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International Journal of Environment ISSN 2091-2854 79 | P a g e
INTERNATIONAL JOURNAL OF ENVIRONMENT
Volume-12, Issue-1, 2023 ISSN 2091-2854
Received: 16 November 2022 Revised: 26 January 2023 Accepted: 11 February 2023
UNDERSTANDING THE Phyllanthus AND Terminalia chebula SPECIES
POPULATION CHANGE, DEPENDENCY AND SUSTAINABILITY: A STUDY IN MALAI
MAHADESHWARA HILLS WILDLIFE SANCTUARY, SOUTHERN INDIA
Harisha R.P1*, Siddappa Setty R1, Ravikanth G2
1Centre for Environment and Development
2Centre for Biodiversity and Conservation, Ashoka Trust for Research in Ecology and the Environment,
Royal Enclave, Srirampura, Jakkur Post, Bangalore 560 064, India
*Corresponding author: hari@atree.org
Abstract
Non-Timber Forest Products (NTFPs) are vital sources of livelihood for forest-dependent communities across
the globe. This study examined the NTFPs species (Phyllanthus emblica, P. indofischeri, and Terminalia
chebula) population change determined by the dependency, disturbances, and accessibility in the dry tropical
forest of Malai Mahadeshwara (MM) Hills wildlife sanctuary. The long-term monitoring population data
were analyzed across three time periods; 2000-01, 2010-11, and 2020-21. The participatory research methods
were used to assess the dependency and accessibility which influence the population structure. The multi-
factor linkage approach was used to identify the significant drivers of population decline. The results indicated
that grazing, fire, hemi-parasite infection, and Lantana invasion influenced the tree population structure and
regeneration of study species. This study has also indicated variations and changes in the interrelationship
among factors that have a significant role in shaping NTFPs species population structure. Multiple factor
analysis determined that grazing, fire, and lantana have significant impacts on population structures,
regeneration, and fruit production of NTFPs species. The study recommended that forest managers should
consider a site-specific adaptive approach and multiple factors models and inclusive management tools
provisioned in recent policies like the Biological Diversity Act -2002 and Forest Rights Act-2006 would hold
great potential for developing sustainable use and co-management practices.
Keywords: Community; Dependency; Forest resources; Sustainability
DOI: https://doi.org/10.3126/ije.v12i1.52444
Copyright ©2023 IJE
This work is licensed under a CC BY-NC which permits use, distribution and reproduction in any medium
provided the original work is properly cited and is not for commercial purposes
International Journal of Environment ISSN 2091-2854 80 | P a g e
1. Introduction
Forests provide a range of socio-economic and environmental benefits that are essential for human life
(Millennium Ecosystem Assessment, 2005; Food and Agriculture Organization, 2014). Globally, more than
a billion people depend directly on NTFPs for their livelihoods (MEA, 2005). Another three billion people
indirectly depend on NTFPs for many economic and social benefits (Agrawal et al., 2013). Thus, NTFPs are
found to be a vital source of livelihood for people from forest fringe communities across the world. Many
studies revealed that NTFPs realize various functions in enhancing human well-being and rural livelihoods,
and it was widely acknowledged globally (Angelsen et al., 2014; Shackleton et al., 2015; Shackleton and
Pullanikkatil, 2018).
In India, NTFPs species contribute 2.7 billion USD per year and provide 55% of the total employment in the
forestry sector (FAO, 2014). These NTFPs are harvested for food, medicines, and handicrafts apart from
subsistence use and as a source of cash income. Past studies have reported that fruit harvesting alone does not
have a significant impact on the NTFPs population, site-specific multiple factors could change the population
structure (Ticktin, 2004; Brummitt and Bachman, 2010; Scoles et al., 2012). Shaanker et al. (2004a),
identified that dependency, degradation, and disturbance factors could contribute to the depletion of NTFPs
resources in tropical forest settings. However, intermittent extraction of NTFPs has influenced stability in the
population structure of NTFPS species (Kodandapani et al., 2004; Shackleton et al., 2005; Varghese et al.,
2015). Moreover, resource accessibility changes dependency on NTFPs and related livelihoods (Marshall et
al., 2006). These changes in local context and livelihoods might have influenced or contributed to changes in
the NTFPs population structure (Endamana et al., 2016). It is also apparent that the contribution of NTFPs to
income varies across ecological settings, seasons, and income levels (Marshall et al., 2006). Considering the
importance of NTFPs in the livelihoods of local communities needs more attention on developing appropriate
policies and evolving strategies for the better management of NTFPs.
Several attempts have been made to understand the dynamics of the NTFPs species population and identify
sustainable harvesting practices (Gaoue and Ticktin, 2010; Ravikanth and Setty, 2017). Most of the NTFPs
studies have exclusively focused on assessing the impacts of harvesting and other anthropogenic factors on
regeneration, productivity, and population structure as well as on genetic diversity (Murali et al., 1996;
Padmini et al., 2001; Ganesan and Setty, 2004; Sinha et al., 2005; Ramesha et al., 2007; Ravikanth et al.,
2009). Further, studies also recommended developing effective conservation strategies, sustaining the
livelihoods of dependent communities and ecology of NTFPs species using anthropogenic factors at multiple
scales, times, disciplines, and linkages (Peres et al., 2003; Schmidt et al., 2011; Ticktin et al., 2012). The
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attributes such as harvest practices, dependency dynamics, and species response patterns are crucial to evolve
an ecologically sustainable practice (Ticktin and Shackleton, 2011).
This study is intended to contribute to the knowledge of understanding the interrelationship between the
dependencies; accessibility change and derived disturbance which influenced the species' response toward
developing sustainable practices. It helps in developing strategies for finding nature-based solutions to address
and manage forest resources. Henceforth, this study formulates the hypothesis that a combination of fruit
harvest frequency, derived disturbances and accessibility play a crucial role in reshaping the population
structure, recruitment rate, and fruit productivity.
2. Materials and methods
2.1. Study site
The study was conducted in Malai Mahadeshwara (MM) Hills which is located in the Chamarajanagara
district of Karnataka in South India. It lies between latitudes 11˚ 55˚N and 12˚ 13˚N, and longitudes 77˚ 30˚
and 77˚ 47˚E with Cauvery Wildlife Sanctuary to its North-East and the Biligiri Rangaswamy Temple Tiger
Reserve (BRT) to its South-West (Figure 1). The sanctuary covers an area of 906 km², hills and valleys are
roofed with extensive forests with a chain of continuous mountain peaks (elevations ranging from 600-1380
m) and mosaic habitat. The climate of MM Hills is moderate throughout the year, hot summer and cold winter.
It receives rain from the southwest monsoon between May-August and from the northeast monsoon between
September-November with a pronounced dry period between January and March. The mean annual
temperature is 35.3˚C and varies between 24˚C in winter to 42˚C in summer (Shaanker et al., 2004a).
The forest harbors rich fauna and flora used by local people in traditional healthcare, cultural, and religious
systems. It has unlike forest types such as dry deciduous (64.34%), scrub woodland (20.50%), and scattered
patches of moist deciduous and riparian forest (2.47%) (Aravind et al., 2010). The sanctuary has been invaded
by Lantana camara which brought significant change to local biodiversity and livelihoods.
Since the 1990s, the NTFPS has played a critical role in the livelihoods of local communities in this region as
there has been an increased demand for forest products in the local markets (Shaanker et al.,
2004b). Phyllanthus emblica, P. indofischeri, and Terminalia chebula are common NTFPS species and fruits
have been harvested as a source of income (Rist et al., 2008; Shaanker et al., 2004a). Phyllanthus species are
widely known as Indian gooseberry, and Amla (in Hindi). Phyllanthus fruits have been used extensively to
make pickles, jams, and herbal drinks; in Ayurvedic medicine and cosmetics. Terminalia chebula is
commonly known as black myrobalan and it is also extensively used in Ayurveda medicine and cosmetics
industries. The populations of T. chebula and P. emblica overlap in forest habitats. However, the populations
of Phyllanthus species are spatially segregated; P. indofischeri occurs at lower elevations (<900m), in scrub
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forests, and P. emblica occurs at higher elevations (>800m) in dry deciduous forests (Setty et al.,
2008). T.chebula is slow-growing shade-tolerant species and has poor seed germination capacity in natural
conditions (Anitha et al., 2010).
There are 31 settlements (villages) scattered within and periphery of MM Hills forest. About eight villages
constitute a homogenous community called Soligas; whereas another 23 villages are constituted
heterogeneous communities called Soligas and Bedagampana. Soligas are the indigenous tribes living in MM
Hills forest for centuries and they have historically been engaged in hunting, NTFP collection, and shifting
cultivation for their livelihood (Harisha et al., 2015). Shifting cultivation and hunting were banned in 1972
under the Wildlife Protection Act, following which the Soligas sedentarized in settlements called 'podu' and
continued settled agriculture (Murali et al., 1996). Most of the villages were notified as revenue villages in
1913. They received titles to their cultivable land ranging in size from 0.5 to 2 hectares under Forest Rights
Act 2006.
Figure 1: Location of the study area, MM Hills, Karnataka, India
2.2. Assessment of population structure and regeneration
The regeneration capacity of a species is reflected in the population size- class distribution; as the assessment
of population structure is convenient and provides information on species population status. Twenty transects
measuring 10m X 100m sample plots were laid for each species in the M M Hills forests in 2000-01 and
monitored annually till the year 2020-21. Sample plots were used to estimate the population structure of P.
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emblica, P. indofischeri, and T. chebula. Sample plot locations are marked based on NTFPs resource maps
developed by harvesters during participatory resource appraisal (PRA) exercises. All the individuals of the
study species in the sample plots were tagged permanently with aluminium tags bearing a unique number
(seedlings to adult trees). The diameter at breast height (DBH) of each tree/sapling (measuring above 1 cm
DBH) was recorded at 1.3 meters height.
The density of Lantana camara was estimated by counting the number of individual clumps in five sample
quadrates measuring 5mX5m within each of the 20 plots. Similarly, grazing intensity was assessed by
counting the dung and pellets within the belt transect measuring two-meter wide and 100 meters long, within
each of the 20 plots. The forest fire incidents were recorded during the study periods and the extent of the area
burned was also mapped in all the plots.
2.3. Assessment of fruit productivity and extraction
All reproductive adults >5cm DBH stem in the transect were identified and estimated fruit production before
harvest to understand the stock of fruits and after harvest to understand the level of extraction. One month
before the fruit harvest, the number of fruits was estimated. The fruit production of the tree was estimated
visually by counting five fruiting branches which were selected randomly out of the total fruiting branches
from each tree and also the number of fruiting branches was estimated. Later average fruits per branch were
calculated for five selected branches and multiplied by the total fruiting branches. . It has been repeated during
the post-harvest to understand the level of fruit extraction by the community. The number of hemi-parasite-
infected branches was counted and branch cut was also recorded before and after the fruit harvest to
understand the level of the tree getting damaged while harvesting the fruits.
2.4. Assessment of dependency and accessibility
The dependency and accessibility of the local community have been assessed by conducting a socio-
economic survey in 2000-01, 2010-11, and 2020-21. The four forest villagers who have traditionally been
involved in NTFPS collection were selected based on proximity to the road (to the transect points) and the
number of people harvesting NTFPS based on the frequency of harvest. Semi-structured interviews were
conducted for 92 households from four villages which were 10% of the total households per village. Villages
such as Anehoa(228 hh), Kumbudukki(232), Gorasane(242), and Kokkubare(218) were located close to the
sample plots and were selected for the household interview. The systematic sampling methods were used to
select households for the interview by considering the family size (number of people in their family) and based
on their occupation (NTFPS collection, farming, daily wage, and others) to draw reliable information. Both
men and women participated in the household survey.
The household survey was conducted from October to November, before fruit harvest, and after harvest from
April to May. The interviews were 1-3 hours long in the local language (Kannada) and information was
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validated by revisiting the households. The household survey captured the socio-economic profile, source of
income, and livelihood change. The community perception of the present status of NTFPS resources,
dependency, and accessibility change under the forest protection regime was also documented.
2.5. Focus group discussion
Most of the participants were NTFPs harvesters and were members of the cooperative society called Large
Scale Adivasi- Multi-Purpose Societies (LAMPS). The community leaders who have been involved in the
Forest Rights Act (FRA) 2006 implementation, and served as Forest Department employees for a long time
in the study area have also participated. During the FGD, participants prepared resource maps and harvesting
timelines for the NTFPS species. They also shared the constraints involved in NTFPS collection, the impacts
of policy change on resources, and their livelihoods.
Annual fruit harvest estimate accessed from LAMPS for the past 20 years (from 2000-01 to 2020-21).
LAMPs are the nodal agency for marketing, monitoring, and management of NTFPs resources. It consists of
a management committee and an executive committee formed by Soliga community representatives and
forest department officials.
2.6. Data analysis
We categorized the data collected across three study periods into; indicators of population
structure (population density, regeneration rate, and annual fruit yield); disturbance variables (grazing
intensity, lantana density, fire frequency, branch cut, and hemi-parasite infection); Dependency
variables (LAMPs data on the quantity of fruit collected and income from NTFPS-which is a direct indicator
of dependency); accessibility change (tenure, policy change and community perceptions on forest regime
which had a significant impact on NTFPs accessibility).
Understanding the variation of variables across study periods: The population structure of study species
that recruit regularly was characterized by reverse J curves (Lykke, 1998; Wright et al., 2003). The size class
distribution was estimated for the study species by using population monitoring data obtained from three study
periods. Repeated measure MANOVA statistics was used to evaluate the variance of mean values of
population structure, disturbance, and dependency indicators across study periods.
Determine the relationship between indicators and variables: We used multiple regression analysis to
determine the relationship between population structure, disturbance, dependency, and accessibility variables.
The data was also used to determine the significant relationship between and within variables. The study used
population density, recruitment rate, and fruit yield as response variables to determine the impact of
disturbances, dependency, and accessibility.
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Characterizing the variables which determine the population structure
Data on variables of dependency, disturbance, accessibility, and indicators of population structure were
averaged across sampled points. The binomial distribution model was employed for seedling density and fruit
production to check the distribution of data points. A generalized Linear Model (GLM) was employed to
explore the link to disturbance, dependency, and indicators of the population structure of the study species.
The GLM was employed for three study periods separately. The analysis started with an initial model of eight
variables to reduce the co-linearity. The variables used were lantana density, grazing intensity, fire frequency,
hemi-parasite infection, branch cut, and extraction rate versus population density, regeneration, and fruit
production.
To assess the overall effect of each predictor variable on population structure and productivity, we have
calculated AICc (second-order Akaike information criteria, used for small sample size), differences in AICc,
and AICc weights for each model in the two model sets (T. chebula and Phyllanthus spp.) using the R package
AICc moving. We calculated the mean and 95% confidence interval for the regression coefficient of each
predictor variable by averaging coefficients across all models, weighted by the AICc weight of each model.
A backward selection based on the Akaike Information Criterion (AIC) was operated to determine the best
combination of factors that have a significant impact on population density, regeneration, and fruit production.
Data analyses have been carried out using Microsoft Excel and R (version 3.3.1).
3. Results
3.1. Understanding the factors across study periods
3.1.1 Indicators of population structure
Size class distribution: The population (>5cm DBH.) density of species varied across the study periods. In
2000-01(P.emblica-20.8/ha.; P. indofischeri -16.9/ha.; T.chebula-43.2/ha.); In 2010-11 (P.emblica-
18.4/ha.; P.indofischeri-15.5/ha.; T.chebula-41.2/ha.); and in 2020-21 (P.emblica-20.1/ha.; P. indofischeri -
16.2/ha.; T.chebula-42.6/ha.). All three species (P. emblica and P. indofischeri and T. chebula) showed three
different size-class distributions (including seedlings, saplings, and adults) across the study period. In the year
2000-01, for all three species, the relative frequency of seedlings and saplings was lower than in 2010-11 and
2020-21 resulting in an unstable population structure. However, in 2010-11, all three species showed a
sporadic size-class distribution. In 2020-21 all three species showed a reverse j-shaped curve (stable, self-
maintaining population) which indicates that the population is stable compared to previous study periods (Fig.
2).
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Figure 2: Size class distributions of species across study periods
Regeneration: The seedling density of Phyllanthus species was lower in 2000-01(89/ha.) compared to 2010-
11 (114/ha.) and (205/ha.) in 2020-21. Similarly, the percentage of seedling survival rate increased from 1.8,
2.1, and 8.9 in 2000-01, 2010-11, and 2020-21 respectively. The seedling density of T. chebula was43/ha.,
64/ha. and 104/ha.in 2000-01, 2010-11, and 2020-21 respectively. The percentage of seedling survival rate
increased from 1.3, 1.8, and 6.3 in 2000-01, and 2010-11 to 2020-21 respectively.
Fruit production: The percentage of fruiting trees across study periods was 35.6% for P. emblica, 31% for P.
indofischeri, and 54.6% for T. chebula. There were no significant differences in the percentage of fruiting trees
for all the species across the study periods. Similarly, the annual fruit production for P. emblica was 135.5
kg./ha., 158kg./ha., and 206.72 kg/ha in 2000-01, 2010-11, and 2020-21 respectively; similarly for P.
indofischeri, it was 115.5 kg./ ha., 114kg./ha., and 161.72 kg/ha., in 2000-01, 2010-11 and 2020-21
respectively; In case of T. chebula it was 235.5 kg./ha., 260.4 kg./ha. and 270.72 kg/ha. in 2000-01, 2010-11,
and 2020-21 respectively.
3.1.2. Dependency variables
Annual fruit harvesting: LAMPS is a tribal co-operative society constituted by a cooperative and the forest
department that purchases fruits/NTFPs buy fruits from harvesters and sells them in the open market with or
without value addition. The fruit harvested by the harvesters in 2000-01 was 67.5 tons, which decreased to
0.00
20.00
40.00
60.00
515 25 35 45
Relative frequency(%)
Size class(DBH in cm.)
Phyllanthus indofischeri
2000-01
2010-11
2020-21
0
20
40
60
80
515 25 35 45 >50
Relative frequency(%)
Size class(DBH in cm.)
Phyllanthus emblica
2000-01
2010-11
2020-21
0
20
40
60
515 25 35 45 >50
Relative frequency(%)
Size class(DBH in cm.)
Terminalia chebula
2000-01
2010-11
2020-21
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4.1 tons in 2020-21 in the case of Phyllanthus. In the case of T. chebula, fruit collection was 13.7 tons in 2000-
01 and it reduced to 3.1 tons in 2020-21 (Fig. 3).
Figure 3: Phyllanthus and Terminalia fruit harvested (source: LAMPS)
NTFPs dependency and income change: Fruit collection season is from November to February, Soliga
community from 31 villages in and around forest area collect and sell them to LAMPS. The number of
households involved in harvesting from each village ranged from 30% to 55% and usually, both men and
women participated in fruit harvest.
The percentage of income from fruit/NTFPS collection in 2000-01 was 43 % per household per capita,
whereas in 2010-11 it reduced to 21% and it further reduced to 13 % in 2020-21. Fruit harvesting was mainly
determined by availability, accessibility, and market linkage. The younger generation has been migrating to
nearby towns for jobs and migratory income has increased by two fold in recent years. The percentage of
people migration increased from 18% to 51% in 20 years that’s very significant (Fig. 4).
Figure 4: NTFPs dependency and income change from the past two decades.
R² = 0.5755
R² = 0.3211
0
10
20
30
40
50
60
70
80
Fruit harvested (Tons)
Amla Arale Expon. (Amla) Expon. (Arale)
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3.1.3. Disturbance variables
Grazing intensity: The forest area was wide open for grazing until the year 2013 when the forest department
declared MM Hills as a wildlife sanctuary. There were 19 cattle sheds spread across the MM Hills forest area
before being declared a wildlife sanctuary. The forest department put a restriction on cattle grazing and banned
cattle sheds inside the wildlife sanctuary. The grazing intensity (the number of cattle and goats) inside the
forest reduced drastically from 2013 onwards. It was indicated indirectly in terms of pellet density change
which was recorded during the study periods. Similarly, the cattle and goat populations in the forest villages
were also reduced. According to the official record, the cattle population reduced from 6500 (in 2000-01) to
3400 (in 2020-21), and the goat population from 4900 (in 2000-01) to 2750 (in 2020-21).
Lantana density: There has been a major change in forest structure due to the invasion of L. camara. This
species spread in the landscape after the mass bamboo felling in the 1970s and development activities in the
region. In the first period (2000-01) the density of lantana was very high (6342/ha.) compared to 2010-11
(4850/ha.) and 2020-21 (3930/ha.). Local people started harvesting lantana from the forest in 2003-04 to make
furniture and utility products, which resulted in a gradual decrease in lantana density.
Fire frequency: Incidents of forest fire got reduced after the declaration of the wildlife sanctuary. In 2000-01,
thirteen sample plots were partially burnt whereas in 2010-11 only six sampled plots were partially burnt and
in 2020-21 only three sampled plots got burned. The incidents of fire reduced across forest areas in general.
In 2000-01, 1.25 hectares of forest area got burned, it was reduced to 0 .84 hectares in 2010-11, and in 2020-
21 it was only 0.38 hectare got burned.
Branch cut and hemi-parasite infection: Branch cut and hemi-parasite infection were found only
in Phyllanthus species. In T. chebula no hemi-parasite infection but branch cuttings were recorded. Therefore,
we considered Phyllanthus species (P.emblica and P.indofischeri) for the analysis. Branch cutting was much
more common in 2000-01 and 2010-11. It is because of open access to use resources in the forest, grazing
livestock, and harvesting fruits. The rate of the branch cut correlated with hemi-parasite infection. Fruit
harvesters have been removing hemi-parasite-infected branches during fruit harvest to check the further
spread of hemi-parasite and grazers to feed their goats. In 2000-01 the frequency of branch cutting was 63%
for P.embica and 59.3% for P.indofischeri. Whereas in 2020-21 branch cutting was less (11%
for P.embica and 6.1% for P.indofischeri). Similarly, hemi-parasite infection was very high in 2000-01(36%
for P.embica and 9.3% for P.indofischeri compared to 2020-21 (23.3% for P.embica and 4.6 %
for P.indofischeri.
3.1.4. Tenure and accessibility change
Since 2013 the MM Hills forest was a reserve forest that offered open access to resource use and collection of
NTFPs in the landscape. Also, the forest opened to cattle sheds inside the forest and another livestock grazing
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in 2013, MM Hills became a wildlife sanctuary which resulted in a ban on grazing, restriction on NTFPS
collection, and other produce collected by the community from the forest (Table 1). Also, the forest
department initiated strict patrolling and improved management practices (for example, issuing NTFPs
collection passes, and timings), which reduced incidents of forest fire, a reduction in grazing, and a significant
reduction in fruit collection.
Table 1: Showing changes in tenure rights across study periods
Attributes
2000-01
2010-11
2020-21
Forest protection status
Reserve
Reserve
Wildlife sanctuary
Using forest for grazing, firewood etc.
Open
Open
Ban
Accessibility fruit harvest
No restriction
No restriction
Regularised
Forest Right Act 2006 implementation
No policy exist
Started claiming
rights
Rights over NTFPs
collection received
3.2. Variables and indicators variations across study periods
The study hypothesized that variables such as population structure, disturbance, dependency, and accessibility
change significantly vary across study periods (2000-01, 2010-11, and 2020-21). The analysis of variance
(ANOVA) between study periods revealed that variables such as population density and percentage of fruiting
trees had not changed significantly. Whereas, regeneration (F=18.52, P=>0.01 for Phyllanthus and F=7.47,
P=>0.01 for T. chebula) rate, fruit production (F=5.52, P=>0.05 Phyllanthus and F=6.37, P=>0.05 for T.
chebula) have seen significant change. Disturbance variables such as grazing intensity observed significant
change (measured indirectly by counting pellets and dung) (F=12.52, P=>0.01 for pellets and F=9.43,
P=>0.01 for dung). Similarly, lantana density (F=8.45, P=>0.05) and fire frequency (F=8.43, P=>0.05) have
changed significantly between study periods (Appendix 1).
3.3. Understanding the variable's relationship with indicators of population structure.
Listed variables such as fruit extraction, hemi-parasite infection, branch cut, lantana density, grazing intensity,
fire frequency, the quantity of fruit harvest, and income were used for multiple regression analysis. Dependent
variables such as tree density, regeneration rate, and fruit productivity were tested with variables to evaluate
the relationship and characterize the impacts on population structure.
In 2000-01 grazing intensity, hemi-parasite infection, branch cut, and forest fire had significant negative
impacts on the population density of Phyllanthus species, in the case of T. chebula, grazing intensity and fire
frequency had significant negative impacts on tree population density. Similar trends were seen in 2010-11 as
well for all three species (Appendix 2). Whereas, in 2020-21 all of the variables had a significant positive
relationship with the population density of Phyllanthus species and T. chebula.
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The regeneration rate was negatively affected by grazing and forest fire for all the species during 2000-01 and
2010-11. Whereas in 2020-21 fruit productivity had a significant positive impact on the regeneration rate
(Supplementary file 1). In 2000-01 and 2010-11, the fruit productivity of Phyllanthus species was negatively
affected by hemi-parasite, branch cut, and grazing, in the case of T. chebula grazing had significant negative
impacts (Appendix 2).
Fruit production in the case of T. chebula factors such as grazing (r² =-0.561, P=<0.05) had a significant
negative impact on fruit production. Whereas, adult tree density (r²=0.403, P=<0.05) has a positive
relationship with fruit production. While branch cut showed a significant negative impact on P.
indofischeri fruit production, in the case of P. emblica, hemi-parasite infection significantly affected fruit
productivity. Adult tree density (r²=0.413, P=<0.05) also had a positive relationship with fruit production in
both the species of Phyllanthus (Appendix 2). The branch cut reduced fruit production
for Phyllanthus species.
3.4. Characterizing the roles of the variable on indicators of population structure
The model selection was based on coefficient value, associated standard error, and corresponding p-value of
the covariates in the model that influence species population density, regeneration, and fruit productivity.
Based on theoretical as well as field knowledge, there were seven models with different combinations of
variables/covariates were used to test against the population density, regeneration, and fruit productivity. The
detailed model selection procedure for covariates which influenced population density, regeneration, and fruit
production for all three species across the study periods was provided in (Appendix 3a,3b,3c). In the
supplementary file three best models with covariates influencing population density, regeneration, and fruit
production of study species was listed.
Population density: In 2000-01 and 2010-11 the variables such as grazing, fire, and hemi-parasite had a
significantly negative influence on the population density of P.emblica. In 2020-21, lantana density and hemi-
parasite had a significant negative influence on population density. Similarly, the P.indoficheri population
structure had a significant negative association with variables such as branch cut; grazing, and fire in 2000-01
and 2010-11. But in 2020-21 lantana density and forest fire had a significant negative influence on population
density. In the case of T. chebula, grazing and fire had a significant negative role in shaping population
structure in 2000-01, whereas in 2010-11 variables like grazing, lantana density, and fire had a significant
impact on population density. But In 2020-21 only lantana density and fire had a significant impact on the
population density of T. chebula (Table 2).
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Table 2: Summary of the best model with covariates that influence population density of the study species.
Species
Year
Covariates
wi
AIC
AICc
Δ AICc
K
Coefficients
SE
p
PE
2000-01
HP+GI+FF
0.92
108.72
106.83
0.16
4
HP:-0.064
GI: -0.299
FF:-0.183
0.005
0.008
0.064
<0.05*
<0.05*
<0.05*
2010-11
HP+GI+FF
0.91
112.43
108.32
0.01
4
HP:-0.064
GI: -0.299
FF:-0.192
0.008
0.069
0.031
<0.05*
>0.05
>0.05
2020-21
HP+LD
0.83
102.41
103.16
0.06
3
HP:-0.064
LD:-0.076
0.041
0.231
<0.05*
<0.05*
PI
2000-01
BC+GI+FF
0.89
117.86
113.91
0.12
5
BC:-0.001
GI: -0.258
FF: -0.076
0.006
0.064
0.008
>0.05
<0.05*
<0.05*
2010-11
BC+GI+FF
0.79
111.92
106.03
0.04
4
BC:-0.064
GI: -0.103
FF:-0.214
0.086
0.067
0.041
<0.05*
>0.05
<0.05*
2020-21
LD+BC
0.86
105.32
105.89
0.05
3
LD:-0.319
BC:-0.031
0.031
0.072
<0.05*
>0.05
TC
2000-01
GI+FF
0.94
104.14
102.65
0.10
4
GI: -0.010
FF: -0.038
0.004
0.007
<0.05*
<0.05*
2010-11
GI+ LD+ FF
0.86
119.82
114.68
0.23
4
GI:- 0.062
LD: -0.246
FF:-0.031
0.013
0.092
0.021
<0.05
<0.05*
<0.05*
2020-21
LD+FF
0.91
102.34
104.31
0.04
3
LD:-0.049
FF:-0.031
0.012
0.031
<0.05*
>0.05
TC=Terminalia chebula; PE=Phyllanthus emblica; PI=P.indofischeri; LD=Lantana density; GI=Grazing
intensity; FF=Fire frequency; TD=Tree density; FT=Fruiting trees; BC=Branch cuts; AIC=Akaike
Information Criterion; AICc= AIC corrected for small-sample bias; Δ AICc= difference in AICc values
between each model and the model with the lowest AICc; wi=AICc model weight; K=Number of parameters
estimated by the model; SE=Standar Error.
Regeneration: Hemi-parasite, branch cut, and lantana have affected the regeneration of P.emblica.
Similarly, the regeneration of P.indoficheri is affected by grazing, fire, and lantana. In the case of T.
chebula variables like grazing, fire, and lantana had a significant impact on both seedlings and saplings across
International Journal of Environment ISSN 2091-2854 92 | P a g e
study periods (Table 3). This indicated that seedlings and saplings were more sensitive to grazing and fire
during drought years which caused high mortality in seedlings and saplings.
Lantana density had a positive association with seedling density but the sapling survival rate was poor across
study periods and species. It is shown that lantana influences regeneration by controlling soil erosion and
increasing soil moisture and nutrition. But transforming from seedling to sapling differed between vegetation
type and presence and absence of L. camara. The positive association between lantana and seedling density
in the dry deciduous forest for all the study species was observed. It indicated that unregulated disturbance
other than fruit harvest in the forest had greater impacts on population structure. Dry tropical tree species are
more prone to physical damage by branch cut, grazing, and fire during drought years (2000-01 and 2007-08).
Table 3: Summary of the best model with covariates that influence regeneration of the study species.
Species
Year
Covariates
wi
AIC
AICc
Δ AICc
K
Coefficients
SE
P-value
PE
2000-01
GI+FF
0.94
102.15
101.5
0.06
3
GI: -0.299
FF:-0.021
0.064
0.041
<0.05*
<0.05
2010-11
GI+FF
0.91
108.43
105.91
0.13
3
GI: -0.113
FF: -0.057
RF: 0.021
0.008
0.069
0.031
<0.05*
>0.05
>0.05
2020-21
FF+LD
0.89
119.82
114.68
0.23
3
RF: 0.031
LD: 0.311
0.013
0.092
<0.05*
>0.05
PI
2000-01
GI+FF+LD
0.93
123.81
120.41
1.62
4
GI: -0.001
FF:-0.051
LD: 0.258
0.006
0.064
0.052
>0.05
<0.05*
>0.05*
2010-11
GI+LD+FF
0.97
113.92
108.31
0.61
4
GI: -0.064
LD: 0.103
FF:-0.031
RF: 0.052
0.086
0.067
0.030
0.051
<0.05*
>0.05
<0.05*
>0.05
2020-21
FF+LD
0.88
117.41
114.81
0.41
3
RF: 0.031
LD: 0.046
0.431
0.836
<0.05*
>0.05
TC
2000-01
GI+FF
0.94
104.14
107.32
0.061
3
GI: -0.010
FF: -0.038
0.004
0.007
<0.05*
<0.05*
2010-11
GI+FF
0.86
116.82
119.45
0.089
3
GI:- 0.062
FF:- 0.246
RF: 0.062
0.013
0.092
0.032
<0.05*
<0.05*
<0.05*
2020-21
FF+LD
0.91
109.56
113.84
0.082
3
RF: 0.021
LD: 0.039
0.084
0.073
<0.05*
>0.05
TC=Terminalia chebula; PE=Phyllanthus emblica; PI=P.indofischeri; LD=Lantana density; GI=Grazing
intensity; FF=Fire frequency; TD=Tree density; FT=Fruiting trees; BC=Branch cuts; AIC=Akaike
International Journal of Environment ISSN 2091-2854 93 | P a g e
Information Criterion; AICc= AIC corrected for small-sample bias; Δ AICc= difference in AICc values
between each model and the model with the lowest AICc;
wi=AICc model weight; K=Number of parameters estimated by the model; SE=Standard Error
Fruit productivity: Hemi-parasite and branch cuts have affected the fruit productivity of P.emblica.
Similarly, the fruit production of P.indoficheri is affected by grazing, and branch cutting. In the case
of T.chebula variables like grazing, branch cut, and density of fruiting trees had a significant impact on fruit
production (Table 4).
Table 4: Summary of the best model with covariates that influence fruit productivity of the study species.
Species
Year
Covariates
wi
AIC
AICc
Δ AICc
K
Coefficients
SE
P-value
PE
2000-01
HP+BC
0.92
102.53
104.32
0.14
3
HP: -0.064
BC: -0.076
RF: -0.299
0.005
0.008
0.064
<0.05*
<0.05*
<0.05*
2010-11
HP+BC
0.91
118.31
119.69
0.01
3
HP: 0.113
BC: 0.057
RF: 0.142
0.008
0.069
0.042
<0.05*
>0.05
<0.05*
2020-21
HP
0.95
111.83
112.19
0.31
2
HP: 0.041
RF: 0.524
0.052
0.034
<0.05*
<0.05*
2000-01
BC+GI
0.89
121.04
121.89
0.51
3
BC: -0.001
RF: -0.258
GI: -0.076
0.006
0.064
0.008
>0.05
<0.05*
<0.05*
2010-11
BC+GI
0.97
106.32
107.82
0.02
3
BC: 0.064
RF: 0.103
GI: 0.210
0.086
0.067
0.083
<0.05*
>0.05
<0.05*
2020-21
RF+FT
0.88
114.02
115.19
0.43
3
RF: 0.412
RD: 0.023
FT: 0.028
0.042
0.039
0.105
<0.05
>0.05*
<0.05*
TC
2000-01
GI+BC
0.94
104.14
106.43
0.08
3
GI: -0.010
RF: -0.038
0.004
0.007
<0.05*
<0.05
2010-11
GI+BC
0.86
112.82
114.94
0.21
3
GI: 0.062
RF: 0.246
0.013
0.092
<0.05*
<0.05
2020-21
FT
0.93
108.5
109.43
0.05
2
RF: 0.531
RD: 0.217
FT: 0.071
0.712
0.372
0.619
>0.05*
>0.05
<0.05
TC=Terminalia chebula; PE=Phyllanthus emblica; PI=P.indofischeri; HP=Hemi parasite; GI=Grazing
intensity; FF=Fire frequency; BC= Branch cut; RF= Rainfall; RD= Rainy days; FT=Fruiting trees;
BC=Branch cuts; AIC=Akaike Information Criterion; AICc= AIC corrected for small-sample bias; Δ AICc=
International Journal of Environment ISSN 2091-2854 94 | P a g e
difference in AICc values between each model and the model with the lowest AICc; wi=AICc model
weight; K=Number of parameters estimated by the model; SE=Standard Error.
4. Discussion
4.1. Understanding the indicators across the study period
The life strategy of a species largely depends on adding more new individuals to its population. Long-term
monitoring of the regeneration history (the frequency and abundance of seedling establishment) was one of
the methods used across the globe to understand population change (Charles,1998). Results demonstrate that
the size class distribution of the study species was stable (reverse ‘J’ shaped curve) in 2020-21 compared to
previous years. A significant decrease in disturbances, low human dependency, and restrictions on fruit
harvest improved regeneration and reduced mortality in 2020-21.
In contrast, in 2000-01 and 2010-11 species population showed very poor seedling and sampling
establishment and the distribution curve reflects that regeneration is severely limited. Grazing, fire, and branch
cut were high in 2000-01 and 2010-11 compared to 2020-21. Frequent drought and high accessibility (open
to access resource) during the period from 2000-01 to 2010-11 in the forest were practiced. These findings
were consistent with population studies elsewhere in the tropical deciduous forest (Kodandapani et al., 2004;
Mandle et al., 2012; Varghese et al., 2015). Results suggest that grazing, branch cut, and the fire was the main
contributors to decreased population density and poor regeneration. These drivers were common across
tropical forests with similar forest habitats and climatic conditions like in India (Sinha et al., 2005; Ticktin et
al., 2012). The study suggests that fruit harvest alone did not have a significant impact on population structure,
regeneration, and fruit production.
Similarly, fruit productivity was low in 2000-01 and 2010-11 compared to 2020-21, which indicates that
combined effects of disturbance factors such as hemi-parasite infection, branch cut, and grazing resulted in
low fruit productivity in Phyllanthus species.
4.2. Evaluating the relationship between the indicators
The study found a significant relationship between changes in size class distribution and variables. The
population structure of Phyllanthus species was shaped by hemi-parasite, grazing, and fire, in the MM Hills
forest. Hemi-parasite is impacting mainly the tree population and significantly increases tree mortality.
Grazing and fire had affected the seedling establishment and sampling transformation into an adult. A high
rate of seedling and sapling mortality was happening in areas where the high intensity of grazing and frequent
fire occurred. Several studies have reported similar impacts in many tropical forest studies elsewhere in India
(Kodandapani et al., 2004; Tiktin et al., 2012).
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In the case of T. chebula, rainfall, grazing, and fire played a role in seedlings and saplings
development. T.chebula also has significant constraints in the fruit set due to predation apart from
microclimatic conditions that were vital for seed germination (Varghese et al., 2015). Lantana density alone
does not have a significant impact on the tree density of Phyllanthus and T.chebula. However, in fire-prone
areas, Lantana density had a significant impact on the seedling and sapling density of both Phyllanthus and T.
chebula species. High grazing usually had low lantana density, which also had a low chance of fire but poor
regeneration of seedlings and a low rate of sapling establishment for both species. The areas such as low-
grazing, medium lantana density, and less fire-prone areas showed good regeneration of seedlings and a high
rate of sapling establishment for both species. Areas with high-density lantana with low grazing were prone
to high fire and had poor regeneration and sapling establishment. These patterns revealed that in tropical
forests elsewhere in India, South Asia, and Africa (Shackleton et al., 2005; MEA, 2005). During drought
years, grazing in fire-prone areas was subjected to a major change in population structure as recorded during
study periods 2000-01 and 2007-08. On the contrary, the low-grazing, less frequent fire, and medium lantana
density have supported the regeneration, seedling, and sapling density in 2014-15 for both Phyllanthus and T.
chebula species.
The change in fruit productivity was determined by rainfall, rainy days and hemi-parasite, and branch cut in
the case of Phyllanthus species. In the case of T. chebula, rainfall and rainy days, and grazing determined fruit
productivity. These findings were consistent with fruit productivity surveys elsewhere in the Western Ghats
(Murali et al., 1996; Mandle et al., 2012). The disturbance factors such as grazing and branch cut were the
common factors that limited fruit production across study species. The infestation by hemi-parasite was an
additional factor that played a significant role in limiting fruit production in the case of Phyllanthus species.
Previous studies reported significant impacts of hemiparasites on fruit production (Shaanker et al., 2004b;
Setty et al., 2008; Mandle et al., 2012). The adult tree density and the number of fruiting trees were the sources
of stocks for fruit productivity for most of the NTFPs species. Several studies have shown that fruit harvest
had a less significant impact on fruit productivity (Murali et al., 1996; Mandle et al., 2012; Varghese et al.,
2015) similar to our study results. Fruit harvest had less impact on the flowering and fruiting phenology of
woody species unless populations were subjected to other stressors like continuous drought, hemi-parasite,
fire, grazing, and lantana invasion.
4.3. Dependency and harvester’s perspective
The change in fruit productivity and regeneration was discussed during the focused group discussions with
harvesters. The harvesters observed a high rate of adult mortality due to hemi-parasite infection, prominent in
the case of P. emblica. They had been practicing branch cutting to avoid further infection of hemi-parasite
during fruit harvest (Rist et al., 2008). They also observed a high rate of fruit abortion and fruit predation in
International Journal of Environment ISSN 2091-2854 96 | P a g e
the case of T. chebula. Research studies also reported that the low fruit set rate is due to poor pollination and a
high rate of immature fruit abscission (Varghese et al., 2015). All three species had high fruit predation and
were severely affected by grazing and fire and lantana invasion (Shaanker et al., 2004a; Ticktin et al., 2012).
The focus group discussion revealed that their dependency on NTFPs species had reduced drastically since
the last decade. The reasons for the reduction in dependency were; a) change in resource accessibility and
imposition of rules and regulations which inhibit people from NTFPS collection b) migration to nearby towns
and cities to work as laborers facilitated by increased transportation facilities c) volatile markets for fruits
harvested from forest and d) socio-economic welfare schemes from the government such as MNREGA,
public distribution systems, etc. These reasons were drivers of livelihood change among forest-dependent
communities across India (Gadgil et al., 1995; Lele and Rao, 1996; Rajindra, 2015).
The combination of fruit abortion, fruit predation, hemi-parasite, fire, grazing, lantana invasion, and drought
has played a significant role in shaping the population structure of study species. Harvesters also mentioned
that hemi-parasite infections have increased over the years. Usually, the harvesters remove the hemiparasites
on Phyllanthus trees during fruit harvesting. The fruit harvesting decrease in recent years led to high infestation
and increased mortality of trees.
4.4. Accessibility
The community also revealed that more than 80% of people, below the age of 35 migrate as laborers to granite
quarries and cities for construction work. Around 45% of the women migrate seasonally to work in the coffee
estates. About 60% of people, above the age of 50 years do farming and local labor work. The NTFPS
cooperative societies (LAMPS) and forest department staff played an important role in the collection of
NTFPs (Lele and Rao, 1996). Lack of accessibility and a volatile market also influenced the reduction of
NTFPs collection.
The People's perceptions of policy implications, livelihoods, and resource accessibility in the forest have been
recorded. About 82.6% of people perceived that wildlife sanctuary status largely affected their livelihoods and
resource accessibility compared to other conservation policies in the region (Lele and Rao, 1996). Similarly,
62% of people felt that the Wildlife Protection Act of 1972 impacted their livelihoods. However, 96.7% of
people mentioned that the Forest Rights Act 2006 would positively affect their livelihood (Roshni et al., 2019).
About 86% were disappointed with the delay in implementation, especially for NTFPS accessibility rights
(community forest rights). The Declaration of wildlife sanctuary resulted in the banning of NTFPS harvest,
increased patrolling, and fire preventive measures restriction on human activity (grazing, fuelwood collection,
and NTFPS collection) inside the wildlife sanctuary.
International Journal of Environment ISSN 2091-2854 97 | P a g e
4.5. Characterizing the potential indicators
The NTFPs species in the tropical forest are subject to multiple disturbances, and the magnitude of individual
and combined effects had been poorly studied. Our results illustrate that untangling the effects of common
drivers was critical for developing effective management strategies. The mix and match between indicators
played a significant role in shaping the population structure, regeneration, and fruit production of the study
species. As previous studies suggest translating as many indicators/drivers as possible into models using a
multi-model inference-based approach which would be more helpful for understanding the population
structure. In this study, we used disturbance, dependency, and accessibility variables in general linear model
(GLM) analysis along with population structure variables. Moreover, Akaike Information Criterion (AIC)
was used to determine the combination of factors, which might have had an impact on the population
structure. This would provide useful insights for developing management and conservation strategies for
NTFPs species.
The combination of the availability of adult trees and an increase in fruit trees would lead to higher
regeneration. Similar patterns were observed across the country with similar climatic and habitat conditions
(Sundaram, 2011; Varghese et al., 2015). During the study period, fire intensity was recorded from 2000 to
2020. Fire season was correlated with drought year generally; dry deciduous forests, sites close to villages,
and cattle camps were identified as fire-prone areas. Because of intensive grazing and frequent fire,
consecutive droughts from 2001 to 2003 and in 2012 and 2013 resulted in poor regeneration and low seedling
establishment for the species, in general.
Specifically, hemi-parasite control is essential as it harms the tree population of Phyllanthus species and
increases adult mortality (Rist et al., 2008). Moreover, the decline of the Phyllanthus species population from
2000-01 to 2010-11 was unable to explain by the invasion of Lantana camara alone (Sundaram, 2011).
Instead, hemi-parasite, fire, and grazing along continuous drought could be impacted. The causes of this
mortality are consistent across plots and other parts of the forest (Ganesan and Setty, 2004). Moreover, the
long-term monitoring revealed the indirect effects of lantana on the NTFPs population (Sundaram, 2011).
Recent studies suggest that the population dynamics of fruit harvesting species may be perplexed with other
drivers, but these indicators have not been considered (Schmidt et al., 2011 Sundaram, 2011; Gooden et al.,
2009). For instance, effective management requires addressing the drivers affecting most of the population's
decline. As studies revealed low regeneration even in low lantana density areas (Schmidt et al., 2011).
Similarly, hemi-parasite was found to be a major threat to Phyllanthus species in both BRT and MM Hills
forest and it affected other species as well (Rist et al., 2008; Setty et al., 2008; Shaanker et al., 2004a).
The pattern of regeneration in T. chebula was determined by the low level of genetic diversity, due to high
levels of inbreeding (Ravikanth and Setty, 2017; Shaanker et al., 2004b; Anitha et al., 2010). In addition, a
International Journal of Environment ISSN 2091-2854 98 | P a g e
low level of germination and a high proportion of seeds with a lack of embryos and pollinator deficiency has
been proposed as possible contributors to low regeneration in the case of T. chebula (Varghese et al., 2015)
however, these need a further vigorous investigation.
4.6. Implications for forest management and conservation
The sustainability of NTFPs species in tropical forests subjected to multiple stressors and the importance of
identifying potential factors which have a greater impact on the population is yet to be answered clearly.
However, fruit harvest was blamed for observed declines in the study species and state policy imposed a ban
on fruit harvest in protected areas. Prohibiting the fruit harvest was ineffective and would not improve the
status of these populations; instead, it resulted in negative economic repercussions for harvesters (Sandemose,
2009). Therefore, it is necessary to direct inclusive conservation policies to frame action plans to reduce the
disturbance identified. For example, effective control of grazing, and monitoring the distribution of invasive
species like lantana could aid the regeneration of the species.
Moreover, effective control strategies for hemi-parasite infection and branch-cut could be the participatory
monitoring and adaptive management strategy. It was emphasized in the Forest Rights Act 2006 to ensure the
sustainable use of resources and the regeneration of the study species (Roshni et al., 2019). Promoting
indigenous control methods for hemi-parasite infection, controlled grazing, and an enterprise-based Lantana
Craft Center(LCC) model would help in the management of lantana invasion and better management.
Lantana density could be a prime factor in reducing fire intensity and grazing during drought years. A better
understanding of the interrelationships between disturbances, dependency, and accessibility could shape the
population structure, regeneration, and fruit production and develop effective forest management policy. The
use of a long-term, multidisciplinary and inclusive approach allowed us to determine the combination of
factors/indicators that should be the focus that could help in developing management strategies.
It is important to assume that unpredictable climatic factors could change the whole scenario and affect the
population structure. Moreover, forest fire and developmental activities in and around natural forests could
impact the health of the forest and the stability of the NTFPs species population. The poor implementation of
inclusive policies such as the Forest Rights Act 2006 could affect the resource sustainability in the forests.
5. Conclusions
The study found that the impact of disturbances, dependency, and accessibility on shaping the NTFPs species
population was very significant. Our results suggested that invasion of L. camara, grazing, hemi-parasite, and
branch cuts are major drivers impacting the population structure of both Phyllanthus species and Terminalia
chebula. The cumulative effect of hemi-parasite infection, grazing, and branch cut had a significant negative
impact on fruit production and regeneration of study species. However, with changes in dependency,
International Journal of Environment ISSN 2091-2854 99 | P a g e
accessibility, and disturbance due to protected area status, the adult tree density, and fruiting trees would have
a significant positive impact on regeneration.
Control of grazing, hemi-parasite spread, and reduction in branch cut would be the better strategy to retain a
stable tree population and regeneration. The multi-factor linkage would evolve to identify drivers of
population decline and formulate effective conservation policy. NTFPs species and the population are at risk
due to multiple disturbances, and it is essential to have inclusive management strategies and practices.
Moreover, site-based adaptive management and participatory resource monitoring approach as provisioned
in the recent Forest Rights Act-2006 would hold great potential for developing sustainable use and co-
management practices in the study area.
Authorship contribution
RPH and SSR contributed to the study conception, design, conceptualization, methodology, software,
investigation, and data curation. The first draft of the manuscript was written by RPH. Material preparation,
data collection, and analysis were performed by RPH. RPH, SSR, and RKG supervised and validated the
work. All authors commented on previous versions of the manuscript, and read and approved the final
manuscript.
Declaration
The authors declare that they have no conflict of interest.
Acknowledgment
This work was supported by Ford Foundation and Dorabji Tata Trust grants to ATREE and by NSF grant
OISE03-52827 to TT. The work was partly supported by USAID and DBT, India. We thank the Soliga
community, T. Ganesh, C. Trauernicht, and K.S. Bawa for their contributions; the Karnataka State Forest
Department for granting permission for this research; and O. Gaoue, L. Mandle, I. Schmidt, and two
anonymous reviewers for valuable comments on previous drafts.
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