ECOHYDROLOGY

Ecohydrol. 1, 149 160 (2008)

Published online in Wiley InterScience
(www.interscience.wiley.com) DOI: 10.1002/eco.14

Ecosystem impacts of disturbance in a dry tropical forest

in southern India
Vishal K. Mehta,1 * Patrick J. Sullivan,2 M. Todd Walter,3 Jagdish Krishnaswamy4
and Stephen D. DeGloria1
1

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

Received 15 December 2007; Accepted 6 April 2008

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.

Prasad, 2004). A majority of these studies has separately

focussed on impacts on either vegetation (Shankar et al.,
1998b; Madhusudan, 2000; Sagar et al., 2003; e.g. Kumar
and Shahabuddin, 2005), soils (e.g. Sahani and Behera,
2001) or wildlife (e.g. Madhusudan, 2004). Few studies have attempted an integrated ecosystem functions
approach to the impacts of forest disturbance, one that
elucidates the linkages and feedbacks between, for example, changes in vegetation structure and composition, soil
impacts, nutrient cycling and hydrology. Research that
addresses the multi-functionality of ecosystems (Hector
and Bagchi, 2007) and the linkages of ecosystem functioning to services they underpin (Balvanera et al., 2006)
is especially important given that ecological change can
alter the ecosystem services currently enjoyed, and can
increase the vulnerability of people and ecosystems to
further changes (Carpenter et al., 2006a).
To address this critical issue, a research campaign
was conducted in a deciduous forest in Bandipur
National Park,(BNP) India, focussing on forest disturbance impacts on three aspects of forest ecosystem
function, namely, vegetation ecology (this paper), soils
(Mehta et al., 2008, this issue) and watershed-scale
hydrology (Krishnaswamy et al., in preparation). In this
paper, we (i) investigate the impact of disturbance on
vegetation structure, diversity and composition in BNP;

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V. K. MEHTA ET AL.

(ii) integrate impacts on vegetation with impacts on soil

processes presented earlier (Mehta et al., 2008, this issue)
by means of ordination analysis; and (iii) based on (i) and
(ii) discuss the feedbacks and implications for services
currently enjoyed by local populations.
STUDY AREA
The study area is the BNP (Longitudes 76 120 76 530 E,
Latitudes 11 350 11 580 N), in the southern state of Karnataka. The park is approximately 874 km2 in area and is
located leeward of the Western Ghats mountain range and
biodiversity hotspot (Figure 1 in Mehta et al., 2008, this
issue). It is home to 17 critically endangered, endangered
and vulnerable plant and animal species (2001 IUCN
Red List, www.redlist.org), and is one of the Project
Tiger conservation areas in the country. BNP lies in
the sub-humid transitional zone between the humid highelevation backslopes of the Western Ghats and the semiarid Deccan plateau interiors (Bourgeon, 1989 p3756).
Elevations in BNP range from a low of 700 m on the
western border, to a maximum of 1450 m at Gopalswamy
Betta in the central portion of the park. Topography
is gently undulating. The climate is tropical savanna,
hot, seasonally dry (IMD, 1984). Average annual rainfall
ranges from 900 to 1200 mm within the park boundaries
(Pascal, 1982). Rainfall is trimodal, occurring as convectional pre-monsoon storms (MarchMay), the southwest
monsoon (JuneSeptember) and the northeast monsoon
(OctoberDecember), with the highest rainfall in October (Devidas and Puyravaud, 1995). Mean minimum and
maximum daily temperatures are 15 and 35 C respectively. There is a prolonged dry season from January to

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).

Figure 1. Research watersheds: D1Hediyala; D2Muntipur; P3Hebhalla; P4Soreda.

Copyright 2008 John Wiley & Sons, Ltd.

Ecohydrol. 1, 149160 (2008)

DOI: 10.1002/eco

ECOSYSTEM IMPACTS OF FOREST DISTURBANCE

Unlike many other protected areas in the country

there are no human settlements within the boundaries of BNP (Madhusudan, 2005). All anthropogenic
biomass pressuresprimarily livestock-grazing and fuelwood removalon the park originate from the villages
along the reserves northern flank. Over 100 000 cattle
from nearly 180 villages graze in the buffer zone of
Bandipurs northern fringes (Lal et al., 1994). The current
density of livestock is estimated at 236 animals per square
kilometre (Madhusudan, 2005). Livestock and wild herbivores compete for grazing resources in BNP; recovery of wild herbivores following livestock decline has
been documented by Madhusudan (2004). Cattle grazing
and fuelwood collection occurs within protected forest
areas, while small-scale farming is practised outside park
boundaries. Until recently, this use of the forest could be
qualified as subsistence use, with local village communities as consumers. However, in a study on the impact of
grazing on vegetation in BNP, Madhusudan (2005) documented the intensification of grazing practices since the
1990s, linked to the export of cattle dung as manure to
a neighbouring coffee growing district. This demand for
dung from a non-local source, in turn driven by global
coffee markets, has converted cattle dung from a local
subsistence agriculture resource to a commercial export
to neighbouring coffee estates. As a result, the pressure
on the northern boundaries of the park has substantially
increased in the last decade.
MATERIALS AND METHODS
Field sampling
Sample locations were nested within four watersheds in
BNP that had been previously identified for a study on
the impacts of forest degradation on rainfall/stream-flow
response (Krishnaswamy et al., 2006). A 1 km 1 km
grid was laid over the entire park, and five grids were
randomly selected within each of the four watersheds.
Soil and vegetation sampling were conducted at ten locations within each grid for a total of 200 sample locations
(sites), between November 2005 and March 2006. Using
a topographical sheet [Survey of India (SOI); 1 : 50 000
scale] of the area, and a GPS unit, a four-wheel-drive
jeep was used to drive to each selected grid on existing trails. Each site was selected at random distances on
either side of the trails that intersected any part of the
selected grid. Among other wildlife, elephants or signs of
elephants were encountered at 150 of the 200 locations,
and a tiger was spotted on two occasions. Considerations
of safety prevented us from venturing more than 500 m
away from the access trails within the forest. Figure 1
shows the Digital Elevation Model (DEM) of the research
watersheds, with the selected grids and sites overlaid.
The DEM (90-m resolution) source for this dataset was
the global land cover facility (http://www.landcover.org).
Research watersheds spanned the elevation and slope
ranges typical of the entire park, and within each watershed selected grids spanned low and high elevations
Copyright 2008 John Wiley & Sons, Ltd.

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
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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 ;

where pi is the proportional abundance of species i.

Since only counts, but not the individual DBH of individuals <10 cm DBH was measured, we used frequency
of occurrence as a measure of the dominant species.
Effects of disturbance on diversity and composition
were analysed in two steps. Sites were first clustered
using K-means clustering using the FDI indicator variables, into two groups (degraded and protected).
K-means clustering is a non-hierarchical method that produces a single partition optimizing within group homogeneity (Legendre and Legendre, 1998). Differences in
diversity between the two groups were analysed using
two sample, unequal, variance t-tests. Community composition differences were assessed by calculating the relative frequencies of the most dominant species within
each cluster group, supported by ordination.
Ordination. Partial canonical correspondence analysis,
(pCCA), (Ter Braak, 1986; Palmer, 1993) was used to
further elucidate the relationship of vegetation sites and
species composition with disturbance (FDI) and soil gradients. Additional variables were projected post facto
onto the ordination space to elucidate soilvegetation
linkages with disturbance. CCA is a method of direct gradient analysis that allows a direct comparison between
community and environmental data matrices (Legendre
and Legendre, 1998). The ordination plot was constrained
by FDI and soil CEC after conditioning out possible
effects of elevation. Among the soil data gathered for
this research, CEC was chosen in this analysis as it is
a key property describing nutrient availability for plant
growth, and was shown to be negatively influenced by
disturbance [Mehta et al. (2008, this issue)]. Significance
of the CCA model, each CCA axis and constraint terms
(FDI and CEC) were assessed separately using permutation tests. Three variablestree heights, soil clay and
grass coverwere projected post facto onto the ordination space to assess their possible associations with
community data. The pCCA was performed on the mean
values of community and environmental data, i.e. based
on grid-means of all species and environmental variables.
Additionally, presenceabsence transformation was performed, and all species that had a mean occurrence of
<5 were removed in order to better interpret the major
structure of the dataset.
Copyright 2008 John Wiley & Sons, Ltd.

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.
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ECOSYSTEM IMPACTS OF FOREST DISTURBANCE

Table I. Species with frequency greater than 10%.

ID
sp1
sp3
sp2
sp5
t1
t36
t3
t4
t2
t6
t23
t8

Species

Frequency (%)

Understory (non-tree) species

Chromolaena odorata
135 (675)
Canthium parviflorum
87 (435)
Lantana camara
72 (360)
Gymnosporia emarginata
27 (135)
Tree species
Anogeissus latifolia
147 (735)
Emblica officinalis
77 (385)
Tectona grandis
45 (225)
Terminalia crenulata
37 (185)
Terminalia alata
26 (130)
Acacia chundra
26 (130)
Careya arborea
24 (120)
Dalbergia latifolia
22 (110)

Table II. Summary of plot characteristics (n D 200 plots).

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

Tree height (m)

Understory height (m)
Canopy cover
CEC (cmol/kg)a
% Soil organic carbona
% Claya

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

Figure 2. Species richness and total density. Dotted line is a lowess

smooth.
Copyright 2008 John Wiley & Sons, Ltd.

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

soil properties used in Figure 7 are included.

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.
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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).

Table III. Size class distribution of dominant tree species.

DBH (cm)

<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

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ECOSYSTEM IMPACTS OF FOREST DISTURBANCE

(a)

(b)

Figure 5. DBH classes for three dominant tree species. (a) Northern watersheds; (b) southern watersheds.

Disturbance and diversity

Species diversity H0 across the 200 sites ranged from 0
to 2836 with a mean of 175 and was approximately
normally distributed overall. These estimates are comparable to H0 estimated in previous studies in other parts of
BNP (Elouard and Krishnan, 1999; Madhusudan, 2000).
Significant differences emerged between degraded and
protected watersheds. Figure 6 shows the boxplots of
H0 by degradation class. Using a two-sample, unequal,

variance t-test, diversity H0 was significantly higher in

the degraded watersheds than in the protected watersheds
(t D 453, p < 0001, n D 100 plots each). In Figure 6(a)
all 47 species were included. Further insight is obtained
by computing H0 separately for tree and non-tree (understory) species. Figure 6(b) and (c) shows boxplots of H0
for tree and understory species respectively. There was no
significant difference in diversity of tree species between
the two watersheds. However, in the case of understory

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.

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V. K. MEHTA ET AL.

species, species diversity was significantly higher in the

degraded watersheds (class 1) than the protected watersheds (w D 6698, p < 0001, Wilcoxon rank-sum test).
The greater overall diversity in degraded forest is a consequence of greater understory diversity compared to the
protected forest.
Disturbance and vegetation composition
Table IV presents a two-way table of relative frequencies
of species present in degraded and protected watersheds.
Only those species that had a frequency of at least 10%
in either watershed class are included. These relative
frequencies are combined with the pCCA results given
below to elucidate composition differences.
CCA results. Figure 7 shows the pCCA ordination plot.
Since mean data (means calculated from 10 samples per
grid) and species with frequency >5 were used, there are
20 sites (corresponding to the 20 sample grids) and 21
species in the ordination plot. The 20 sites are colourcoded by watershed as in previous figures. Directions of
the biplot vectors (FDI and CEC) allow interpretation
of the CCA axes. CCA1 is interpreted as a degradation

Table IV. Relative frequencies of species within degradation

class.
IDa

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

spunderstory species; ttree species.

1degraded; 2protected.

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.

Ecohydrol. 1, 149160 (2008)

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ECOSYSTEM IMPACTS OF FOREST DISTURBANCE

gradient (increasing in the negative CCA1 direction).

CCA2 is correlated with soil CEC. Table V reports the
model summary and significance tests based on 1000
permutations. The constrained axes (CCA and CCA2)
together account for 23% of the total variability (inertia or
mean square contingency coefficient) after conditioning
out the effect of elevation. The overall model tested
using permutation tests for the eigenvalues is significant
(Table V part b). On the basis of permutations tests
for significance of individual axes and terms, CCA1 is
highly significant. However CCA2 (correlated to CEC)
is not significant at the 10% level. This means that after
conditioning out the elevation effect, CEC is a weak
gradient compared to disturbance (FDI) in explaining the
major variation in the community data. Nevertheless, it
is retained in the ordination plot because CEC was found
to be highly significant when all 200 plots were used in
the pCCA (not shown). Only the grid-mean data results
are reported in Table V.
The ordination plot confirms that degradation level
(FDI or CCA1) is a very good discriminator between
sites that are in the northern degraded watersheds and
sites in the southern protected watersheds. All grids in
the degraded watersheds (black and red open symbols)
are located in the left half of Figure 7, while all grids
in the protected watersheds are located in the right
half. Projection of sites (species), onto any vector in
the plot, allows a comparison between sites (species)
along that vector. In the case of the degraded watersheds,
projection of sites onto CEC, clay and tree height vectors
allows further discrimination within the degraded sites.
Sites in watershed D1 have less soil CEC, average
tree heights and soil clay than those in watershed D2;
Table V. pCCA results.
pCCA model results
a. Model summary

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

hence D1 can be rated as more degraded than D2.

This internal discrimination does not appear between
the two protected watersheds P3 and P4. The directions
of (post facto fitted) clay, tree height and SOC vectors
show them to be closely related to each other and
to CEC. Projections of the sites onto these vectors
confirm earlier findings that the protected sites also
exhibit greater average tree heights (this paper), soil
clay contents and organic carbon (Mehta et al., 2008,
this issue) compared to the degraded sites. Reduced
clay content in disturbed sites is attributed to increased
vulnerability of soils to erosion upon reduction of canopy
cover with increasing disturbance (Figure 3(c)), with
consequent selective removal of finer clay particles.
CCA1 being a degradation gradient, species composition differences emerge on either side of the CCA1 origin
with degraded sites to the left (CCA1 <0) more populated by species that are also to the left. Hence, as also
shown in Table IV earlier, the degraded sites are more
commonly populated by tree species t17 and t6 than the
protected sites, with tree species t3 and t8 more prevalent in protected sites. As for shrub species, sp5 and sp7
are more prevalent in degraded sites compared to protected sites, whereas sp11 is more prevalent in protected
sites. Those species that do not show discrimination along
the degradation gradient appear closest to the origin of
the pCCA plot. Projecting the levels of the grass factor
variable, it is evident that the most degraded sites have
short Themeda spp. cover (G1); moderately degraded
sites have tall Themeda spp. cover (G2); moderately protected sites have short Cymbopogon spp. cover (G3) and
well protected sites have tall Cymbopogon spp. cover
(G4).
On the basis of the pCCA results and Table IV,
the following community characteristics and differences
between protected and watersheds are summarized. The
dominant trees in degraded watersheds are A. latifolia,
E. officinalis, Acacia chundra and Terminalia spp. The
understory is dominated by the weeds C. odorata, L.
camara and native Canthium parviflorum; which are
almost equally dominant.
The dominant trees in protected watersheds are A. latifolia, E. officinalis, T. grandis and Terminalia spp. The
understory is dominated by C. odorata, C. parviflorum
and L. camara, with C. odorata more widespread than
the others. The contrasts between degraded and protected
forests are as follows. In tree species, (i) degraded
forests had less widespread teak, (ii) degraded forests
had several small woody tree species in moderate
frequencyChloroxylon swietenia, Careya arborea,
Erythroxylon monogynum, Ziziphus xylopyrus
compared to protected forests in which these species
were almost absent, (iii) protected forests had a moderate frequency of large woody treesDalbergia latifolia and Schrebera swietenioides compared to degraded
forest which had practically none. In understory species,
(i) Gymnosporia emarginata and Ziziphus oenoplia were
moderately widespread in degraded forests, compared to
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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.

DISCUSSION AND CONCLUSIONS

In BNP, disturbance quantified as a combined impact
of grazing, fuelwood/fodder extraction and fire are significantly related to forest structure, composition and
diversity. Tree heights, densities and diameters were
less in more degraded sites. There was no significant
relationship in understory heights and shrub densities
with disturbance. A reduced recruitment of teak and an
increased mortality of A. latifolia in bigger size classes
are evident in degraded watersheds. Species composition and diversity differences exist. Degraded forests
had greater diversity, attributed to greater diversity in
shrub species. Species assemblages in degraded forests
showed a greater presence and density of small woody
tree species, and greater numbers of understory species.
The increasing dominance and diversity of small woody
species, along with shrubaceous species that bear thorny
structures (e.g. G. emarginata, Z. oenoplia) has been
reported in other dry tropical forests in India (Pandey
and Singh, 1991; Shankar et al., 1998b; Kumar and Shahabuddin, 2005; Madhusudan, 2005). Invasives, C. odorata and L. camara, are widespread and abundant throughout the forest, indicating that protected and degraded
classifications are relativeprotected southern forested
areas are also disturbed, but less so than northern areas.
A. latifolia (family Combretaceae) is widespread throughout the forest, and teak individuals as well as Terminalia
spp. are also present in degraded areas (albeit not abundantly). This supports the published vegetation maps we
used as baseline information as well as local and forest
department accounts that the entire study area was once
covered with dry deciduous woodland to savanna woodland forests of the A. latifoliaT. grandisTerminalia
spp. type.
In dry tropical forests of the Western Ghats, intense
overgrazing and lopping has been theorized to convert
the tree savanna into clump thickets, with the selection of thorny and unpalatable species (Pascal, 1988
p291293). This study provides field evidence in support of this theory in BNP. The removal of dominant tree
species for timber/fuelwood opens the canopy allowing
more species to compete. Following continual biomass
extraction and further aridification of the soils and microclimate, unpalatable, small, woody and thorny species
become more dominant.
Feedbacks and implications for services
Field evidence shows that forest degradation in BNP has
substantial impact on the entire forest ecosystem, through
its impact on forest structure, diversity and composition
(this paper); and soil physical, nutrient and hydraulic
characteristics Mehta et al. (2008, this issue). Nutrient
Copyright 2008 John Wiley & Sons, Ltd.

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)
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ECOSYSTEM IMPACTS OF FOREST DISTURBANCE

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

This research was financially supported by grants from

the International Foundation for Science (IFS, Sweden);
Ford Foundation with UNESCO; Cornell Universitys
Center for the Environment and Bradfield Awards. We
thank the Karnataka Forest Department for research
permissions, and the Ashoka Trust for Research in
Ecology and the Environment (ATREE) Bangalore, for
research and logistic support. International travel support
was provided by Cornell Universitys Einaudi Center
for International Studies and the South Asia Program.
We are grateful for assistance in field logistics from
N. Samba Kumar and the Centre for Wildlife Studies,
Bangalore. Special thanks to local field assistant, Mr
Dasappa, and to Kiran M.C., Kiran Yadav, Rakesh K.N.
and Dr Pradeep Joshi for research assistance. We are
grateful to Dr R. Ganesan for help with taxonomy and
local uses of vegetation inventoried. We express thanks to
Dr Hugh Gauch Jr for ordination advice and constructive
comment on the manuscript.
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