Remote Sensing of Environment 113 (2009) 857867

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Remote Sensing of Environment

j o u r n a l h o m e p a g e : w w w. e l s ev i e r. c o m / l o c a t e / r s e

Quantifying and mapping biodiversity and ecosystem services: Utility of a

multi-season NDVI based Mahalanobis distance surrogate
Jagdish Krishnaswamy a,b,, Kamaljit S. Bawa a,c, K.N. Ganeshaiah a,d, M.C. Kiran a
a

Ashoka Trust for Research in Ecology and the Environment, No 659, 5th A Main Road, Hebbal, Bangalore, 560024, India
Stockholm Resilience Centre, Stockholm University, SE-106 91 Stockholm, Sweden
Department of Biology, University of Massachusetts, Boston, MA 02125, USA
d
Department of Genetics and Plant Breeding, University of Agricultural Sciences, GKVK, Bangalore, 560065, India
b
c

a r t i c l e

i n f o

Article history:
Received 7 April 2008
Received in revised form 24 December 2008
Accepted 26 December 2008
Keywords:
Mapping biodiversity and ecosystem services
Biodiversity surrogate
Tropical forest
Remote sensing

a b s t r a c t
There is an urgent need for techniques to rapidly and periodically measure biodiversity and ecosystem
services over large landscapes. Conventional vegetation classication and mapping approaches are based on
discrete arbitrary classes which do not capture gradual changes in forest type (and corresponding biodiversity and ecosystem services values) from site to site. We developed a simple multi-date NDVI based
Mahalanobis distance measure (called eco-climatic distance) that quanties forest type variability across a
moisture gradient for complex tropical forested landscapes on a single ecologically interpretable, continuous
scale. This Mahalanobis distance, unlike other distance measures takes into account the variability in the
reference class and shared information amongst bands as it is based on the covariance matrix, and therefore
is most useful to summarize ecological distance of a pixel to a reference class in multi-band remotely sensed
space In this study we successfully apply this measure as a surrogate for tree biodiversity and ecosystem
services at two nested scales for the Western Ghats Bio-diversity hotspot. Data from over 500 tree-plots and
forest type maps was used to test the ability of this remotely sensed distance to be a surrogate for abundance
based tree-species compositional turn-over and as a continuous measure of forest type and ecosystem
services. Our results suggest a strong but scale dependant relationship between the remotely-sensed distance measure and oristic distance between plots. The multi-date NDVI distance measure emerges as very
good quantitative surrogate for forest type and is a useful complement to existing forest classication
systems. This surrogate quanties forest type variability on a single, continuous quantitative scale and has
important applications in conservation planning and mapping and monitoring of hydrologic and carbon
storage and sequestration services.
2008 Elsevier Inc. All rights reserved.

1. Introduction
Continuing biodiversity and ecosystem services loss urgently
requires techniques to rapidly assess and monitor changes in biodiversity and ecosystem services (Balvanera et al., 2001; Menon & Bawa,1997;
Margules & Pressey, 2000; Ramesh et al., 1997; Stork & Samways, 1995).
Assessment of changes in biodiversity at a large spatial scale often
depends upon the use of surrogate information to detect trends in
general patterns because direct information on changes in the distribution of individual species or species assemblages is hard to collect
and is time consuming (Ferrier, 2002; Faithe et al., 2004). Remotely
sensed imagery combined with limited sampling on the ground has the
potential to ascertain spatial and temporal variation in biodiversity over
large spatial scales (Couteron et al., 2005; Nagendra, 2001; Rosenzweig
& Abramsky, 1993; Tuomisto et al., 2003). However, the potential of
Corresponding author. Ashoka Trust for Research in Ecology and the Environment,
No 659, 5th A Main Road, Hebbal, Bangalore, 560024, India.
E-mail address: jagdish.krishnaswamy@gmail.com (J. Krishnaswamy).
0034-4257/$ see front matter 2008 Elsevier Inc. All rights reserved.
doi:10.1016/j.rse.2008.12.011

remotely sensed imagery in mapping various components of biodiversity has not yet been fully utilized or demonstrated effectively in a
rigorous manner (Couteron et al., 2005; Lillesaeter, 1982; Nagendra &
Gadgil, 1999; Nagendra, 2001; Sanchez-Azofeifa & Rivard, 2001) with a
few notable exceptions usually in a single type of forest and at a single
spatial scale (e.g. Rocchini, 2007; Tuomisto et al., 2003).
Existing vegetation and habitat classication and mapping
approaches have several severe limitations. Forests are not mosaics of
discrete categories of vegetation types, but are a continuously changing
terrain of biological diversity and corresponding ecosystem services. In
addition, seasonal tropical forests have varying degrees of deciduousness and tree-density which inuences a range of ecological and hydroecological processes. Every site or pixel should potentially have a distinct
value on an easily interpretable ecological or ecosystem service distance
scale. Earlier methods of mapping or classication did not reect this.
However, fuzzy, sub-pixel unmixing, and other soft classication methods have the capability to reect the variability (Brown, 1997; Foody,
1996a,b; Lobell & Asner, 2004) and increasingly the utility of a continuous measure based on remote sensing is becoming apparent (e.g.

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J. Krishnaswamy et al. / Remote Sensing of Environment 113 (2009) 857867

Asner et al., 2005; Carlson et al., 2007; Puzzolo et al., 2003; Rocchini,
2007; Tuomisto et al., 2003). There is thus an increasing need and
interest in linking biodiversity across a landscape to a well-dened and
interpretable remotely sensed continuous distance measure that is related to eco-climatic variability, and takes into account the variability of
the reference class. In this study, the rst set of criteria is met by adopting
the Mahalanobis distance measure and the second criteria by using
multi-season Normalized Difference Vegetation Index (NDVI) data for
dening the eco-climatic distance.
The Mahalanobis distance is a distance in multi-dimensional space
from a point to the centroid of a multi-dimensional variable sample. It
is superior to other distance measures since it takes into account the
variability of the reference class and the covariance matrix accounts for
shared information in correlated variables, both of which are common
feature of multi-band remotely sensed data. Formally, the Mahalanobis

distance from a group of values with mean and covariance matrix for
a multivariate vector to a point or pixel is dened as:
q
DM x = x T 1 x
"
#
$
%
= 1 ; 2 ; 3 ; 4 ; N p ; x = x1 ; x2 ; x3 ; N ; xp

where x is a vector of values in p different bands or dimensions for a

particular pixel and is the mean vector of values for the same bands
or dimensions for the reference class pixels, and is the covariance
matrix for these p bands for all the reference class pixels.
NDVI derived from remotely sensed data is a measure of green
biomass in a habitat and is an alternative. NDVI ranges from 1 to +1.
All surfaces yield an NDVI value, with values b0 being non-vegetated,
0 being equated with water, and values N0 representing vegetation.

Fig. 1. The eco-climatic distance map and location of ground measurement plots for two different landscape scales in the Western Ghats hotspot in India. The scale in reverse also spans
above ground carbon storage in biomass from approximately 50 Mg C ha 1 for very high distance values to over 250 Mg C ha1 at sites with very low distance values, in evergreen forests.

J. Krishnaswamy et al. / Remote Sensing of Environment 113 (2009) 857867

NDVI has been shown to be linearly correlated with fPAR (fraction of

PAR intercepted by plants) and is often transformed accordingly. In a
continuously vegetation forested environment, maxmin NDVI may
well be correlated with maxmin fPAR (Berry & Roderick, 2002). Green
biomass is a measure of productivity, and productivity is known to be
correlated with tree species richness (Bailey et al., 2004; Haggar &
Ewel, 1997; Rosenzweig & Abramsky, 1993; Waide et al., 2000) and a
positive relationship exists between NDVI and (within a habitat)
diversity of tree species in tropical forests in the Western Ghats of
southwest India (Bawa et al., 2002). In the Western Ghats and adjacent
areas the diversity of forest ecosystems, habitats and tree communities
are related to strong environmental gradients of moisture availability
and length of dry season (Barboni et al., 2003; Gunnel, 1997; Pascal,
1982; Prasad et al., 2005). We reasoned that NDVI values of various
forest types across a moisture gradient should show different levels of
seasonal variation in NDVI. This then enabled us to use multi-date
NDVI (23.5 m spatial resolution) within a Classication and Regression
Tree (CART) framework to classify vegetation of a 540 km2 area into
various types or categories (Krishnaswamy et al., 2004).
We then asked whether we can dene a single quantitative measure
using multi-season NDVI to predict patterns of biodiversity posed the
following specic questions:
1. Is there a way of using RS data to arrange forest types along the
moisture gradient, and dene a continuous measure of forest type
that is related to changes in biodiversity and ecosystem services?
2. Is such a map related to spatial turn-over in biodiversity composition or beta-diversity?
3. Is the relationship between the ecoclimatic map derived from RS
data, and biodiversity compositional turn-over, dependent on spatial
scale (grain and extent)?
In this paper, we explore the utility of this Mahalanobis distance based
approach for mapping tree species compositional turnover between
forest types and sites at ne scales and at multiple landscape scales. This
was motivated by the successful use of this surrogate for some
biodiversity components (especially endemic trees and amphibians ) at
very coarse spatial scales (1:25,000) in a major conservation planning
exercise for the entire Western Ghats biodiversity hotspot (Das et al.,
2006). Potentially this distance measure could complement existing
forest type maps with discrete classes and could be used for a variety of
applications in mapping biodiversity and ecosystem services.
We now rigorously test its relationship with actual ground data on
tree species composition from two different and independent sources
(Bawa et al., 2002; Shivraj et al., 2000) at two different spatial scales
because it is necessary to analyze the effect of scale on patterns of beta
diversity or on techniques to predict beta diversity (Fraser, 1998; Koleff
& Gaston, 2002; Lennon et al., 2001). We also explore its utility in
quantifying and mapping ecosystem services.
2. Methods
2.1. Study area
This study was conducted in the Western Ghats Biodiversity hotspot
(Myers et al., 2000; Das et al., 2006). The study used (1) tree plot data,
and remotely sensed data at two landscape scales from a very diverse
part of the hotspot (Fig. 1) as well as comparisons of remotely sensed
distance images with existing forest type maps for two protected areas
in the Western Ghats.
2.1.1. Nested landscape component
The tree-plots based component was done at two landscape scales
(Fig. 1). At each scale we used independently collected and archived
ground data (Bawa et al., 2002; Shivraj et al., 2000) and processed
corresponding, remotely sensed, data of the appropriate spatial resolution. Each scale is dened by the extent of the study area and spatial

859

resolution of the tree-plot data (plot size) and pixel size of the remotely
sensed data. Each scale is also dened by distinct forest types that are
spread across a strong gradient of moisture availability but share common species (or abundances of species). The nested ne scale component
(landscape scale 1) was located in the Biligiri Rangaswamy Temple (BRT)
Wildlife Sanctuary, in Karnataka, described in detail later in this section.
At the broad landscape scale (landscape scale 2) our study used
independent ground and remotely sensed data for a 25,000 km2 region
(Latitude 11 32 48.31 N to 13 2 21.83 N and Longitude 75 7 38.98
E to 77 48 8.41 E) spread over the Western Ghats hotspot. Various
forest types such as evergreen, moist deciduous, dry deciduous, scrub
forests and grass lands occur in this area which span a rainfall gradient
from over 2000 mm to 500 mm yr 1. The natural forests in this area are
spread over 15,223 km2 of which 3423 km2 is evergreen forests. The
elevation ranges from 70 m to 2205 m. 3828 km2 of area is under
protection (Rodgers et al., 2002). Approximately six million people
inhabit this landscape (Census, 2001).
The comparison of the continuous remotely sensed measure to
existing, independently forest type classication was done for two
important protected areas in the Western Ghats bio-diversity hotspot:
BRT Wildlife sanctuary, and Bhadra Wildlife Sanctuary and Tiger Reserve (Fig. 2). These two protected areas are described briey below:
Biligiri Rangaswamy Temple (BRT) Wildlife Sanctuary, (latitude 11 43
to 12 08 N and longitude 77 to 77 16 E), approximately 540 km2 in size
and elevation from 600 to 1800 m. The major forest types as per the
Champion and Seth (1968) classication in the sanctuary are evergreen,
grassland, moist deciduous, dry deciduous and scrub forests. This corresponds approximately to a rainfall gradient from 1800 to 500 mm yr 1. The
reserve has an indigenous Soliga tribal community with a long history
of past shifting cultivation as well as high levels of current dependence
on forest product extraction.
Bhadra Wildlife Sanctuary was expanded and made into the present wildlife sanctuary covering an area of 492 km2 in 1974 and is now
a tiger reserve. Bhadra Tiger Reserve is located in Karnataka, India
(Longitude 75 29 E to 75 47 E and Latitude 13 22 to 13 47 N) with
the altitude ranging from 670 m to 1870 m. Bhadra receives an annual
rainfall varying spatially from less than 1000 to 2500 mm. The
vegetation in Bhadra Reserve is primarily moist deciduous forest with
bamboo and patches of dry deciduous occurring on the northern
fringes. At the higher elevation in Bababudangiri, Coffee plantations
dominate, but there are patches of a third type of forest known as the
Shola wet evergreen forests and some grassland patches occur.
2.2. Sources of data
2.2.1. Tree-plot data
The ground data used was independently obtained for each of the
two landscape scales. For landscape scale 1, data from an earlier study
(Murali et al., 1998) was used. For this study, a network of plots was
established in and adjacent to the BRT sanctuary. The study area was
divided into 2 2 km grid cells and at the centre of each cell, an
80 5 m plot was laid (Bawa et al., 2002; Murali et al., 1998). All trees
greater than 10 cm diameter at breast height (DBH) were enumerated
for each cell. The number of cells or plots that fell within the sanctuary
was 125. The total number of tree species was 127 across eight forest
types, (Ramesh & Menon, 1997).
For landscape scale 2, we used ground data from the network of 0.1 ha
plots established by the Forest Survey of India (Shivraj et al., 2000). These
plots are spread over an area of 15,000 km2 within landscape scale 2. All
tree species greater than 10 cm DBH were recorded. The Forest Maps of
South India (Pascal et al., 1982, 1992) were used to classify the vegetation
of landscape scale 2 into 18 types. At each scale the vegetation type to
which each plot belonged was determined by its location on these forest
maps. The forest types are basically a function of bio-climate and span a
strong gradient of moisture availability mainly related to rainfall and

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Fig. 2. Comparison of eco-climatic distance maps with conventionally classied forest bio-climatic maps for two protected areas (BRT and Bhadra) with diverse forest types and
rainfall regimes.

length of dry season (Barboni et al., 2003; Champion & Seth,1968; Pascal,
1982; Prasad et al., 2005). Only plots for which recorded locations could
be veried using meta-data such as topographic sheet number and site
description and which fell within a distinct forest vegetation type were
nally used. A total of 464 plots with 320 species were included in the
study.
In summary, the density of plots is about 1 per 3.94 km2 for the ner
scale and 1 per 32.33 km2 for the coarser, broad scale. The ne scale had

regularly spaced plots, whereas the broad scale had irregularly spaced
plots (Fig. 1).
2.2.2. Remotely sensed data
The Indian Remote Sensing (IRS) satellite's LISS III images which had
23.5 m spatial resolution acquired on 2nd February 1998, 16th November
1998 and 15th April 1999 were used for the landscape scale 1. For
landscape scale 2, IRS WiFS images (188 m spatial resolution) acquired on

J. Krishnaswamy et al. / Remote Sensing of Environment 113 (2009) 857867

14th December 2000, 15th January 2001 and 23rd March 2003 were
used. These dates approximately span the period with maximum seasonal variation in eco-physiological and eco-climatic conditions. Due to
cloud cover and large area coverage, it was difcult to get enough cloud
free images for same year.
2.2.3. Climate data
The annual potential evapotranspiration (PET) at 30 min resolution
was derived from the Global data set (Ahn & Tateishi, 1994). Rainfall
data for every 5 km resolution pixel was obtained from a kriged layer
of rainfall generated from secondary rainfall data from stations in and
around the Western Ghats available from the National Institute of
Hydrology, Belgaum and Water Resources Development Organization,
Bangalore.
2.3. Data processing and analyses
To summarize, the ne scale study covered an area of about 500 km2,
plot size of 0.04 ha and a pixel resolution of 23.5 m whereas the
corresponding values are 15,000 km2, 0.1 ha and 188 m for the broader
scale study.
2.3.1. Eco-climatic distance
At each spatial scale, a new RS based distance measure called ecoclimatic distance was used to compute the distance in NDVI space for
each pixel, tree-plot or group of tree plots pooled by forest type/moisture
regime from the reference forest type class. We dened a Mahalanobis
distance to the reference class in multi-band space using the four NDVI
derived layers.
In this study the reference class vector and the pixel vector x
consists of four values (p = 4) based on three multi-date NDVI:
1. Mean of NDVI from three dates (1.end of wet-season, 2. beginning
of dry season, 3. late dry season)
2. Coefcient of variation (CV) of NDVI from the three dates
3. NDVI difference 12
4. NDVI difference 23
These four layers derived from multi-date NDVI span the period of
maximum phenological and eco-climatic variation. The rst band 1
above is a surrogate for tree density and canopy foliage biomass, and 2
above is an excellent surrogate for degree of deciduousness as related to
length of dry season, plant available moisture availability and evapotranspiration (Barboni et al., 2003; Krishnaswamy et al., 2004; Prasad
et al., 2005) and 3 and 4 are related to intra-seasonal phenological and
plant available moisture trends (Krishnaswamy et al., 2004).
Rainfall or moisture is the principal environmental inuence on
forest type in the Western Ghats (Barboni et al., 2003; Gunnel, 1997;
Pascal, 1982; Prasad et al., 2005) and we dened the reference forest
type class at one end of the rainfall/moisture gradient: evergreen forest
type. The evergreen forest type is at one end of the principal environmental gradient, is relatively stable with respect to NDVI over the dry
season, has the highest evapotranspiration, and is also the most diverse
in tree-species, and so it should be a good candidate for a reference
forest/eco-service type. The metric is based on the energy absorbed/
reected by photosynthesising foliage in the forest canopies. Plant
photosynthesis is constrained by seasonal uxes in water availability. So,
the connection with climate is that rainfall is the dominant input to the
soil water balance and the availability of water to plants. So, the metric is
more of a plant-ecophysiology distance or a plant phenology metric
as the link with climate is inferred. In addition, as this metric is related to
leaf area index, canopy cover, tree density and degree of deciduousness,
it is also a measure of an evapotranspiration and soil erosion protection
gradient.
Several pixels from this class were identied and chosen for each
landscape scale. The values of the four NDVI layers or variables for
these reference pixels dene the mean vector and covariance matrix

861

for the reference class. An S-PLUS (S-Plus, 2000) macro was written to
compute the distance from each pixel to the reference class in multi
dimensional space.
2.3.2. Forest type classication comparison
Digitized Forest type maps for BRT and Bhadra were processed
from the French Institute Bio-Climatic Map sheets. For each forest type
identied in the map, the corresponding eco-climatic pixel data were
used to generate a Box and Whiskers plot for each site.
2.3.3. Analyses of tree-plot data pooled and averaged by bio-climatic/
moisture regime/forest type
The remotely sensed eco-climatic distance was averaged by bioclimatic or moisture regime/forest type using the tree-plot locations. In
the case of the landscape scale 1 (ne), the mean value for a 7 7 window
placed around each plot was used whereas in the landscape scale 2
(broad), the exact value of the eco-climatic distance measure corresponding to the WiFS pixel (0.4 ha) within which the plot (0.1 ha) was located
was used directly. The window was used for the ner scale study rather
than the original pixel resolution to ensure that the smaller plot would be
located within a particular pixel window given the geo-referencing error.
We nally compared the map of eco-climatic distance with the
most detailed and widely used existing maps of vegetation and habitat
types produced by the French Institute at Pondicherry, India at both
landscape scales.
2.3.4. Tree species compositional distance
At each landscape scale, data from the plots within each distinct
forest type were pooled.
A macro was written in S-Plus software that computed the mean
number of trees for each species for a given vegetation type. Only those
vegetation types that had at least ve plots corresponding to it were
selected. There were four such vegetation types for landscape scale 1, 15
types respectively for landscape scale 2. We used the BrayCurtis or
Czekanowski's coefcient or Srenson (abundance) (Bray & Curtis, 1957;
EstimateS, 2000). We computed compositional distances to the reference class in EstimateS 6.0 software (EstimateS, 2000). The reference
plots were selected from the dense evergreen forest type. There were 23
reference plots for landscape scale 1 and 50 plots for landscape scale 2.
Each distance measure was computed as 1-similarity.
2.3.5. Regressions
The log transformed eco-climatic distance of each vegetation type to
the reference evergreen class was regressed against the tree species
compositional distance for those classes. The independent variable is the
log transformed eco-climatic distance and the dependant variable was
the tree species compositional distance of a vegetation type to the reference evergreen class. Regressions were done for each landscape scale. We
tested the eco-climatic distance with all four different measures of
dissimilarity distances as described above.
Image processing software Idrisi Kilimanjaro (Idrisi, 2003) and
Erdas 8.3.1 were used to process the satellite images.
2.3.6. Tree-plot level analyses
The analyses described above were done by averaging the tree-plot
data within a bio climatic/rainfall regime type. Another set of analyses
(see below) was done using the unpooled tree-plot data at both
landscape scales.
2.3.7. Quantilequantile plots
The pair-wise BrayCurtis tree-species compositional or oristic
distances were plotted against the corresponding pair-wise log (ecoclimatic) distances for both landscape scales. As these pair-wise distances are not independent of each other, and the number of data points
is very large, conventional statistical methods (e.g. regression, linear or
non-linear with specic parametric distributional assumptions for the

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errors) are not possible. We used the graphical, non-parametric

quantilequantile plots to summarize the ability of the remotely sensed
surrogate to predict oristic compositional distances between the treeplots. Quantilequantile are one the most powerful techniques that are
widely used to compare the shapes of distributions of two variables with
respect to each other (Cleveland, 1994; Kuczmarski & Rosenbaum, 1999;
Marden, 1998; Wilk & Gnanadesikan, 1968). Quantile plots are useful for
large sized, noisy data sets for two reasons: (1) simple patterns in
quantile plots have simple interpretations in terms of distributional
shape; and (2) the plots are often more robust and more revealing than
numerical measures of distributional shape. We computed quantiles in
the range 01 at intervals of 0.05 for the oristic BrayCurtis distances
and the remotely sensed distances and plotted these against each other,
overlayed on the scatter-plot.
2.3.8. Mantel's analyses using pair-wise distance matrices
The simple Mantel test (Mantel, 1967; Mantel & Valand, 1970) was
used to test the hypothesis that the pair-wise oristic distances among
sites are linearly independent of the pair-wise remotely sensed
distances among the same sites and to quantify the linear correlation
between the two distance matrices. In our study, the simple Mantel
correlation addresses the basic ecological question, are sites or bioclimatic regimes that are similar in the remotely sensed eco-climatic
distance values also similar in their tree-species composition? A test of
signicance is evaluated via (1000 in this study) permutations of one
of the matrix because the elements of a distance matrix are not
independent (Manly 1997; Legendre & Fortin 1989). We tested in
particular whether distances in the RS measure between sites or sites
pooled by broad classes pre-dened across an environmental gradient
is a good surrogate for the oristic distance between them. Using the
environmental gradient level-averaged data, a dissimilarity matrix of
tree species composition between forest types was generated for each
landscape scale. The Mantel's tests were conducted using Ecodist-R
(see Acknowledgements).
2.3.9. Quantifying and mapping ecosystem services
The Zhang et al. (2001) model predicts evapotranspiration (ET) as a
function of precipitation (P), E0 (potential evapotranspiration) and a
plant-available moisture index, w.
0

1 + w EP0
B
C
ET = @
" #1 AP:
1 + w EP0 + EP0
NDVI is a very good surrogate for Leaf Area Index (LAI), which is
strongly related to available soil moisture at rooting depth (Donohue
et al., 2007). Variability of soil moisture from site to site as summarized
in the parameter w in the above model can therefore be calibrated with
NDVI or eco-climatic distance using ground information on vegetated
sites with extreme low and high values of w (0.5 to 2, Zhang et al.,
2001). Using this calibration, and spatially explicit data on rainfall and
potential evapotranspiration, we could predict and map green-water
(evapotranspiration) and blue-water (rainfall-evapotranspiration)
uxes. Subtracting predicted evapotranspiration from rainfall gives the
blue-water ux for every pixel.

Table 1
Mantel's correlation coefcients between remotely sensed eco-climatic distance and
tree-species compositional distance
Region and landscape scale
Western Ghats (Broad)
BRT (Fine)

Fig. 3. Tree-species compositional distance between tree-plots in relation to eco-climatic

distance between tree-plots at (a) landscape scale 1(500 km2) and (b) landscape scale 2
(15,000 km2). The quantilequantile plots are overlayed on the scatter-plots.

The eco-climatic distance measure we have dened is related to

tree-density, LAI and degree of deciduousness, so it is a direct measure
of carbon storage. Available data from published studies on above
ground biomass storage in Western Ghats forest types indicate that
the carbon storage potential captured by the range of the eco-climatic
distance metric (Fig. 1) is approximately 50 to over 250 Mg C ha 1
(Chave et al., 2008; Madugundu et al., 2008; Rai & Proctor, 1986).
3. Results

Mantel rm coefcients
Pooled and averaged

Un-pooled tree-plots

0.70 (n = 16)
0.63(n = 5)

0.10 (n = 464)
0.0085 (n = 125)

Signicant values (p b0.05) are marked and number of samples, n are indicated.

3.1. Eco-climatic distance and forest type classication

The eco-climatic for the entire landscape is shown in Fig. 1. Even a
simple visual comparison of the RS quantitative scale with the broad

J. Krishnaswamy et al. / Remote Sensing of Environment 113 (2009) 857867

863

Fig. 4. The scatter plot of the eco-climatic distance measure against the tree-species compositional distance to reference evergreen forest at two landscape scales. The eco climatic distance
for different vegetation types to the reference evergreen class is solely derived from remotely sensed data and shown on a log scale. The tree-species compositional distance to the reference
evergreen class is calculated from measurements from ground plots using the BrayCurtis/Sorenson abundance based index and is pooled by forest type (bio-climatic regime) and averaged.
Each data point represents a pooled average for a particular forest type. These plots indicate that oristic distance of different forest types to evergreen forest is correlated with remotely
sensed distance of that forest type to evergreen forest.

forest types derived from the Champion and Seth and the French Institute
classication system for two protected areas reveals a close correspondence (Fig. 2). The eco-climatic distance measure effectively captures the
continuum of bio-climatic variability across environmental gradients
including eco-tones. In addition, it captures variability across the
landscape in two ecosystem functions related to ecosystem services,
evapotranspiration and carbon storage and sequestration.

3.2. Relationship between remotely sensed distances and oristic distance

3.2.1. Tree-plots pooled by bio-climatic/forest type regime
Pair-wise tree-species compositional distance between predened bio-climatic regimes/forest types was highly correlated with
the corresponding remotely sensed distance measure at both landscape scales: (Table 1, ne scale 500 km2, rm = 0.63, p 0.05 and

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broad, 15,000 km2, rm =0.70, p b 0.05). These results indicates that:

(1) Tree species oristic differences between forest types over a
moisture gradient can be captured by the RS distances averaged by
bio-climatic/forest type. (2) Relative distance to evergreen reference
bio-climatic class is a good measure of oristic distance between
forest types (3) The remotely sensed distance measure can be a good
surrogate for use in forest type or bio-climatic mapping and can
express broad changes in tree-species composition and vegetation
type within a single quantitative, continuous scale.
3.2.2. Tree-plots (unpooled)
The scatter-plot of the pair-wise remotely sensed surrogate
dissimilarity with oristic dissimilarity shows a fairly typical pattern
representative of spectral distance decay in a tropical landscape with a
large amount of scatter and a large number of pairs of plots that are at
dissimilarity = 1. A few plots at the larger landscape scale with extremely small number of trees of the same species, and usually located
close to each other, had extremely small BrayCurtis distances with
respect to each other in the dissimilarity matrix. These were rounded
off to zero distances in the scatter-plot, and were retained in the main
plot. However these plots were dissimilar with respect to the remotely
sensed distance measure, because of the differences in the non-tree
ora, mainly the under-storey and invasive species shrubs.
The quantilequantile plots and comparisons between the two
landscape scales, are strongly supportive of a non-linear and scale dependant relationship between the remotely sensed distance measure
and the corresponding oristic distances between tree-plots (Fig. 3). The
plots are supportive of the apriori hypothesis that a remotely sensed
surrogate that is dened over a moisture and LAI gradient will be effective in predicting oristic distances between sites.
At the larger landscape scale, the saturation level of the oristic
distance measure is achieved at relatively low levels on the corresponding remotely sensed distance measure, compared to the smaller landscape scale (Fig. 3b).
Tree oristic distance between plots was positively correlated with
remotely sensed eco climatic distance between the plots (rm =0.10, p 0.001,
broad scale, n=464), although there was no signicant correlation at the
ner landscape scale (Table 1), a result that is not supported by the
quantilequantile plots. We attribute this to the fact that Mantel's tests are
based on a linear correlation whereas we have a clearly non-linear relationship indicated by the quantilequantile plots (Fig. 3).
3.3. Ordinary regressions

climatic distance suggests that the most important environmental

gradient inuencing forest type is moisture availability and bio-climate
at both landscape scales. It is to be noted that after reaching a threshold
canopy cover and biomass, NDVI no longer increases as has been identied in numerous studies throughout the tropics (Asner et al., 2003;
Lillesaeter, 1982) and this is a shortcoming of NDVI based distance
measures. This technique is therefore most successfully applicable to
regions with a distinct dry season and a mosaic of deciduous and evergreen forest types rather than an area with only dense, moist evergreen
forest types.
Turn-over in tree-species composition (beta-diversity) in disturbed
and fragmented landscapes such as the Western Ghats is determined
through a complex set of processes including species traits (e.g. dispersal
mechanism) and characteristics of the bio-physical and socio-ecological
landscape (e.g., environmental dissimilarity, topographic factors, geographical proximity). Geographic variation in beta-diversity reects past
and present differences in environment, human-use, ecological interactions, and biogeographic history (McKnight et al., 2007; Seidler &
Plotkin, 2006). RS distance rather than spatial distance may be powerfully used in such highly interspersed landscapes. Methods based on
distance decay alone do not necessarily account for environmental
heterogeneity, especially in heavily fragmented landscapes (Palmer,
2005). Moreover, at local scale, spatial distance alone may not be high
enough to reveal patterns of beta-diversity.
In our study, At the ner scale, sites are relatively spatially proximate
to each other, the forests are largely deciduous and dominated by winddispersed and therefore tree species composition are likely to more
similar even between sites that are different in their eco-climatic distance measure. At the larger scale, sampled tree-plots are more spatially
distanced, and thus tree-species composition differences between sites
are likely to be more different even for sites with similar eco-climatic
distance attributes.
At both scales, anthropogenic disturbance tends to complicate the
ability to predict rates tree-species compositional turn-over between
sites across an environmental and geographical gradient, and this shows
up as considerable scatter in tree-species compositional dissimilarity for
a given lag in eco-climatic distance.
Thus we recommend that spatial proximity and remotely sensed
eco-climatic dissimilarity between sites taken together would be good
covariates to predict turn-over in tree-species composition between
sites, but this model would be weaker in human-transformed landscapes. We also recommend caution in use of Mantel's tests in testing
surrogates for biodiversity without checking for non-linearity using
quantilequantile plots.

The remotely sensed eco-climatic distance averaged across forest

type is closely and signicantly correlated to tree species compositional
distance averaged across forest type to reference evergreen forest type,
at both landscape scales (Fig. 4). Linear regression models to predict
BrayCurtis/Sorenson tree compositional distance to reference evergreen forest using the remotely sensed measures were highly signicant
(r 2 = 0.86, p b 0.00001) respectively.
4. Discussion
The remotely sensed distance measure we have dened is a good,
robust and easily interpretable complement to conventional classied
digital image based classication. Our results show that tropical forest
types across a moisture gradient show variability in structural/
physiognomic classication that captures the gradients in tree oristic
composition as well. Our results conrm and support early concepts
about the relationship between tropical forest species composition and
structure (e.g. Webb et al., 1970). In addition, the inferred relations
between physical environment and tropical forest structural types,
inclusive of canopy foliage attributes were quantitatively established by
earlier studies (Mackey, 1994, 1993; Webb, 1968). The strong relationships between oristic distance to reference evergreen forest and eco-

Fig. 5. Examples of habitat or niche of three large mammals from the Western Ghats (Chital
deer (Axis axis), Sambar deer (Cervus unicolor) and Lion-Tailed Macaque (Macaca silenus)
and Western Ghats endemic bird species dened on the remotely sensed eco-climatic
distance measure. The range for each species was dened approximately using locational
information available from published studies.

J. Krishnaswamy et al. / Remote Sensing of Environment 113 (2009) 857867

5. Conclusion
Our results are noteworthy for three reasons.
First, the remotely sensed eco-climatic distance successfully quanties habitat and forest variability, with low values corresponding to
more moist, denser, more evergreen forest habitats with high evapotranspiration, and high carbon storage and higher values to more open
dry deciduous and scrub habitats with low evapotranspiration and
lower carbon storage (Figs. 1 and 2). Thus it is has quantied forest type
and ecosystem services on a single continuous numerical scale.
Furthermore we have already shown in a previous study (Das et al.,
2006) and further demonstrated it in this paper, that the RS based metric

865

is a good approximate surrogate for various components of biodiversity

at broad to very broad scales.
Second, we can rigorously quantify the bio-climatic niche of species
(e.g. Fig. 5) as this multi-date NDVI measure integrates leaf area index,
canopy cover, tree-density, degree of deciduousness and plant available moisture availability within a single metric.
Third, this remotely sensed surrogate has important applications
in quantifying and mapping of ecosystem services especially hydrologic and carbon services for large landscapes. Hydrologic uxes are
now divided into blue (surface and ground-water) and green water
(evapotranspiration) ux based ecosystem services (Gordon et al.,
2008; Rockstrm et al., 1999). As evapotranspiration is increasingly

Fig. 6. Spatially explicit (25 km2 resolution) (a) green-water and (b) blue-water uxes in mm per year in a part of the Western Ghats predicted by the Zhang, Walker and Dawes model
using the eco-climatic distance based prediction of plant-available moisture index. Protected areas are overlayed. An ideal measure of net hydrologic service would be a weighted
average of blue and green water uxes, adjusted for rainfall. The scale for green-water uxes above also approximately covers above ground carbon storage from less than
approximately 50 Mg C ha 1 to over 250 Mg C ha 1.

866

J. Krishnaswamy et al. / Remote Sensing of Environment 113 (2009) 857867

being considered a necessary eco-physiological process that sustains

other ecosystem goods and services (e.g. biomass, ood and overlandow control) and part of evapotranspiration is recycled as rainfall at
various spatial scales, and not all of blue-water ux is always benecial (e.g. ood water in the wet-season), a more appropriate measure
of a net hydrologic service would be a suitably weighted average of
green and blue water uxes, adjusted for rainfall (Fig. 6). In addition,
as the eco-climatic distance measure we have dened is related to
tree-density, LAI and degree of deciduousness, so it is a direct measure
of carbon storage (Fig, 1 and Fig. 6) and sequestration as well.
Overall, we have demonstrated that we can use multi-date
remotely sensed data to quantify forest type variability on a single
numerical scale and use this approach to detect broad scale patterns of
bio-diversity and ecosystem services that can be used in conservation
planning. Although our techniques would be most suitable for regions
where tree species are relatively diverse in their phenological
responses, advancements in satellite sensors, spatial and temporal
resolution, the utility of these techniques will be enhanced for other
regions. Our approach has the potential of further improvement as
satellite data improves in spectral and temporal resolution. The low
cost, simple techniques based on remotely sensed imagery may allow
us to map patterns of species assemblages and ecosystem services over
large spatial scales and detect changes in biodiversity and ecosystem
services over time due to land-cover change, global climate change and
other human impacts.
Acknowledgements
This study was supported by grants from the Ford Foundation, New
Delhi, Dorabji Tata Trust, Mumbai, and the Ministry of Environment and
Forests, Government of India, Save the Tiger Fund, Natural Environment
Research Council, UK, and the Stockholm Resilience Centre. This paper is
number 3 in the Suri Sehgal Centre for Conservation Science, ATREE
series. We thank B. Shivraj who arranged for access to the data from the
Forest Survey of India's network of plots in the Western Ghats and
Murali and S. Setty who helped generate the BRT eld data. N. Barve and
Bipin C. spent a considerable amount of effort in converting the FSI data
into a spatial format. S. Davande helped with image processing. A. Das
and M. Irfan Ullah provided support. Robert Chandran provided useful
inputs with interpretation. Harini Nagendra commented on an earlier
draft. We thank Dean Urban for sending the Splus modules developed by
Sarah Goslee and him which enabled us to do the Mantel's tests. We
thank two anonymous reviewers and the editor for critical inputs that
considerably enhanced the quality of the manuscript.
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