Trees, Forests and People 11 (2023) 100371

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Trees, Forests and People

journal homepage: www.sciencedirect.com/journal/trees-forests-and-people

Tree community structure, carbon stocks and regeneration status of

disturbed lowland tropical rain forests of Assam, India☆
Manish Kuntal Buragohain a, c, Ashaq Ahmad Dar a, c, Kanda Naveen Babu a, b,
Narayanaswamy Parthasarathy a, *
a
Department of Ecology and Environmental Sciences, School of life Sciences, Pondicherry University, Puducherry, 605 014, India
b
Department of Ecology, French Institute of Pondicherry, St. Louis Street, Puducherry, 605001, India

A R T I C L E I N F O A B S T R A C T

Keywords: Tropical rain forests are global hotspots of biodiversity and key climate change regulators. Despite the decades of
Carbon loss conservation efforts, tropical forests are increasingly under continuous threat from human activities. Therefore,
Conservation understanding the impact of anthropogenic disturbances on biodiversity is necessary for better conservation and
Disturbance
management. This study focused on a tropical rainforest located in Kakoi reserve, Assam, in the northeastern
Eastern himalaya
Himalayas, aiming to compare tree species composition, diversity, carbon stocks, and regenerating status in two
Species diversity
Tropical rainforest forest sites experiencing disturbance regimes. A total of 55 tree species representing 36 genera and 26 families
were recorded in the study area. The tree species richness was slightly higher in moderately disturbed (MD)
Dirgha than in the highly disturbed (HD) Kakoi-Rajgarh forest. Tree density and basal area were 582 and 446
individuals ha− 1, 38.43, and 32.63 m2 ha− 1, respectively, in MD-Dirgha and HD-Kakoi-Rajgarh. Forest stand
structure followed a typical reverse J-shaped trend in both sites, with a lesser density of large girth-class (>330
cm girth at breast height) individuals in HD-Kakoi-Rajgarh, reflecting the influence of past onsite disturbances.
The analysis of the regeneration strata depicted that seedling and sapling density was 1.8 and 1.4 times lower in
HD-Kakoi-Rajgarh compared to MD-Dirgha. Above-ground biomass and total carbon stocks were maximum in
MD-Dirgha, followed by HD-Kakoi-Rajgarh. About 63.64% of the recorded tree species, including Critically
Endangered Vatica lanceifolia are under IUCN Red List Categories, thus stressing the need for conservation. Our
findings suggest that conserving species diversity and carbon reserves in tropical forests would principally
depend on minimizing forest loss and degradation.

1. Introduction other environmental changes, which has resulted in a significant

The importance of tropical ecosystems for global biodiversity and
their critical role in sustaining essential ecological functions have been
invariably emphasized (Alkama and Cescatti, 2016) and acknowledged
in international agreements (UNFCCC, 2015). Tropical forests being
complex and heterogenous habitats are rapidly fragmented (Laurance
et al., 2011), degraded, and cleared for development, creating a severe
danger to global biodiversity and ecosystem services (Barlow et al.,
2016). The floristic composition and distribution across tropics are
determined by topography, local climatic regime, and edaphic features
(Qin et al., 2019). Over 50% of the natural tropical forest cover has
already been destroyed (FAO, 2010) as a result of land-use practices and
biodiversity loss (Gibson et al., 2011) and substantial greenhouse gas
emissions (Metz et al., 2007). Deforestation alone is estimated to have
degraded more than 20% of the total area between 2000 and 2005, with
a net loss of 8 million ha per year (Asner et al., 2009). These
human-caused disruptions consequently impact the local species pool
and capacity to store/sequester carbon. Under the climate change
perspective, the carbon reserves in forests have reduced by 11 Gt glob­
ally during the last 25 years (FAO, 2015). To address these aforemen­
tioned global challenges at the regional scale, investigating floristic
diversity and drivers governing regional patterns is essential for main­
taining and sustainable use of biodiversity (Devictor et al., 2010).
Disturbances (natural as well as human-driven) have an overall

☆
This article is part of a special issue entitled: “Tropical Forestry: Conservation, Management and Utilization” published at the journal Trees, Forests and People.
* Corresponding author.
E-mail address: nparthasarathypu@gmail.com (N. Parthasarathy).
c
Both authors contributed equally.

https://doi.org/10.1016/j.tfp.2023.100371
Received 5 November 2022; Received in revised form 3 January 2023; Accepted 4 January 2023
Available online 5 January 2023
2666-7193/© 2023 The Author(s). Published by Elsevier B.V. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-
nc-nd/4.0/).
M.K. Buragohain et al.
impact on forest species abundance, diversity, structure, and regenera­
tion patterns in general (Dutta and Devi, 2013). Regeneration strata in
forest ecosystems facilitate the establishment and development of
indigenous flora, improving the stability, adaptability, and species het­
erogeneity (Liira et al., 2011; Danková and Saniga, 2013). The recovery
rate for structural characteristics such as canopy structure, tree height,
and basal cover differed depending on intensity, frequency, and scale of
disturbance that occurred (de Carvalho et al., 2017). Following distur­
bances that severely impact soil, as well as above-ground biota, the
regeneration of the forest is substantially delayed. By far, the Neotropics
has received greater attention in research on tropical forest regeneration
over the last 20 years, that has skewed our knowledge of seedling
regeneration patterns in rainforests. Limited research comes from the
tropics of Asia or, to a greater extent, Africa, two extremely diverse
tropical biomass areas with very distinct biogeographies (Ashton et al.,
2018).
Tropical forests play a key role in global carbon balance, contrib­
uting 70% to global live biomass accounting for ~250 Pg carbon (Pan
et al., 2011). In tropical forests, carbon stocks are governed by
human-driven activities, characteristics of the tree community, and
environmental factors (Pyles et al., 2022). Chronic human disturbances
cause abiotic and biotic changes that influence the forest structure and
hence the ability to store carbon (Osuri et al., 2014). These variations
include tree density and basal area reduction, which lowers carbon
storage, as well as decreased abundances of high wood-density tree
species, which decreases biomass per unit volume (de Paula et al.,
2011). Gross tropical deforestation outflows are estimated to be 2.9 Gt C
year− 1, with human-induced disturbances as the major drivers in trop­
ical forests contributing 8% to 15% to GHG emissions (Houghton et al.,
2015). Despite regional variations, emissions from logging and burning
wood for fuel make up the majority of emissions caused by forest
degradation (Köhl et al., 2015). The estimated annual CO2 emissions due
to forest degradation are 2.1 Gt C, of which 53% were attributable to the
extraction of timber, 30% to firewood, and 17% to forest fires (Pearson
et al., 2017). Even though forest degradation was included in interna­
tional climate change mitigation policies such as reducing emissions
from deforestation and forest degradation (REDD+) mechanism in 2008
(UNFCCC, 2008), the magnitude and longevity of biomass loss in forests
with varied disturbance regimes are still mostly obscure (de Andrade
et al., 2017). By encouraging the management and regeneration of for­
ests, REDD+ seeks to increase global carbon sinks (UNFCC, 2018). The
1.5–2 ◦ C objective set out by the Paris Agreement cannot be achieved
without REDD+ and conservation initiatives aiming at enhancing forest
carbon reserves in developing nations and sustainable management of
forested areas (Paris Agreement, 2015). Such climate-focused initiatives
provide side advantages for tropical forest biodiversity protection
(Strassburg et al., 2010; Gilroy et al., 2014).
Mountain ecosystems have an important role in the formation of
mountain hotspots, as acknowledged by the Convention on Biological
Diversity (CBD) long ago (CBD, 2004). In fact, Sustainable Development
Goal (SDG) 15 Target 4 of the 2030 agenda incorporates protection of
mountain biodiversity as a goal, making it a global objective (Makino
et al., 2020). The easternmost (northeast) region of India is
well-acknowledged for its ecological diversity. Northeast states repre­
sent 25% of the country’s forest area, constituting over 17 million ha.
With 36% forest cover, Assam is one of the hilly states with great
ecological significance in the Himalayan biodiversity hotspot (Forest
Survey of India, 2021). Tropical forests of Assam support considerable
species richness, likely due to prolonged isolation and difficult terrain.
While human demand to exploit their resources for economic progress is
growing (Borah and Garkoti, 2011), understanding how diversity and
ecosystems will adapt is imperative in the context of the environmental
changes being triggered by climate change and human-induced
2
Trees, Forests and People 11 (2023) 100371
constraints. The plant diversity in northeast Assam has been well
documented, but scientific evidence on disturbances influencing plant
diversity in the northeast tropical rainforest has been incomprehensively
investigated (Borah et al., 2014; Gogoi et al., 2022). Therefore, this
forest consequently necessitates an urgent investigation to comprehend
the extent and/or effect of human disturbance on their
physico-ecological health, which will additionally be beneficial in
developing practical approaches for the sustainability of this forest be­
sides forests facing similar challenges. In this research, we investigated
species composition, regeneration strata, and productivity in differently
disturbed tropical rainforest sites of Kokoi reserve, Assam. We hypoth­
esized that anthropogenic interventions would result in floristic differ­
entiation and limit biomass and carbon reserves in the highly disturbed
(HD-) forest site (Kakoi-Rajgarh) compared to the moderately disturbed
(MD-) site (Dirgha).
2. Methodology
2.1. Study area
The present study was conducted in a lowland rain forest of Kakoi
reserve forest, one among the four reserve forests of Lakhimpur district
of Assam state; it is primarily composed of semi-evergreen forest. At
~150 elevation, the forest lies between 27◦ 18′ 51′′ and 27◦ 25′ 17′′ N and
93◦ 59′ 09′′ and 94◦ 09′ 32′′ E, with a surface area of about 4415.03 ha on
moderate slope (12–16◦ ) facing northern aspect (Fig. 1; Table 1). The
undulating terrain is characterized by the deposits of pebbles and
boulders of gneissic and quartzitic rocks in a matrix of fine sand, silt, and
clay. The soil in Kakoi reserve forest is predominantly alluvial with a
coarse loamy texture. The average monthly rainfall in Kakoi reserve
forest is 3200 mm (https://worldclim.org/). Typically, rainfall occurs
for about 125 days a year, with July being the wettest month (Statistical
Handbook of Assam, 2014). The average annual temperature of the
study area is 23.8 ◦ C. The relative humidity is within the range of 74 to
89%, with an average of 81%.
2.2. Plot stratification and sampling design
A reconnaissance survey was conducted during winter season in the
month of February 2021 to get an overview of accessibility, terrain,
species composition, disturbance and overall structure of Kakoi Reserve
Forest. Five beets (an administrative sub-unit of a forest range) namely
Diju, Joyhing, Kakoi Rajgarh, Dirgha, and Boginodi were identified.
Based on the reconnaissance survey (by exploring all the five beets), the
entire reserve forest was divided into two blocks [A (Heavily disturbed) -
Diju, Joyhing and Kakoi Rajgarh and B (Moderately disturbed) - Dirgha,
and Boginodi] considering the disturbance impact. Two beets (Diju and
Joyhing) had heavy elephant movements, making it impossible to
consider for establishing the plot. Hence Kakoi-Rajgarh was considered
from block A and Digha from block B for the final inventory. Disturbance
parameters were estimated within a 300 m × 300 m area demarcated at
each block, starting from the southern boundary. Five disturbance pa­
rameters, viz., fuel wood collection, timber logging, forest fire, wind
throw, and elephant felling, were identified and quantitatively assessed
by counting the trees with cut branches, logged trees, trees with fire
marks, wind-fell trees, and trees broken by elephants. In addition, can­
opy cover percentage an important predictor of disturbance (Bartels
et al., 2016), was retrieved for both the sites (Fig. A.1) from high-­
resolution (30 m) global forest cover maps (Hansen et al., 2013). The
disturbance percentage (%) of each parameter (excluding canopy cover
(%)) was calculated in relation to the total observation of all disturbance
parameters (Eq. (1)).
M.K. Buragohain et al. Trees, Forests and People 11 (2023) 100371

Table 1
Topography and climate of the studied tropical rainforest sites of Kakoi reserve, Assam.
Plots Mean Annual Temperature ( ◦ C) Mean Annual Precipitation (mm) Slope (o) Aspect (o) Elevation (m)

Dirgha 23.5 3234 16 N 156

Kakoi Rajgarh 23.6 3219 12 N 153

Number of observations of a disturbance parameter

Disturbance percentage (%) = × 100 (1)
Total number of observations of all disturbance parameters

extrapolation sampling curves were generated using the Hill numbers

Based on the nature and degree of disturbance, Dirgha (27◦ 22′ 23′′ N,
94 05′ 08′′ E) was designated as moderately disturbed, while Kakoi-
◦
Rajgarh (27◦ 20′ 42′′ N, 94◦ 03′ 08′′ E) was assigned as a highly
disturbed area (Table 2).
Vegetation sampling and measurements were conducted in the study
sites (Dirgha and Kakoi-Rajgarh) using standard quadrat method (Curtis
and MacIntosh, 1950). Two 100 m × 100 m (1-ha) sample plots (one at
each site) were laid with in the 300 m × 300 m compartment and further
subdivided into 10 m × 10 m size subsample quadrats to facilitate the
vegetation assessment (Fig. 1). Within each sub-quadrat, all trees with a
girth size of ≥30 cm at breast height (1.3 m above ground level) were
identified and measured. In this study, adult trees were defined as
woody plants with GBH ≥ 30 cm, saplings as woody plants with GBH ≥
3 cm to < 30 cm, and seedlings as woody plants with GBH < 3 cm and a
height ≥ 20 cm. The seedlings having <20 cm height were considered
ephemeral and not counted (Dutta and Devi, 2013). For buttressed trees,
girth measurements were taken above the point of measurement. In the
case of multi-stems, the girths were measured independently. Tree
seedlings and saplings were counted from twenty-five nested 5 m × 5 m
sub-subsample quadrats laid in alternative 10 m × 10 m subsample
quadrats and rows (Fig. 1). All the species were identified in the field
with the help of published literature on local and regional flora, viz.,
Flora of Assam (Kanjilal et al., 1934–1940), Plant diversity of Assam: a
checklist of Angiosperms and Gymnosperms (Barooah and Ahmed,
2014) and subsequently confirmed with experts. Taxonomic details of
the identified tree species were updated following The World Flora
Online (WFO; http://www.worldfloraonline.org/; Accessed on: 08 Jun
2022).
2.3. Data analyses
The vegetation data of the quadrats were quantitatively analyzed for
density, frequency, and abundance (Curtis and McIntosh, 1950). The
important value index (IVI) was calculated using equations provided by
Curtis and McIntosh (1950) in "BiodiversityR" library (https://CRAN.R-p
roject.org/package=BiodiversityR). Diversity indices (Shannon-Wiener,
Simpson, and Pielou evenness) were computed using "vegan" package
(https://CRAN.R-project.org/package=vegan). Rarefaction and
Table 2
Quantitative assessment of disturbances recorded in tropical rain forest sites of
Kakoi reserve, Assam.
Disturbance Dirgha Kakoi-Rajgarh
Forest fire (trees with fire marks;%) 5 (0.08) 0 (0.00)
Wind-felling (trees broken by wind;%) 12 (0.19) 2 (0.01)
Elephant felling (trees broken by elephants;%) 34 (0.53) 12 (0.06)
Lopping (trees with cut branches;%) 11 (0.17) 107 (0.53)
Stumps (%) 2 (0.03) 80 (0.40)
Canopy cover (%) 75 – 97 45 – 85
Total 64 201
family based on abundance-incidence data for diversity comparisons
(Chao et al., 2014). The bootstrap method was used to determine the
95% probability level with 200 randomizations. These investigations
were conducted in "iNEXT" library (https://CRAN.R-project.org/p
ackage=iNEXT). The abundance/frequency (A/F) ratio was used to
determine and classify the species dispersion patterns, which were
divided into three categories: regular/uniform (<0.025), random dis­
tribution (0.025–0.050), and contagious/clumped (>0.050; Whitford,
1949). The conservation and endemic status of the recorded species
were retrieved from IUCN Red List database (https://www.iucnredlist.
org/) and available key literature (Chaudhuri, 2007; Borah et al.,
2020). All the statistical analyses were worked out in R software (R Core
Team, 2021). The status of regeneration of species was determined
based on the density of seedlings, saplings, and adults of tree species as
good, fair, poor, no, and new (Table 3; modified from Khan et al., 1987;
Shankar, 2001; Khumbongmayum et al., 2006).
Using the DBH along with wood density acquired from the global
database (Zanne et al., 2009), individual-level above-ground biomass
(AGB) was computed following Chave et al. (2005). Below-ground
biomass (BGB) was estimated by multiplying AGB with 0.26 (Ravin­
dranath and Ostwald, 2008), which was combined with AGB to estimate
the total biomass in each plot. Carbon Stock (CS) of trees was computed
to be 50% of the total biomass (IPCC, 2005; Eq. (2)).
(AGBest + BGBest )
CSest = (2)
2
3. Results
3.1. Tree species composition and diversity
A total enumeration of 1028 tree individuals accounting 55 species
belonging to 36 genera and 26 families, were recorded from moderately
disturbed (MD–) Dirgha and highly disturbed (HD–) Kakoi-Rajgarh
forest sites in Kakoi reserve forest. Dirgha site accounted for all the lis­
ted species in Kakoi reserve forest, while seven species, which occurred
in Dirgha, were completely absent from Kakoi-Rajgarh, indicating
greater species richness in MD–Dirgha than HD–Kakoi-Rajgarh study
site (Table 4; Table A.1, Supplementary Material). Species contribution
by families was disproportionate, with seven families accounting for
more than half (54.55%) of the species, while the remaining belonged to
19 families, 13 of which were monotypic, representing single species.
The taxonomically diverse plant families included Lauraceae and Mag­
noliaceae, with six species (10.91%) each. The families, Meliaceae,
Lamiaceae, Phyllanthaceae and Anacardiaceae contributed to the major
floristic differentiation between the sites (Table A.1, Supplementary
Material).
The H′ between the sites ranged from 3.55 (in MD–Dirgha) to 3.68
(in HD–Kakoi-Rajgarh), suggesting both sites were almost equally
diverse. The D was 0.05 and 0.03 in Dirgha and Kakoi-Rajgarh, while 1-

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M.K. Buragohain et al. Trees, Forests and People 11 (2023) 100371

Fig. 1. Map showing the location of the two study plots (Plot A – Dirgha; Plot B – Kakoi-Rajgarh) and the sampling design in tropical rainforests of Kakoi Reserve,
Assam, India.

1.62–31.20 and 0.81–44.21 in HD–Kakoi-Rajgarh and MD–Dirgha,

Table 3
respectively (Table S3). Four dominant and co-dominant tree species,
Regeneration status of the of recorded tree species in two tropical rainforest sites
Castanopsis indica (31.20 IVI), Mesua ferrea (17.85 IVI), Magnolia griffithii
of Kakoi reserve, Assam.
(15.24 IVI), and Magnolia pealiana (10.21 IVI), together shared
Population trend Regeneration status approximately 25% of the cumulative IVI in HD–Kakoi-Rajgarh, while
Seedling Sapling Adults the corresponding figure for MD–Dirgha was much greater (37%),
✓ > ✓ > ✓ Good contributed by Mesua ferrea (44.21 IVI), Castanopsis indica (26.34 IVI),
✓ > or ≤ ✓ ≤ ✓ Fair Magnolia griffithii (21.89 IVI), and Mesua assamica (17.29 IVI).
✓ ≤ ✓ > ✓
⨯ ✓ <, > or = ✓ Poor
⨯ ⨯ ✓ No 3.2. Tree density and basal area
✓ or/and ✓ ⨯ New
The MD-Dirgha exhibited 1.3 times greater density than HD-Kakoi-
Rajgarh (Table 4). The five tree species with the highest density were
Table 4 Mesua ferrea, Castanopsis indica, Magnolia griffithii, Mesua assamica and
Phytosociological and productivity attributes in two tropical rainforest sites of Canarium resiniferum across the study sites. Specifically, Mesua ferrea
Kakoi reserve, Assam. (84 stem ha− 1), followed by Castanopsis indica (59 stems ha− 1) and
Attribute MD-Dirgha HD-Kakoi-Rajgarh Magnolia griffithii (43 stems ha− 1), accounted more than 31% of tree
Tree diversity and composition
density. In Kakoi-Rajgarh, Castanopsis indica shared more than 9.4% of
Number of species/genera/families 55/36/26 48/30/24 total tree density at the site, followed by Mesua ferrea (6.1%) > Magnolia
Shannon diversity (H′ ) 3.55 3.68 griffithii (5.4%) > Magnolia pealiana (3.6%). The cumulative basal area
Simpson dominance (D) 0.05 0.03 across the two sites was 71.06 m2, with MD-Dirgha being the major
Simpson diversity (1-D) 0.95 0.97
contributor (54%). Mesua ferrea (7.04 m2 ha‒1) followed by Mesua
Pielou evenness (J) 0.89 0.95
Structure assamica (3.18 m2 ha‒1) and Castanopsis indica (3.02 m2 ha‒1) represent
Density (Ind. ha− 1) 582 446 highest basal area. In HD-Kakoi-Rajgarh, Castanopsis indica (13%)
Basal area (m2 ha− 1) 38.43 32.63 accounted for the highest basal area, followed by Mesua ferrea (6%),
Biomass and carbon stocks Magnolia griffithii (4.6%), and Canarium resiniferum (3.6%). In both the
Aboveground biomass (AGB; Mg ha− 1) 371.78 285.47
Belowground biomass (BGB; Mg ha− 1) 96.66 74.22
sites Lagerstroemia speciosa exhibit lower basal area. By family, Calo­
Total biomass (TB; Mg ha− 1) 468.45 359.69 phyllaceae in MD-Dirgha, in contrast to Fagaceae in HD-Kakoi-Rajgarh
Carbon stocks (TC; Mg ha− 1) 234.22 179.85 stocked the maximum basal area accounting 26.5% and 18.1% within
respective sites (Table A.2, Supplementary Material).

D was 0.95 and 0.97 for the same sites. Similarly, J was recorded to be
0.89 in Dirgha and 0.95 in Kakoi-Rajgarh (Table 4). However, HD–Ka­
koi-Rajgarh had the highest (1 = 39.60) first-order diversity (Shannon
exponential), whereas MD–Dirgha had the least (1 = 34.79). Similar
trends were observed for second order 2 (Inverse of Simpson), i.e.,
HD–Kakoi-Rajgarh (2 = 31.52) > MD–Dirgha (2 = 20.52; Fig. 2).
The importance value index (IVI) of tree species is in the range of
3.3. Forest stand structure and regeneration status
The distribution of tree density and basal area varied among the girth
classes between the two study sites (Fig. 3). The density of threshold
girth class (30–60 cm) contributed 49.6% (HD-Kakoi-Rajgarh) to 56.2%
(MD-Dirgha) of the total recorded tree density in the two sites. The
population structure corresponds to the reverse J-shaped trend in both

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M.K. Buragohain et al. Trees, Forests and People 11 (2023) 100371

Fig. 2. Observed (solid lines) and expected (extrapolated) curves of taxonomic diversity in HD–Kakoi-Rajgarh and MD–Dirgha, the two study sites of disturbed
tropical rainforests of Kakoi Reserve, Assam. Symbols •, ▴ and ■ represent Species richness (0), Shannon exponential (1) and Simpson’s reverse (2).

Fig. 3. Distribution of tree density and basal area across girth classes in two tropical rainforest sites of Kakoi reserve, Assam.

Fig. 4. Regeneration status by species percentage of recorded tree species in two tropical rainforest sites of Kakoi reserve, Assam.

sites, with a marginal increase in the 180–200 cm class in HD-Kakoi-
Rajgarh. However, the allocation of basal area among girth classes fol­
lowed an asymmetric trend with a maximum contribution (14.2% and
18%) from 180 to 210 cm and >330 cm individuals in HD-Kakoi-Rajgarh
and MD-Dirgha forest sites.
The regeneration potential of the sites was verified by analyzing the
adult tree density with regenerating seedlings and sapling populations of
species. Seedling density in HD-Kakoi-Rajgarh site (8352 Ind. ha− 1) was
1.8 times lower than MD-Dirgha site (15,120 Ind. ha− 1). Similarly,
sapling density was 1.4 times lower in HD-Kakoi-Rajgarh (4224 Ind.
ha− 1) than MD-Dirgha site (6000 Ind. ha− 1). The recorded seedling
densities in the studied forest sites were greater than saplings and adults,
thus, signifying an overall “Good” regeneration potential. Accordingly,
29.09% of species in MD-Dirgha and 37.50% in HD-Kakoi-Rajgarh,
forest sites showed “Good” regeneration. Also, 10.90% in MD-Dirgha
and 14.58% of species in HD-Kakoi-Rajgarh showed “Fair” regenera­
tion (Table A.1, Supplementary Material). About 25.25% and 29.17%
species in MD-Dirgha and HD-Kakoi-Rajgarh have “No” regeneration, i.

5
M.K. Buragohain et al.
e., no seedlings and saplings, indicating inhibited natural regeneration
(Fig. 4). The dominant tree species, M. ferrea presented best regenera­
tion potential in MD-Dirgha site, where it had excellent seedling density
(6160 Ind. ha− 1) with maximum sapling and adult densities of 1504 Ind.
ha− 1 and 84 Ind. ha− 1. In contrast, C. indica showed the highest regen­
eration potential in HD-Kakoi-Rajgarh, with 42 adult trees, 816 saplings,
and 2992 seedlings.
3.4. Distribution patterns (abundance/frequency ratio)
M. ferrea manifested the highest frequency of 55% in MD-Dirgha,
while Mallotus paniculatus, Lagerstroemia speciosa, and Bridelia retusa
had the lowest frequency of 3% each. In HD-Kakoi-Rajgarh, occurrence
of C. indica was frequent (33%), while G. arborea had the lowest fre­
quency of 2% (Table A.2, Supplementary Material). Abundance to fre­
quency ratio for the tree species indicated that none of the species across
the studied sites exhibit regular distribution. Majority of the species in
MD-Dirgha and HD-Kakoi-Rajgarh displayed contagious distribution
(94.55% and 97.92%) followed by random distribution (5.45% and
2.08%) patterns (Table A.2, Supplementary Material).
3.5. Forest biomass and carbon stock
The results revealed maximum cumulative tree biomass for MD-
Dirgha (468.45 Mg ha− 1) and minimum (359.69 Mg ha− 1) for HD-
Kakoi-Rajgarh with an average value of 414.07 Mg ha− 1. The AGB
and cabon stocks ranged from 371.78 Mg ha‒1 to 285.47 Mg ha− 1 and
234.22 Mg ha− 1 and 179.84 Mg ha− 1 in MD-Dirgha and HD-Kakoi-
Rajgarh (Table 4). The dominant species in MD-Dirgha and HD-Kakoi-
Rajgarh contributed more than 13% of total carbon (27.57% and
13.52%, respectively). The biomass storage capability of M. ferrea
(129.15 Mg ha− 1) was highest among 55 tree species, with a carbon
content of 64.58 Mg ha− 1 in MD-Dirgha. Mesua assamica with 24.79 Mg
ha− 1 and Terminalia chebula with 20.44 Mg ha− 1 carbon were co-
dominant species with respect to carbon storage. In HD-Kakoi-
Rajgarh, Castanopsis indica contributed the highest to the carbon stor­
age (24.319 Mg ha− 1), while the lowest carbon stocks of 0.35 Mg ha− 1
were contributed by Litsea salicifolia. Carbon storage by girth class
depicted that ≥330 cm and 180–21 cm in MD-Dirgha and HD-Kakoi-
Rajgarh stocked the maximum (64.29 Mg ha− 1 and 29.02 Mg C ha− 1)
Fig. 5. IUCN status (LC: Least concern; NE: Not Evaluated; EN: Endangered;
DD: Data Deficient; CR: Critically Endangered; VU: Vulnerable) of recorded tree
species in two tropical rainforest sites of Kakoi reserve, Assam.
6
Trees, Forests and People 11 (2023) 100371
carbon. However, girth classes with lowest carbon storage were
270–300 cm and ≥330 cm in MD-Dirgha and HD-Kakoi-Rajgarh.
3.6. Endemism and conservation status of tree species
The IUCN status of Kakoi reserve species revealed that 20 species
were classified into Not Evaluated (NE) category, and the remaining
63.64% species are included under Red List categories. Among them
Vatica lanceifolia (Roxb.) Blume is Critically Endangered (CR); one
species Magnolia pealiana King is Endangered (EN); one species Elaeo­
carpus prunifolius Wall. ex Müll.Berol. is Vulnerable (VU), two species
Hydnocarpus kurzii (King) Warb. and Magnolia griffithii Hook.f. &
Thomson falls under Data Deficient (DD) category, and 30 species are
Least Concern (LC; Fig. 5). In addition, two endemic species with
restricted distribution to Assam (Mesua assamica (King & Prain) Kos­
term.) and the northeast region including Assam (Phoebe goalparensis
Hutch.) were also recorded.
4. Discussion
Minimizing the decline of irreplaceable biodiversity and carbon re­
serves of tropical forests and ensuring the carbon-storing capability
represent a two-fold challenge as the human footprint in tropics con­
tinues to grow (Watson et al., 2018; Osuri et al., 2020). The present
study recorded 55 tree species in Kakoi reserve forest, with low diversity
in disturbed site. The overall diversity is almost comparable to the
number of species recorded in other tropical semi-evergreen and ever­
green forests of the region (Akash et al., 2019;Baidya et al., 2022;
Thakur et al., 2022). Moreover, the recorded species richness was within
the range of 20–223 species ha− 1 exhibited by tropical rainforests
worldwide (Whitmore, 1984). Considering the continuous
socio-ecological trends, sustaining biodiversity and ecosystem services
for long-term human well-being is a major challenge, especially in
mountain forests where there is considerable demographic stress and
consistent land-use shift (Gogoi and Sahoo, 2018). The Kakoi reserve
forest provides a number of widely preferred ecosystem services to forest
fringe human populations, which has led to extensive exploitation of
forest areas. In particular, the forest harbors valuable timber-yielding
species and species of medicinal, fodder, fiber, oil, and agroforestry
importance used by fringe communities (Fig. 6). In the Himalaya, forest
resources play a significant role in the livelihood security of distant and
poor rural populations (Chakraborty et al., 2018). The high commercial
demand leading to over-exploitation of the recorded species could be
responsible for reduced species richness in places considered to be
highly disturbed (Kakoi-Rajgarh). The challenge with this disturbance is
that forests are not given ample time to recuperate, since human
Fig. 6. Resource value of recorded tree species in two tropical rainforest sites of
Kakoi reserve forest, Assam.
M.K. Buragohain et al.
activities rarely cease (Singh, 1998). Lower diversity of woody species
may also be caused by increased road accessibility that disrupts moun­
tain ecosystems directly via habitat destruction (Foggin, 2016). There­
fore, lesser species diversity is usually a characteristic of heavily
disturbed forest ecosystems (Dumbrell et al., 2008; Hughes, 2012), as
reported in our study from HD-Kakoi-Rajgarh.
In forest ecosystems, diversity indices characterize the species di­
versity and density patterns which help us to acknowledge the local
processes driving species composition (Dar and Parthasarathy, 2022).
Most diversity indicators for the Kakoi reserve forest were greater in
MD-Dirgha, but lesser in HD-Kakoi-Rajgarh (Table 4). The rationale for
this outcome was that MD-Dirgha experienced lesser anthropogenic
stress, and the HD-Kakoi-Rajgarh site suffer strong human threats. The
results corroborate with the findings of Pandey et al. (2020) in Central
Himalaya, Deb et al. (2020) in tropical forests of Tripura, Ao et al.
(2021) in sub-tropical forests of Nagaland and other global tropical
ecosystems (Muhati et al., 2018; Bentsi-Enchill et al., 2022). Changes in
dominance (based on IVI) in the examined sites represented the most
evident floristic distinction caused by disturbance. M. ferrea is dominant
tree species in the MD-Dirgha as reported in other studies in undisturbed
forest types (Borah et al., 2014); on the other hand, C. indica has
dominated in disturbed site, i.e., HD-Kakoi-Rajgarh. The higher IVI
values and dominance of M. ferrea and C. indica, as well as other
co-dominant tree species, suggest that these species have a stronger
potential and competitive power to survive and outcompete other tree
species in the community. These tree species may have been more
competent depending on their morpho-physio-phenological traits
(Chazdon et al., 2010; Osazuwa-Peters et al., 2015). Similarly, domi­
nance differentiation by the degree of disturbance is also prominent at
family level.
The role of disturbance events in altering the community structure is
well-recognized (Fakhry et al., 2020). The elephants in the entire reserve
forest feed mainly on elephant apple (Dillenia indica fruits), which is
available in most of the areas of reserve forest. However, compared to
Dirgha site, Kakoi Rajgarh forest fringes are highly populated, and
human access to the forest is frequent, which limits the elephant
movements. As a result, there are lesser elephant fellings in Kakoi Raj­
garh compared to Dirgha. We noticed a reduced tree density at the site
(HD-Kakoi-Rajgarh) with a higher number of cut stems. Our findings are
consistent with previous research reporting lower tree density and basal
area in disturbed stands in contrast to undisturbed forests (Sahoo et al.,
2020; Chapagain et al., 2021; Bentsi-Enchill et al., 2022). The illegal tree
felling and fuelwood collection by the local community in
HD-Kakoi-Rajgarh could be the reason for lower basal area of this site.
The recorded tree density (446–582 ha− 1) in our study lies within this
range besides comparable to records from Bhuban hills, Assam (Borah
et al., 2014; 344–794 ind. ha− 1), lowland tropical rainforest, Assam
(Gogoi and Sahoo, 2018; 235–645 ind. ha− 1), subtropical broad-leaved
forest, Meghalaya (Singh et al., 2020; 276–868 ind. ha− 1) and tropical
forests of Dibru‑Saikhowa biosphere reserve, Assam (Joshi, 2020;
410–830 ind. ha− 1).
The distribution of tree individuals by diameter class is a key indi­
cator of alterations in stand structure and species composition in forest
ecosystems (Newbery and Gartlan, 1996). The present study recorded
lower basal area in higher girth classes in HD-Kakoi-Rajgarh compared
to MD-Dirgha, which have likely resulted from lesser numbers of larger
individuals (Gonzales and Zak, 1996), owing to selective removal of
large girth class trees. The study also revealed the declining tree density
trend with girth class, indicating the role of disturbance in affecting the
successional trend of the community, which is congruent with other
tropical wet evergreen forests in northeast India (Borogayary et al.,
2018).
Existing regenerating strata in forest ecosystems facilitate in antici­
pating the projected species composition. The lower density of regen­
erating strata at HD-Kakoi-Rajgarh may be the result of fire and other
unregulated human activities, which would have reduced the
7
Trees, Forests and People 11 (2023) 100371
reproductive individuals of several species that could have dispersed
adequate seeds for regeneration. The results revealed that 25.45% (MD-
Dirgha) and 29.17% (HD-Kakoi-Rajgarh) “not-regenerating” species
which indicates their sensitivity to existing disturbances and may go
extinct locally in future. Human disturbance may have altered seed and
fruit dispersal syndromes, germination, and regeneration processes in
several species, as seen by reduced seedling densities in HD-Kakoi-
Rajgarh; thus, certain species failed to regenerate naturally. Further­
more, juvenile individuals are more susceptible to environmental stress,
which may have accounted for reduced seedling density in HD-Kakoi-
Rajgarh. The strong regeneration ability of C. indica among the species
at HD-Kakoi-Rajgarh may be favored by disturbance and opportunistic
traits. Eilu and Obua (2005) and Rentch et al. (2010) reported that forest
disturbances frequently involve an uneven distribution, mechanical
injury, and higher seedling mortality of regenerating strata. However,
MD-Dirgha was dominated by M. ferrea, which has extensive regenera­
tion potential, thereby resulting in the suppressed or reduced potential
for regeneration of other species. Other variables, including light, can­
opy cover, soil nutrients, and moisture, may also impact species’ ability
to regenerate (Iqbal et al., 2012). Due to the lack of M. ferrea mother
trees, which allowed other species to flourish and increase in number,
HD-Kakoi-Rajgarh (18) had a higher percentage of species displaying
strong regeneration than MD-Dirgha (16). Even though the phenomenon
of plant regrowth on degraded sites is well characterized (Liu et al.,
2018), it is crucial to support restoration with a deeper knowledge of
vegetation recovery and shifting nutrient dynamics, given the unpre­
dictable disturbances. Most forest restoration efforts to recover
degraded areas primarily concentrate on planting trees which might
affect the species regeneration and recovery. Therefore, identifying in­
dicator species is crucial for eco-restoration.
The mean AGB and total carbon were 328.62 Mg ha‒1 and 207.04 Mg
ha across the study plots, with a maximum and minimum corre­
‒1
sponding to MD-Dirgha and HD-Kakoi-Rajgarh. The recorded estimates
were comparable to other tropical studies (Terakunpisut et al., 2007;
Joshi and Dhyani, 2019; Joshi, 2020). This indicates that anthropogenic
activities had unfavorable impact on carbon sequestration through
changes to the structure and composition of the forest. This was
confirmed by lower biomass and carbon stocks in HD-Kakoi-Rajgarh
than MD-Dirgha. Additionally, tree logging, fall, and transfer of logged
wood might harm non-target trees (often small trees), leading to tree
mortality, reduction in biomass altogether, and impeding the regener­
ation of disturbed areas (Putz et al., 2012). Similar findings in tropical
forests were reported by Gibbs et al. (2007) and Pan et al. (2011),
concluding that disturbances tend to reduce biomass and carbon stock of
forests, lower their potential, and also account for 15% of total carbon
emissions.
Besides stand characteristics, tree size class distribution predomi­
nantly influences the biomass and carbon storage in forest ecosystems
with large girth tress as major contributors (Clark et al., 2000; Baishya
et al., 2009). The differences in size class heterogeneity result from
various local and regional forces. In MD-Dirgha, 27.45%, whereas in
HD-Kakoi-Rajgarh 0.01% biomass and carbon stocks were represented
by trees >330 cm GBH. This suggests that selective cutting of large trees
in HD-Kakoi-Rajgarh was predominantly responsible for the lower
contribution of higher GBH classes to biomass and carbon stores. The
ability of tropical forests to store more carbon has also been linked to
species diversity (Liu et al., 2018). The greater species richness and
correspondingly better carbon storage in the MD-Dirgha support the
potential advantage of species diversity on carbon sequestration (Liu
et al., 2018; Wondimu et al., 2021). On the other hand,
HD-Kakoi-Rajgarh, with a substantial contribution from young in­
dividuals to biomass stocks, indicates the regenerative forest with sig­
nificant future carbon sequestration ability, provided management
interventions are put in action.
M.K. Buragohain et al.
5. Conclusion
Our study provides unequivocal evidence of the implications of
disturbance on tree species composition, regeneration, stand structure,
and biomass/carbon in a tropical rainforest in Eastern Himalaya. The
findings revealed that Kakoi reserve forest is considerably rich in
floristic diversity dominated by Mesua ferrea and Castanopsis indica. The
higher tree density, basal area, biomass, and carbon stock in MD-Dirgha
compared to HD-Kakoi-Rajgarh supports our hypothesis of disturbance-
induced species loss and their ability to take in atmospheric CO2. Despite
the persistent efforts of forest officials to minimize the overexploitation
of resources, illegal felling of trees for timber and fuelwood in the forest
are on the rise, which can have deleterious effects on the floristic di­
versity and stand structure of the forest. We conclude that lesser
recruitment and absence/lower number of large girth-class trees due to
overexploitation could seriously hamper the forest longevity if vulner­
able areas are not protected, restored, and managed sustainably. The
data suggest that disruptive and unsustainable resource extraction
methods jeopardize sustainable forest management. The insights
emphasize that the efforts for forest conservation are key elements for
ecosystem services, such as co-benefits between biomass/carbon stock
and biodiversity under REDD+, and are positively associated with
higher carbon stocks, particularly in forests that have experienced little
to moderate disturbance. To protect biodiversity and sustain critical
ecological services, we recommend to design adaptive management
techniques that encourage biodiversity conservation; rehabilitating/
restoring forest via natural or assisted regeneration; and intensify in-situ
conservation of focal species (vulnerable, threatened, and endemic),
including strict regulations for the safety of surviving mother trees.
Further investigations are encouraged to analyze the effect of distur­
bances on tree recruitment, mortality, growth patterns, and carbon
sequestration through long-term monitoring.
Consent for publication
Not applicable.
Ethics approval and consent to participate
Not applicable.
Funding
This work has received no funding.
Declaration of Competing Interest
The authors declare that they have no competing interests.
Data availability
Data will be made available on request.
Acknowledgements
Authors thank Principal Chief Conservator of Forests (PCCF) for
granting permission to work in the Reserve Forest. Thanks to Mr. Sukur
Ali, Beat Officer, and his team for assistance during the fieldwork. MKB
is indebted to Dr. Chittaranjan Borah and Dr. Budhadev Basumatary,
Department of Botany, North Lakhimpur College, Mr. Mantu Mahanta
and Mrs. Jully Gohain Neog, Head of the Department of Botany Pandu
College (Guwahati) and LTK College (Lakhimpur), and Rain Forest
Research Institute, Jorhat, Assam for their help in identification of tree
species.
8
Trees, Forests and People 11 (2023) 100371
Supplementary materials
Supplementary material associated with this article can be found, in
the online version, at doi:10.1016/j.tfp.2023.100371.
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