Article

Plant Community Composition and Carbon Stocks of a

Community Reserve Forest in North-East India
Aosanen Ao 1 , Sapu Changkija 2 , Francis Q. Brearley 3, * and Shri Kant Tripathi 1

1 Department of Forestry, Mizoram University, Aizawl 796004, Mizoram, India

2 Department of Genetics and Plant Breeding, Nagaland University, Medziphema 797106, Nagaland, India
3 Department of Natural Sciences, Manchester Metropolitan University, Manchester M1 5GD, UK
* Correspondence: [email protected]

Abstract: Anthropogenic activities are altering the structure and functioning of forests and their
services to society. However, we know little about the degree to which such activities are changing
the health of forests through edge effects in fragmented forests in different regions of the world.
The present study was carried out in Minkong Community Reserve Forest of Nagaland (North-east
India) with the aim to determine the effects of anthropogenic activities on floristic composition and
diversity, population structure, and biomass and carbon (C) stocks in the core zone (CZ) and buffer
zone (BZ) of the forest. We established 15 plots of 0.04 ha each in the two forest zones. We identified
31 trees, 18 shrubs, and 22 herbs in the CZ, and 22 trees, 25 shrubs, and 24 herbs in the BZ; tree species
diversity was greater in the CZ whereas the diversity of shrubs and herbs was greater in the BZ. The
values for tree density and basal area in the CZ and BZ were 303 and 197 individuals ha−1 and 32.6
and 22.2 m2 ha−1 , respectively; in contrast, the shrub and herb density increased in the BZ (4470 and
50,200 individuals ha−1 ) compared to that of the CZ (2530 and 35,500 individuals ha−1 ). The total
stand biomass (including that below-ground) was 327 Mg ha−1 in the CZ and 224 Mg ha−1 in the BZ.
Similarly, the total ecosystem C stocks in the CZ and BZ were 224 Mg C ha−1 and 173 Mg C ha−1 ,
indicating that the overall ecosystem C pool including soil in the CZ was approximately 30% greater
than the BZ. These results show how fragmentation and anthropogenic disturbance can reduce
forest diversity and C stocks and that community forest management can play a role in conserving
biodiversity and act as an ecosystem management tool to mitigate climate change.

Citation: Ao, A.; Changkija, S.; Keywords: carbon; community forestry; diversity; Minkong; Nagaland; species composition
Brearley, F.Q.; Tripathi, S.K. Plant
Community Composition and
Carbon Stocks of a Community
Reserve Forest in North-East India. 1. Introduction
Forests 2023, 14, 245. https:// Globally, forests play vital roles in conserving biodiversity, biogeochemical cycling
doi.org/10.3390/f14020245
including storing a large amount of carbon (C) in the vegetation and soil, and sustaining
Received: 11 December 2022 peoples’ livelihoods [1]. Tropical forests contribute significantly to the global terrestrial C
Revised: 12 January 2023 stocks and harbour half of the world’s biodiversity whilst supporting human populations
Accepted: 18 January 2023 by providing a wide range of goods and services. However, these forests are regularly
Published: 28 January 2023 over-exploited leading to the loss of forest cover (>10 million ha year−1 ), thereby losing
their potential for ecological service provision [2,3]. Tropical forests, with appropriate man-
agement, can provide economic benefits through ecotourism, non-wood forest products,
a sustainable timber source, and also through emerging C financing mechanisms [4,5].
Copyright: © 2023 by the authors.
Tropical moist forests are characterized by the high density of understory trees, high species
Licensee MDPI, Basel, Switzerland.
diversity, and a high degree of endemism [6]. Habitat fragmentation due to anthropogenic
This article is an open access article
activities (e.g., agriculture, road construction, urbanization, etc.) has become one of the
distributed under the terms and
most pervasive and conspicuous forms of disturbance in tropical rain forests [7–9]. Studies
conditions of the Creative Commons
on tropical forest fragmentation have shown edge effects to influence changes in the abiotic
Attribution (CC BY) license (https://
environment impacting upon forest structure and composition. Studies on a range of
creativecommons.org/licenses/by/
4.0/).
tropical forests reported the influence of fragmentation on ecosystem processes resulting

Forests 2023, 14, 245. https://doi.org/10.3390/f14020245 https://www.mdpi.com/journal/forests

Forests 2023, 14, 245 2 of 14

in increasing biomass loss, tree mortality rates, and ultimately changes to the population
structure of the forest [10–13].
Community forestry plays an important role in forest conservation, where the local
people are involved in the management and protection of the forest [14,15]. Community
forestry is practiced over the world but it is mainly dominant in Asia and has had a long
history in India [16]. The Indian state of Nagaland, geographically located in the north-east
of the country within the Himalaya and Indo-Burma biodiversity hotspots, is endowed
with diverse vegetation cover that supports the predominantly tribal communities in
their livelihoods. The constitution of India has provided special customary rights to the
population of the state for the traditional use of natural resources, and thus over 85% of
the natural habitats are owned by individuals or clans through the co-ordination of village
councils and other traditional institutions [17]. Since most of the economic activities in
rural areas are dependent on natural resources, the over-exploitation of forest resources in
recent years is posing a threat to the region’s biodiversity.
Minkong Community Reserve Forest is managed by local communities within a
mountainous region of Mokokchung district in the state of Nagaland surrounded by three
villages, namely, Chuchuyimpang, Longmisa, and Sungratsu. The forest, which is owned
by the local communities, is traditionally managed by laying down certain customary laws
such as restricting the cutting of trees, gathering of natural resources, or hunting of wild
animals from the forest without permission from the local village councils. Heavy fines are
levied upon the defaulters to act as a deterrent and help protect the forest.
In this study, we look at the floristic and structural attributes of a community-conserved
sub-tropical humid forest and we hypothesize that human disturbance reduces species di-
versity and C stocks. We also examine the role of edge effects through not only biophysical
disturbances but also increased human interference on the forest composition. Therefore,
the present study aims to: (i) assess the vegetation community composition (trees, shrubs,
and herbs), diversity, and species distribution patterns; (ii) estimate tree biomass and
ecosystem C stocks; and (iii) determine how these differ between the core zone and buffer
zone of the Minkong Community Reserve Forest of Nagaland in North-east India.

2. Materials and Methods

2.1. Study Area
The study was located in Minkong Community Reserve Forest, having a total area
of 2.75 km2 located at 26◦ 210 N and 94◦ 330 E and about 1400 m above mean sea level
in Mokokchung district, Nagaland, North-east India (Figure 1). The vegetation is sub-
tropical evergreen forest [18]. The forest area, which was initially owned by the state
government until the 1980s, was handed over to the surrounding village communities
which later declared it a Community Reserve Forest with the aim to conserve and protect
the biodiversity inhabiting it due to the high level of anthropogenic disturbance at that
time. About 40% of the forest area is considered secondary/fallow forests from shifting
cultivation practices by the Naga agriculturalists. While the core zone (CZ) is situated in
the centre of the reserve forest without any disturbance, the buffer zone (BZ) surrounds
this zone and is located about 2 km from the human settlements. The forest soil has, on
average, a pH of 5.8 [19] and 3.4% soil organic carbon (SOC) [20].

2.2. Vegetation Sampling

For a detailed quantitative analysis of vegetation, the forest was divided into two
zones—the core zone (CZ) in the centre of the reserve forest without disturbance and the
buffer zone (BZ) present towards the periphery. Tracts of the forest were selected in each
zone for the measurement of species diversity and C stocks. In each zone, 15 quadrats each
measuring 20 m × 20 m (0.04 ha) were laid randomly for tree species (DBH ≥ 10 cm). Within
each 20 m × 20 m plot were nested two 5 m × 5 m subplots for shrubs and four 1 m × 1 m
subplots for herbs (including epiphytes, lithophytes, and climbers). The plant species
were photographed and identified, and herbarium vouchers were prepared following
 Forests 2023, 14, 245 3 of 14

Jain and Rao [21]. Vouchers were deposited in the Herbarium, School of Agricultural
Sciences and Rural Development, Nagaland University, Medziphema, Nagaland, India.
Plant community attributes such as frequency, density, and basal area were analyzed.
The importance value index (IVI) of each individual species was calculated to study the
dominance of the species. Tree species were classified into different diameter classes to
study the population structure of the forest. The Shannon–Weiner diversity index [22] of
each lifeform in each quadrat and Sørensen’s similarity index [23] between the CZ and
Forests 2023, 14, x FOR PEER REVIEW 3 of 16
BZ for each of the lifeforms were also calculated. Comparisons between the two zones
were made with t-tests using Minitab 19.2, whilst a detrended correspondence analysis
ordination was conducted on the three groups of lifeforms (i.e., trees, shrubs, and herbs)
using Canoco 5.15 [24].

Figure 1. Map of the study site and villages surrounding the Minkong Community Reserve Forest,
Figure 1. Map district,
Mokokchung of the study site and
Nagaland, villages surrounding the Minkong Community Reserve Forest,
India.
Mokokchung district, Nagaland, India.
2.3. Soil Sampling and Analysis
2.2. Vegetation Sampling
Soil samples were collected from two random locations from each 0.1 ha sampling plot
at two
Fordepths (0 to 25
a detailed cm and 25 toanalysis
quantitative 50 cm). Soil
of samples collected
vegetation, from five
the forest wassampling
dividedplots
into two
were composited to represent one sample with a total of three samples from 15 sampling
zones—the core zone (CZ) in the centre of the reserve forest without disturbance and the
plots of 0.04 ha from each of the two zones. Soil bulk density was determined using a
buffer zone (BZ) present towards the periphery. Tracts of the forest were selected in each
stainless-steel tube of known inner volume within which soils were oven-dried at 40 ◦ C to
zone for the measurement of species diversity and C stocks. In each zone, 15 quadrats
constant weight and then passed through a sieve of 2 mm. Soil organic carbon (SOC) was
each measuring
analyzed using the20 m × 20 m
method of(0.04 ha)and
Walkley were laid[25].
Black randomly for tree species (DBH ≥ 10 cm).
Within each 20 m × 20 m plot were nested two 5 m × 5 m subplots for shrubs and four 1 m
× 12.4.
mEstimation
subplots offorPlant Biomass
herbs and Carbon
(including Stock lithophytes, and climbers). The plant spe-
epiphytes,
cies were photographed and identified, and
Total above-ground biomass of trees herbarium
(AGB vouchers using
est ) was estimated were prepared following
the allometric
equation
Jain and Raodeveloped from trees
[21]. Vouchers in North-east
were depositedIndia by Herbarium,
in the Nath et al. [26]:
School of Agricultural Sci-
ences and Rural Development, Nagaland University, Medziphema, Nagaland, India.
AGBest = 0.32 (D2 × Hδ) 0.75 × 1.34 (1)
Plant community attributes such as frequency, density, and basal area were analyzed. The
importance
where D isvalue index
the DBH, H (IVI)
denotesof each individual
the height of thespecies was
tree, and calculated
δ is to study
wood-specific the dom-
gravity
inance of the
obtained fromspecies.
the ICRAFTreedatabase
species[27].
were classified into different diameter classes to study
Below-ground
the population biomass
structure wasforest.
of the estimated
Theusing the allometric diversity
Shannon–Weiner equation developed
index [22]byof each
Cairns et
lifeform inal. [28]:quadrat and Sørensen’s similarity index [23] between the CZ and BZ for
each
each of the lifeforms were BGBalso exp [−1.085Comparisons
est =calculated. + 0.9256 × ln (AGB)]
between the two zones were (2) made
with t-tests using Minitab 19.2, whilst a detrended correspondence analysis ordination
was conducted on the three groups of lifeforms (i.e., trees, shrubs, and herbs) using
Canoco 5.15 [24].
Forests 2023, 14, 245 4 of 14

The vegetation C stock was estimated assuming 45.6% C content of dry biomass [29]. Total
above-ground biomass of shrubs was estimated using equation developed by Ali et al. [30]:

AGB = exp (-3.5 + 1.65 × ln (CD) + 0.842 × ln (H) (3)

where CD is the collar diameter and H denotes the height of the shrub.
The forest floor mass was collected by laying a quadrat of 50 cm × 50 cm in each
sampling plot of 0.04 ha. Litter was collected twice in a year (dry and wet seasons) and the
mean litter mass of the two seasons was presented. Herb biomass was estimated using the
harvest plot method on the basis of fresh/dry weight. All the above- and below-ground
vegetation was harvested inside the herb plot after which roots were separated from the
shoots and the fresh weight was recorded in the field. About 100 g of a fresh sample of
roots and shoots of the herbs was then oven-dried (60 ◦ C) to constant weight. Soil organic
carbon (SOC) stock (0 to 50 cm) was estimated as the product of the SOC content, bulk
density, and soil depth.

SOC stock (Mg C ha−1 ) = SOC × BD × SD × 100 (4)

where SOC is soil organic carbon content (%), BD is bulk density (g cm3 ), and SD is
soil depth in metres (m). The total ecosystem C stock was calculated by summing total
vegetation C stock and SOC stock.

3. Results
The present study showed that species composition varied significantly between the
CZ and BZ with the DCA ordinations showing a clear separation of the plots (Figure 2).
The total number of tree species recorded in the CZ was 31, which was reduced to 22 species
in the BZ. However, the total species richness of shrub and herb species was greater (25 and
24 species) in the BZ and lower (18 and 22 species) in the CZ (Table 1). The dominant families
of trees, shrubs, and herbs were: Lauraceae and Moraceae, Rosaceae and Urticaceae, and
Leguminosae and Zingiberaceae, respectively. Structural attributes also varied between
the two zones. The tree density and basal area were greater (303 individuals ha−1 and
32.6 m2 ha−1 ) in the CZ which declined to 197 individuals−1 ha−1 and 22.2 m2 ha−1 in the BZ.
However, the density of shrub and herb species increased (4470 and 50,200 individuals−1 ha−1 )
in the BZ compared to the CZ (2530 and 35,500 individuals−1 ha−1 ) (Table 1).

Table 1. Structural attributes and diversity (mean ± standard error) of trees, shrubs, and herbs in
core and buffer zones of Minkong Community Reserve Forest, Nagaland, India.

Parameter Core Zone Buffer Zone p-Value

No. of species Trees 9.47 ± 0.98 6.33 ± 0.55 0.01
Shrubs 7.60 ± 0.60 12.1 ± 0.70 <0.001
Herbs 6.60 ± 0.43 10.5 ± 0.85 0.001
Density (ind. ha−1 ) Trees 303 ± 2.20 197 ± 1.10 0.01
Shrubs 2530 ± 21.5 4470 ± 14.8 <0.001
Herbs 35,500 ± 132 50,200 ± 267 0.004
Basal area (m2 ha−1 ) Trees 32.6 ± 5.71 22.2 ± 4.30 0.09
Shannon–Wiener diversity index (H’) Trees 2.11 ± 0.10 0.72 ± 0.10 0.01
Shrubs 1.85 ± 0.10 2.39 ± 0.05 <0.001
Herbs 1.73 ± 0.06 2.19 ± 0.07 <0.001

The Shannon’s diversity index (H’) for tree species was greater (2.11) in the CZ and
lower (0.72) in the BZ, whereas the H’ for shrub and herb species showed the opposite
pattern (Table 1). Based on IVI values, the most important tree species in the CZ were Betula
alnoides (15.4), Choerospondias axillaris (15.3), Garcinia pedunculata (15.3), and Lithocarpus
pachyphyllus (15.0). On the other hand, the most important tree species in the BZ were:
Lithocarpus elegans (31.6), Morus macroura var. macroura (22.4), and Castanopsis indica (20.5)
 Forests 2023, 14, 245 5 of 14

(Table 2). The most important shrub species in the CZ were Clerodendrum glandulosum
(21.7), Eriosolena involucrata (16.9), Millettia pachycarpa (15.7), and Boehmeria japonica (15.5);
in contrast, Osbeckia nepalensis (13.7), Dendrocnide sinuata (11.3), and Rubus lucens (11.3)
were the most important species in the BZ (Table 2). Moreover, the most important herb
species in the CZ were Girardinia diversifolia (22.3), Smilax aspera (17.6), and Commelina
benghalensis (16.1); in the BZ, Commelina benghalensis (12.6) was the most important species,
along with Curculigo orchioides (12.6) Hellenia speciosa (12.4), and Gomphostemma strobilinum
(12.1) (Table 2). The dominance–diversity curves followed a log-normal distribution pattern
which indicates that different plant life forms (trees, shrubs, and herbs) showed high
equitability and low dominance of species in both zones (Figure 3). The greatest similarity
Forests 2023, 14, x FOR PEER REVIEW (Sørenson) (73%) between the CZ and BZ was recorded for the herb community, followed 5 of 16
by shrubs (46%) and trees (26%); the overall community had 47% similarity between the
two zones.

Detrendedcorrespondence
Figure2.2.Detrended
Figure correspondenceanalysis
analysisordination
ordinationofof(a)(a)tree,
tree,(b)
(b)shrub,
shrub,and
and(c)(c)herb
herbspecies
species
ininthe
thecore
core(filled
(filledcircles)
circles)and
andbuffer
buffer(open
(opencircles)
circles)zones
zonesininMinkong
MinkongCommunityCommunityForest
ForestReserve,
Reserve,
Nagaland,
Nagaland,India.
India.TheThecomparative
comparativesize
sizeofofthe
thecircles
circlesrepresents
representsthe thespecies
speciesrichness
richnessofofeach
eachplot.
plot.

Table 1. Structural attributes and diversity (mean ± standard error) of trees, shrubs, and herbs in
core and buffer zones of Minkong Community Reserve Forest, Nagaland, India.

Parameter Core Zone Buffer Zone p-Value

No. of species Trees 9.47 ± 0.98 6.33 ± 0.55 0.01
Shrubs 7.60 ± 0.60 12.1 ± 0.70 <0.001
Forests 2023, 14, 245 6 of 14

Table 2. Density (individuals ha−1 ), basal area (m2 ha−1 ), and important value index (IVI) of trees,
shrubs, and herbs in core and buffer zone of Minkong Community Reserve Forest, Nagaland, India.

Core Zone Buffer Zone

Species Family Basal Basal
Density IVI Density IVI
Area Area
Trees
Acer thomsonii Miq. Sapindaceae 13.3 1.16 13.6
Actinodaphne obovata (Nees) Blume Lauraceae 6.67 0.52 8.94
Aglaia spectabilis (Miq.) S.S.Jain & S.Bennet Meliaceae 8.33 0.81 8.05
Alnus nepalensis D.Don Betulaceae 10.0 1.86 12.5
Bauhinia purpurea L. Fabaceae 8.33 0.66 11.5
Beilschmiedia roxburghiana Nees Lauraceae 8.33 0.55 9.92
Betula alnoides Buch.-Ham. ex. D.Don Betulaceae 10.0 2.54 15.4
Bischofia javanica Blume Phyllanthaceae 11.7 1.44 11.8 8.33 0.81 13.2
Brassaiopsis hainla (Buch.-Ham.) Seem. Araliaceae 3.33 0.36 3.61
Callicarpa arborea Roxb. Lamiaceae 8.33 0.69 7.71
Castanopsis indica (Roxb. ex Lindl.) A.DC. Fagaceae 15.0 1.05 12.4 10.0 2.72 20.5
Castanopsis tribuloides (Sm.) A.DC. Fagaceae 8.33 0.49 9.64
Cephalotaxus griffithii Hook.f. Taxaceae 11.7 1.29 9.21
Choerospondias axillaris (Roxb.) B.L.Burtt &
Anacardiaceae 15.0 1.30 15.3
A.W.Hill
Cinnamomum sulphuratum Nees Lauraceae 10.0 0.44 8.20
Cinnamomum verum J.Presl. Lauraceae 15.0 0.74 10.8 6.67 0.32 6.95
Dalrympelea pomifera Roxb. Staphyleaceae 10.0 0.74 12.7
Elaeocarpus tectorius (Lour.) Poir. Elaeocarpaceae 11.7 1.31 11.4
Engelhardia spicata Lechen ex Blume Juglandaceae 8.33 0.70 12.7
Eurya cerasifolia (D.Don) Kobuski Pentaphylacaceae 8.33 0.48 10.7
Ficus hispida L.f. Moraceae 5.00 0.37 4.90 8.33 0.20 9.41
Ficus neriifolia Sm. Moraceae 5.00 0.25 6.85
Ficus semicordata Buch.-Ham. ex Sm. Moraceae 16.7 0.87 13.8 3.33 0.20 4.71
Garcinia pedunculata Roxb. ex Buch.-Ham. Clusiaceae 16.7 1.58 15.3
Grewia serrulata DC. Malvaceae 3.33 0.31 3.48
Hovenia dulcis Thunb. Rhamnaceae 8.33 1.03 8.73
Ilex sp. Aquifoliaceae 3.33 0.10 2.83
Ilex dipyrena Wall. Aquifoliaceae 10.0 1.13 15.5
Juglans regia L. Juglandaceae 11.7 1.49 12.7
Lithocarpus elegans (Blume) Hatus. ex Soepadmo Fagaceae 10.0 1.90 12.0 16.7 3.01 31.6
Lithocarpus pachyphyllus (Kurz) Rehder Fagaceae 16.7 1.02 15.0
Macaranga denticulata (Blume) Müll.Arg. Euphorbiaceae 11.7 0.75 15.7
Macropanax dispermus (Blume) Kuntze Araliaceae 10.0 0.96 8.38
Mallotus nepalensis Müll.Arg. Euphorbiaceae 8.33 0.45 6.24
Morus macroura var. macroura Miq. Moraceae 13.3 2.28 22.4
Myrica esculenta Buch.-Ham. ex D.Don Myricaceae 6.67 0.44 6.37
Oreocnide integrifolia (Gaudich.) Miq. Urticaceae 3.33 0.20 3.13
Prunus napaulensis (Ser.) Steud. Rosaceae 5.00 2.04 14.9
Quercus lamellosa Sm. Fagaceae 5.00 2.52 11.5
Quercus serrata Murray Fagaceae 13.3 1.11 11.3 8.33 0.83 12.2
Sloanea dasycarpa (Benth.) Hemsl. Elaeocarpaceae 11.7 0.62 16.2
Sterculia lanceolata var. coccinea (Jack) Phengklai Sterculiaceae 11.7 0.71 15.5
Terminalia myriocarpa Van Heurck & Müll.Arg. Combretaceae 10.0 1.33 10.2
Toona ciliata M.Roem. Meliaceae 5.00 1.66 8.16
Trema cannabina Lour. Cannabaceae 6.67 0.30 5.94
Xerospermum noronhianum (Blume) Blume Sapindaceae 8.33 2.21 18.5
Shrubs
Abutilon indicum (L.) Sweet Malvaceae 120 3.78
Agapetes macrantha (Hook.) Benth. & Hook.f. Ericaceae 160 6.86
Amomum dealbatum Roxb. Zingiberaceae 147 8.20
Boehmeria japonica (L.f.) Miq. Urticaceae 213 15.5 200 10.5
Forests 2023, 14, 245 7 of 14

Table 2. Cont.

Core Zone Buffer Zone

Species Family Basal Basal
Density IVI Density IVI
Area Area
Breynia retusa (Dennst.) Alston Phyllanthaceae 173 6.61
Calamus rotang L. Arecaceae 93.3 7.22
Camellia oleifera C.Abel Theaceae 173 7.16
Chromolaena odorata (L.) R.M.King & H.Rob. Asteraceae 160 9.85 240 9.20
Clerodendrum glandulosum Lindl. Lamiaceae 280 21.7 107 4.57
Crotalaria juncea L. Fabaceae 120 8.28
Croton caudatus Geiseler Euphorbiaceae 147 12.0
Debregeasia longifolia (Burm.f.) Wedd. Urticaceae 173 7.71
Dendrocnide sinuata (Blume) Chew. Urticaceae 133 10.6 227 11.6
Deutzia compacta Craib Hydrangeaceae 80.0 7.58
Eriosolena involucrata (Wall.) Tiegh. Thymelaeaceae 160 16.9
Grona heterocarpos (L.) H.Ohashi & K.Ohashi Fabaceae 213 9.69
Hibiscus sabdariffa L. Malvaceae 93.3 4.28
Leucosceptrum canum Sm. Lamiaceae 187 11.8 200 8.30
Maesa indica (Roxb.) Sweet Primulaceae 147 9.33
Melastoma malabathricum L. Melastomataceae 120 11.8 240 9.75
Millettia pachycarpa Benth. Fabaceae 173 15.7
Mussaenda roxburghii Hook.f. Rubiaceae 93.3 8.99 227 9.45
Neillia thyrsiflora D.Don Rosaceae 133 7.36
Osbeckia nepalensis Hook. Melastomataceae 293 13.7
Oxalis acetosella L. Oxalidaceae 133 5.72
Oxyspora paniculata (D.Don) DC. Melastomataceae 200 8.30
Rhaphiolepis bengalensis (Roxb.) B.B.Liu & J.Wen Rosaceae 93.3 8.11 173 7.71
Rubus ellipticus Sm. Rosaceae 213 9.69
Rubus efferatus Craib Rosaceae 40.0 4.23 133 5.72
Rubus lucens Focke Rosaceae 187 11.3
Securinega sp. Phyllanthaceae 240 9.75
Senegalia pennata (L.) Maslin Fabaceae 120 10.1 66.7 3.13
Uraria oblonga (Wall. ex Benth.) H.Ohashi &
Fabaceae 173 10.4
K.Ohashi
Herbs
Arisaema concinnum Schott Araceae 667 3.80 2170 10.7
Bidens pilosa L. Asteraceae 1330 7.60 1670 6.49
Commelina benghalensis L. Commelinaceae 4000 16.1 3170 12.6
Curculigo orchioides Gaertn. Hypoxidaceae 2500 12.6
Cymbidium aloifolium (L.) Sw. Orchidaceae 1830 9.97 2170 8.75
Dendrobium lituiflorum Lindl. Orchidaceae 2330 11.0
Dioscorea glabra Roxb. Dioscoreaceae 100 4.74 2500 10.1
Drymaria cordata (L.) Willd. ex Schult. Caryophyllaceae 1670 7.58
Elatostema sessile J.R.Forst. & G.Forst. Urticaceae 2330 12.3 1170 4.86
Entada rheedei Spreng. Fabaceae 1170 5.21 1000 5.16
Erythropalum scandens Blume Olacaceae 833 3.31
Fagopyrum cymosum (Trevir.) Meisn. Polygonaceae 2670 9.11
Fragaria nilgerrensis Schltdl. ex J.Gay Rosaceae 1670 7.58 2330 8.45
Girardinia diversifolia (Link) Friis Urticaceae 4170 22.3 2170 6.85
Gnetum latifolium Blume Gnetaceae 1500 9.99 1830 6.19
Gomphostemma strobilinum Wall. ex Benth. Lamiaceae 1670 9.50 3830 12.1
Hedychium coronarium J.Koenig Zingiberaceae 1330 6.45
Hellenia speciosa (J.Koenig) S.R.Dutta Costaceae 1500 12.9 3670 12.4
Mucuna macrocarpa Wall. Fabaceae 1330 8.56 2330 10.4
Oxalis corniculata L. Oxalidaceae 833 5.23
Papilionanthe teres (Roxb.) Schltr. Orchidaceae 1500 13.8
Phanera vahlii (Wight & Arn.) Benth. Fabaceae 2000 12.4 1330 4.56
Poranopsis paniculata (Roxb.) Roberty Convolvulaceae 2330 9.46 2830 10.1
Forests 2023, 14, 245 8 of 14

Table 2. Cont.

Core Zone Buffer Zone

Species Family Basal Basal
Density IVI Density IVI
Area Area
Saccharum longesetosum (Andersson) V.Naray.
Poaceae 1000 3.89
ex Bor
Smilax
Forests 2023, 14, x FOR PEERaspera L.
REVIEW Smilacaceae 2170 17.6 1170 10 5.49
of 16
Forests 2023, 14, x FOR PEER REVIEW 10 of 16
Spatholobus parviflorus (Roxb. ex G.Don) Kuntze Fabaceae 2330 11.6
Tetrastigma eucostaphylum (Dennst.) Alston Vitaceae 667 2.59
Tinospora sinensis (Lour.) Merr. Menispermaceae 2000 7.78

Figure 3.
Figure 3. Dominance–diversity
Dominance–diversity curve
curvefor
fortrees,
trees,shrubs,
shrubs,and
andherbs
herbsinincore
core(left)
(left)and
andbuffer
buffer(right)
(right)
Figurein3.Minkong
zones Dominance–diversity
Community curveReserve,
Forest for trees, shrubs, and
Nagaland, herbs in core (left) and buffer (right)
India.
zones in Minkong Community Forest Reserve, Nagaland, India.
zones in Minkong Community Forest Reserve, Nagaland, India.
While tree
While treedensity
densityand andbasal
basalarea
areawere
werelower
lowerin inthe
theBZ BZasascompared
comparedtotothe theCZ CZ(Table
(Table1),
1), the While
tree tree
density density
and and
basal basal
area area
showed werethe
the tree density and basal area showed the greatest occurrences lower in
greatest the BZ as
occurrences compared
in the 10 to the CZ
10 to 20 cmDBH
to 20 cm (Table
DBH
1), the
range in
range tree
in bothdensity
both zones and basal area showed
zones (Figure 4) with the number the greatest
number of occurrences
of individuals
individualsdeclining in the
decliningwith10 to 20 cm
withincreasing DBH
increasing
range
DBH in
DBH in both
in both zones (Figure 4) with
both zones; there were proportionally the number
proportionally fewer of individuals
fewertrees
treesin inthe
the20declining
20toto30
30DBH with
DBHclass increasing
classininthe
the
BZDBH
BZ in both to
compared
compared zones;
tothe there
theCZ.
CZ.The were
The proportionally
overall
overall plant
plant fewer
biomass
biomass and trees
and C in
C stock the
stock
were20greater
wereto greater
30 DBH class
in
in the CZ in
the CZthe
than
inBZ
than
the compared
in
BZ.theOutBZ. tothe
of thetotal
Out CZ.
of theThetotal
biomassoverall
(327plant
biomassMg ha biomass
−1 )Mg
(327 and
ha
in the −1) C stock
CZ,inthethetreewere
CZ, greater
the
biomass tree in the CZ
biomass
contributed
thanand in the
contributed
98% theBZ.
98% Outandofthe
remaining the
was total
remainingbiomass
contributed was(327 Mg haherb,
bycontributed
shrub, −1 ) in the
byand CZ,
floorthe
shrub, treeand
herb,
mass. biomass
floor contrib-
Similarly, mass.
of the
utedbiomass
total 98% of
Similarly, andthethe
(224 total ha−1 ) in
remaining
Mgbiomass wasthecontributed
(224 Mg
BZ, ha −1) in by
trees theshrub,
BZ, trees
contributed herb, and
Thefloor
contributed
97%. mass.
97%.
overall Similarly,
The overallof
distribution of
the total
distribution biomass
of tree (224 Mg
biomass ha −1) in the BZ, trees contributed 97%. The overall distribution
among different girth classes showed
tree biomass among different girth classes showed the greatest biomass in the trees of 10 the greatest biomass in
toof30
the treecmbiomass
trees DBH toamong
of 10 which 30 cm different
DBH which
contributed girth
79% classes
contributed
in the CZshowed
79%
and in the
71% thegreatest
inCZ
theand biomass
BZ. 71%
Thein in the
the
maximum BZ.trees
treeof
The
10 to 30 of
maximum
biomass cmtree
DBH
50% which
biomass
and 58%ofcontributed
50%contributed
was and 58% 79% in
was the
theCZ
bycontributedand
10 to 2071%
by
cmthein10thetoBZ.
diameter Thediameter
20 class
cm maximum
in the CZ tree
class
and
biomass
in
BZ, the CZof and
respectively50%BZ, and 58% 5).was contributed
respectively
(Figure (Figure 5). by the 10 to 20 cm diameter class in the CZ and
BZ, respectively (Figure 5).

Figure 4.
Figure 4. Contribution
Contribution ofof tree
tree stand
stand density
density and
and basal
basalarea
areabased
basedon
ondiameter
diameterclass
classdistributions
distributionsinin
the core (a) and buffer (b) zones in Minkong Community Forest Reserve, Nagaland,
the core (a) and buffer (b) zones in Minkong Community Forest Reserve, Nagaland, India. India.

Figure 4. Contribution of tree stand density and basal area based on diameter class distributions in
the core (a) and buffer (b) zones in Minkong Community Forest Reserve, Nagaland, India.
Forests
Forests 2023,
2023, 14,
14, x245
FOR PEER REVIEW 11 9ofof16
14

Treebiomass
Figure5.5.Tree
Figure biomassdistribution
distributionin
invarious
variousDBH
DBHclasses
classesfor
forthe
thecore
coreand
andbuffer
bufferzones
zonesin
inMinkong
Minkong
CommunityForest
Community Forest Reserve,
Reserve, Nagaland,
Nagaland, India.
India.

The total C contributed by different tree species to AGB C varied between the species
The total C contributed by different tree species to AGB C varied between the species
and was directly associated with the tree basal area. In the CZ, the contribution of major
and was directly associated with the tree basal area. In the CZ, the contribution of major
tree species (i.e., Betula alnoides, Bischofia javanica, Cephalotaxus grifithii, Lithocarpus elegans,
tree species (i.e., Betula alnoides, Bischofia javanica, Cephalotaxus grifithii, Lithocarpus elegans,
Quercus lamellosa, Terminalia myriocarpa, and Toona ciliata) was around 40% of the total
Quercus lamellosa, Terminalia myriocarpa, and Toona ciliata) was around 40% of the total
AGB and C stocks, with a similar contribution of major tree species (i.e., Castanopsis indica,
AGB and C stocks, with a similar contribution of major tree species (i.e., Castanopsis indica,
Ilex dipyrena, Lithocarpus elegans, Prunus nepaulensis, Quercus serrata, and Xerospermum
Ilex dipyrena, Lithocarpus elegans, Prunus nepaulensis, Quercus serrata, and Xerospermum no-
noronhianum) in the BZ. The total tree biomass (AGB + BGB) was estimated at 322 Mg ha−−11
ronhianum) in the BZ. The total tree biomass (AGB + BGB) was estimated at 322 Mg ha
and 218 Mg ha−1 in the CZ and BZ, respectively (Table 3). The total understory biomass
and 218 Mg ha−1 in the CZ and BZ, respectively (Table 3). The total understory biomass
(shrubs + herbs) was 1.90 and 2.94 Mg ha−1 in the CZ and BZ.
(shrubs + herbs) was 1.90 and 2.94 Mg ha−1 in the CZ and BZ.
Table 3. Total biomass (Mg ha−1 ) (mean ± standard error) in core and buffer zones of Minkong
Table 3. Total biomass (Mg ha−1) (mean ± standard error) in core and buffer zones of Minkong Com-
Community Reserve Forest, Nagaland, India).
munity Reserve Forest, Nagaland, India).
Ecosystem Component
Ecosystem Component Core Zone Core Zone Buffer Zone
Buffer Zone
Above-ground tree biomass
Above-ground tree biomass 263 ± 8.4 263 ± 8.4 ± 11± 11
177 177
Below-ground tree biomass 58.7 ± 3.4 40.7 ± 4.1
Below-ground tree biomass 58.7 ± 3.4 40.7 ± 4.1
Shrub biomass 1.20 ± 1.2 2.01 ± 0.3
Shrub biomass
Herb biomass 0.70 ± 0.6 1.20 ± 1.2 0.932.01
± 0.4± 0.3
FloorHerb
mass biomass 3.3 ± 0.8 0.70 ± 0.6 3.4 0.93
± 0.5± 0.4
Total stand Floor
biomassmass 327 ± 13.3 3.3 ± 0.8 224 ± 18.7
3.4 ± 0.5
Total stand biomass 327 ± 13.3 224 ± 18.7
The total SOC stock (0 to 50 cm) was not different between the CZ (75 ± 4.51 Mg C ha−1 )
and The total
the BZ (71SOC stock
± 3.45 −1cm)
Mg(0Ctoha50 was4).
) (Table notThe
different
total Cbetween
stored inthe
all CZ (75 ± 4.51
ecosystem Mg C
compart-
ha −1
ments,) and the BZ (71
including ± 3.45
SOC, Mg Ca ha
showed sum of 224 Mg C ha−1 inCCZ
−1) (Table 4). The total stored in all
and 173 Mg C ha−1 incom-
ecosystem BZ.
partments, including SOC, showed a sum of 224 Mg C ha−1 in CZ and 173 Mg C ha−1 in
Table 4. Distribution of carbon stocks (Mg C ha−1 ) (mean ± standard error) in different ecosystem
BZ.
compartments of core and buffer zone of Minkong Community Forest Reserve, Nagaland, India.
Table 4. Distribution of carbon stocks (Mg C ha−1) (mean ± standard error) in different ecosystem
Ecosystemof
compartments Compartment Core Zone
core and buffer zone of Minkong Community Forest Reserve,Buffer Zone India.
Nagaland,
Above-ground tree biomass 120 ± 0.31 80.7 ± 0.43
Ecosystem Compartment Core Zone Buffer Zone
Below-ground tree biomass 26.8 ± 1.45 18.6 ± 2.10
Above-ground
Shrub biomass tree biomass 0.54 ± 1.16 120 ± 0.31 0.92 80.7 ± 0.43
± 0.52
Below-ground
Herb biomass tree biomass 0.31 ± 1.2026.8 ± 1.45 0.42 18.6 ± 2.10
± 0.45
Floor massbiomass
Shrub 1.50 ± 0.120.54 ± 1.16 ± 0.15
1.55 0.92 ± 0.52
Total biomass carbon 149 ± 3.40 102 ± 1.21
Herb biomass 0.31 ± 1.20 0.42 ± 0.45
Soil organic carbon 75 ± 4.51 71 ± 3.45
Floor mass 1.50 ± 0.12 1.55 ± 0.15
Forests 2023, 14, 245 10 of 14

4. Discussion
Community reserve forests can play an important role in biodiversity conservation
and provide socio-ecological benefits for regional populations [14,15,31–33]; however,
overexploitation of these forests, combined with ecological edge effects, can considerably
affect their structure and function. In the present study, the CZ and BZ of a community
reserve forest in North-east India were studied with respect to their species composition,
diversity, biomass, and C stocks. The greater tree species richness in the CZ compared
to the BZ may be because of the protection provided by the traditional communities [33]
and less exploitation, whereas the lower density and species richness of trees in the BZ,
being more closely located to human settlements, is likely due to anthropogenic pressures
(e.g., extraction of timber and natural resources, forest fire, and agriculture expansion)
along with further environmental changes brought about by the opening of the canopy.
The diversity indices in the present study showed the changing pattern of the diversity
of various lifeforms in the forest community. The tree community was more diverse than
the shrub and herb communities, although this could be due to the greater area sampled
for trees. High values of the Shannon–Wiener diversity index (H’) and low dominance
are characteristic features of an old-growth forest [34] which was also supported by our
study. Additionally, changes in species composition have been shown at the edges of
forest fragments with a move towards more early successional species and a decline in the
abundance of old-growth specialists [35]. In the BZ, we found, for example, that Macaranga
denticulata, a typical secondary forest tree species, was an abundant tree but it was not
present in the CZ. In contrast, a greater diversity and density of undergrowth plants (shrubs
and herbs) in the BZ may be attributed to a more open canopy due to anthropogenic and
natural disturbances allowing more sunlight to reach the forest floor and thus creating
favorable microsite conditions enhancing the growth of understory vegetation [36]. The
composition of the shrub community in the BZ was clearly distinct from that in the CZ,
whereas there was more overlap in the herb community. Similar observations have been
reported from other natural undisturbed forests in India [37–40]. This is supported by the
similarity measures (Sørenson) which are the most instinctive and common methods for
comparing two or more sites with respect to their species overlap [41]. The low similarity
between the CZ and BZ may also be because of succession brought about by disturbance,
leading to contrasting micro-environmental conditions [42,43]. Different plant life forms
(trees, shrubs, and herbs) recorded from the CZ and BZ showed similar characteristics of
dominance–diversity curves with high equitability and low dominance of species. The
log-normal distribution pattern in both sites indicates high species richness and equitable
distribution of natural resources among the species in the forest ecosystem [44,45]. The
reduced proportion of trees in the 20 to 30 cm DBH category in the BZ suggests that this size
category is preferentially used by the local communities. In other studies of disturbed forest,
larger trees are found at lower proportions leading to changes in the slope of a scaling
relationship between stem diameter and density [46,47]. Our results present an interesting
comparison to this as large trees seem to be retained in the BZ with medium-sized trees
appearing to be preferentially removed.
The estimated above-ground biomass for trees in both zones ranged from 177 to
263 Mg ha−1 with a mean of 220 Mg ha−1 which falls well within the reported range (133
to 262 Mg ha−1 ) for various natural forests of North-east India [48–55]. Decreased biomass
in the BZ may be attributed to lower tree density and size due to various anthropogenic
disturbances associated with edge effects [56]. Tree species composition also affects the
total biomass and C stock of forests [51,57,58] and the change in species composition in
the BZ plots will influence total forest biomass if lighter-wooded species become more
dominant. Below-ground biomass contributed about 20% to the total biomass stock in both
zones and plays an important role in below-ground C storage; this value is an estimate
based on above-ground biomass so it would be valuable to directly measure root biomass
in these forests for a more accurate estimate, e.g., [59,60]. Furthermore, in both the zones,
tree biomass distribution across diameter classes showed the greatest biomass allocation
Forests 2023, 14, 245 11 of 14

in smallest classes (10 to 20 cm DBH) which may be due to the occurrence of greater tree
density and basal area of the young trees and the lack of particularly large trees due to
the montane elevation. A similar study was also conducted in a sacred natural forest of
Manipur, North-east India which reported the greatest biomass in the smaller girth class
of 30 to 60 cm (equivalent to 10–20 cm DBH) [55]. The greatest familial contribution to
total biomass C stocks in both the CZ and BZ was by the Fagaceae, followed by Betulaceae,
Sapindaceae, Moraceae, Euphorbiaceae, and Meliaceae, which contributed about 70% of
the total biomass among the six families.
There was no difference in the SOC stock between the BZ (71 Mg C ha−1 ) compared
to the CZ (75 Mg C ha−1 ). It is known that forest conversion to other land-uses such as
shifting cultivation or plantations can have a clear effect on SOC stocks with reductions
regularly seen [61,62]. It appears that, currently, the low-intensity disturbance in the buffer
zone was not severe enough to influence SOC stocks. However, changes in litter input
and soil disturbance in the future may influence soil changes, so additional monitoring
would be valuable. Determining long-term C sequestration rates and how they may be
influenced by disturbance, tree growth, and mortality rates combined with litterfall and
other measures of primary production, e.g., [13,59], would be an important next step.

5. Conclusions
We show here that whilst community reserve forests can be valuable for conserving
tropical biodiversity, edge effects brought about by human usage of the forest lead to
reductions in tree species richness but an increase in shrub and, particularly, herb species
richness with a clear impact on the community composition of all three groups of plant
life-forms with edge-affected (buffer-zone) plots having a distinct composition to those
in the core zone. Total biomass and C stock were both high as compared to other Indian
natural forests indicating the high potential of the site to store C in the plant biomass
and soil, and the buffer zone can provide similar ecosystem services to that of the core
zone if adequate protection is provided. However, the buffer zone can act as an ex situ
conservation ground for the valuable local plant species and protect the ecosystem services
at a regional level by buffering the human disturbances. We recommend the extension of
the buffer zone or the creation of a peripheral zone around Minkong Community Reserve
Forest, perhaps in the form of a plantation forest or agroforest, that may help in conserving
local species and ecosystem services on a sustained basis. Such an effort may provide
livelihood opportunities to the local tribal communities by conserving regional diversity
and mitigating climate change.

Author Contributions: Conceptualization, A.A. and S.K.T.; Data curation, A.A.; Formal analysis,
A.A., F.Q.B. and S.K.T.; Methodology, A.A. and S.K.T.; Resources, A.A., S.K.T. and S.C.; Writing—
draft, A.A., F.Q.B. and S.K.T.; Supervision, S.K.T., S.C. and F.Q.B.; Validation, A.A., S.K.T. and S.C.;
Writing—review & editing, A.A., F.Q.B. and S.K.T. All authors have read and agreed to the published
version of the manuscript.
Funding: This research was partly supported by a fellowship to A.A. from the Ministry of Tribal
Affairs, Government of India (202021-NFST-NAG-00620).
Institutional Review Board Statement: Not applicable.
Informed Consent Statement: Not applicable.
Data Availability Statement: Data on species composition is available from the corresponding author.
Acknowledgments: The authors would like to thank the Nagaland Forest Department and the
villagers living near the Minkong Community Reserve Forest for granting us permission to collect
the data and study the forest vegetation. Our sincere thanks to local residents for extending help as
local guides during the field survey.
Conflicts of Interest: This article has no conflict of interest to any person or organization.
Forests 2023, 14, 245 12 of 14

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