Kumar et al.

Carbon Research (2025) 4:6

https://doi.org/10.1007/s44246-024-00158-5

ORIGINAL ARTICLE Open Access

Tree diversity, carbon sequestration

and production potential of Oryza sativa L.
in traditional agroforestry systems of Garhwal
Himalaya, India
Sachin Kumar1, Sandeep Kumar1, Vinod Prasad Khanduri1* , Bhupendra Singh1, Reena Joshi1,
Manoj Kumar Riyal1, Deepa Rawat1 and Kewat Sanjay Kumar2,3

Abstract
Agroforestry is an alternative land use practice that holds promise for societal benefits and the attainment of eco-
system sustainability. The objectives of this study were to evaluate the tree diversity, carbon sequestration, soil
carbon pool, oxygen production and rice productivity under traditional agroforestry systems at different elevations
in the Garhwal Himalayan region of India. Tree diversity, carbon sequestration and oxygen production were quantified
by field measurements (using 0.04 ha quadrats) and subsequent calculations. Rice productivity was assessed using
grain yield, straw yield and biological yield, while soil properties were analyzed in the laboratory using standard meth-
ods. Results of the study showed that tree diversity was higher at the 1200–1600 m elevation and had a maximum
Shannon Diversity Index (1.29) and Simpson Diversity Index (0.69). The 1600–2000 m elevation stored more carbon
(34.43 Mg ­ha−1) and total oxygen production (91.79 Mg ­ha−1). Among the agroforestry trees, Quercus leucotrichophora,
Melia azedarach and Prunus cerasoides showed the highest carbon storage and total oxygen production. Elevation
and soil depth were found to affect the soil properties. The agroforestry systems had higher soil organic carbon
and lower bulk density than sole cropping systems. Compared to the agroforestry system, the monoculture produced
more rice (Oryza sativa). The study shows that traditional agroforestry is a valuable tool for carbon sequestration
and soil improvement, albeit with potential compromises in crop productivity. It emphasises the need for tailored
management approaches to harness the ecological and environmental benefits of agroforestry in the Himalayas.
This study draws attention to the potential of traditional agroforestry in the Garhwal Himalaya for carbon seques-
tration, climate change mitigation and soil quality improvement which provides a reference for striking a balance
between the ecological advantages of agroforestry and the socio-economic considerations of local communities.
However, it also underlines the importance of considering trade-offs between environmental benefits and crop yields
when implementing such agroforestry systems.

*Correspondence:
Vinod Prasad Khanduri
[email protected]
Full list of author information is available at the end of the article

© The Author(s) 2025. Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which
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Kumar et al. Carbon Research (2025) 4:6 Page 2 of 18

Highlights
• Elevation plays an important role for tree diversity, net carbon sequestration and net oxygen production in Himalaya.
• Compared to sole cropping system, agroforestry systems had higher soil pH and soil organic carbon,
along with lower bulk density.
• Production was high in sole cropping system while agroforestry had ecological benefits like carbon storage and soil
improvement.
Keywords Below-ground carbon, Carbon stock, Elevational gradients, Importance Value Index, Oxygen, Soil
properties

Graphical Abstract

1 Introduction (Jose 2019). Traditional agroforestry is a collection of

The increased demand for food, fodder, and timber centuries-old agroforestry practices that have been
is primarily responsible for the over-exploitation of used globally with varying levels of structure, function,
forests. Therefore, agroforestry is considered as an socioeconomic characteristics, and ecological services
alternative land use practice for the benefits of soci- (Ghale et al. 2022).
ety and has the potential to achieve ecosystem sus- In the Himalayas, elevation significantly influences
tainability (Russo 2022). Indian National Agroforestry tree diversity and carbon storage. With increasing ele-
Policy (2014) defines agroforestry as the use of trees in vation, tree diversity tends to decrease due to harsher
croplands and rural landscapes, with or without live- climatic conditions and limited soil nutrients. This
stock, to improve the productivity, profitability, diver- trend of decreasing tree species diversity with increas-
sity, and sustainability of ecosystems. A pan-European ing elevation has also been observed in other studies
study by Rois-Diaz et al. (2017), acknowledges that in the Himalayas (Wani et al. 2022). However, despite
"agroforestry is a new term for something exceedingly their lower species richness, forests and agroforestry
huge and old". However, a more recent definition of systems at high elevations play a crucial role in carbon
this land use system as "agroforestry" is that it encom- sequestration and thus contribute to climate regula-
passes productivity, additional ecosystem services, tion and ecological balance in this fragile mountain
and environmental advantages in a new and larger way region. The varied topography of Garhwal Himalaya
Kumar et al. Carbon Research (2025) 4:6 Page 3 of 18

changes the species diversity, composition, and struc- holds significant importance and is commonly cultivated
ture along the elevations, even over short distances. in rotation with other crops like wheat (Kumar et al.
Tree species present in the agroforestry systems pro- 2018). Its cultivation within the Himalayan agroforestry
vide fuel, fodder, fruit, and fibers to the rural com- framework fosters a balanced interaction that promotes
munity up to a greater extent. In the Himalayas of sustainability and biodiversity in the fragile Himalayan
Uttarakhand, traditional agroforestry systems are environment that is a resulting benefit to both the ecosys-
mainly dominated by several multipurpose and other tem and the economy. At lower elevations, rice is often
fruit tree species (Kumar et al. 2021), that provide the intercropped with trees like Grewia optiva, Celtis austra-
key ecosystem services including carbon sequestra- lis and Morus alba, while at higher elevations it witnesses
tion and oxygen production. integration with species like Quercus leucotrichophora
Plant biomass and soil carbon pools are considered (Khybri et al. 1992). However, this study focuses on rice
as the most important carbon sink of the terrestrial eco- cultivation under traditional agroforestry systems of Utt-
system. Agroforestry systems increase overall biomass arakhand Himalayas.
through the accumulation of dead wood, litter and plant Cultivating woody perennials in agroforestry systems is
material, thus promoting the growth of trees, shrubs and considered as a most effective means for mitigating cli-
crops and also sequester significant amounts of carbon. mate change. Hence, the assessment of tree biomass and
Studies have shown that agroforestry outperforms mono- carbon sequestration (under climate change mitigation
cropping in various aspects, including different economic strategies) are the most crucial variables to be studied
and ecosystem services (Castle et al. 2022). Soil con- generally to elicit their value, as well as the ecosystem ser-
tains the prime terrestrial carbon pool and plays a deci- vices provided by an agroforestry system (Watson et al.
sive role in the exhaustive carbon balance (Scharlemann 2000). Traditional agroforestry in Garhwal Himalaya fea-
et al. 2014). The depletion of soil organic carbon (SOC) turing diverse tree species intercropped with food crops,
degrades soil quality and reduces biomass productivity; which positively influences ecological and agronomic
this decomposition is foreseeable due to the expected factors. We hypothesized that greater tree diversity in
global warming. these systems will link to higher carbon sequestration
Forests and trees are the main sources of oxygen and and more oxygen production. The anticipated synergistic
important reservoirs of carbon dioxide. The potential for interactions between trees and crops are expected to pro-
oxygen production in an agroforestry system depends on mote a more sustainable and resilient agricultural ecosys-
plant growth and biomass, which is influenced by tree tem, contributing to soil and water conservation, thereby
species and density, soil fertility, climatic conditions, benefiting environmental preservation and enhancing
and the implemented management practices (Albrecht overall crop productivity (Mehta et al. 2024). Therefore,
and Kandji 2003). In general, the potential for producing the present study was aimed to evaluate (1) tree diversity,
oxygen in agroforestry systems with a high tree density is carbon sequestration in tree biomass, soil organic carbon
greater than that of systems with a low tree density, espe- stock, and oxygen production at different elevations, and
cially when the fast-growing species are included (Nath (2) the productivity of Oryza sativa under the traditional
et al. 2015). In addition, some species are more tolerant agroforestry systems in different elevations of Garhwal
of adverse environmental conditions, such as drought Himalaya.
or low nutrient availability, which can affect their ability
to produce oxygen (Seleiman et al. 2021; Keethika and 2 Materials and methods
Chavan 2022; Zhang et al. 2022). 2.1 Experimental sites
Intercropping between trees and crops at various The study survey was conducted in the farmers’ fields of
elevations in the Himalayas highlights the potential for the mid-hills region of Uttarkashi district, Uttarakhand,
enhancing biodiversity, improving soil fertility, and opti- India. Nine villages (three in each elevation range) were
mizing land uses, thereby also enhancing the rural live- selected in the northwest region of Uttarkashi district
lihoods of people (Arunachalam et al. 2019; Ghale et al. lying between 30°47′09.18" N to 30°55′12.50" N latitude
2022). Rice (Oryza sativa L.) is the world’s most impor- and 78°01′30.50" E to 78°09′24.37" E longitude in the ele-
tant agricultural produce, with nearly 40% of the global vational ranges of 800–1200 m asl, 1200–1600 m asl and
population relying on it as their primary staple food. 1600–2000 m asl. The selected area is renowned for rice
Rice cultivation within diverse agroforestry systems is cultivation in the Uttarakhand Himalayas. The geograph-
already underway in various regions worldwide. Tomar ical details like elevation and coordinates of each village
et al. (2013) and Wijayanto and Briliawan (2022) have were measured using GPS (Garmin, vista) and given in
documented the response of rice crops to various tree Fig. 1a.
species. In the Himalayan agroforestry systems, rice
Kumar et al. Carbon Research (2025) 4:6 Page 4 of 18

Fig. 1 a The location map of the study sites; b overview of traditional agroforestry system in the study area

2.2 Sampling design π D2
The tree diversity, carbon stock, and capacity of oxy- Basal area = (1)
4
gen released by traditional agroforestry systems were
assessed by laying out 10 quadrats/sample plots of
0.04 ha (20 × 20 m) in the agroforestry systems of each 2.4 Trees diversity
village. Based on soil control sections, soil samples were The Importance Value Index (IVI) demonstrates the
collected in 15 cm increments from three fixed depths, importance and dominance of species in the study area
i.e., 0–15 cm, 15–30 cm, and 30–45 cm, from random by combining relative density, frequency, and dominance
spots within a 0.04-hectare quadrat. The productivity of percentages according to the methods of Curtis (1959).
rice was estimated during the Kharif season. In agrofor- Species richness was determined by counting the spe-
estry systems, the random selection of 5 sample plots of cies in the agroforestry systems. Menhinick’s richness
1 × 1 m (0.0001 ha) was laid out in 0.04 ha sample plots, index was computed following Whittaker (1977), the
while in a sole cropping system (control condition), 5 Shannon–Wiener index was determined according to the
sample plots of 0.0001 ha were randomly laid out for esti- approach outlined by Shannon and Wiener (1963), Simp-
mating the productivity of rice (Oryza sativa L.). son’s diversity index was calculated as Simpson’s (1949),
and Pielou’s Evenness index, which indicates the relative
2.3 Trees enumeration dominance of each species in the area, was determined as
Individual species having ≥ 10 cm diameter (31.41 cm per Pielou (1966). Additionally, the Whittaker β-diversity
girth) at breast height were considered individual trees index was computed following Whittaker’s (1972).
(FSI 2021). The tree girth at breast height (1.37 m) was
estimated using measuring tape and converted in diam-
2.5 Tree volume and carbon stock
eter using the formula, i.e., D = G/3.14, and expressed in
Tree volume was measured following non-destructive
centimetres (cm). The total height of the tree was meas-
methods based on recorded height and diameter values
ured by using a Ravi multimetre in all sample plots and
with the existing volume equations of a single tree and
expressed in meters. The tree basal area was derived
the general equation for tree volume from the Forest Sur-
from diameter using the following formula (Eq. [1]) and
vey of India (FSI 1996) for the Yamuna catchment in the
expressed in m­ 2, where D = Tree diameter and π = Pi is
Garhwal Himalaya. The volume of each tree was calcu-
constant (3.14).
lated within the sample plots to determine the growing
Kumar et al. Carbon Research (2025) 4:6 Page 5 of 18

stock volume density (GSVD), which was then converted Net O2 production Mg ha−1 yr−1 = Net carbon sequestration × 32/12
to ­m3 ­ha−1 by multiplying a value of 25. Above-ground (5)
tree biomass was determined by applying a biomass
2.7 Crop productivity
expansion factor for broadleaf trees (Brown et al. 1999) to
the GSVD of the sample plot. Below-ground tree biomass Rice productivity (q ­ha−1) was assessed using grain (eco-
was calculated using a regression equation by Cairns nomical) yield, straw yield and biological (grain + straw)
et al. (1997). yield. The harvested rice was then separated into grain
Using the default carbon fraction value of 0.50, as and straw components. The harvest index of rice was
per the IPCC (2000), the above-ground (bole and estimated by the following equation (Eq. [6]).
crown) biomass, below-ground (root) biomass, and
total (above + below ground) biomass values were
Economical Yield
converted into above-ground carbon stock (AGCS), Harvest Index = × 100 (6)
below-ground carbon (BGCS), and total tree carbon Biological Yield
stocks (TCS) (Eq. [2]).

Total tree carbon stock = Total tree biomass × Carbon % (i.e., 0.50)
(2)

After arriving at the carbon stock of each tree, the 2.8 Soil properties analysis
above ground net carbon sequestration (AGNCS) The soil analysis was carried out in the soil science lab-
potential, below ground net carbon sequestration oratory of the College of Forestry (VCSG UUHF), Ran-
(BGNCS) and net carbon sequestration (NCS) in terms ichauri. Soil pH was measured according to Jackson
of ­CO2 (eq.) was calculated by using the following for- (1967). The soil bulk density (g c­ c−1) was measured by
mula (Eq. [3]): the core tube sampler method. The modified Walkley
and Black method was utilised to analyse the organic car-
bon content of the soil (Walkley and Black 1934) and the
CO2 (eq.) = (Carbon stock × 44) / 12 (3) Brown (2004) formula was used to calculate soil organic
The above equation represents the quantity of total carbon stock (Mg ­ha−1).
­CO2 sequestered in terms of C ­ O2 (eq.) of the tree during
its entire lifespan. To obtain a yearly C sequestration rate, 2.9 Statistic analysis
the overall ­CO2 equivalent by the tree was divided by its One-way ANOVA (Analysis of Variance) was used to
age (Eq. [4]). The information on the age of the tree was study the tree’s carbon stock and oxygen production
collected from the local farmers. statistically, while the 3-factorial ANOVA analysis was
applied to analyse the soil parameters. The degree of the
linear associations between the various parameters in
Net C sequestration = CO2 (eq.) / Age of the tree
both agroforestry and sole cropping systems was ascer-
(4)
tained following the Pearson correlation coefficient,
using IBM Corporation’s SPSS software.
2.6 Oxygen released by trees
The net oxygen production (NOP) of trees was calculated
by subtracting the oxygen absorbed during respiration 3 Results
from the oxygen produced during photosynthesis. If a tree 3.1 Tree density, diversity and Importance Value Index (IVI)
absorbs more C ­ O2 during photosynthesis than it releases The highest tree density (95.19 trees ­ha−1) was observed
during respiration, it sequesters carbon, which indi- at the middle elevation range, and the upper elevation
cates the growth of trees and net oxygen production. The has the lowest tree density with 86.84 trees ­ha−1. Grewia
total oxygen production (TOP) including above ground optiva (41.75), Celtis australis (33.4), and Quercus leu-
(AGOP), below ground (BGOP) of trees was estimated by cotrichophora (55.11) had the highest numbers of indi-
the total carbon sequestration, while the net oxygen pro- viduals at lower, middle, and upper elevation ranges,
duction, including above-ground (AGNOP) and below- respectively. Middle elevation exhibited the highest fre-
ground (BGNOP) contributions of trees was estimated quency of species, followed by lower and upper eleva-
by the net carbon sequestration of trees according to the tional ranges. At lower elevations (800–1200 m), 5 tree
formula (Eq. [5]) of Nowak and Dwyer (2007). species were recorded in the traditional agroforestry sys-
tem while six tree species were recorded at middle and
Kumar et al. Carbon Research (2025) 4:6 Page 6 of 18

upper elevations. The study reveals varying basal areas ranges compared to the lower elevation (1.04) range.
of tree species across different elevations, reflecting their The Plieou Index of Evenness indicates differences in the
distribution, growth and dominance. At lower elevations, evenness of tree species among elevation ranges, with the
the total basal area is 3.83 ­m2 ­ha−1, with P. cerasoides middle elevation having the highest evenness (0.90), fol-
and P. pashia being notable basal area. Middle elevations lowed by the lower elevation range (0.88) while the upper
have a total basal area of 3.78 m ­ 2 ­ha−1, showing a more elevation shows the lowest evenness (0.70). Upper eleva-
even growth and distribution of species. The upper eleva- tion had the highest beta diversity (2.24), followed by
tion shows a marked increase to the total basal area of lower (1.96) and middle (1.61) elevation ranges (Fig. 2).
4.98 ­m2 ­ha−1, dominated by Q. leucotrichophora (3.62
­m2 ­ha−1), indicating significant species variation with ele- 3.2 Tree carbon stock and net carbon sequestration
vation (Table 1). The carbon stock of the trees varied between different
The results of tree diversity parameters show that the elevations (Table 2). The upper elevation had the highest
Simpson Diversity Index values differ slightly among ele- AGCS and BGCS, i.e., 26.71 Mg ­ha−1 and 7.72 Mg ­ha−1,
vation ranges, with upper elevation showing lower diver- respectively, while the 800–1200 m elevational range had
sity (0.53) compared to lower (0.67) and middle (0.69) a minimum above and below ground carbon stock. The
elevations. The upper elevation shows the highest Men- TCS was highest at the upper elevation (34.43 Mg ­ha−1),
hinick’s Index, i.e., 1.04, and minimum in the lower eleva- followed by the middle elevation (33.80 Mg ­ha−1) and the
tion (0.95). Marglef ’s index values indicate higher overall lower elevation (30.49 Mg ­ha−1). Among the elevations,
diversity in the upper (1.16) and middle elevation (1.13) the middle elevation showed the highest mean value
of AGNCS (8.34 Mg ­ha−1 ­yr−1), followed by the upper
elevation (7.10 Mg ­ha−1 ­yr−1) and the lower elevation
Table 1 Average number of tree density, frequency, tree basal (7.08 Mg ­ha−1 ­yr−1). Similarly, the middle elevation had
area and Importance Value Index (IVI) in traditional agroforestry the highest mean value of BGNCS (2.40 Mg ­ha−1 ­yr−1),
systems at different elevational ranges followed by the upper elevation (2.05 Mg ­ ha−1 ­yr−1)
−1 −1
and the lower elevation (1.96 Mg ­ ha ­yr ). Over-
Tree species Density Frequency Basal area IVI
(number (m2 ha−1) all, net carbon sequestration followed a similar pat-
of trees tern, with the middle elevation recording the highest
ha–1) mean value (10.74 Mg ­ha−1 ­yr−1), followed by the upper
Lower elevation (800–1200 m) elevation (9.15 Mg ­ha−1 ­yr−1) and the lower elevation
Grewia optiva 41.75 0.73 0.97 106.65 (9.05 Mg ­ha−1 ­yr−1).
Pyrus pashia 25.05 0.60 1.10 86.41 At 800–1200 m elevation, Prunus cerasoides showed
Melia azedarach 8.35 0.20 0.49 56.90 the highest total and net carbon sequestration rates,
Prunus cerasoides 8.35 0.33 1.21 31.50 i.e., 8.94 Mg ­ha−1 and 2.65 Mg ­ ha−1 ­yr−1 respec-
Celtis australis 6.68 0.20 0.06 18.54 tively, followed by P. pashia and G. optiva, while C.
Total 90.18 2.07 3.83 300.00 australis sequestered the least total and net carbon
Middle elevation (1200–1600 m)
(3.15 Mg ­ha−1 and 0.94 Mg ­ ha−1 ­yr−1, respectively)
Celtis australis 33.40 0.87 0.54 110.06
(Table 2). At the middle elevation range (1200–1600 m),
Melia azedarach 30.06 0.73 0.91 81.87
the highest value of carbon stock (8.59 Mg ­ha−1) and
Grewia optiva 20.04 0.60 0.58 58.91
net carbon sequestration (2.73 Mg ­ ha−1 ­yr−1) was
Pyrus pashia 6.68 0.27 0.39 27.38
observed for M. azedarach among all species. P. pashia
Quercus leu- 3.34 0.13 0.23 14.37
exhibited the lowest TCS (4.00 Mg ­ ha−1) and NCS
−1 −1
cotrichophora (1.27 Mg ­ha ­yr ). Similarly, carbon stock and seques-
Prunus cerasoides 1.67 0.07 0.13 7.41 tration were also recorded as maximum for M. azedar-
Total 95.19 2.67 3.78 300.00 ach and minimum for P. pashia. At the upper elevation
Upper elevation (1600–2000 m) range (1600–2000 m), among all tree species, Q. leu-
Quercus leu- 55.11 0.87 3.62 182.04 cotrichophora sequesters the highest amount of carbon
cotrichophora and possesses the maximum carbon stock, whereas C.
Grewia optiva 11.69 0.33 0.45 39.84 australis exhibits the lowest total and net carbon stock,
Celtis australis 10.02 0.33 0.21 33.10 i.e., 3.14 Mg ­ha−1 and 0.83 Mg ­ha−1 ­yr−1, respectively.
Melia azedarach 3.34 0.13 0.26 15.82 Overall, the upper elevation range revealed the maxi-
Prunus cerasoides 3.34 0.13 0.24 14.72 mum total carbon stocks (34.43 Mg ­ha−1) and the mid-
Pyrus pashia 3.34 0.13 0.20 14.47 dle elevation range proclaimed the highest net carbon
Total 86.84 1.93 4.98 300.00
Kumar et al. Carbon Research (2025) 4:6 Page 7 of 18

Fig. 2 Tree diversity parameters in traditional agroforestry systems at different elevational ranges

sequestration (10.74 Mg ­ha−1 ­yr−1) among the three At lower elevations, P. cerasoides exhibits the high-
elevation ranges. est total oxygen production, i.e., 23.85 Mg ­ha−1. Con-
versely, at middle and upper elevations, M. azedarach
(22.91 Mg ­ha−1) and Q. leucotrichophora (38.40 Mg ­ha−1)
3.3 Trees oxygen production demonstrated the highest oxygen production, both
Upper elevation exhibited the highest mean value aboveground and belowground, as well as the highest
of above-ground oxygen production (AGOP), net oxygen production. At the upper elevation range
i.e., 71.22 Mg ­ha−1, followed by middle elevation (1600–2000 m asl), Q. leucotrichophora exhibits the high-
(69.96 Mg ­ha−1), and lower elevation (63.67 Mg ­ha−1). est NOP (10.21 Mg ­ha−1 ­yr−1) values among all species.
The highest belowground oxygen production (BGOP) Across all elevation ranges, above-ground oxygen pro-
and net oxygen production (NOP) were observed at duction (AGOP), and above-ground net oxygen produc-
the upper elevational range, followed by the middle and tion (AGNOP) consistently surpasses the below-ground
then the lower elevation. Across all elevation ranges, oxygen production (BGOP) and below-ground net oxy-
there was a general trend of increasing oxygen pro- gen production (BGNOP) (Table 3).
duction, represented by the Total Oxygen Production
(TOP) values. As a result, the TOP was also highest in 3.4 Soil properties
the upper elevation range (91.79 Mg ­ ha−1), reflecting 3.4.1 Soil pH
larger oxygen production, followed by middle elevation In the lower and middle elevational ranges, the tradi-
(90.12 Mg ­ha−1), and lower elevation (81.31 Mg ­ha−1) tional agroforestry system exhibited higher soil pH val-
ranges. The highest value of AGNOP was recorded in the ues compared to the sole agricultural cropping system.
middle elevation range (22.23 Mg ­ha−1 ­yr−1), followed by Conversely, the sole cropping system had higher soil pH
upper elevation (18.93 Mg ­ha−1 ­yr−1) and lower elevation values than the agroforestry system at higher elevations.
(18.88 Mg ­ha−1 ­yr−1) ranges. The data reveals distinct In the studied agroforestry, soil pH ranged between 5.06
trends in above and belowground net oxygen production to 5.34, 6.14 to 6.43 and 5.94 to 6.00, respectively, in the
across the elevations. The net oxygen production (NOP) lower, middle and upper elevations; while 4.74 to 4.90,
follows a similar pattern as of net carbon sequestration, 5.04 to 6.23, and 5.44 to 6.20, respectively, in the sole
with the highest mean value recorded in the middle agricultural cropping system (Table 4). At lower and mid-
elevation range (28.64 Mg ­ha−1 ­yr−1), followed by upper dle elevation ranges, the agroforestry system exhibited
elevation (24.38 Mg ­ha−1 ­yr−1) and Lower Elevation higher soil pH values compared to the rice-cultivated
(24.40 Mg ­ha−1 ­yr−1) (Table 3). fields. Conversely, at the upper elevation range, the sole
cropping system showed higher soil pH values compared
Kumar et al. Carbon Research (2025) 4:6 Page 8 of 18

Table 2 Carbon stock (Mg ­ha−1) and net carbon sequestration (Mg ­ha−1 ­yr−1) of each tree species in studied traditional agroforestry
systems (Mean ± SD)
Tree species AGCS BGCS TCS AGNCS BGNCS NCS

Lower elevation (800–1200 m asl)

Pyrus pashia 5.09 ± 0.62 1.49 ± 0.17 6.58 ± 0.79 1.51 ± 0.20 0.44 ± 0.06 1.95 ± 0.25
Prunus cerasoides 6.98 ± 0.18 1.96 ± 0.79 8.94 ± 0.19 2.07 ± 1.14 0.58 ± 1.07 2.65 ± 0.66
Grewia optiva 4.98 ± 1.01 1.46 ± 1.75 6.43 ± 1.76 1.48 ± 1.92 0.43 ± 0.53 1.91 ± 0.64
Melia azedarach 4.14 ± 0.23 1.24 ± 0.14 5.38 ± 0.25 1.23 ± 0.17 0.37 ± 0.18 1.60 ± 0.15
Celtis australis 2.68 ± 0.11 0.47 ± 0.28 3.15 ± 0.27 0.80 ± 0.23 0.14 ± 0.49 0.94 ± 0.50
Total 23.87 6.62 30.49 7.08 1.96 9.05
Middle elevation (1200–1600 m asl)
Melia azedarach 6.73 ± 0.56 1.86 ± 0.15 8.59 ± 0.7 2.14 ± 0.40 0.59 ± 0.11 2.73 ± 0.23
Celtis australis 4.37 ± 0.53 1.27 ± 0.12 5.64 ± 0.11 1.39 ± 0.39 0.40 ± 0.12 1.79 ± 0.25
Grewia optiva 3.55 ± 0.11 1.06 ± 0.24 4.61 ± 0.32 1.13 ± 0.61 0.34 ± 0.15 1.46 ± 0.33
Quercus leucotrichophora 3.71 ± 0.34 1.10 ± 0.15 4.80 ± 0.15 1.18 ± 0.83 0.35 ± 0.39 1.53 ± 0.35
Pyrus pashia 3.07 ± 0.15 0.93 ± 0.18 4.00 ± 0.07 0.98 ± 0.06 0.30 ± 0.10 1.27 ± 0.07
Prunus cerasoides 4.81 ± 0.04 1.34 ± 0.20 6.15 ± 0.15 1.53 ± 0.21 0.42 ± 0.24 1.95 ± 0.15
Total 26.23 7.56 33.80 8.34 2.40 10.74
Upper elevation (1600–2000 m asl)
Quercus leucotrichophora 11.43 ± 0.56 2.97 ± 0.15 14.40 ± 0.70 3.04 ± 0.40 0.79 ± 0.11 3.83 ± 0.50
Celtis australis 2.39 ± 0.61 0.75 ± 0.18 3.14 ± 0.56 0.64 ± 0.21 0.20 ± 0.23 0.83 ± 0.97
Grewia optiva 4.23 ± 0.18 1.24 ± 0.19 5.47 ± 0.37 1.13 ± 0.13 0.33 ± 0.25 1.45 ± 0.10
Prunus cerasoides 2.38 ± 0.79 0.82 ± 0.20 3.20 ± 0.39 0.63 ± 0.12 0.22 ± 0.08 0.85 ± 0.35
Melia azedarach 2.73 ± 0.74 0.88 ± 0.49 3.61 ± 1.24 0.72 ± 0.04 0.23 ± 0.03 0.95 ± 0.21
Pyrus pashia 3.55 ± 0.06 1.06 ± 0.06 4.61 ± 0.10 0.94 ± 0.20 0.28 ± 0.35 1.22 ± 0.59
Total 24.71 7.72 34.43 7.10 2.05 9.13
Source of variation F ratio
Elevation 7.230** 10.287** 8.197** 27.864** 34.012** 37.527**
Village 1.790NS 1.970NS 1.873NS 40.299** 50.613** 54.576**
Elevation × Village 0.810NS 0.342NS 0.704NS 33.946** 40.416** 45.491**
AGCS Above-ground carbon stock, BGCS Below-ground carbon stock, TCS Total carbon stock, AGNCS Above-ground net carbon sequestration, BGNCS Below-ground
net carbon sequestration, NCS Net carbon sequestration, SD Standard deviation, NS Non-significant
**
Significant at 0.01 level

to the studied agroforestry system. In the sole cropping 3.4.2 Soil bulk density
system, at the lower elevation range, pH increased with Bulk density (BD) of soil was recorded maximum at
depth, implying higher pH values at greater depths within 30–45 cm depth and decreased with elevation in the
this range. At the middle elevation range, pH decreased studied agroforestry and sole cropping systems. Under
with increasing depth, indicating a decline in pH values the agroforestry systems, the bulk density was slightly
at deeper levels within this range. There was no consist- higher at deeper depths (30–45 cm) compared to shal-
ent pattern of soil pH with increasing depth in the agro- lower depths (0–15 cm). For the sole cropping system,
forestry system, indicating a varying influence on pH at the bulk density generally tended to increase slightly from
different depths. In the agroforestry system, the maxi- shallower to deeper depths, although the differences were
mum pH was recorded at the middle elevation range with not substantial. Agroforestry soils showed slightly lower
a depth of 15–30 cm (6.43). In contrast, the sole cropping bulk density compared to sole cropping soils across the
system recorded its maximum pH (6.23) at 0–15 cm soil elevational ranges. In the agroforestry system, the soil
depth in the middle elevation range. The lowest pH for bulk density reached its highest value of 1.28 g ­cc−1 at the
both systems was observed at the lower elevation range, lower elevation, specifically in the 30–45 cm depth range.
with the agroforestry system having a depth of 15–30 cm Similarly, in the sole cropping system, the maximum bulk
(50.6) and in the sole cropping system at a shallower density of 1.31 g ­cc−1 was also observed at the lower ele-
depth of 0–15 cm (4.74). vation within 30–45 cm depth range (Table 4).
Kumar et al. Carbon Research (2025) 4:6 Page 9 of 18

Table 3 Total oxygen production (Mg ­ha−1), net oxygen production (Mg ­ha−1 ­yr−1) of each tree species in traditional agroforestry
systems at different elevational ranges (Mean ± SD)
Tree species AGOP BGOP TOP AGNOP BGNOP NOP

Lower elevation (800–1200 m asl)

Pyrus pashia 13.58 ± 1.21 3.97 ± 1.47 17.55 ± 2.12 4.03 ± 1.20 1.18 ± 1.12 5.21 ± 1.26
Prunus cerasoides 18.61 ± 2.26 5.24 ± 2.69 23.85 ± 2.11 5.52 ± 1.24 1.55 ± 0.85 7.07 ± 2.13
Grewia optiva 13.27 ± 1.31 3.89 ± 1.36 17.16 ± 2.41 3.94 ± 1.32 1.15 ± 0.89 5.09 ± 2.36
Melia azedarach 11.04 ± 1.75 3.30 ± 2.58 14.35 ± 2.36 3.28 ± 1.14 0.98 ± 1.10 4.25 ± 2.14
Celtis australis 7.16 ± 1.88 1.25 ± 1.54 8.41 ± 2.54 2.12 ± 1.10 0.37 ± 0.88 2.49 ± 1.85
Total 63.67 17.64 81.31 18.88 5.23 24.12
Middle elevation (1200–1600 m asl)
Melia azedarach 17.95 ± 1.45 4.97 ± 1.23 22.91 ± 1.23 5.70 ± 1.25 1.58 ± 0.84 7.28 ± 2.55
Celtis australis 11.65 ± 1.12 3.39 ± 0.85 15.05 ± 2.12 3.70 ± 1.65 1.08 ± 0.87 4.78 ± 1.84
Grewia optiva 9.47 ± 1.32 2.82 ± 0.89 12.29 ± 2.36 3.01 ± 0.89 0.90 ± 0.59 3.91 ± 1.42
Quercus leucotrichophora 9.88 ± 1.01 2.93 ± 1.10 12.81 ± 2.12 3.14 ± 1.05 0.93 ± 0.88 4.07 ± 1.63
Pyrus pashia 8.19 ± 1.22 2.48 ± 0.87 10.67 ± 2.10 2.60 ± 1.23 0.79 ± 0.45 3.39 ± 1.25
Prunus cerasoides 12.82 ± 1.32 3.56 ± 1.12 16.39 ± 2.41 4.07 ± 1.08 1.13 ± 0.65 5.21 ± 2.12
Total 69.96 20.16 90.12 22.23 6.41 28.64
Upper elevation (1600–2000 m asl)
Quercus leucotrichophora 30.47 ± 2.10 7.93 ± 2.12 38.40 ± 2.30 8.10 ± 1.26 2.11 ± 0.26 10.21 ± 2.32
Celtis australis 6.38 ± 1.32 1.99 ± 1.10 8.37 ± 2.10 1.70 ± 1.20 0.53 ± 0.55 2.23 ± 2.10
Grewia optiva 11.29 ± 1.54 3.30 ± 1.32 14.59 ± 2.14 3.00 ± 1.34 0.88 ± 0.84 3.88 ± 2.14
Prunus cerasoides 6.34 ± 1.15 2.18 ± 1.14 8.52 ± 1.85 1.69 ± 1.54 0.58 ± 0.65 2.27 ± 1.84
Melia azedarach 7.28 ± 1.07 2.35 ± 0.97 9.63 ± 1.26 1.92 ± 1.13 0.61 ± 0.59 2.52 ± 1.62
Pyrus pashia 9.46 ± 1.65 2.82 ± 1.02 12.28 ± 2.05 2.52 ± 1.20 0.75 ± 0.26 3.27 ± 1.15
Total 71.22 20.57 91.79 18.93 5.46 24.38
Source of variation F ratio
Elevation 7.226** 10.287** 8.197** 37.527** 34.018** 37.527**
Village 1.789 NS 1.970NS 1.873NS 54.576** 50.613** 54.576**
Elevation × Village 0.809NS 0.342NS 0.704NS 45.491** 40.416** 45.491**
AGOP Above-ground oxygen production, BGOP Below-ground oxygen production, TOP Total oxygen production, AGNOP Above-ground net oxygen production,
BGNOP Below-ground net oxygen production, NOP Net oxygen production, NS Non-significant
**
Significant at 0.01 level

3.4.3 Soil organic carbon percentage 30–45 cm. Similarly, in the sole cropping system, the
The percentage of SOC increased with the studied ele- minimum soil organic carbon percentage of 1.35% was
vational ranges for both agroforestry and single crop- observed at the lower elevation within 30–45 cm depth
ping systems at all three depths. At a depth of 30–45 cm, range (Table 4).
more variation in the percentage of soil organic carbon
was appeared, but there was a trend towards higher val- 3.4.4 Soil organic carbon stock
ues in the studied traditional agroforestry as compared The soil organic carbon stock (SOCS) increases with
to the rice cultivated fields, especially at the upper eleva- elevation at each depth. In general, SOCS was highest at
tions. In agroforestry, the highest percentage of SOC the upper elevations and lowest at the lower elevations
(3.67%) was recorded in the upper elevations, especially in both agroforestry and sole cropping systems (Table 4).
in 0–15 cm depth range. Similarly, the highest percent- In every elevational range, the amount of organic carbon
age of SOC (2.73%) was observed in the elevation range stock in the soil tended to decrease with increasing soil
of 1600–2000 m for the rice cultivated area at 0–15 cm depth. In agroforestry, SOCS was generally higher at all
depth. Conversely, the lowest percentage of soil organic three elevations and soil depths than in the monoculture
carbon (2.13%) was observed in the agroforestry system system. In the agroforestry system, the highest SOCS
at the lower elevation, specifically in the depth range of
Kumar et al. Carbon Research (2025) 4:6 Page 10 of 18

Table 4 Soil properties at different soil depths under traditional agroforestry and sole cropping systems at different elevational ranges
(Mean ± SD)
Elevation range Agroforestry system Sole cropping system
0–15 cm 15–30 cm 30–45 cm 0–15 cm 15–30 cm 30–45 cm

Soil pH
Lower elevation 5.19 ± 0.17 5.06 ± 0.12 5.34 ± 0.09 4.74 ± 0.06 4.84 ± 0.09 4.90 ± 0.16
Middle elevation 6.19 ± 0.09 6.43 ± 0.03 6.14 ± 0.05 6.23 ± 0.14 5.78 ± 0.12 5.04 ± 0.07
Upper elevation 6.00 ± 0.13 5.94 ± 0.04 5.97 ± 0.13 6.15 ± 0.11 5.44 ± 0.09 6.20 ± 0.05
Soil bulk density (g cc−1)
Lower elevation 1.10 ± 0.12 1.17 ± 0.15 1.28 ± 0.09 1.06 ± 0.02 1.18 ± 0.04 1.31 ± 0.11
Middle elevation 1.04 ± 0.08 1.12 ± 0.03 1.22 ± 0.05 1.11 ± 0.14 1.17 ± 0.12 1.29 ± 0.05
Upper elevation 1.01 ± 0.06 1.07 ± 0.04 1.16 ± 0.12 1.08 ± 0.06 1.15 ± 0.09 1.21 ± 0.07
Soil organic carbon (%)
Lower elevation 2.52 ± 0.57 2.32 ± 0.73 2.13 ± 0.67 1.93 ± 0.78 1.53 ± 0.22 1.35 ± 0.15
Middle elevation 3.50 ± 0.21 2.71 ± 0.60 2.32 ± 0.50 2.32 ± 0.70 2.12 ± 0.63 1.73 ± 0.52
Upper elevation 3.67 ± 0.43 3.50 ± 0.44 3.15 ± 0.54 2.73 ± 0.52 2.52 ± 0.52 2.20 ± 0.51
Soil organic carbon stock (Mg ha−1)
Lower elevation 41.58 ± 0.42 40.72 ± 0.33 40.90 ± 0.22 30.69 ± 0.30 27.08 ± 0.36 26.53 ± 0.65
Middle elevation 54.60 ± 0.35 45.53 ± 0.40 42.46 ± 0.16 38.63 ± 0.40 37.21 ± 0.26 33.48 ± 0.56
Upper elevation 55.60 ± 0.74 56.18 ± 0.23 54.81 ± 0.32 44.23 ± 0.25 43.47 ± 0.32 39.93 ± 0.27
Source of variation Degree of freedom F calculated
pH BD SOC SOCS
Elevation 2 98.303** 8.638** 21.748** 15.594**
System 1 26.283** 9.667** 44.126** 34.130**
Soil depth 2 2.825* 54.934** 8.709** 1.544NS
Elevation × System 2 5.881** 2.422NS 0.325NS 0.088NS
NS NS
Elevation × Soil depth 4 7.875** 0.849 0.406 0.366NS
NS NS
System × Soil depth 2 3.555* 0.133 0.112 0.035NS
NS NS
Elevation × System × Soil depth 4 5.159** 0.410 0.414 0.286NS
BD Bulk density, SOC Soil organic carbon, SOCS Soil organic carbon stock, NS Non-significant
*
Significant at 0.05 level
**
Significant at 0.01 level

(56.18 Mg ­ha−1) was found in the 1600–2000 m elevation the agroforestry system was observed in the lower eleva-
range at a depth of 15–30 cm. Conversely, the highest tions, especially at the depth of 15–30 cm with a value
SOCS (44.23 Mg ­ha−1) was observed in the sole crop- of 40.72 Mg ­ha−1. In the sole cropping system, the lowest
ping system at the surface soil depth (0–15 cm) in the SOCS of 26.53 Mg ­ha−1 was recorded at the 800–1200 m
upper elevation. The lowest stock of organic carbon in elevation within the 30–45 cm depth.

Table 5 Productivity of rice (q ­ha−1) under traditional agroforestry system at different elevational ranges (Mean ± SD)
Elevation range Grain yield Straw yield Biological yield Harvest index (%)

Traditional agroforestry system

Lower elevation 17.49 ± 0.64 25.62 ± 4.23 43.10 ± 3.64 40.57 ± 4.21
Middle elevation 16.63 ± 2.01 22.34 ± 0.19 42.33 ± 0.79 39.28 ± 3.12
Upper elevation 11.48 ± 2.54 20.28 ± 0.39 39.45 ± 2.32 29.11 ± 3.26
Sole cropping system (control condition)
Lower elevation 21.05 ± 1.61 28.56 ± 0.24 49.20 ± 1.52 42.78 ± 1.35
Middle elevation 19.31 ± 0.90 26.71 ± 1.25 46.05 ± 0.54 41.93 ± 3.18
Upper elevation 13.71 ± 1.38 25.25 ± 0.73 42.96 ± 1.42 31.91 ± 2.08
Kumar et al. Carbon Research (2025) 4:6 Page 11 of 18

3.5 Productivity of Oryza sativa decreased by increasing elevational range, with the mid-
The grain yield in the sole rice cultivated land is higher dle elevation recorded as 39.28% and the upper eleva-
at all elevations than in the agroforestry system. Among tion showing the lowest index of 29.11%. At the lower
the elevational ranges, the grain yield was maximum at elevation, the sole cropping system achieved the maxi-
800–1200 m elevation (21.05 q ­ha−1) in the sole crop- mum harvest index of 42.78%. Similarly, as the elevation
ping system, while it was 17.49 q h ­ a−1 in the traditional increased, the harvest index declined, with the mid-
agroforestry (Table 5). In agroforestry and monoculture dle elevation displaying 41.93% and the upper elevation
systems, grain, straw and biological yields of rice were 31.91%. The grain and biological yield in agroforestry sys-
decreasing with increasing elevations. In the agroforestry tem at lower elevation showed maximum yield reduction
system, grain yields vary across elevations with values of of 16.91% and 12.4% respectively, while a reduction of
17.49 q h­ a−1, 16.63 q h
­ a−1, and 11.48 q h­ a−1 in the lower, 19.68% in straw yield had been recorded maximum at the
middle, and upper elevational ranges, respectively. Com- upper elevation range as compared to that of sole crop-
paratively, in rice-cultivated fields, slightly higher yields ping system (Fig. 3).
of 21.05 q ­ha−1, 19.31 q ­ha−1, and 13.71 q ­ha−1, respec-
tively, were recorded for the corresponding elevations. 3.6 Correlation between different soil parameters,
Similar to grain yield, straw yield and biological yield is vegetation, and crop yield
also higher in the sole cropping system as compared to A significant negative Pearson correlation was found in
that of agroforestry system at all elevations. Overall, the the agroforestry system between grain yield and basal
data suggest that the lower elevation favours higher yields area, Whittaker β diversity and trees ­ha−1, and Men-
in agroforestry and sole rice fields. Additionally, the sole hinik’s index and straw production. Additionally, a strong
cropping system generally outperforms the agroforestry positive connection was observed between elevation
system in terms of crop productivity across all elevations. and Menhinick’s index, and Simpson’s diversity index to
The values of the harvest index were reported higher in Plieou’s index of evenness, TCS and TOP. In the case of
the rice field than in the agroforestry system. The highest average soil properties of all the depths, BD shows a sig-
harvest index of 40.57% was observed within the agro- nificant negative correlation with elevation, SOC, SOCS,
forestry system at the lower elevation, which gradually TCS, and TOP while Menhinick’s index with BD. A

Fig. 3 Yield reduction % of rice (Oryza sativa) in agroforestry system as compared to that of sole cropping system in different elevation ranges
Kumar et al. Carbon Research (2025) 4:6 Page 12 of 18

Fig. 4 Pearson correlation coefficient between different soil parameters, vegetation, and crop yield under agroforestry systems with average soil
properties. BD = Soil bulk density, SOC = Soil organic carbon, SOCS = Soil organic carbon stock, H’ = Shannon Diversity Index, D = Simpson diversity
index; MeI = Menhinick’s index; MI = Marglef’s index, PI = Plieou index of evenness, βD = Whittaker β diversity, BA = Basal area (trees), TCS = Total
carbon stock (trees), TOP = Total oxygen production (trees), GY = Grain yield (rice), SY = Straw yield (rice), ** = Correlation is significant at the 0.01 level,
* = Correlation is significant at the 0.05 level

significant positive correlation was found between SOC a noteworthy inverse relationship between SOCS
and SOCS. The soil pH shows no significant relation with and straw yield was found. Grain yield and bulk den-
any of the studied characters (Fig. 4). sity were shown to be significantly correlated at a
In the sole cropping system, elevation range and depth of 30–45 cm, while SOCS and straw yield were
grain yield are negatively correlated. SOC shows a sig- found to be significantly negatively correlated. There
nificant positive correlation with elevation at a depth was a strong negative correlation between SOCS and
of 0–15 cm, while SOCS reveals a significant inverse rice straw yield and a significant positive correlation
correlation with rice straw yield. SOC has a positive between SOC and SOCS for average soil parameters
correlation with SOCS and a negative correlation with at all depths. SOC also had a strong negative cor-
straw yield at a soil depth of 15–30 cm. Furthermore, relation with straw yield and a positive correlation

Fig. 5 Pearson correlation coefficient between different soil parameters and crop yield under sole cropping systems at (a) 0–15 cm soil depth; (b)
15–30 cm soil depth; (c) 30–45 cm soil depth and (d) average soil properties of all three studied soil depth. BD = Soil bulk density (g ­cc−3), SOC = Soil
organic carbon (%), SOCS = Soil organic carbon stock (Mg ­ha−1), GY = Grain yield of rice (q h
­ a−1), SY = Straw yield of rice (q ­ha−1), ** = Correlation
is significant at the 0.01 level, * = Correlation is significant at the 0.05 level
Kumar et al. Carbon Research (2025) 4:6 Page 13 of 18

with elevation. Similar to the agroforestry system, pH tree species in traditional agroforestry along the vari-
showed no significant relation with any of the studied ous elevations of the Uttarakhand Himalaya was also
characters (Fig. 5). reported by Kumar et al. (2021) and Singh et al. (2024).
Diversity is a function of the relative distribution of
4 Discussion individuals among species and is a key aspect of ecosys-
Tree-crop components of agroforestry systems interact tems. It is influenced by various factors including long-
in terms of responses such as changes in soil fertility, term community stability and evolutionary time as well
microclimate changes, and resource availability (nutri- as microclimate and macroclimate heterogeneity (Sahoo
ent, light, and water) and utilization (Rao et al. 1997). et al. 2021). The middle elevation stood out in terms of
Ecological interactions between trees and crops are diversity, with the highest values observed for Shannon,
beneficial for three important reasons: (i) trees have Simpson Diversity Index, and Plieou Index of evenness,
a beneficial impact on soil fertility through greater due to the occurrence of maximum number of trees in
production of organic matter and nutrient recycling this elevational range. The calculated values of diversity
(Young 1986). Trees act as nutrient pumps, i.e., extract- parameters, i.e., Simpson’s and Shannon’s diversity index,
ing nutrients from deeper depths and making them Menhinick’s index, Plieou index of evenness, Whittaker
available to shallow-rooted crops in the process of recy- β diversity, and Marglef ’s index, are align with or are
cling (Yamoah et al. 1986). (ii) A tree-crop combination comparable to findings from other studies, including
increases the production of biomass, as more water those by Deb et al. (2008), Gairola et al. (2011) and Sahoo
and nutrients can be absorbed due to the different et al. (2021). Maximum tree density occurred at the mid-
rooting depths (Huxley et al. 1989). (iii) The presence dle elevation which can be attributed to various factors
of trees serves as a defence buffer against soil erosion like soil type, moisture availability, historical land-use
and is also used as shelter and windbreaks (Wiersum practices, and encouragement of agroforestry practices
1984). Several workers have observed positive interac- in regions where people reside in abundance. This range
tions where the tree-crop components of agroforestry might offer more suitable habitats for a broader range of
resulted in greater economic returns per hectare (Wise species, leading to higher tree density. The abundance of
and Cacho 2005). Negative interactions are not uncom- trees in the upper elevation range may be influenced by
mon as well (Fikreyesus et al. 2011); trees and crops several ecological factors, including climate, temperature,
both compete for light where the tree crown causing moisture, precipitation, and soil quality. These conditions
shade that harms the productivity of the annual crops might favour specific tree species in this elevation range,
(Srinivasan et al. 1990). resulting in a higher number of trees. The recorded val-
ues of tree diversity from the present study were less than
4.1 Tree diversity and Importance Value Index (IVI) those reported by Gairola et al. (2011) for certain aspects
The study recorded a total of six tree species in agrofor- of basal area and tree density. These differences could be
estry systems of the study area, and distribution of their attributed to site-specific variations, local ecological con-
dominance across different elevations revealed interest- ditions, and the composition of tree species in the study
ing patterns. At lower, middle, and upper elevations, G. area.
optiva, M. azedarach, and Q. leucotrichophora exhibited
the highest IVI, respectively. This suggests that these spe- 4.2 Tree carbon stock
cies might have the highest combined influence regard- Agroforestry systems present a valuable opportu-
ing the maximum basal area, frequency, and density nity for sequestering atmospheric carbon, contrib-
within their respective elevational ranges. Ecological uting to mitigating climate change impacts (Paudel
studies including phytosociology and community struc- et al. 2017). In this study, the traditional agroforestry
ture are largely confined to the Q. leucotrichophora-based system at lower elevations sequestered a relatively
agroforestry system which is broadly distributed between smaller amount of carbon than at medium and higher
1400–2200 m elevational ranges in Uttarakhand Himala- elevational ranges. This can be explained by the fact
yas (Singh and Rawat 2012; Nautiyal et al. 2020; Kumar that the agroforestry systems at lower elevations have
et al. 2021). The distribution range of G. optiva dominant smaller tree basal areas and diameters than those at
traditional agroforestry system varies between 500m middle and higher elevations that exhibit vigorous
and 2500 m (Semwal et al. 2002; Murasing et al. 2022), growth. This finding is in consistent with the results
while in Northwest Himalaya it is common in mid-hills of Sahoo et al. (2021), who highlighted a positive cor-
and foothills areas. A more or less similar association of relation among tree diameter, basal area, and carbon
sequestration potential in ecosystems. Agroforestry
Kumar et al. Carbon Research (2025) 4:6 Page 14 of 18

systems at the elevation of 1500–2000 m in the Garh- 11.57 Mg ­ha−1 ­yr−1) in forest ecosystems at elevations of
wal Himalaya sequestered more carbon than those at 700–2200 m in the Garhwal Himalayan region, which is
higher elevations of 2000–2500 m (Kumar et al. 2021). approximately at par with the findings of present inves-
This observation is likely influenced by number of tigation. The differences in net oxygen production can be
tree species, tree density, its diameter and total basal attributed to several factors, including location, tree spe-
area. In the present study, the carbon stock values for cies, net carbon sequestration, tree age and diameter. Of
traditional agroforestry systems are aligned with the all the tree species studied, Q. leucotrichophora had the
findings of Kumar et al. (2021), who have reported the highest oxygen production potential as it is widely dis-
carbon sequestration values of 36.94 Mg h ­ a−1 at an tributed in the study sites and also had the largest basal
elevation of 1400 - 1800 m and 38.84 mg h ­ a−1 at 1800 area and the highest carbon storage potential among all
- 2200 m in Quercus leucotrichophora dominated tra- the tree species in the study sites.
ditional agrisilviculture systems in the Tehri Garhwal
district of Uttarakhand Himalaya. At different eleva- 4.4 Soil properties
tions, P. pashia, M. azederach and Q. leucotrichophora 4.4.1 Soil pH and bulk density
have the highest carbon stocks due to their domi- The soil bulk density is an indicator of soil pore space and
nance in the respective elevational ranges. Studies by compaction that depends on soil texture, structure, poros-
Khan et al. (2020), Kumar et al. (2021) and Sahoo et al. ity, and organic matter content as well as the freezing and
(2021) show the positive correlation of carbon stock thawing process, plant roots, and soil micro-organisms
with tree diameter and basal area. Other studies (Man- (Unger 1991; Chen et al. 1998; Goss and Ulery 2022). In
dal et al. 2016; Khan et al. 2020) emphasise how car- the present study, soil bulk density ranged between 1.06 to
bon stock varies with tree size and species. Consistent 1.31 g ­cc−1 at lower elevations, 1.04 to 1.29 g ­cc−1 at mid-
results are noted by Kumar et al. (2021) and Singh et al. dle elevations, and 1.01 to 1.21 g ­cc−1 at upper elevations.
(2024). Moreover, net carbon sequestration is more The soil at higher elevations has a coarser organic matter
similar in the middle and higher elevations than in the structure and has enriched the gaps with organic carbon, as
800–1200 m elevation. This could be linked to factors shown by the lower soil bulk density value at higher eleva-
such as tree age and diameter (Kanime et al. 2013). tions (Kumar et al. 2021). Soil bulk density was higher at
lower elevations than at middle and upper elevations, which
is consistent with the findings of Mishra et al. (2018) and
4.3 Trees oxygen production Kumar et al. (2021). The lower value of soil bulk density at
Plants are the main source of oxygen and are important the 1600–2000 m elevational range can be explained by the
reservoir of carbon dioxide. In the present study, total high SOC and SOCS, as appeared by the inverse relation-
oxygen production (TOP) was maximum in the upper ship with soil bulk density. The bulk density also varied with
sites (101.42 Mg ­ha−1), indicating overall higher oxygen soil depth, as the lowest (1.01 g ­cc−1) was recorded in the
generation in both above and below-ground tree bio- top layer (0–15 cm), which gradually increased with soil
mass, followed by the middle sites (90.12 Mg ­ha−1) and depth and the highest (1.31 g ­cc−1) was observed at soil
the lower sites (81.31 Mg ­ha−1). The varying oxygen pro- depth of 30–45 cm. The subsurface soil has less organic
duction potential among trees at different elevations matter; aggregation, pore space, and root penetration are
may stem from factors like tree density, biomass, diam- more compacted than the surface layer (USDA 2008). The
eter, age, health, architecture, and maintenance practices, lower bulk density of soil observed in the 0–15 cm soil layer
including agroforestry systems. The comparison of find- is attributed to the accumulation of more organic matter in
ings with earlier research indicates that the oxygen pro- the topsoil compared to the subsoil (Manpoong et al. 2021).
duction values in this study are lower than those reported Studies in Northeast India (Manpoong et al. 2020) and the
by Keerthika and Chavan (2022). In this study, overall Garhwal Himalayas (Kumar et al. 2021) consistently show
net oxygen production (total of both above and below that soil bulk density tends to increase with soil depth.
ground) ranged from 24.12 to 28.64 Mg ­ha−1 ­yr−1, with Compared to traditional agroforestry systems, sole cropping
different tree species in the agroforestry system. Singh systems have a relatively higher bulk density. This difference
et al. (2024) found that O­ 2 production ranged from 50.56 could be attributed to several factors including increased
to 72.10 Mg ­ha−1 (3.00 to 4.02 Mg ­ha−1 ­yr−1) in tradi- soil compaction, more intense crop and tree root systems,
tional agroforestry and 58.18 to 264.10 Mg ­ha−1 (2.32 to and greater addition of soil organic matter from increased
litter decomposition in agroforestry systems. Abera and
Kumar et al. Carbon Research (2025) 4:6 Page 15 of 18

Bekele (2019) also reported that the lower soil bulk density and Sharma (2014) who have reported that SOCS ranged
in agroforestry was due to higher fine roots in tree vicinity from 25.92 to 51.71 Mg C h ­ a−1 in different agroforestry
and greater accumulation of organic matter under tree can- practices of Uttarakhand. The studied agroforestry
opy by litter fall. Such an inverse relationship between bulk system had greater SOC and SOCS due to increased
density and SOC has also been demonstrated by Mhawish accumulation of leaf litter and plant root dynamics in
(2015). tree-based agroforestry systems. The agroforestry soils
Agroforestry systems typically include trees and crops, had a higher microbial biomass and enzymatic activ-
which can lead to increased organic matter input through ity than conventional systems (Araujo et al. 2012). This
litterfall and root exudates. Trees often have deep and could contribute to an accelerated decomposition rate
extensive root systems that can reach deeper soil layers, and increased storage of organic carbon in the agrofor-
influencing pH through nutrient uptake, organic matter estry land. In complex land-use systems such as agro-
decomposition, and mineral weathering. In the present forestry, returning more litter and/or cutting biomass to
study, the higher soil pH values observed in the agrofor- the soil and root degradation enhances the abundance
estry system, especially in the middle elevation range, of organic matter (Rahangdale and Pathak 2016). The
might be due to the cumulative effect of tree roots on soil amount of SOC and carbon stock typically decreases
processes and pH regulation. In the sole cropping sys- with soil depth in agricultural fields and agroforestry
tem, the decline in pH with increasing depth at the mid- systems. The fact that leaf litter continuously accumu-
dle elevation range might be related to soil acidification lates in the uppermost layer, which is further decom-
processes. Leaching of base cations (e.g., calcium and posed and enriched, can explain the increased SOC and
magnesium) due to intensive agricultural practices and SOCS in the uppermost soil layer (0–15 cm), which is
nutrient uptake can lead to soil acidification, especially at also supported by the findings of Babu et al. (2020) and
deeper levels where buffering capacity is reduced. Ferti- Kumar et al. (2021).
lizer application, irrigation, and other management prac-
tices can impact soil pH. Rana et al. (2020) reported the 4.5 Productivity of rice under the agroforestry system
pH value ranged 6.17–6.21 in G. optiva in Tehri Garhwal The yield of rice varied with concerning elevation and
(Uttarakhand). They have found acidic soil pH in all three site and also with sole cropping system (control condi-
elevations. Kar et al. (2019) also observed the soil pH of tion) and traditional agroforestry system. In the present
6.80 to 6.89 in G. optiva–based agroforestry in the middle study, grain yield under the agroforestry system ranged
hills of Himachal Pradesh. 11.48–17.49 q ­ha−1 and in the control condition it ranged
13.71–21.05 q ­ha−1. Sati and Kumar (2023) reported an
4.4.2 Soil organic carbon and soil organic carbon stock 11.38 q ­ha−1 average yield of rice in the Garhwal Himala-
In the present study, the percentage of SOC tends to yan region of India. It was observed that rice productivity
decrease from the topsoil to subsurface soil in studied was lower in agroforestry conditions as compared to the
agroforestry and the sole cropping systems. The high- control due to various factors such as tree crop interac-
est SOC content in the uppermost soil layers (0–15 cm) tion, the allelopathic effect of trees on rice, and compe-
reflects the highest root density and increased biologi- tition for sunlight, water or soil moisture, and nutrients
cal activity (Ghosh et al. 2020; Soinne et al. 2024). SOC among trees and rice crops. Higher crop yield in sole
decreased with higher soil depth, which is probably due cropping (control) system for all the elevational range is
to the accumulation of more organic matter from vegeta- due to the absence of trees which resulted in no tree-crop
tion in the topsoil (Mishra et al. 2018). In addition, the interaction, and less competition for light, space, and
agroforestry system generally has higher SOC content nutrients (Kumar et al. 2021). In the Garhwal Himalaya,
than the sole cropping system at different elevations and various tree-crop interactions have been observed in dif-
soil depths, which can be attributed to the accumulation ferent agroforestry systems. These interactions, such as
of litter, residues, debris, and other plant parts from culti- the allelopathic effects of tree leaves, bark, and weeds on
vated crops and trees, as well as changes in the microcli- agricultural crops have been reported by Bhatt and Singh
mate (Kumar et al. 2021). The soil organic matter content (2009) and Padu et al. (2023). These effects pose a threat
in the present study is also consistent with the study of to seed germination, root and shoot length, dry weight,
Lepcha and Devi (2020). nutrient levels, and chlorophyll content of the crops.
It has been observed that soil organic carbon stock Bhatt and Singh (2009) also reported that the different
(SOCS) is correlated with both bulk density and SOC agroforestry tree species inhibit the growth and develop-
content. The value of SOCS in the reported agroforestry ment of Oriza sativa due to allelopathic effects. Recorded
systems ranged between 40.72 Mg C ­ha−1 and 55.60 Mg rice productivity in this investigation was in line with the
C ­ha−1, which is in line with the observation of Gupta
Kumar et al. Carbon Research (2025) 4:6 Page 16 of 18

productivity estimated by Chauhan et al. (2012), Verma for their valuable comments and suggestions which have immensely
improved the quality of the manuscript.
and Rana (2014), and Newaj et al. (2020). The findings
of the current study on grain and straw yield as well as Authors’ contributions
the rice (Oryza sativa) harvest index were similar to the All authors contributed to the study’s conception and design. Conceiving
and designing the research was performed by Vinod Prasad Khanduri and
results found by Tripathi et al. (2005) and Verma and Bhupendra Singh. Material preparation, data collection, and analysis were
Rana (2014). Rice yield was reported to be higher at lower performed by Sachin Kumar. The first draft of the manuscript was written by
elevations than at mid and upper elevations. This obser- Sachin Kumar and Sandeep Kumar. Revising and rewriting the manuscript
was performed by Sandeep Kumar, Vinod Prasad Khanduri, Bhupendra Singh,
vation is well supported by the findings of Hairman- Reena Joshi, Manoj Kumar Riyal, Deepa Rawat and Kewat Sanjay Kumar and all
sis et al. (2017), Liu et al. (2014) and Sanou et al. (2012) authors commented on previous versions of the manuscript. All authors read
and emphasizes the influence of environmental factors and approved the final manuscript.
on rice yields at different elevations. Nishad et al. (2018) Funding
also reported that the slow physiological process of rice is This research received no external funding.
involved in lower grain production and yield due to direct
Data availability
exposure to cold temperatures at higher elevations, as the The datasets used or analyzed during the current study are available from the
earth’s temperature normally falls with increasing eleva- corresponding author upon reasonable request.
tion. Joshi et al. (2018) reported that the mean annual
temperature lapse rate in Western Himalaya is − 0.53 °C Declarations
per 100 m elevation increase. This observation is at par
Competing interests
with the findings of Sanou et al. (2012), Liu et al. (2014) The authors declare no competing interests.
and Hairmansis et al. (2017), further emphasizing the
influence of environmental factors on rice yield across Author details
1
College of Forestry, Veer Chandra Singh Garhwali Uttarakhand Univer-
different elevations. sity of Horticulture and Forestry, Ranichauri, Uttarakhand 249 199, India.
2
Department of Forestry, Mizoram University, Aizawl, Mizoram 796 004, India.
3
Department of Botany, University of Allahabad, Prayagraj, Uttar Pradesh 211
5 Conclusion 002, India.
The study indicates an elevation-dependent variation in
tree density, diversity, carbon sequestration, and oxygen Received: 1 April 2024 Revised: 17 September 2024 Accepted: 7 October
2024
production capacity of the traditional agroforestry sys-
tem in the Himalayas. This elevation-dependent trend
extends to carbon stock and net carbon sequestration,
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