Journal of Asia-Pacific Biodiversity xxx (xxxx) xxx

Contents lists available at ScienceDirect

Journal of Asia-Pacific Biodiversity

journal homepage: http://www.elsevier.com/locate/japb

Original Article

Variation in Species Composition, Structural Diversity, and Regeneration

Along Disturbances in Tropical Dry Forest of Northern India
Anil Sharma, Sanoj Kumar Patel, Gopal Shankar Singh*
Institute of Environment and Sustainable Development Banaras Hindu University, Varanasi, India

a r t i c l e i n f o a b s t r a c t

Article history: Understanding disturbances is crucial for preventing forest degradation and resolving vegetation loss
Received 30 September 2022 which remain to be a major concern. The present study was to evaluate the vegetation dynamics of
Received in revised form different disturbed forest stands, based on cumulative disturbance index score in Vindhyan highlands,
21 November 2022
India. Our findings demonstrated that the mean tree density was greater in least disturbed (56.24
1.44
Accepted 21 November 2022
Available online xxx
hae1) than in highly disturbed (53.28
2.67 hae1) and moderately disturbed (50.27
0.94 hae1) forest
stands. Shannon, Simpson, Evenness, Margalef, and b-diversity indices differed significantly amongst the
three disturbed stands. The regeneration potential of tree species was found to be good (35.08%), fair
Keywords:
Biodiversity indices
(16.42%), and poor to no regeneration (48.5%) across the three stands, in which Diospyros melanoxylon,
Dispersion Syzygium heyneanum, and Holarrhena antidysenterica were shown to have strong regeneration capacity.
Disturbance gradients Principal component analysis revealed that the first component (pc1) explained 45.3% variance, while
Regeneration potential the second component (pc2) elucidated 20.1% variance in both disturbances and life-form wise distri-
Species diversity bution with a cumulative score of 65.4%. Our findings would be helpful to understand how different
disturbanceeregeneration combinations influence the successful regeneration of tree seedlings as well
as how variations in compositional diversity will impact the vegetation dynamics and restoration of plant
community in dry tropics.
Ó 2022 National Science Museum of Korea (NSMK) and Korea National Arboretum (KNA), Publishing
Services by Elsevier. This is an open access article under the CC BY-NC-ND license (http://
creativecommons.org/licenses/by-nc-nd/4.0/).

Introduction The regeneration profile of a forest is determined by the dis-

Tropical forests are biodiversity hotspots with their various bi-
omes (Sullivan et al. 2017) and are primarily concerned with life
sustainability on the planet (Baraloto et al. 2013). Despite consti-
tuting only 7% of the land surface, it is recognized to be home to
almost half of the world’s species (Cusack et al. 2016). Tropical
forests, which have a large standing biomass and higher produc-
tivity, are generally described to be the world’s largest carbon sink
(Raj and Jhariya 2021b). It is significant because they provide a
variety of ecosystem services, including species protection, soil
erosion avoidance, and habitat preservation for plants and animals
(Armenteras et al. 2009). Despite their importance, tropical forests
are constantly subjected to anthropogenic pressures, resulting in
forest fragmentation and biodiversity loss in many regions of the
globe (Chow et al. 2013; Osuri et al. 2017).
* Corresponding author. Tel.: þ91 94 5053 0681
E-mail address: gopalsingh.bhu@gmail.com (GS Singh)
Peer review under responsibility of National Science Museum of Korea (NSMK) and
Korea National Arboretum (KNA).
tribution of individuals of each species in population structure,
which is based on tree density, frequency, basal area, and species
significance value (Tesfaye et al. 2002; Shibru and Balcha 2004).
The population structure and dynamics of seedlings and saplings
can be used to determine the regeneration potential of a species
(Duchok et al. 2005; Bhadouria et al. 2017). Furthermore, under-
standing previous disturbance incidence, current regeneration
scenario, and future projections of the population require sufficient
knowledge of the population structure of a species (Subashree et al.
2021). Disturbance is a key factor in determining the species
composition of forests (De Marzo et al. 2022; Sakai et al. 2022).
Disturbances immediately result in the loss of tree cover, which has
a knock-on effect on other living forms, diminishing the diversity of
forest species (Campbell et al. 2017). Consequently, the declining
tree species in a forest can have a negative impact on the structure
and function as well as the forest’s ecosystem services (Tekalign
et al. 2017; Raj and Jhariya 2021a; Raj and Jhariya 2021b). Accord-
ing to studies conducted in many regions of the world, the bio-
logical diversity of plants and other aspects of forest vegetation
such as density, basal area, and species diversity decreases as the

https://doi.org/10.1016/j.japb.2022.11.004
pISSN2287-884X eISSN2287-9544/Ó 2022 National Science Museum of Korea (NSMK) and Korea National Arboretum (KNA), Publishing Services by Elsevier. This is an open
access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).

Please cite this article as: Sharma A et al., Variation in Species Composition, Structural Diversity, and Regeneration Along Disturbances in
Tropical Dry Forest of Northern India, Journal of Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2022.11.004
2 A Sharma et al. / Journal of Asia-Pacific Biodiversity xxx (xxxx) xxx
amount of disturbance increases (Pretto et al. 2010; Mayor et al.
2012; Cardelús et al. 2019; Thakrey et al. 2022).
India is one of the world’s 18 mega-diverse countries, and it is
ranked the 10th largest country in geographical area (329 million
ha). It has 47,513 floral species (Singh and Das 2013), 28% of which
are endemic, accounting for 11.4% of the total flora on earth
(Arisdason and Lakshminarasimhan 2017). Several studies have
been previously conducted in India to better understand the spe-
cies composition, dispersion, and diversity along disturbance gra-
dients in the Himalayas (Pokhriyal et al. 2012; Gautam et al. 2016;
Malik et al. 2016), North Eastern India (Dutta and Devi 2013;
Majumdar and Datta 2015), Western Ghats (Anitha et al. 2009;
Jayakumar and Nair 2013; Murthy et al. 2016), Eastern Ghats region
(Basha and Rao 2017), Central and North India (Sagar et al. 2003;
Sagar and Singh 2006; Kala 2015). The tropical dry forests of
Northern India are ecologically rich having a wide range of floristic
compositions. It also has a diverse range of economic and medicinal
plant species that the locals rely on to meet their basic re-
quirements and ensure their livelihood (Sharma et al. 2021; Patel
et al. 2022). The tropical forests of the Vindhyan highlands are
fragmenting at a faster rate (Sagar and Singh 2006; Singh et al.
2017), like other tropical forests across the world. The dry decidu-
ous forest of the study area is under a great deal of anthropogenic
strain from fuel and fodder collection, habitat modification, road
construction, cattle grazing, cement, mining and thermal power
industries (Jha and Singh 1990; Sagar et al. 2003; Goparaju et al.
2010; Singh et al. 2017). It is essential to comprehend the vegeta-
tion dynamics in tropical dry forests for the development of forest
conservation strategies, although, this forest type has received less
attention.
However, the earlier studies were primarily concerned with the
phytosociological characteristics of tree species, (Sagar et al. 2003;
Sagar and Singh 2006). Only a small number of studies have been
done on other living forms such as shrubs, herbs (Singh et al. 2017),
seedlings and saplings (Tripathi and Raghubanshi 2014) of tree
which are more crucial for regeneration. The high conservation
priority and the existing dearth of understanding regarding the
impact of disturbance in this threatened ecosystem lead us to
compare the status of different disturbed forest stands. Therefore,
the following concerns were addressed: (1) determine the variation
in diversity, structure, and distribution pattern of trees, shrubs, and
herbs species, (2) how different disturbanceeregeneration combi-
nations influence the successful regeneration status in disturbed
forest stands, (3) to investigate the inter-relationships between
phytosociological attributes in various distinct layers along distur-
bances. Moreover, the current study suggests a framework for
developing suitable conservation priorities and restoration strate-
gies in the context of biodiversity conservation.
Material and methods
Study area
The study area is the Dudhi Tehsil in Sonbhadra district, India,
which is a section of the Vindhyan highlands that is topographically
a rocky plateau with a gently sloping landscape and a valley at the
bottom. The terrain is undulating and characterized by hillocks,
escarpments, and plateau like landforms. The study was carried out
in three villages belonging namely Gardarva (240 15’ 55’’ N and 830
10’ 45’’E), Saudih (240 5’ 54’’ N and 830 07’ 09’’E) and Rajkhar (240
13’ 57’’ N and 830 11’ 56’’E) at altitudes ranging from 313 to 483 m
above mean sea level (Figure 1). The soils are reddish-dark gray,
sandy loam-textured, residual ultisols, and deficient in nutrients
(Singh et al. 1989). The region has a tropical monsoon climate, with
rainy seasons (late JuneeOctober), winter (NovembereFebruary),
Please cite this article as: Sharma A et al., Variation in Species Composition, Structural Diversity, and Regeneration Along Disturbances in
Tropical Dry Forest of Northern India, Journal of Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2022.11.004
and summer (Marchemid-June) throughout the year. During the
research period, the temperature varied from 16 to 32 C for both
the year 2016 and 2017 (Figure 2). Moreover, rainfall also showed
variable trend for different years during the study period with 1344
mm rainfall for the year 2016, whereas 667 mm rainfall for the year
2017. In each yearly cycle, the region experiences nine months of
dry period and remaining three months of humid weather. Rainfall
is erratic and unpredictable, falling mostly during the three months
of rainy season. Furthermore, likelihood of rain in the winter is
hazy, while the weather from March to mid-June is mostly dry with
infrequent precipitation. This results in moisture stress for the
entire year.
Sampling protocol
The three study sites were chosen for field sampling based on
observations in the field and interactions with forest authorities.
The sites were graded in order of how much disturbance they face.
The cumulative disturbance index (CDI) is an index that aggregated
various individual disturbances into a single cumulative score
which was used to characterize disturbance across sites based on
disturbance criteria and indicators such as road, market, grazing,
fuelwood collection, non-timber forest products (NTFPs), leaf-litter
collection, cut stumps, and lopping. The disturbance regimes and
their estimated relative impacts on each of the tree locations were
characterized (Table 1). Sites with a closer proximity to the market
and to the road are subjected to greater utilization pressure and
disturbance. Thus, the site that is farthest from the highway and the
market is given a score of 1 impact, while 2 for medium and 3
impact for nearer one (Sahoo et al. 2020; Patel et al. 2022). Grazing
pressure is quantified in a forest by observing grazing livestock,
browsing patterns, and the presence of grazing animal faeces (Silori
2004; Chaturvedi et al. 2012). Data on fuelwood, NTFPs, and leaf-
litter collection were gathered through interviews and informal
conversations among the inhabitants (Ogra and Badola 2008), and
disturbances were noted based on the information provided by
them. Field observation and visual estimate were also used to
identify cut stumps and lopping (Patel et al. 2022).
Thus, the calculated score of disturbances viz grazing, fuelwood,
NTFPs, and leaf-litter had given 1 impact for low value of distur-
bance and the succeeding, 2 impact for medium and 3 for
maximum score (Table 1) (Sagar et al. 2003; Dhyani et al. 2019). The
overall CDI score of a site was determined by the sum of all factors
assigned to different criteria and indicators. The CDI score varied
from one to twenty-four, with 1 being the lowest and 24 being the
highest score. The total value received was divided into three
ranges in order to divide the range of scores into three classes
(Table 1). Villages are located on the forest fringes in all three forest
sites (Gardarva, Saudih, and Rajkhar), where inhabitants traveled a
distance of 1e12 km to harvest NTFPs, fuelwood, leaf-litter, and
animal grazing. The sample plots have been categorized as least
disturbed (LD), moderately disturbed (MD), and highly disturbed
(HD) categories based on the above-mentioned disturbances of
surrounding villages and the level of human-induced disturbances.
The LD forest stands (0.3 ha  5) are within 6 km of Gardarva
settlement and have experienced the least amount of anthropo-
genic stress in terms of resource exploitation, grazing, and other
repercussions. The sample plots (0.3 ha  5) of MD are located 4 km
from the Saudih village with medium degree of disturbances,
whereas the sample plots (0.3 ha  5) of HD are located more than
2.5 km from the Rajkhar village.
The study was carried out between October 2016 and
September 2018 in three disturbed forest stands to better un-
derstand the diversity of various taxonomic groups such as trees,
shrubs, herbs, saplings, and seedlings in Sonbhadra district,
 A Sharma et al. / Journal of Asia-Pacific Biodiversity xxx (xxxx) xxx 3

Figure 1. Geographical location of Dudhi Tehsil, Sonbhadra district showing three study villages (GRDV Gardarva, SAUD Saudih, and RJKH Rajkhar) in Vindhyan highlands, India.

Vindhyan highlands, India. The random sampling was used to
acquire the most illustrative composition of vegetation. A pre-
liminary survey was also carried out to understand the forest
composition, dominating species, and their common allies. The
nested quadrat approach was used to perform vegetation survey
where sampling was conducted in the peak growing season of
plants (JulyeSeptember and DecembereFebruary). In each of the
five sample plots, a hundred quadrats were put on each site in
order to observe the trees and forty quadrats for the saplings from
the 10  10 m size quadrat. Within the same quadrat, a 5  5 m
size of forty quadrat were also set for the counting of seedlings
and shrubs. Again, forty quadrats of 50  50 cm were spread out
for the measurement of herbs. The most frequent variable used to
quantify tree size is diameter at breast height (DBH), which may
be used as a proxy for tree age. Individuals > 30 cm dbh were
classified as trees, whereas those < 30 cm but > 10 cm were
classified as saplings. The seedlings and shrubs (<10 cm dbh) were
estimated at the collar height above ground level. During the
study, each plant species has its own herbarium specimens which
was photographed in the field. Vernacular names, books and lit-
eratures, published field inventories and floras were all used to
identify plant species (Khare 2008; Joshi 2019). The current
approved names for the identified species were allocated using
online database by means of The World Flora Online (www.
worldfloraonline.org).
Assessment of compositional and structural diversity
Following procedures were adapted for calculating community
metrics including frequency, density, basal area, and importance
value index (IVI) for each species (Curtis 1959; Mishra 1968;
Mueller-Dombois and Ellenberg 1974). Various diversity indices
were used to calculate the species diversity in a given community
such as Shannon and Wiener (1963), Simpson’s concentration of
dominance (CD) index (Simpson 1949), Pielou’s evenness index
(Pielou 1975), Margalef index (Margalef 1958), and Beta diversity
index (Whittaker, 1975).
Abundance to frequency ratio
The plant species distribution pattern was calculated using the
abundance to frequency (A/F) ratio, which was introduced by
Whitford (1949) as a degree of contagiousness. The ratio denotes
regular (0.025), random (0.025e0.05), and contagious (>0.05)
dispersions (Curtis and Cotton 1956).

Please cite this article as: Sharma A et al., Variation in Species Composition, Structural Diversity, and Regeneration Along Disturbances in
Tropical Dry Forest of Northern India, Journal of Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2022.11.004
4 A Sharma et al. / Journal of Asia-Pacific Biodiversity xxx (xxxx) xxx

Figure 2. Climatograph showing rainfall and temperature pattern (2016-2017) of Dudhi tehsil in Sonbhadra District, Vindhyan highlands, India.

Regeneration status ‘Multi-co-linearity’ was decreased by eliminating parameters with a

For better analysis and understanding of the population struc-
ture, all individuals of trees were divided into three categories:
saplings, seedlings, and trees. The regeneration status was deter-
mined by counting the number of saplings, seedlings, and trees in
each of the following categories: (a) ’good’ if seedling > sapling >
tree; (b) ’fair’ if seedling > sapling < tree; (c) ’poor’ if a species only
sustains as saplings, but not as seedlings (saplings might be smaller,
larger or equal to trees); (d) ’none’ if a species is not available as a
sapling or seedling stages, but is present as a tree (Ballabha et al.
2013).
Statistical analysis
Various species diversity indices were computed using Paleon-
tological Statistics (PAST) software, version 3.18 (Hammer et al.
2001). Individual density and diversity indices over different
disturbance gradients were assessed using a one-way analysis of
variance (ANOVA) approach in statistical package for social sciences
(SPSS) version 23. The distribution of plant species in various
families at different disturbance sites was determined using heat
map clustering. The inter-relationship between multiple layers of
tiered vegetation was investigated using cluster analysis and
dendrogram for IVI values along different disturbance parameters.
To analyze the association between species and phytosociological
variables along three disturbance sites, we further utilized the
‘FactoMiner’ package in R software (Le et al. 2008) to perform
principal component analysis (PCA), with the ‘FactoExtra’ package
being used for visual representation (Kassambara and Mundt 2017).
Before analysis, the data were standardized by ‘FactoMiner,’ and
correlation coefficient greater than
0.7 (Verma et al. 2020).
Results
Species composition, richness, and structural diversity
The tree layer comprises of 45 species under 40 genera and 22
families. In terms of disturbance, 37, 33, and 28 tree species were
recorded to occur in LD, MD, and HD forest stands of 15 transects
(2.7 ha) of Sonbhadra district, Vindhyan highlands, India (Figure 3).
Under varying disturbance gradients, a heat map is a visual rep-
resentation of the abundance of individual species in different
families (Figure 4). According to the diversity and species richness
analysis, the total species encompass 41 families with Fabaceae (31
species) followed by Apocynaceae (15 species), Malvaceae (12
species), and Anacardiaceae (10 species).
The density of individual trees does not differ significantly
across the three sites, according to a one-way ANOVA (p ¼ 0.112
Tukey’s honestly significant difference (HSD) test). LD forest stands
had a higher mean tree density (56.24
1.44 hae1) than HD
(53.28
2.67 hae1) and MD (50.27
0.94 hae1) forest stands
(Figure 5). LD stands had a total basal area of 18.53 m2 hae1, 16.11
m2 hae1 in MD stands, and 13.54 m2 hae1 in HD stands. The IVI for
different tree species varied greatly in different forest stands.
Bombax ceiba (25.16), Madhuca longifolia (23.61), and Butea mono-
sperma (17.33) were the most common trees in HD forest stands.
Diospyros melanoxylon (20.85) had the highest IVI in MD stands
closely followed by S. robusta (19.42), H. binata (15.36), and Adina
cordifolia (14.64) (see Appendix A). According to the findings of
species distribution pattern, 78.95% of tree species were dispersed

Table 1. Characterization of disturbance parameters through cumulative disturbance index (CDI) across the different forest stands of Vindhyan highlands, India.

Disturbance parameters

Road Market Grazing Fuelwood collection NTFPs collection Leaf-litter collection Cut stumps Lopping CDI CDI
LD 1 1 1 1 2 1 1 1 9 LD
MD 2 2 3 2 1 3 2 3 18 MD
HD 3 3 2 3 2 2 3 3 21 HD

1-Least disturbed, 2-moderately disturbed, 3- highly disturbed.

Please cite this article as: Sharma A et al., Variation in Species Composition, Structural Diversity, and Regeneration Along Disturbances in
Tropical Dry Forest of Northern India, Journal of Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2022.11.004
 A Sharma et al. / Journal of Asia-Pacific Biodiversity xxx (xxxx) xxx 5

of species displayed a random distribution pattern with some

Figure 3. Taxonomic composition of plant species in least disturbed (LD), moderately
disturbed (MD), and highly disturbed (HD) forest stands of Vindhyan highlands, India.
contagiously and 21.05% species were randomly distributed in LD
stand followed by MD (72.72% contagious, 24.25% random, and
3.03% regular) and HD (42.85% contagious and 57.15% random)
forest stands (Figure 6).
On the other hand, individual density for saplings, seedlings,
shrubs, and herbs differed significantly (p < 0.05) across the three
forest stands. Shorea robusta was the prevalent sapling species with
the highest IVI in all three human-impacted forest stands. The IVI
values in HD forests varied from 12.18 to 22.79, respectively, in
which S. robusta had the highest IVI value, whereas Pongamia pin-
nata had the lowest. The IVI of individual species differed from 8.42
to 18.15 in the MD stand whereas, IVI varied from 9.11 to 17.40 at
the LD stand (see Appendix B). A/F ratio indicated that the majority
contagious distribution. The IVI of seedling species ranging from
10.94 to 38.70. The IVI value in MD stands ranged from 11.38 to
35.57, with Diospyros melanoxylon having the highest IVI in both HD
and MD stands. Whereas, the IVI value ranged from 7.88 to 29.88
which is highest for S. robusta at LD stands (see Appendix C). A/F
ratio of seedling depicted that the maximum species dispersed
randomly to contagious.
IVI of individual shrub density varied from 23.32 to 52.36 for MD
stands, 20.15 to 48.32 for LD, and 18.59 to 41.06 for HD forest
patches. Woodfordia fruticosa had the highest IVI value in the MD
and HD stands followed by Lantana camara in the LD stand (see
Appendix D). According to A/F ratio, majority of the shrubs were
dispersed in a random to regular pattern. The IVI of herb layer
varied from 8.88 to 27.94 in the HD stand, 10.04 to 27.66 in the MD
stand, and 8.42 to 25.09 in the LD stand, respectively (see Appendix
E). Cynodon dactylon had the highest IVI of all the stands. Herb
species displayed a random distribution in this stratum followed by
a regular to contagious distribution.
Diversity indices
The species richness and health of a specific region at a specific
time are measured by diversity indices. Except for seedlings (H’ p ¼
0.086; b, p ¼ 0.041), the Shannon-Wiener and beta diversity index
are substantially different (p < 0.05) across the three sites for trees,
saplings, shrubs, and herbs (Table 2). Simpson’s index was differed
significantly for saplings, shrubs, and herbs (p < 0.05) with the
exception of trees and seedlings (CD, p ¼ 0.175 and 0.451)

Figure 4. Heat map clustering showing the distribution of species individual in various families on the basis of disturbance gradient in least disturbed (LD), moderately disturbed
(MD), and highly disturbed (HD) forest stands of Vindhyan highlands. The extent of individual species is shown from green to red color on the basis of z-score.

Please cite this article as: Sharma A et al., Variation in Species Composition, Structural Diversity, and Regeneration Along Disturbances in
Tropical Dry Forest of Northern India, Journal of Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2022.11.004
6 A Sharma et al. / Journal of Asia-Pacific Biodiversity xxx (xxxx) xxx

Figure 5. Mean density (individuals hae1) of trees, saplings, seedlings, shrubs, and herbs species in least disturbed (LD), moderately disturbed (MD), and highly disturbed (HD)
forest stands of Vindhyan highlands, India.

Figure 6. Distribution pattern of plant species through abundance to frequency (A/F) ratio in least disturbed (LD), moderately disturbed (MD), and highly disturbed (HD) forest
stands of Vindhyan highlands, India.

Please cite this article as: Sharma A et al., Variation in Species Composition, Structural Diversity, and Regeneration Along Disturbances in
Tropical Dry Forest of Northern India, Journal of Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2022.11.004
 A Sharma et al. / Journal of Asia-Pacific Biodiversity xxx (xxxx) xxx 7

Table 2. Species composition and diversity indices in least disturbed (LD), moderately disturbed (MD), and highly disturbed (HD) forest stands of Vindhyan highlands, India.

Variables LD MD HD F p-value Total mean

Species diversity (Shannon-Wiener index)

Trees 3.42
0.03a 3.33
0.02b 3.20
0.01b 19.39 0.000* 3.32
0.02
Saplings 3.18
0.01a 3.10
0.00b 2.82
0.00c 340.57 0.000* 3.03
0.04
Seedlings 2.63
0.02a 2.57
0.02a 2.55
0.02a 3.03 0.086 2.58
0.01
Shrubs 2.18
0.01a 2.06
0.02b 2.29
0.01c 33.71 0.000* 2.18
0.02
Herbs 3.15
0.00a 3.03
0.01a 3.04
0.01b 33.51 0.000* 3.07
0.01
Simpson’s index (CD)
Trees 0.96
0.00a 0.95
0.00a 0.95
0.00a 2.02 0.175 0.95
0.00
Saplings 0.95
0.00a 0.95
0.00b 0.93
0.00b 23.94 0.000* 0.94
0.00
Seedlings 0.91
0.00a 0.90
0.00a 0.90
0.00a .851 0.451 0.91
0.00
Shrubs 0.87
0.00a 0.85
0.00ab 0.88
0.00b 9.75 0.003* 0.87
0.00
Herbs 0.95
0.00a 0.94
0.00a 0.94
0.00b 6.62 0.012* 0.94
0.00
Evenness index (Pielou’s)
Trees 0.83
0.02a 0.85
0.02a 0.88
0.01a 1.21 0.332 0.85
0.01
Saplings 0.93
0.01a 0.92
0.00a 0.93
0.00a .561 0.585 0.93
0.00
Seedlings 0.77
0.01a 0.82
0.02ab 0.85
0.01b 4.26 0.040* 0.81
0.01
Shrubs 0.88
0.01a 0.87
0.02a 0.90
0.01a .516 0.610 0.88
0.00
Herbs 0.90
0.00a 0.90
0.01a 0.87
0.01a 1.81 0.204 0.89
0.00
Margalef index
Trees 5.97
0.02a 5.51
0.01b 4.74
0.04c 425.47 0.000* 5.40
0.13
Saplings 4.66
0.01a 4.50
0.03b 3.58
0.04c 349.90 0.000* 4.25
0.12
Seedlings 3.37
0.01a 3.19
0.04a 3.20
0.02b 8.44 0.005* 3.25
0.02
Shrubs 1.84
0.02a 1.74
0.02a 2.15
0.03b 53.38 0.000* 1.91
0.04
Herbs 4.16
0.01a 3.84
0.03b 4.11
0.04b 28.37 0.000* 4.04
0.04
B-diversity
Trees 3.69
0.07a 3.90
0.07b 3.05
.007b 33.74 0.000* 3.55
0.10
Saplings 2.13
0.04a 2.46
0.06b 2.45
0.07b 8.47 0.005* 2.35
0.05
Seedlings 2.39
0.08a 2.36
0.07a 2.63
0.06a 4.20 0.041 2.46
0.05
Shrubs 2.11
0.05a 1.78
0.09a 1.70
0.08b 7.72 0.007* 1.86
0.63
Herbs 1.24
0.02a 1.41
0.04b 1.49
0.03b 9.99 0.003* 1.38
0.03

Superscripts (a, b) following means within each row differ significantly (p < 0.05) based on Tukey’s HSD test; Significant p values are shown by *.

respectively. Evenness index did not change significantly except
seedlings; however, Margalef’s index differed considerably (p <
0.05) across all the sites. When comparing the Shannon index
values of the tree species, the LD site (3.42
0.03) had the highest
value followed by MD (3.33
0.02) and HD (3.20
0.01) site. For the
tree layer, Simpson’s (CD) index values varied from 0.95
0.00 to
0.96
0.00, evenness index from 0.83
0.02 to 0.88
0.01, Margalef’s
index from 4.74
0.04 to 5.97
0.02, and beta diversity from
3.05
0.07 to 3.90
0.07, respectively.
For shrub layer, Shannon index ranged from 2.06
0.02 to
2.29
0.01, Simpson’s index varied from 0.85
0.00 to 0.88
0.00,
evenness from 0.87
0.02 to 0.90
0.01, Margalef’s index from
1.74
0.02 to 2.15
0.03, and beta diversity from 1.70
0.08 to
2.11
0.05, respectively, in different forest stands (Table 2). According
Figure 7. Regeneration status of tree species in least disturbed (LD), moderately
disturbed (MD), and highly disturbed (HD) forest stands of Vindhyan highlands, India.
to the diversity indices of the herb layer, Shannon index, Simpson’s
index, and Margalef’s index values were all greater at LD site. The
forest stands of HD site recorded the highest value of beta diversity,
whereas in case of evenness, both the LD and MD sites possessed
higher value.
Regeneration status
The regeneration potential of a species is determined by the
density of seedlings and saplings. The general pattern of regener-
ation (overall seedling, sapling, and tree density) demonstrated
that the LD stand (430.55
11.41 seedlings/ha) had the highest
seedling density followed by MD stand (345.00
21.95 seedlings/
ha) and HD stand (262.67
10.07 seedlings/ha). LD stand shows the
population pattern of the stand as seedlings > saplings > trees,
while similar pattern was also followed by MD and HD stand (see
Figure 5).
In the LD stand, only 31.58% species had good regeneration
potential, 31.58% had poor regeneration, 21.05% species were not
regenerating, and rest of the 15.79% species had fair regeneration
status. In the MD stand, 34.38% species had good as well as poor
regeneration, 15.62% had fair, and 15.62% species were not regen-
erating. On the other hand, in HD stands, 39.29% species have good
regeneration potential, whereas 21.43% species have poor and no
regeneration occurs. In nature, 17.85% species have a fair regener-
ation potential (Figure 7). Diospyros melanoxylon, Syzygium
heyneanum, and Holarrhena antidysenterica reflected good regen-
eration potential in LD site and S. robusta and D. melanoxylon
showed good regeneration in case of MD stands. On the other hand,
Ziziphus nummularia, Butea monosperma, and Madhuca longifolia
showed strongly good regeneration potential for HD forest stands.

Please cite this article as: Sharma A et al., Variation in Species Composition, Structural Diversity, and Regeneration Along Disturbances in
Tropical Dry Forest of Northern India, Journal of Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2022.11.004
8 A Sharma et al. / Journal of Asia-Pacific Biodiversity xxx (xxxx) xxx
Discussion
Human-induced disturbances have a significant influence on
the structure and composition of tropical forests (Kumar and Ram
2005; Zhu et al., 2007), although the amount of the impact is
dependent on the severity and form of the disturbances (Chown
2010). It has been objectively demonstrated that anthropogenic
disruptions have a significant impact on biodiversity.
Variation in species composition and diversity
The first hypothesis unveiled by the present study is that
species richness decreased as the disturbance increased from LD
to HD stands. With increasing disturbance, the number of various
layers (trees, saplings, seedlings, shrubs, and herbs) of species
declined, with most accumulation in LD forests. The tree species
found in this study (37, 33, and 28) was lesser than the species
found in three stands (77, 72, and 62 species) of tropical moist
deciduous forests in Odisha (Sahoo et al. 2020) and four plots (72,
71, 70, and 62 species) of tropical forests of Andhra Pradesh (Naidu
and Kumar 2016). The higher species richness in the LD stand is
likely due to the succession process, which tends to promote
species diversity (Yam and Tripathi 2016) while also providing
protection by local people and forest management committees.
On the other hand, anthropogenic disturbances like lopping,
extraction pressure, and other activities may be blamed for the
loss of species richness in MD and HD stands. Fabaceae family has
been recognized as the most prominent family with thirty-one
species, which is also found in Sal forests of Ranchi, Jharkhand
with 17 species (Kumar and Saikia 2020) and a tropical dry forest
in south Ecuador, accounting for 20% of the total species (Graefe
et al. 2020). The presence of leguminous species in Fabaceae
family demonstrated their vast ecological amplitude, and as a
result, these plants are well adapted toward nutrient constraint,
which promotes the establishment of higher-N fixer species
(Singh et al. 2017). Beside these, dryness also encourages the
growth and survival of N-fixer plant species in tropical dry forests
(Vargas et al. 2015).
Species composition, degree of disturbances, and successional
strategies of the stands may all contribute to variances in the basal
area of tree layers (LaFrankie et al. 2006). The basal area of trees in
the present study LD (13 m2/ha), MD (16 m2/ha), and HD (18 m2/ha)
is greater than the value (1.31e13.78 m2/ha) reported by Sagar and
Singh (2006) in their previous study. Other research have found
that evergreen forests in southern India have basal areas between
10.79 to 20.44 m2/ha (Parthasarathy and Sethi 1997).
IVI analysis reveals the social standing of a species and can be
identified as patterns of affiliation of dominating species in a
community. Bombax ceiba (IVI, 25.16) and Madhuca longifolia (IVI,
23.61) were the most common tree species in the HD stand fol-
lowed by Hardwickia binata (IVI, 23.88) and Lagerstroemia parviflora
(IVI, 20.59) dominated the LD stand, whereas Diospyros melanox-
ylon (IVI, 20.85) and Shorea robusta (IVI, 19.42) showed a clear
dominance in MD stand. Kumar and Saikia (2020) reported tree
species with higher IVI values such as Diospyrous melanoxyon (IVI,
13.04) in Sal forests of Ranchi, Jharkhand and tropical moist de-
ciduous forests (IVI, 12.20) of Odisha, India (Sahoo et al. 2020),
respectively. Higher IVI values suggest good regeneration potential,
flexibility, and wide ecological amplitude.
The highest IVI value was recorded by Woodfordia fruticosa in
the shrub layers in MD and LD stands (IVI, 52.36 and 44.08). The
growth and development of tree species may be hampered by
Please cite this article as: Sharma A et al., Variation in Species Composition, Structural Diversity, and Regeneration Along Disturbances in
Tropical Dry Forest of Northern India, Journal of Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2022.11.004
biotic and anthropogenic interferences. As a result, successional
change is hindered, which encourages the growth of alternative
vegetation communities such as non-native and shrub species
(Jhariya 2014; Jhariya 2017). Cynodon dactylon has propagated and
spread abundantly across all sites in the herb layer. The maximum
IVI of C. dactylon was also reported in open grazing areas (IVI, 15.82)
and enclosure areas (IVI, 7.42) in the Harishin rangelands, eastern
Ethiopia (Hailu 2017). The IVI value varied significantly among
disturbances, which might be attributable to environmental factors
like temperature and moisture as well as anthropogenic activities
such as grazing pressure and gathering livelihood goods around the
respective sites (Oraon et al. 2014).
Species distribution pattern
In the case of trees, an analysis of dispersion patterns revealed
that the majority of species exhibited contagious distribution,
albeit a few species had random dispersion; nonetheless, no
species with regular distribution was found. In case of shrubs and
herbs, random and regular distribution patterns were prevalent in
all three stands. According to Odum (1971), contagious distribu-
tion is frequent in nature and is caused by modest but consider-
able differences in environmental circumstances, unequal
distribution of nutrients, or other resources in the environment
which reflects species habitat preferences. It should also be
emphasized that random distribution shows homogenous nature
of species found exclusively in relatively consistent environments,
whereas regular distribution can only be found when individuals
compete fiercely (Joshi et al. 2022). Contagious distribution has
been seen to varied degrees in the tropical forests of India (Singh
et al., 2016; Kumar et al. 2017).
Diversity indices
Diversity indices are an important tool for determining the
structure of a community. Shannon-Wiener diversity index (H’) for
Indian forests was observed to range from 0.81 to 4.10 (Pandey
2000; Sahu et al. 2012). The values for trees (3.20
0.01 to
3.42
0.03) ranging from HD to LD stands, and shrubs (2.06
0.02 to
2.29
0.01) and herbs (3.03
0.01 to 3.15
0.00) ranging from MD to
HD stands, respectively, lies within the range reported for tropical
forests. Simpson’s CD values for tree species varied from 0.95 to
0.96, which is higher than the values (0.28 to 0.62) were recorded
by Mandal and Joshi (2014) in Doon valley, Western Himalaya.
Kumar and Saikia (2020) observed CD values for shrubs (1.10) and
herbs species (0.08), which was lower than shrubs (0.85 to 0.88)
and higher than herbs (0.94 to 0.95) at all sites. This discrepancy
might be explained by the varied amounts of disturbance that exist
in these forests.
Evenness in the present study (0.83 to 0.88 for trees) is higher
than the range of sacred forests (0.31 to 0.81) and community
forests (0.13 to 0.75) of Meghalaya (Suchiang et al. 2020). Evenness
displays equal or similar representation of species and less even
plant communities as shown in the LD and MD stands and in other
form of shrubs and herbs. This finding is also in line with Gotore
et al. (2021) in Miombo woodland, Zimbabwe. Margalef index for
trees is within the range 4.54e23.41 for tropical forests as sup-
plemented by previous studies (Kumar et al. 2010; Sathish et al.
2013). The b-diversity value of trees were higher than those pre-
viously reported by Sagar et al. (2003). There was a significant
decline in shrub, herb, and tree species richness, when collated to
the study provided by Gautam et al. (2006). This drop might be
 A Sharma et al. / Journal of Asia-Pacific Biodiversity xxx (xxxx) xxx 9

attributed to an increase in anthropogenic activities in forest
regions.
Regeneration status
The seedlings and saplings in the LD stand was higher than in
other disturbed forest stands for most species. In the present study,
35.08% of species displayed good regeneration, 16.42% exhibited
fair regeneration, and rest of the 48.5% species showed poor to no
regeneration across the three sites. Similarly, the regeneration in
degraded sal forests of North-eastern Uttar Pradesh was insuffi-
cient, with just 18.6% of woody species depicting good regenera-
tion, 31.6% showing fair, and the remaining species highlighting
poor regeneration (Pandey and Shukla 2001).
Anthropogenic disturbances such as fuelwood collection, cut-
ting, lopping, and grazing were shown to be the primary cause of
poor regeneration in this study. Overgrazing devastates the ground
flora and prevents the regeneration of prominent tree species such
as D. melanoxylon, S. robusta, and L. parviflora. There may be addi-
tional causes for the poor regeneration status of these kind of trees,
such as low biotic potential, which affects fruiting or seed germi-
nation, or effective seedling to sapling stage conservation (Sarkar
and Devi 2014). The survival, growth, and establishment of the
tree seedlings have been recognized to be controlled by in-
teractions among the numerous resources, including light and
water availability, nutrient addition, herbivory, and grass compe-
tition along disturbances (Tripathi and Raghubanshi 2014;
Bhadouriya et al. 2016, 2017). Drought is another important cause
of poor seedling survival, with the sprouting of newly distributed
seeds. It used to be high in June and July, with a peak in recruitment
in August and a decline in seedling population following the rainy
season, as well as increased fatality during the dry winter (Kumar
and Saikia 2021).
Relationship between species and distinct stratified layers of
vegetation
The following is a list of several codes (TR Tree; SHR Shrub; H
Herb) that were used to assess the affinity between distinct levels
of graded vegetation in three disturbed forest stands. For this
purpose, IVI measurements were used to create a cluster analysis
and a dendrogram (Table 3). The cluster analysis of phytosocio-
logical features of distinct stratified layers of vegetation in LD
stands is depicted in the (Appendixes FeH).

Table 3. List of different codes used for various stratified layers of vegetation in Vindhyan highlands, India.

Tree species code Botanical name Shrub species code Botanical name Herb species code Botanical name

TR1 Acacia auriculiformis SHR 1 Abution indicum H1 Achyranthes aspera

TR2 Acacia catechu SHR2 Calotropis procera H2 Ageratum conyzoides
TR3 Acacia nilotica SHR3 Desmodium gangeticum H3 Alternanthera sessilis
TR4 Adina cordifolia SHR4 Helicteres isora H4 Andrographis paniculata
TR5 Aegle marmelos SHR5 Lantana camara H5 Argemone mexicana
TR6 Albizia lebbeck SHR6 Lawsonia inermis H6- Asphodelus tenuifolius
TR7 Alstonia scholaris SHR7 Martynia annua H7 Bacopa monnieri
TR8 Anogeissus latifolia SHR8 Nerium indicum H8 Boerhavia diffusa
TR9 Azadirachta indica SHR9 Opuntia dillenii H9 Cannabis sativa
TR10 Bauhinia variegata SHR10 Randia uliginosa H10 Cassia tora
TR11 Bombax ceiba SHR 11 Ricinus communis H11 Catharanthus roseus
TR12 Boswellia serrata SHR12 Tabernaemontana divaricata H12 Cynodon dactylon
TR13 Bridelia retusa SHR13 Tephrosia purpurea H13 Cyperus longus
TR14 Buchanania lanzan SHR14 Thevetia peruviana H14 Cyperus rotundus
TR15 Butea monosperma SHR15 Woodfordia fruticosa H15 Datura innoxia
TR16 Cassia fistula H16 Eclipta alba
TR17 Diospyros melanoxylon H17 Euphorbia hirta
TR18 Emblica officinalis H18 Evolvulus alsinoides
TR19 Elaeodendron glaucum H19 Hyptis suaveolens
TR20 Gardenia turgida H20 Malvestrum coromandelianum
TR21 Gmelina arborea H21 Mimosa pudica
TR22 Hardwickia binata H22 Ocimum tenuiflorum
TR23 Holarrhena antidysenterica H23 Oxalis corniculata
TR24 Lagerstroemia parviflora H24 Physalis minima
TR25 Lannea coromandelica H25 Plumbago zeylanica
TR26 Madhuca longifolia H26 Rauvolfia serpentina
TR27 Mangifera indica H27 Rumex dentatus
TR28 Miliusa tomentosa H28 Sida acuta
TR29 Mitragyana parviflora H29 Solanum nigrum
TR30 Moringa oleifera H30 Tagetes erecta
TR31 Nyctanthes arbor-tristis H31 Tribulus terrestris
TR32 Oroxylum indicum
TR33 Pongamia pinnata
TR34 Prosopis juliflora
TR35 Pterocarpus marsupium
TR36 Semecarpus anacardium
TR37 Shorea robusta
TR38 Soymida febrifuga
TR39 Syzygium heyneanum
TR40 Tectona grandis
TR41 Terminalia bellirica
TR42 Terminalia chebula
TR43 Terminelia tomentosa
TR44 Ziziphus nummularia
TR45 Ziziphus xylopyrus

Abbreviations TR, Tree; SHR, Shrub; H, Herb.

Please cite this article as: Sharma A et al., Variation in Species Composition, Structural Diversity, and Regeneration Along Disturbances in
Tropical Dry Forest of Northern India, Journal of Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2022.11.004
10 A Sharma et al. / Journal of Asia-Pacific Biodiversity xxx (xxxx) xxx

Figure 8. Principal component analysis (PCA) biplot of different phytosociological variables (IVI ¼ Importance value index; A/F¼Abundance to frequency; BA¼Basal area) in least
disturbed (LD), moderately disturbed (MD), and highly disturbed (HD) forest stands of Vindhyan highlands, India.

Figure 9. Principal component analysis (PCA) biplot of different life-forms (TR¼ Trees; SAP¼ Saplings; SL¼ Seedlings; SH¼ Shrubs; HR¼ Herbs) in least disturbed (LD), moderately
disturbed (MD) and highly disturbed (HD) forest stands of Vindhyan highlands, India.

As a result, the findings envisaged that the inter-relationships
among IVI of tree, shrub, and herb species in LD and MD stands
had substantial relationships between tree and herb species. On the
other hand, tree and shrub species revealed the interdependence of
multiple stratified layers for vegetation forms in HD stands. The
reciprocal process between the multistrata of plants must be pro-
tected for disturbed environments that are under threat. Some of
the species demonstrate the importance of the region in terms of
associated ecological functions which are critical for species sur-
vival (Kujur et al. 2021; Sharma et al. 2021; Patel et al. 2022).

Please cite this article as: Sharma A et al., Variation in Species Composition, Structural Diversity, and Regeneration Along Disturbances in
Tropical Dry Forest of Northern India, Journal of Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2022.11.004
 A Sharma et al. / Journal of Asia-Pacific Biodiversity xxx (xxxx) xxx
Cluster analysis indicated a high relationship between L. parviflora
and H. suaveolens in LD stands. Shorea robusta and Cassia tora,
H. binata and B. diffusa were shown to have a strong relationship in
MD stands, but B. ceiba and C. procera, M. longifolia and O. dilenii
were significantly associated in HD stands. These species should be
given special attention in terms of conservation and management
as well as encouraging the plantation in the disturbed region.
The phytosociological data obtained during the present inves-
tigation were analyzed through PCA. The Pc1 accounted for 45.3%
variance whereas Pc2 had 20.1% variations in both disturbed forest
stands and life-form wise distribution with cumulative score of
65.4%. The PCA result of the phytosociological data about vegeta-
tion along different disturbed forest stands and association of
various life-forms has shown the dominance on major axis
(Figures 8 and 9). In these results, the first four Pcs explained 97.9%
of the variation in the data. Most of the variables were important
for disturbed forest stands as well as for life forms. Based on the
Eigen value, the elucidations of PCs were based on the most highly
associated variables which were organized into three principal
components. Frequency (0.95), density (0.73), and IVI (0.33) were
found to be significant as Pc1. In the first Pc, according to increased
frequency, density and IVI suggested that these factors are closely
connected and if one increases, the other two tend to rise with it.
These associations offered more species diversity, which is a sign of
more intricate and healthy community structure. Additionally,
greater vegetation density and frequency results in a diversified
system of species, which allows for strong interactions that
enhance system stability (Prescott 2002; Giri et al. 2008). The high
IVI points out that these species have developed specific coping
mechanisms through time to adapt to disturbances. IVI (0.91) and
A/F ratio (0.22) are significant as Pc2, while A/F ratio (e0.48) and
density (e0.21) had a strong negative association in the third Pc.
Thus, understanding the general structure and function of any
ecosystem requires data on the floristic characteristics.
Conclusion
Biodiversity influences critical ecological systems making it vital
to evaluate ecosystem health. The plant species are key elements of
the ecosystem since they shape forest architecture and impact
community composition. An excellent database for forest man-
agement is created by recording the diversity and distribution
patterns. The current study gathered data on plant diversity, stand
structural attributes, and regeneration potential in three disturbed
sites (LD, MD, and HD) of Vindhyan highlands, India. As a result of
human-induced disturbances, the species diversity, structure, and
composition of tropical forests have been substantially altered in
recent decades. In the study area, the LD stand has the highest
species richness in terms of total plant species followed by MD and
HD stands. The patterns of diversity and structure are mostly
determined by the site conditions.
The general population pattern of tree species revealed that
seedlings contributed the most to the total population followed by
saplings and finally trees. It demonstrates that in all forest stands,
around 48.5% of the species had poor to no regenerating in-
dividuals. The species composition will be affected in the imminent
future by the presence of such non-regenerating or poor-
regenerating individuals. Future species composition and associ-
ated ecosystem services may be negatively impacted, if adequate
management measures are not implemented. Therefore, a well-
thought-out management strategy is essential for conservation
and sustainable utilization. The outcomes of this study will aid
forest managers and key officials in developing long-term action
plans for better conservation and management of forests.
Please cite this article as: Sharma A et al., Variation in Species Composition, Structural Diversity, and Regeneration Along Disturbances in
Tropical Dry Forest of Northern India, Journal of Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2022.11.004
11
Declaration of competing interest
The authors declare that they have no known competing
financial interests or personal relationships that could have
appeared to influence the work reported in this paper.
Acknowledgments
The authors would like to acknowledge Dean and Director of
Institute of Environment & Sustainable Development, BHU, Vara-
nasi for providing all the necessary facilities. The authors are also
grateful to the Divisional Forest Officer, Renukoot, Uttar Pradesh,
India for allowing to work in the vicinity of forests. AS would like to
acknowledge University Grants Commission, New Delhi for finan-
cial support in the form of fellowship.
Appendixes A-H. Supplementary data
Supplementary data to this article can be found online at
https://doi.org/10.1016/j.japb.2022.11.004.
References
Anitha K, Joseph S, Ramasamy EV, et al. 2009. Changes in structural attributes of
plant communities along disturbance gradients in a dry deciduous forest of
Western Ghats, India. Environmental Monitoring and Assessment 155:393e405.
Arisdason W, Lakshminarasimhan P. 2017. Available at: http://www.bsienvis.nic.in/
Database/Status_of_Plant_Diversity_in_India_17566.aspx. [Accessed 16 May
2019] https://doi.org/10.1007/978-3-642-54601-3_7.
Armenteras D, Rodríguez N, Retana J. 2009. Are conservation strategies effective in
avoiding the deforestation of the Colombian Guyana Shield? Biological Con-
servation 142:1411e1419.
Ballabha R, Tiwari JK, Tiwari P. 2013. Regeneration of tree species in the sub-tropical
forest of Alaknanda Valley, Garhwal Himalaya, India. Forest Science and Practice
15:89e97.
Baraloto C, Molto Q, Rabaud S, et al. 2013. Rapid simultaneous estimation of
aboveground biomass and tree diversity across Neotropical forests: a compar-
ison of field inventory methods. Biotropica 45:288e298.
Basha SK, Rao BRP. 2017. Biodiversity and population structure of trees of Gund-
labrahmeswaram Wildlife Sanctuary, Andhra Pradesh, India. IOSR Journal of
Pharmacy and Biological Sciences 12:87e100.
Bhadouria R, Singh R, Srivastava P, et al. 2016. Understanding the ecology of tree-
seedling growth in dry tropical environment: a management perspective. En-
ergy, Ecology and Environment 1:296e309.
Bhadouria R, Srivastava P, Singh R, et al. 2017. Tree seedling establishment in dry
tropics: an urgent need of interaction studies. Environment Systems and De-
cisions 37:88e100.
Campbell MJ, Edwards W, Magrach A, et al. 2017. Forest edge disturbance increases
rattan abundance in tropical rain forest fragments. Scientific Reports 7:1e12.
Cardelús CL, Woods CL, Bitew Mekonnen A, et al. 2019. Human disturbance impacts
the integrity of sacred church forests, Ethiopia. PLoS One 14:e0212430.
Chaturvedi RK, Raghubanshi AS, Singh JS. 2012. Effect of grazing and harvesting on
diversity, recruitment and carbon accumulation of juvenile trees in tropical dry
forests. Forest Ecology and Management 284:152e162.
Chow J, Doria G, Kramer R, et al. 2013. Tropical forests under a changing climate and
innovations in tropical forest management. Tropical Conservation Science 6:
315e324.
Chown SL. 2010. Temporal biodiversity change in transformed landscapes: a
southern African perspective. Philosophical Transactions of the Royal Society B:
Biological Sciences 365:3729e3742.
Curtis JT, Cotton G. 1956. Plant ecology workbook. Laboratory, field and reference
manual. Minnesota: Burgess Publishing Company.
Curtis JT. 1959. The vegetation of Wisconsin: an ordination of plant communities.
Madison: University of Wisconsin Press.
Cusack DF, Karpman J, Ashdown D, et al. 2016. Global change effects on humid
tropical forests: Evidence for biogeochemical and biodiversity shifts at an
ecosystem scale. Reviews of Geophysics 54:523e610.
De Marzo T, Gasparri NI, Lambin EF, et al. 2022. Agents of Forest Disturbance in the
Argentine Dry Chaco. Remote Sensing 14:1758.
Dhyani S, Maikhuri RK, Dhyani D. 2019. Impact of anthropogenic interferences on
species composition, regeneration and stand quality in moist temperate forests
of Central Himalaya. Tropical Ecology 60:539e551.
Duchok R, Kent K, Khumbongmayum AD, et al. 2005. Population structure and
regeneration status of medicinal tree Illicium griffithii in relation to disturbance
gradients in temperate broad-leaved forest of Arunachal Pradesh. Current Sci-
ence:673e676.
12 A Sharma et al. / Journal of Asia-Pacific Biodiversity xxx (xxxx) xxx
Dutta G, Devi A. 2013. Plant diversity, population structure, and regeneration status
in disturbed tropical forests in Assam, northeast India. Journal of Forestry
Research 24:715e720.
Gautam MK, Manhas RK, Tripathi AK. 2016. Patterns of diversity and regeneration in
unmanaged moist deciduous forests in response to disturbance in Shiwalik
Himalayas, India. Journal of Asia-Pacific Biodiversity 9:144e151.
Gautam MK, Tripathi AK, Manhas RK. 2006. Changes in the Scenario of Dominance
and Diversity in Shorea robusta Gaertn. F. (Sal) Forests of Lachchhiwala, Doon
Valley, India. Indian Forester 132:1645e1652.
Giri D, Tewari A, Rawat YS. 2008. Vegetation Analysis in mixed Banj
(Q. leucotricophora, A. Camus) Tiloj-oak (Q. floribunda Lindl.) forest in Nainital
catchment. Indian Journal of Forestry 31:167e174.
Goparaju L, Jha CS. 2010. Spatial dynamics of species diversity in fragmented plant
communities of a Vindhyan dry tropical forest in India. Tropical Ecology 51:55e
65.
Gotore T, Ndagurwa HG, Kativu S, et al. 2021. Woody plant assemblage
and the structure of miombo woodland along a disturbance gradient in
Hurungwe, Zambezi Valley, Zimbabwe. Journal of Forestry Research 32:1867e
1877.
Graefe S, Rodrigo R, Cueva E, et al. 2020. Impact of disturbance on forest structure
and tree species composition in a tropical dry forest of South Ecuador. Eco-
tropica 22:202002.
Hailu H. 2017. Analysis of vegetation phytosociological characteristics and soil
physico-chemical conditions in Harishin Rangelands of Eastern Ethiopia. Land
6:4.
Hammer O, Harper DAT, Ryan PD. 2001. PAST: Paleontological statistics software
package for education and data analysis. Palaeontologia Electronica 4:1e9.
Jayakumar R, Nair KKN. 2013. Species diversity and tree regeneration patterns in
tropical forests of the Western Ghats, India. International Scholarly Research
Notices 2013:890862.
Jha CS, Singh JS. 1990. Composition and dynamics of dry tropical forest relation to
soil texture. Journal of Vegetation Science 1:609e614.
Jhariya MK. 2014. Effect of forest fire on microbial biomass, storage and sequestration
of carbon in a tropical deciduous forest of Chhattisgarh. Raipur, Chhattisgarh:
Indira Gandhi Krishi Vishwavidyalaya. PhD thesis.
Jhariya MK. 2017. Vegetation ecology and carbon sequestration potential of shrubs
in tropics of Chhattisgarh, India. Environmental Monitoring and Assessment 189:
1e15.
Joshi MC. 2019. Hand book of Indian medicinal plants. Jodhpur, Rajasthan: Scientific
Publishers.
Joshi VC, Bisht D, Sundriyal RC, et al. 2022. Species richness, diversity, structure, and
distribution patterns across dominating forest communities of low and mid-
hills in the Central Himalaya. Geology, Ecology, and Landscapes:1e11.
Kala CP. 2015. Forest structure and anthropogenic pressures in the Pachmarhi
biosphere reserve of India. Journal of Forestry Research 26:867e874.
Kassambara A, Mundt F. 2017. Factoextra: extract and visualize the results of multi-
variate data analyses76; 2017. p. 2.
Khare CP. 2008. Indian medicinal plant: an illustrated dictionary. Berlin: Springer-
Verlag.
Kujur E, Jhariya MK, Yadav DK, et al. 2021. Phytosociological attributes and regen-
eration potential of riparian vegetation in Northern Chhattisgarh, India. Envi-
ronment, Development and Sustainability 24:2861e2886.
Kumar A, Jhariya MK, Yadav DK, et al. 2017. Vegetation dynamics in Bishrampur
collieries of northern Chhattisgarh, India: eco-restoration and management
perspectives. Environmental Monitoring and Assessment 189:1e29.
Kumar A, Ram J. 2005. Anthropogenic disturbances and plant biodiversity in forests
of Uttaranchal, central Himalaya. Biodiversity and Conservation 14:309e331.
Kumar JN, Kumar RN, Bhoi RK, et al. 2010. Tree species diversity and soil nutrient
status in three sites of tropical dry deciduous forest of western India. Tropical
Ecology:51273e51279.
Kumar R, Saikia P. 2020. Floristic analysis and dominance pattern of sal (Shorea
robusta) forests in Ranchi, Jharkhand, eastern India. Journal of Forestry Research
31:415e427.
Kumar R, Saikia P. 2021. Population structure and regeneration status of Shorea
robusta and associated trees in Sal forests of Ranchi, Eastern India. Tropical
Ecology 62:34e51.
LaFrankie JV, Ashton PS, Chuyong GB, et al. 2006. Contrasting structure and
composition of the understory in species-rich tropical rainforests. Ecology 87:
2298e2305.
Lê S, Josse J, Husson F. 2008. FactoMineR: an R package for multivariate analysis.
Journal of Statistical Software 25:1e18.
Majumdar K, Datta BK. 2015. Vegetation types, dominant compositions, woody
plant diversity and stand structure in Trishna Wildlife Sanctuary of Northeast
India. Journal of Environmental Biology 36:409.
Malik ZA, Pandey R, Bhatt AB. 2016. Anthropogenic disturbances and their impact
on vegetation in Western Himalaya, India. Journal of Mountain Science 13:69e
82.
Mandal G, Joshi SP. 2014. Analysis of vegetation dynamics and phytodiversity from
three dry deciduous forests of Doon Valley, Western Himalaya, India. Journal of
Asia-Pacific Biodiversity 7:292e304.
Margalef R. 1958. Temporal succession and spatial heterogeneity in phytoplankton.
In: Buzzati-Traverso A, editor. Perspectives in marine biology. Berkeley: Univer-
sity of California Press. pp. 323e347.
Mayor SJ, Cahill JF, He F, et al. 2012. Regional boreal biodiversity peaks at inter-
mediate human disturbance. Nature communications 3:1e6.
Please cite this article as: Sharma A et al., Variation in Species Composition, Structural Diversity, and Regeneration Along Disturbances in
Tropical Dry Forest of Northern India, Journal of Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2022.11.004
Mishra R. 1968. Ecological workbook. Calcutta, New Delhi: Oxford and IBH Pub-
lishing Company.
Mueller-Dombois B, Ellenberg H. 1974. Aims and methods of vegetation ecology. New
York: John Wiley and Sons, Inc.
Murthy IK, Bhat S, Sathyanarayan V, et al. 2016. Vegetation structure and
composition of tropical evergreen and deciduous forests in Uttara Kannada
District, Western Ghats under different disturbance regimes. Tropical Ecology
57:77e88.
Naidu MT, Kumar OA. 2016. Tree diversity, stand structure, and community
composition of tropical forests in Eastern Ghats of Andhra Pradesh, India.
Journal of Asia-Pacific Biodiversity 9:328e334.
Odum EP. 1971. Fundamentals of ecology. Philadelphia: Saunders.
Ogra M, Badola R. 2008. Compensating humanewildlife conflict in protected area
communities: ground-level perspectives from Uttarakhand, India. Human
Ecology 36:717e729.
Oraon PR, Singh L, Jhariya MK. 2014. Variations in herbaceous composition of dry
tropics following anthropogenic disturbed environment. Current World Envi-
ronment 9:967.
Osuri AM, Chakravarthy D, Mudappa D, et al. 2017. Successional status, seed
dispersal mode and overstorey species influence tree regeneration in tropical
rain-forest fragments in Western Ghats, India. Journal of Tropical Ecology 33:
270e284.
Pandey SK, Shukla RP. 2001. Regeneration strategy and plant diversity status in
degraded sal forests. Current Science 95:102.
Pandey SK. 2000. Population status and regeneration strategy of some perennial le-
gumes in plantation forests of North-Eastern Uttar Pradesh. Gorakhpur, Uttar
Pradesh: Deen Dayal Upadhyaya Gorakhpur University. PhD thesis.
Parthasarathy N, Sethi P. 1997. Trees and liana species diversity and population
structure in a tropical dry evergreen forest in south India. Tropical Ecology 38.
Patel SK, Sharma A, Singh R, et al. 2022. Diversity and distribution of traditional
home gardens along different disturbances in a dry tropical region, India.
Frontiers in Forests and Global Change 5:822320.
Pielou EC. 1975. Ecological diversity. New York: John Wiley and Sons.
Pokhriyal P, Chauhan DS, Todaria NP. 2012. Effect of altitude and disturbance on
structure and species diversity of forest vegetation in a watershed of central
Himalaya. Tropical Ecology 53:307e315.
Prescott CE. 2002. The influence of the forest canopy on nutrient cycling. Tree
physiology 22:1193e1200.
Pretto F, Celesti-Grapow L, Carli E, et al. 2010. Influence of past land use and current
human disturbance on non-native plant species on small Italian islands. Plant
Ecology 210:225e239.
Raj A, Jhariya MK. 2021a. Site quality and vegetation biomass in the tropical Sal
mixed deciduous forest of Central India. Landscape Ecological Engineering 17:
387e399.
Raj A, Jhariya MK. 2021b. Carbon storage, flux and mitigation potential of tropical
Sal mixed deciduous forest ecosystem in Chhattisgarh, India. Journal of Envi-
ronmental Management 293:112829.
Sagar R, Raghubanshi AS, Singh JS. 2003. Tree species composition, dispersion and
diversity along a disturbance gradient in a dry tropical forest region of India.
Forest Ecology and Management 186:61e71.
Sagar R, Singh JS. 2006. Tree density, basal area and species diversity in a disturbed
dry tropical forest of northern India: implications for conservation. Environ-
mental Conservation 33:256e262.
Sahoo T, Acharya L, Panda PC. 2020. Structure and composition of tree species in
tropical moist deciduous forests of Eastern Ghats of Odisha, India, in response
to human-induced disturbances. Environmental Sustainability 3:69e82.
Sahu SC, Dhal NK, Mohanty RC. 2012. Tree species diversity, distribution and
population structure in a tropical dry deciduous forest of Malyagiri hill ranges,
Eastern Ghats, India. Tropical Ecology 53:163e168.
Sakai A, Arbain A, Sugiarto S, et al. 2022. Composition and diversity of tree species
after fire disturbance in a lowland tropical forest in East Kalimantan, Indonesia.
Biodiversitas Journal of Biological Diversity 23:1576e1587.
Sarkar M, Devi A. 2014. Assessment of diversity, population structure and regen-
eration status of tree species in Hollongapar Gibbon Wildlife Sanctuary, Assam,
Northeast India. Tropical Plant Research 1:26e36.
Sathish BN, Viswanath S, Kushalappa CG, et al. 2013. Comparative assessment of floristic
structure, diversity and regeneration status of tropical rain forests of Western Ghats
of Karnataka, India. Journal of Applied and Natural Science 5:157e164.
Shannon CE, Wiener W. 1963. The mathematical theory of communication. Urbana II:
University of Illinois Press.
Sharma A, Patel SK, Singh GS. 2021. Traditional knowledge of medicinal plants
among three tribal communities of Vindhyan highlands, India: an approach for
their conservation and sustainability. Environmental Sustainability:1e35.
Shibru S, Balcha G. 2004. Composition, Structure and regeneration status of woody
species in Dindin Natural Forest, Southeast Ethiopia: An implication for con-
servation. Ethiopian Journal of Biological Sciences 3:15e35.
Silori CS. 2004. Socio-economic and ecological consequences of the ban on
adventure tourism in Nanda Devi Biosphere Reserve, western Himalaya.
Biodiversity & Conservation 13:2237e2252.
Simpson EH. 1949. Measurement of diversity. Nature 163:688.
Singh S, Malik ZA, Sharma CM. 2016. Tree species richness, diversity, and regen-
eration status in different oak (Quercus spp.) dominated forests of Garhwal
Himalaya, India. Journal of Asia-Pacific Biodiversity 9:293e300.
Singh JS, Raghubanshi AS, Singh RS, et al. 1989. Microbial biomass acts as a source of
plant nutrients in dry tropical forest and savanna. Nature 338:499e500.
 A Sharma et al. / Journal of Asia-Pacific Biodiversity xxx (xxxx) xxx 13

Singh P, Dash SS. 2013. Plant discoveries: new genera, species and new records. Kol-
kata: Botanical Survey of India.
Singh R, Sagar R, Srivastava P, et al. 2017. Herbaceous species diversity and soil
attributes along a forest-savanna-grassland continuum in a dry tropical region.
Ecological Engineering 103:226e235.
Subashree K, Dar JA, Karuppusamy S, et al. 2021. Plant diversity, structure and
regeneration potential in tropical forests of Western Ghats, India. Acta Ecologica
Sinica 41:259e284.
Suchiang BR, Nonghuloo IM, Kharbhih S, et al. 2020. Tree diversity and
community composition in sacred forests are superior than the other com-
munity forests in a human-dominated landscape of Meghalaya. Tropical Ecology
61:84e105.
Sullivan MJ, Talbot J, Lewis SL, et al. 2017. Diversity and carbon storage across the
tropical forest biome. Scientific Reports 7:1e12.
Tekalign M, Van Meerbeek K, Aerts R, et al. 2017. Effects of biodiversity loss and
restoration scenarios on tree-related ecosystem services. International Journal of
Biodiversity Science, Ecosystem Services & Management 13:434e443.
Tesfaye G, Teketay D, Fetene M. 2002. Regeneration of fourteen tree species in
Harenna forest, southeastern Ethiopia. Flora-Morphology, Distribution, Func-
tional Ecology of Plants 197:461e474.
Thakrey M, Singh L, Jhariya MK, et al. 2022. Impact of disturbance on biomass,
carbon, and nitrogen storage in vegetation and on soil properties of tropical dry
deciduous forest in Chhattisgarh, India. Land Degradation & Development 33:
1810e1820.
Tripathi SN, Raghubanshi AS. 2014. Seedling growth of five tropical dry forest tree
species in relation to light and nitrogen gradients. Journal of Plant Ecology 7:
250e263.
Vargas GG, Werden LK, Powers JS. 2015. Explaining Legume Success in Tropical Dry
Forests Based on Seed Germination Niches. Biotropica 47:277e280.
Verma P, Singh R, Bryant C, et al. 2020. Green space indicators in a social-ecological
system: A case study of Varanasi, India. Sustainable Cities and Society 60:102261.
Whitford PB. 1949. Distribution of woodland plants in relation to succession and
clonal growth. Ecology 30:199e208.
Whittaker RH. 1975. Communities and ecosystems. 2nd ed. New York: Macmillan.
Yam G, Tripathi OP. 2016. Tree diversity and community characteristics in Talle
Wildlife Sanctuary, Arunachal Pradesh, Eastern Himalaya, India. Journal of Asia-
Pacific Biodiversity 9:160e165.
Zhu J, Mao Z, Hu L, et al. 2007. Plant diversity of secondary forests in response to
anthropogenic disturbance levels in montane regions of northeastern China.
Journal of Forest Research 12:403e416.

Please cite this article as: Sharma A et al., Variation in Species Composition, Structural Diversity, and Regeneration Along Disturbances in
Tropical Dry Forest of Northern India, Journal of Asia-Pacific Biodiversity, https://doi.org/10.1016/j.japb.2022.11.004