Department of Crop and Soil Sciences, Cornell University, Ithaca, NY 14853, USA
2 Department of Natural Resources, Cornell University, Ithaca, NY 14853, USA
Department of Biological and Environmental Engineering, Cornell University, Ithaca, NY 14853, USA
4 Ashoka Trust for Research in Ecology and the Environment (ATREE), Bangalore-560 024, India
ABSTRACT
Indian forests provide a multitude of services to vast populations. Common human activities including livestock grazing,
fuelwood extraction and burning have the potential to impact forest ecosystem structure and function. The effects of these
activities on vegetation, ecology and soil properties were investigated in Bandipur National Park (BNP) in southern India.
Data were collected from 200 sites in four watersheds within the park. Sample sites spanned a degradation gradient measured
by a field disturbance index (FDI). This paper focusses on the impacts on vegetation structure, diversity and composition, and
integrates impacts on soil. Shrub and tree species were inventoried and evaluated in plots 10-m in diameter. The tree layer
was dominated by AnogeissusEmblicaTectona species. The understory was dominated by invasives Chromolaena odorata
and Lantana camara, and native Gymnosporia emarginata. Vegetation plot heights, canopy cover and tree diameters were
negatively correlated with field disturbance resulting in stunted forest stature in degraded sites. Vegetation composition in
degraded watersheds was dominated by small woody tree species and a greater diversity of shrub species. Ordination analysis
was used to integrate soil data with vegetation and disturbance, revealing that deciduous forest in the park is degrading to scrub
forest along with negative impacts on soil characteristics. Consequences of services currently enjoyed by local populations
are discussed. Copyright 2008 John Wiley & Sons, Ltd.
KEY WORDS
anthropogenic disturbance; ecosystem impacts; tropical dry forest; vegetation and soils; ordination; India
INTRODUCTION
Indian forests have been used by communities over
millennia for a variety of uses and practices (Lele and
Hegde, 1997; Shankar et al., 1998a). Of these, livestock
grazing, fuelwood, fodder extraction, and burning to
promote grasses for fodder are common historic and
continuing uses of the forest (Banerjee, 1995; Bhat et al.,
2001; Saha, 2002; Kodandapani et al., 2004; FAO, 2006).
These activities represent substantial pressures on the
forest resource base. The anthropogenic pressures from
communities residing within and on the fringes of these
reserves continue to be enormous. Over five million
people have been estimated to reside within a countrywide network of 593 wildlife reserves and national parks
(Kothari et al., 1995); (Madhusudan, 2005).
Consequently, livestock grazing, fuelwood, fodder
extraction and burning are recognized as a chronic disturbance (Singh, 1998) that can have substantial impacts
on the entire forest ecosystem (Tilman and Lehman,
2001) including impacts on vegetation, soil and water
resources, fauna and micro-climate. However, quantitative studies on the impacts of forest disturbance in
India were relatively few until recently (Shahabuddin and
* Correspondence to: Dr. Vishal K. Mehta, 133 D St. Suite F, Davis CA,
95616, USA. E-mail: vkm2@cornell.edu
Copyright 2008 John Wiley & Sons, Ltd.
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V. K. MEHTA ET AL.
May (Pascal, 1982). BNP falls in the granitegneiss complex of the Archaean (Peninsular gneiss) group, the chief
rocks being gneisses, granites and charnockites. Soils
in the research watersheds are haplic alfisols of moderate depth with ustic soil moisture regime and are well
drained (Bourgeon, 1989; Shiva Prasad et al., 1998). Surface soils are generally sandy clay loam with underlying
argilic horizons of high to medium-base saturation, and
coloured deep red with iron oxides. Soils are moderately
acidic (pHKCl 53), with a cation exchange capacity
(CEC) of approximately 20 cmol/kg, and base saturation
above 70% (Ferry, 1994 p190193). Mehta et al. (2008,
this issue) report the negative impacts of forest disturbance on CEC, through reductions in soil organic carbon
(SOC) and soil clay content. Soil hydraulics are also negatively impacted by reduced available water capacity, and
a likelihood of increased Hortonian overland flow due to
higher cattle trail density in degraded forest areas.
Vegetation in BNP is classified as dry deciduous
woodland to savanna woodland forests of the Anogeissus latifolia Tectona grandis Terminalia tomentosa
type (Pascal, 1982). Most researchers believe that the
present vegetation represents degraded stages of succession, varying from a savanna woodland to low discontinuous thickets. These clump thickets and stunted
scrub forests can be found extensively along the northern
and eastern boundaries of BNP. Grasses belong predominantly to the Cymbopogon and Themeda spp (Prasad and
Sharatchandra, 1984). Grass heights and cover are noticeably low and sparse in the degraded northern borders of
the park. Vegetation is prone to annual, low-intensity fires
at the end of the dry season (Devidas and Puyravaud,
1995).
151
(Figure 1). Details on watershed characteristics are available elsewhere (Mehta et al., 2008, this issue).
At each of the 200 sites, a 10-m-diameter plot
was marked out. Shrub and tree vegetation of above
one foot in height was inventoried within each plot.
Species counts and heights were noted. Individual tree
diameter at breast height (DBH) was measured when
greater than 10 cm, and noted as below 10 cm otherwise. Average plot tree height, understory height and
canopy cover were recorded. Grasses were not inventoried completelyonly the dominant grass species and
height in each plot were noted. Dominant grasses were
either Cymbopogon spp. or Themeda spp. Grass observations were coded as a factor variable with four levels for
use in ordination as follows: G1 (Themeda spp, <3 ft
high); G2 (Themeda spp, 35 ft); G3 (Cymbopogon
spp, <5 ft; and G4 (Cymbopogon spp, 57 ft). Vegetation heights were measured using a Suunto PM-5 optical
height metre (Suunti Finland, Vantaa Finland). Plot slopes
were measured using an Abney level (Lawrence & Mayo,
Bangalore India). Tree canopy cover was measured using
a densiometer (Forestry Suppliers, Inc., Jackson, MI,
USA). Observations of the field disturbance in the plot
were recorded and combined into a field disturbance
index (FDI). FDI comprised of presence/absence observations of five indicator variables: TTrails (cattle or
jeep); Ccut and/or broken stems; D livestock dung;
PPeople; and FFire. For each plot, FDI was calculated as FDI D T C C C D C P C F. Indicator P was
assigned a value of 1 if people were sighted while sampling. The choice of indicator variables was informed
by literature concerning anthropogenic impacts on vegetation ecology in Indian forests (Shankar et al., 1998b;
Kumar and Shahabuddin, 2005).
Data analysis
All analyses were conducted using the open-source R
statistical software (R Development Core Team, 2006).
Ordination was run in R using the VEGAN community
ecology package (Dixon, 2003; Oksanen et al., 2007).
Vegetation structure. Average tree and understory
heights were calculated for each sampled vegetation plot.
Density for each vegetation plot was calculated as the
sum of all individuals per species per plot. Frequency
was the number of sites in which each species occurred.
The mean density of each species (MD1), when it occurs,
was calculated by dividing the species density by the frequency. A mean density (MD2) was also calculated over
all 200 plots. To study the relationship of tree diameter with disturbance, individual tree species in each plot
were classified into tree and saplings if their DBH
was above or below 10 cm, respectively. This allowed
tree density and sapling density estimation for each plot.
Additionally, DBH frequency distributions for three dominant tree species were tested for differences between
degraded and protected watersheds using Chi-square tests
on contingency tables. Effects of disturbance on vegetation structure were investigated by testing the significance
Ecohydrol. 1, 149 160 (2008)
DOI: 10.1002/eco
152
V. K. MEHTA ET AL.
of Pearsons correlation coefficient (r) between gridmean structure variables and grid-mean FDI. Grid-mean
FDI was used as a continuous variable after ensuring that
it was approximately normally distributed.
Vegetation composition and diversity. Diversity (H0 )
was estimated using the Shannon index, which combines
the number of species (richness) and evenness (Legendre
and Legendre, 1998);
H0 D $6i pi log2 pi ;
RESULTS
Field disturbance index
Cut and/or broken stems(C) were most frequently
observed in 55% of the plots. Jeep or cattle trails (T) were
encountered in 40% of the plots. Signs of fire (F) were
observed in 205% of the plots, cattle or cattle dung (D) in
155% and people sighted (P) in only four of the plots.
The combined disturbance variable FDI ranged from 0
to 4there were no plots in which all five indicators
were observed. The livestock indicator (D) was probably under-counted, because the dung deposited by livestock inside the forest boundaries is removed by herders
and sold to coffee estates in the neighbouring district
(Madhusudan, 2005). Similarly, people sightings (indicator P) were also probably under-counted because of the
evasive behaviour of people who are illegally within the
park. The results were not affected by exclusion of this
variable from the combined FDI disturbance index.
The distribution of FDI from the 50 samples in
each of the four watersheds indicates that the two
northern watersheds (D1 and D2, mean FDI of 226
and 198 respectively) are more disturbed than the two
southern watersheds (P3 and P4, mean FDI of 042 and
066 respectively). This was confirmed with K-means
clustering performed on FDI. Cluster 1 (total D 113)
contained 91 out of the 100 sites sampled in the two
northern watersheds (D1 and D2). Cluster 2 (total D
87) contained 78 of the 100 sites sampled in the two
southern watersheds (P3 and P4). As a result, the two
northern watersheds were grouped together as degraded,
and the two southern watersheds as protected. This
binary classification is used below to assess diversity and
composition differences.
Species inventory, frequency and density
In all, 47 species (excluding grasses) were recorded,
including 35 tree species. Themeda triandra and Cymbopogon citratus were the dominant grass species. As also
documented by Prasad and Hegde (1986), most species
occur at low frequencies. Only 5 species occur in more
than 50 plots, and 40 of the 47 species occur in less than
30 plots. The observed species richness (no. of non-grass
species) per plot ranged from 1 to 8 with a mean of
47. The complete vegetation inventory along with local
uses of each species is listed in (Mehta, 2007, Appendix
B). Except for the invasives and two other species, all
species are used locally for at least one, and most often
for several purposes.
Table I below lists the 12 species, including 8 tree
species, that occurred in more than 10% of the 200
plots. The invasives, Chromolaena odorata and Lantana Camara occurred frequently throughout the forest
understory. Among the tree species, A. latifolia, Emblica
officinalis and T. grandis most commonly made up the
tree layer. Few species occurred at high densities within
a plot. The widespread tree species listed in Table I occur
at low densities (MD1), whereas the weeds, C. odorata
and L. camara, are both widespread as well as abundant.
Ecohydrol. 1, 149160 (2008)
DOI: 10.1002/eco
153
Species
Frequency (%)
MD1
MD2
Variable
91
41
52
31
612
178
187
042
295
204
162
222
208
189
121
15
217
079
037
041
027
025
015
017
The denser the vegetation in a plot, the more the number of species (species richness) in the plot (Figure 2).
Exceptions were plots 12 and 122, where only one
(non-grass) species was found at high density, corresponding to clumped distributions of Schrebera spp. and
Phoenix spp., respectively, found at these sites. Three
tree speciest5 (Eucalyptus spp.), t11 (Schrebera spp.)
and t33 (Phoenix spp.)showed a particularly clumped
distribution, occurring at low frequency but high density.
One of the randomly sampled grids overlay a Eucalyptus
plantation planted by the Forest Department, accounting
for its clumped distribution and the presence of this nonnative tree species in the dataset.
Disturbance and structure
Tree heights, diameter and canopy cover. Table II summarizes the average plot heights and canopy cover
over all 200 plots. Tree heights were on average
95 m (C/$ 39 m sd). Tree heights are significantly
negatively correlated with degradation (r D $07, p <
0005). Figure 3(a) shows the grid-means of tree heights
a Only
Mean (1sd)
95
17
128
176
244
220
(39)
(048)
(99)
(787)
(104)
(694)
Range
CV (%)
27195
0030
00489
3203940
008492
6603840
410
281
773
446
426
315
against FDI. Understory heights showed no relationship with degradation. The understory in most degraded
sites was a mix of thorny shrubs and regenerating trees
<10 cm DBH of short stature, whereas in protected sites
the understory was comprised of either only grass or
weeds at low abundance. Figure 3(c) shows the positive
relationship between tree heights and canopy cover for all
200 plots. As expected, tree heights are positively correlated to canopy cover. Disturbance reduces the canopy
cover by replacing tall stature with short-stature forest
types with lower canopy cover (Figure 3). Figure 4(a)
and (b) compares tree densities (DBH >10 cm) and percentage saplings (percentage of tree individuals <10 cm),
respectively against FDI. Grid-means of tree densities and
percentage saplings are plotted against FDI, respectively.
Both correlations (r D $077 and r D 080 respectively)
are significant at the 5% level. Degraded sites have lower
tree (DBH >10 cm) densities, and are dominated by
saplings (juvenile or regenerating individuals) when tree
species are present. Across all tree species, the stature of
the plots, both in tree height and diameter, is reduced to
more degraded plots. There was no significant relationship between understory density and disturbance.
Size class distribution of dominant tree species. Table
III and Figure 5 display the DBH distribution for the
three tree species that were most dominantA. latifolia,
E. officinalis and T. grandis grouping them according
to degraded (D1 and D2) and protected (P3 and P4) forest classes. A. latifolia is the most abundant tree species
across the gradient. Further, T. grandis (teak), a highly
valued timber species, is more abundant in the protected
forest (62 individuals) than in the degraded forest (11
individuals). E. officinalis is a valued non-timber forest
product (NTFP) for its fruit. Its greater relative abundance
in the protected forest likely reflects more frequent interference to its growth in the degraded forest by lopping of
its branches for fruit. Among these three tree species, the
frequency distribution of DBH classes was significantly
different at the 5% level between the degraded and protected watersheds in the case of A. latifolia ($ 2 D 95,
df D 3; three DBH classes used : <10 cm, 1020 cm,
2040 cm). In the case of E. officinalis and T. grandis,
although the total abundance was greater in protected
watersheds, the size class distribution, (proportional to
the total number of trees within each group) was not significantly different between the groups at the 5% level.
Ecohydrol. 1, 149 160 (2008)
DOI: 10.1002/eco
154
V. K. MEHTA ET AL.
Figure 3. Tree heights and canopy cover. Error bars are 1SE (colour on-line).
Figure 4. Tree density and sapling density against FDI (colour on-line).
<10
1020
2030
3040
4050
5065
More
Total
Class 1degraded
Class 2protected
A. latifolia
E. officinalis
T.grandis
A. latifolia
E. officinalis
T. grandis
242
15
0
0
0
0
0
257
56
7
0
0
0
0
0
63
6
0
1
2
1
1
0
11
86
74
16
1
0
0
0
177
70
16
7
0
0
1
0
94
28
5
14
11
2
2
0
62
155
(a)
(b)
Figure 5. DBH classes for three dominant tree species. (a) Northern watersheds; (b) southern watersheds.
Figure 6. Boxplots of Diversity H0 . 1degraded; 2protected (a) All 47 species; (b) only tree species; (c) only understory species.
Copyright 2008 John Wiley & Sons, Ltd.
156
V. K. MEHTA ET AL.
Species
1b
2b
sp1
sp2
sp3
sp5
sp7
sp11
t1
t2
t3
t4
t5
t6
t7
t8
t11
t16
t17
t21
t23
t34
t36
Chromolaena odorata
Lantana camara
Canthium parviflorum
Gymnosporia emarginata
Ziziphus oenoplia
Phyllanthus reticulatus
Anogeissus latifolia
Terminalia alata
Tectona grandis
Terminalia crenulata
Eucalyptus spp.
Acacia chundra
Ziziphus xylopyrus
Dalbergia latifolia
Schrebera swietenioides
Premna tomentosa
Grewia tiliifolia
Chloroxylon swietenia
Careya arborea
Erythroxylon monogynum
Emblica officinalis
051
054
056
027
012
002
071
01
009
015
01
026
01
005
00
004
01
014
016
015
031
084
018
031
00
001
015
076
016
036
022
00
00
003
017
018
01
003
00
008
00
046
a prefix
b
Figure 7. pCCA ordination plot. Grid-mean data with symbols coded by watershed. Symbolssites; italic textspecies; Solid arrowsconstraint
vectors; dashed arrowsfitted vectors; G1 to G4centroids of grass factor variable (colour on-line).
Copyright 2008 John Wiley & Sons, Ltd.
Inertia
Total (%)
Total
08307
100
Conditional
01242
150
Constrained
01926
232
Unconstrained
05139
619
Eigenvalues for constrained axes
CCA1
CCA2
01408
00518
b. Permutation tests for significance (1000 permutations)
df
$2
F
Pr > F
1. Overall
2
01926
299
<0005
2. Individual terms
FDI
1
0141
438
<0005
CEC
1
0052
162
0109
3. CCA axes
CCA1
1
0141
438
<0005
CCA2
1
0052
162
0131
4. Correlation of fitted vectors with CCA axes
CCA1
CCA2
r2
Pr > r
Clay
0656
$0754
05019 <0005
Tree height
085
$0526
06647 <0005
G: G1
$0922
02803 0299
0044
G2
$04048 $03721
G3
04685
00895
G4
13243 $02137
Copyright 2008 John Wiley & Sons, Ltd.
157
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V. K. MEHTA ET AL.
almost no presence in protected forests; (ii) the herbaceous Phyllanthus reticulatus, was more frequent in protected forests compared to degraded forests.
and water availability are key resources for plant productivity and quality (Tilman and Lehman, 2001; Sankaran
and McNaughton, 2005). The negative impacts on both
represent a negative feedback on regeneration. With the
consequent (possibly irreversible) change in vegetation
composition towards unpalatable species and a reduction in grass cover, the feedback towards continuation
of grazing and forage services currently enjoyed by local
populations is also negative. In their related study, Mehta
et al. (2008, this issue) discuss the negative impacts of
disturbance on hydrological services to downstream agriculture. Evidence points to increased vulnerability to both
the ecosystem as well as people (Carpenter et al., 2006a).
We note that greater diversity in degraded forests illustrates that anthropogenic disturbance can increase biodiversity, but still impair ecosystem functioning. Therefore,
from a management perspective, the use of a diversity
index alone as an indicator of ecosystem health is not
advisable. Further, diversity indices are rarely applied to
the whole community and do not address functional differences among species (Krebs, 1999).
Research results point towards potential impacts on
other aspects of ecosystem function. Reduction in canopy
cover by replacement of tall stature forest by open
shrubaceous vegetation also impacts boundary-layer climatology and rainfall partitioning (Giambelluca, 2002).
Reduced soil carbon in degraded sites suggests losses to
soil carbon sequestration services.
Dry tropical forests account for some 60% of the Indian
forest cover (WRI, 1996) and more than 70% of the Western Ghats forests (Kodandapani et al., 2004). In the past
decade, several studies on floristic structure, composition and impacts of forest disturbance have begun filling
the gap in ecological knowledge that existed previously.
However, inter-disciplinary studies on ecosystem functioning that include energy, water and nutrient cycling
are necessary for a comprehensive understanding on local
and regional ecosystem change impacts in a region that
forms the major watershed for southern India and is experiencing considerable forest cover loss and fragmentation
(Menon and Bawa, 1998; Jha et al., 2000; Amarnath
et al., 2003). Coordinated ecological experimental programmes (e.g. BIODEPTH in European grasslands (Hector and Bagchi, 2007) ), will have to be conducted, along
with a network of long-term environmental monitoring
stations. Efforts are under way to set up a coordinated
network of terrestrial, atmospheric and marine monitoring stations for India called INDOFLUX, along the lines
of FLUXNET (Sundareshwar et al., 2007), that will begin
addressing the environmental data gap. Simultaneously,
ecological and socioeconomic research will need to be
integrated to effectively inform forest management policy
(Carpenter et al., 2006b). Forest management response to
uses and impacts described here call for options that meet
local energy and livelihood needs while preserving forest
ecosystem health. Alternative sources of local energy in
the form of options including biogas and social forestry,
and attempts by the Forest Department in Bandipur to
Ecohydrol. 1, 149160 (2008)
DOI: 10.1002/eco
distribute cooking gas cylinders to villages are conceivably viable options. However, the linkage of grazing and
manure collection to export-oriented coffee markets is
changing the traditional outlook of local forest use as
local subsistence practice. Although fuelwood extraction
may be addressed through alternative energy schemes,
the response to grazing pressures under the existing scenario may well involve a combination of forest protection
and the provision of alternative economic opportunity in
the long term. At the time of writing this, researchers
with a partner institution in India (Lele, pers. comm.) are
analysing the socioeconomic aspects of ecosystem change
in the Bandipur region.
ACKNOWLEDGEMENTS
159
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V. K. MEHTA ET AL.