EXTRACTION OF NON-TIMBER FOREST PRODUCTS IN THE

FORESTS OF BILIGIRI RANGAN HILLS, INDIA. 5. INFLUENCE OF

DISPERSAL MODE ON SPECIES RESPONSE TO ANTHROPOGENIC
PRESSURES 1
K . N . GANESHAIAH, R . U M A SHAANKER, K . S. MURALI, U M A SHANKAR,
AND K . S. B A W A 2

K. N. Ganeshaiah (Department of Genetics and Plant Breeding, University of Agricultural

Sciences, GKVK Campus, Bangalore 560 065, India), R. Uma Shaanker (Department of Crop
Physiology, University of Agricultural Sciences, GKVK Campus, Bangalore 560 065, India),
K. S. Murali (Tata Energy Research Institute, 50/7, Palace Road, Bangalore 560 052, India),
Urea Shankar (Tata Energy Research Institute, Ghoshpara, Hakimpura, Siliguri 734 401,
India) and K. S. Bawa (Department of Biology, University of Massachusetts, Boston, MA
02125, USA). EXTRACTIONOF NoN-TIMBER FOREST PRODUCTS IN THE FORESTS OF BILIGIRIRANGAN
HILLS, INDIA 5. INFLUENCEOF DISPERSAL MODE ON SPECIES RESPONSE TO ANTHROPOGENICPRESSURES. Economic Botany 52(3):316-319. 1998. We examined the response offorest tree species

with different dispersal modes to anthropogenic pressure in a d O, deciduous forest of South

India. The species and their populations were sampled in two forest stands, one in proximity
to a Soliga settlement (greater disturbance) and the other distant to the settlement (lower
disturbance). Our results suggest that the populations of animal dispersed species than those
of wind or passively dispersed species are more vulnerable to human disturbance. In fact wind
dispersed species seem to be facilitated by human disturbances. The proximal site had a higher
representation of understo~ plants and seedling belonging to wind dispersed species than that
of animal dispersed species. We discuss the results in the context of the role of dispersal mode
in shaping species response, and vegetation composition of forest to anthropogenic pressures.

Key Words:

dry deciduous forest; NTFP extraction; wind dispersal; animal dispersal; Soliga.

The response of the forest vegetation to anthropogenic pressures may depend upon the intrinsic features of its flora (Kruckeburg and Rabinowitz 1985). In particular, the reproductive
features and regeneration potential of the constituent species might strongly influence their response to human induced pressures such as harvesting, fire and grazing. For instance, Daniels,
Gadgil, and Joshi (1995) showed that in the
tropical humid forest of western ghats in south
India, species with good coppicing ability were
less vulnerable to anthropogenic disturbance
than those with poor potential for coppicing. In
orchids, floral features, such as large size and
bright colors, were found to predispose species
to extinction following harvesting (Lokesha and

~Received 30 December 1996; accepted 25 January

1998.
2 Correspondence.

Vasudeva 1992). Forests impacted by human

disturbances tend to have a thinner canopy and
relatively drier environment (Daniels, Gadgil,
and Joshi 1995). The openness thus created may
adversely affect the activity of animal dispersers.
On the other hand, unhindered wind movement
in disturbed habitats would facilitate efficient
dispersal of wind dispersed species. Indeed, in
the central Amazonian forest, Gentry (1980,
1982) found that the proportion of wind dispersed species increased and the proportion of
animal dispersed species decreased with the dryness of the forest. Furthermore, Lokesha, Vasudeva, and Yellappa Reddy (1991) showed that
animal dispersed species are more prone to become rare and endangered than wind or passively dispersed species. In this paper, we examine
the change in the composition of forest vegetation with respect to the dispersal modes of species under different degrees of anthropogenic

Economic Botany 52(3) pp. 316-319. 1998

9 1998 by The New York Botanical Garden, Bronx, NY 10458 U.S.A.

1998]

GANESHAIAH

ET AL.: N O N - T I M B E R F O R E S T P R O D U C T S , I N D I A , 5

Y = 2 5 . 8 3 + 2.74 X, R 2 = 41
r = 0 . 6 3 8 , P < 0.05

pressure in a dry deciduous forest of Biligiri

Rangan Hills, India. We offer evidence to suggest that the dispersal mode of a species plays
an important role in shaping its response to anthropogenic pressures.
MATERIALS AND METHODS
This study is based on an earlier work by
Murali et al. 1996, involving the enumeration of
the vegetation at the Biligiri Rangan Temple
(BRT) Wildlife Sanctuary (11-13 ~ N and 77-78 ~
E), Mysore district, India; detailed account of
the site is available in other papers in this series
(Hegde et al. 1996; Ramesh 1989; Uma Shankar
et al. 1996; Uma Shankar et al. 1998; Uma
Shankar, Hegde and Bawa 1998). Murali et al.
(1996) identified two sites in the dry deciduous
forest, namely a "proximal" site which is located in the vicinity of a settlement called Kanneff colony and a "distant" site located away
from the settlement. At each site, they laid five
linear transects, 1000m long and 10m wide. All
species > 1 0 c m dbh were enumerated and their
girths were measured. Within each transect, 10m
10m quadrats were laid at 100m intervals and
individuals > l c m dbh but < 1 0 c m dbh were
enumerated. Within each of these quadrats, one
1 m x 1 m sub-quadrats were laid randomly and
all understory plants, saplings/seedlings of tree
species enumerated. As a measure of the extent
of human disturbances they computed the proportion of cut and broken trees at each of the
transects. They showed that the proximal site
had significantly higher proportion of cut and
broken stems compared to the distant site. The
disturbance index of the transects ranged from
2.6 to 8.81 for the proximal site and 0 to 0.98
for the distant site. Thus, the two sites clearly
appeared to experience different levels of human
pressures.
For this study, we ascribed for each species
enumerated by Murali et al. (1996) at the two
sites, a dispersal mode from among the three
following categories: wind, animal or passive
(including explosive) by examining fruit and
seed features and following Gamble (19151934). The dispersal mode of a few rare species
could not be ascertained, and they were excluded from the analysis. The frequency of species
and of individual stems (->10 cm dbh) belonging
to the three dispersal modes was computed for
each site. The impact of human disturbance on
species with different dispersal modes was an-

317

60

p
[]

>_401

.~
u

~20
~rd

E~

p
~

D
D

10

~ J, ~- ~ + 8 6 fo
DISTURBANCE INDEX

Fig. 1. Relation between percent individual stems

dispersed by wind and disturbance levels of the ten
transects. (D = site distant to human settlement; P =
site proximal to human settlement).

alyzed in two ways: (a) the disturbance index

was correlated to the percentage of species as
well as the percentage of stems with a given
dispersal mode in individual transects and (b) by
estimating the independence of the dispersal
mode with the location of the sites (proximal or
distant to human settlement). For this, contingency tables were set up for the number of species or individual stems (->lcm dbh) belonging
to various dispersal modes at proximal and distant sites (pooled over all five transects in each
regime) and a contingency chi-square test performed (Sokal and Rohlf 1969).
RESULTS AND DISCUSSION
The percent of stems (individuals) of species
dispersed by wind was positively correlated with
the disturbance level of the transect (r = 0.638,
P < 0.047, Fig. 1), while those of animal dispersed species were negatively correlated (r =
-0.591, P < 0.072, Table 1). Passive dispersal
mode did not exhibit any association with the
disturbance index of the transects. Thus, disturbance seems to enhance the frequency of stems
dispersed by wind and decrease those dispersed
by animals. These results are also borne through
the contigency chi-square analysis of the association of the stems of various dispersal modes
and the disturbance levels. The two sites exhibited significant differences in the number of individual stems with different dispersal classes

318
TABLE

1.

ECONOMIC BOTANY

[VOL. 52

TABLE 3.

A S S O C I A T I O N OF DISPERSAL M O D E OF

R E L A T I O N BETWEEN DISPERSAL MODE

( O F SPECIES A N D I N D I V I D U A L STEMS) A N D THE DIS-

UNDERSTORY PLANTS A N D SAPLINGS W I T H A N T H R O -

T U R B A N C E LEVEL OF T H E TRANSECTS.

POGENIC PRESSURE.

Dispersal mode

Dispersal mode

Wind

Animal

Passive

Percent
species

r = 0.456
P < 0.185

r = 0.275
P < 1.000

r = -0.173
P < 1.000

Percent
stems

r = 0.638
P < 0.047

r = -0.591
P < 0.072

r = 0.447
P < 0.195

Site

Wind

Passive

Animal

Distant

31
(49) ~

164
(157)

438
(427)

Proximal

41
(23)

68
(75)

195
(205)

Values in parentheses refer to the expected values. Chi-square = 22.4,

df = 2, P < 0.01.

(Chi-square = 146.6, df = 2, P < 0.001; Table

2). The proximal site had a greater number of
stems of wind dispersed species than the distant
site; nearly 54% of the stems in the proximal
site were wind dispersed, while only 35% of the
stems in the distant site were wind dispersed.
Conversely, a greater percentage of the stems in
the distant site (43%) were animal dispersed
than in the proximal site (22%).
However, on per cent species basis, there was
no significant correlation (Table 1) nor association (Chi-square = 0.26, df = 2, NS) between
the dispersal modes and the disturbance levels.
This could be attributed to the possible immigration of propagules into the sites from the
neighboring forest or to the common background vegetation that existed before the sites
were impacted by the anthropogenic pressures.
The predominance of wind dispersed stems at
the proximal site may have been facilitated by a
more effective dispersal, recruitment and regeneration of propagules due to the relatively open
canopy. In contrast, the reduction in the frequency of stems of animal dispersed species in
the proximal site may be due to poor dispersal
consequent to the interference to animal vectors
or due to the loss of safe sites. In fact, in the

proximal site, there was a greater number of understory plants and saplings/seedlings of wind
dispersed species than expected (Table 3); conversely, there was lesser number of seedlings of
animal dispersed species than expected (Chisquare = 22.43, df = 2, P < 0.001, Table 3).
The number of saplings recruited per individual
tree (->10 cm dbh) of wind dispersed species
was relatively more than that of the species in
the proximal compared to the distant site (Table
4). These differences, however, were not significant because of the high variance contributed
by the wide range of the species constituting a
given dispersal mode.
Thus, there appears to be a strong association
between anthropogenic pressure and the preponderance of species with wind dispersal in communities subjected to human pressures. Animal
dispersed species appear to be more vulnerable
to human impact than wind or passively dispersed species. Wind dispersed species, in fact

TABLE 4.

MEAN

NUMBER OF UNDERSTORY SAP-

LINGS AND SEEDLINGS PER A D U L T TREE OF SPECIES

B E L O N G I N G TO T H E VARIOUS DISPERSAL MODES A T

TABLE 2.
SPECIES

PROXIMAL A N D D I S T A N T SITES.I
A S S O C I A T I O N OF DISPERSAL MODE OF

(ON

I N D I V I D U A L STEM BASIS) W I T H A N -

Site

T H R O P O G E N I C PRESSURES.
Dispersal
mode

Dispersal mode
Site

Wind

Passive

Animal

Distant

576
(705) ~

355
(374)

698
(550)

Proximal

674
(545)

308
(289)

278
(426)

~Values in parentheses refer to expected frequency. Chi-Square =

146.6, df = 2, P < 0.001.

Wind
Animal
Passive

Distant
n

X + SD

9 0.100 + 0.119
15 2.510 + 4.212
4 0.177 + 0.490

Proximal
n

X + SD

11 0.367 + 1.277
16 1.350 + 2.082
2 0.118 + 0.074

~The number of understory saplings or seedlings per adult tree (>10

cm dbh) for each species of a dispersal mode was computed and averaged
over all species of the similar dispersal mode for each of the sites. Two
way analysis of variance for the effect of disturbance regimes on the
recruitment of seedlings per tree was non-significant.
2 n = number of species.

1998]

GANESHAIAH ET AL.: NON-TIMBER FOREST PRODUCTS, INDIA, 5

319

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Volume 15, Plenum Press, New York.
Hedge, R., S. Suryaprakash, L. Achoth and K. S.
Bawa. 1996. Extraction of non-timber forest products in the forests of Biligiri Rangan Hills, India.
1. Contribution to rural income. Economic Botany
50:243-251.
Kruckeberg, A. R., and D. Rabinowitz. 1985. Biological aspects of endemism in higher plants. Annual Review of Ecology and Systematics 16:447479.
Lokesha, R., R. Vasudeva, and A. N. Yellappa Reddy. 1991. Do rare/endangered/threatened plant species of south India have specific reproductive syndromes promoting their extinction? Proceedings of
the Symposium on Rare, Endangered, and Endemic
Plants of the Western Ghats. Trivandrum, Kerala.
Lokesha, R., and R. Vasudeva. 1992. Commercial
exploitation--a threat to Indian orchids? Current
Science 63:740-744.
Murali, K. S., Urea Shankar, R. Urea Shaanker, K.
N. Ganesbaiab, and K. S. Bawa. 1996. Extraction
of non-timber forest products in the forest of Biligiri Rangan Hills, India. 2. Impact of NTFP extraction on regeneration, population structure and
species composition. Economic Botany 50:252269.
Ramesh, B. R. 1989. Vegetation map of the Biligiri
Rangan Hills French Institute, Pondicherry.
Sokal, R. R, and F, J. Rohlf. 1969. Biometry. Freeman, San Francisco.
Uma Shaanker, R., K. N. Ganeshaiah, and T. R.
Radhamani. 1990. Association among the modes
of pollination and seed dispersal----ecologicalfactors and phylogenetic constraints. Evolutionary
ACKNOWLEDGMENTS
Trends in Plants 4:107-111.
This paper represents contribution number 52 of a research program
Uma
Shankar, K. S. Murali, R. Urea Shaunker, K.
in Conservation of Biodiversity and the Environment jointly co-ordinated
N. Ganeshaiah, and K. S. Bawa. 1996. Extraction
by the Tata Energy Research Institute, New Delhi and the University of
Massachusetts, Boston. The program is supported in part by the Macof non-timber forest products in the forests of the
Arthur Foundation. Dr. K. N. Ganeshaiah and Dr. R. Urea Shaanker were
Biligiri Rangan Hills, India. 3. Productivity, extracalso supported by a grant from the Department of Science and Technoltion and prospects of sustainable harvest of Amla,
ogy, Government of India, New Delhi.
Phyllanthus emblica (Euphorbiaceae). Economic
Botany 50:270-279.
LITERATURE C I T E D
Uma Shankar, R. Hedge, and K.S. Bawa. 1998. ExDaniels, R. J. R., M. Gadgil, and N. V. Joshi. 1995.
traction of non- timber forest products in the forests
Impact of human extraction of tropical humid forof Biligiri Rangan Hills, India 6. Fuelwood presests in the Western Ghats in Uttara Kannada, South
sure and management options. Economic Botany
India. Journal of Applied Ecology 32:866-874.
52:320-336.
Gamble, J. S. 1915-1934. Flora of the Presidency of Urea Shankar, K. S. Murali, R. Uma Shaanker, K.
Madras, 1-3, Adlard, London.
N. Ganeshaiah, and K, S. Bawa. 1998. Extraction
Gentry, A. H. 1980. Phytogeographic patterns as evof non-timber forest products in the forests of Biidence for a Choco refuge in biological diversifiligiri Rangan Hills, India. 4. Impact on floristic dication in the tropics. Pages 112-136 in G. Prance,
versity and population structure in a thorn scrub
ed., Columbia University Press, New York.
forest. Economic Botany 52:302-315.

appear to be relatively favored by human disturbance. While our results suggest a possible
role for dispersal mode in the response of species to anthropogenic pressures, it is likely that
other plant features or processes may be similarly important. Urea Shaanker, Ganeshaiah, and
Radhamani (1990) showed that there exists a
strong association between the modes of dispersal and pollination; wind dispersed species
tend to be wind pollinated and animal dispersed
species tend to be insect or animal pollinated. It
is therefore likely that the response of species to
anthropogenic pressures is also influenced by
their pollination mode. Wind pollinated and
wind dispersed species may be more resilient to
anthropogenic disturbances in the forest compared to species that are insect pollinated and
animal dispersed.
It can also be argued that disturbance should
favor early successional species that are likely
to have wind dispersed propagules. While this
might be true, our results seem to suggest that
wind dispersed species might gradually replace
species with other modes of dispersal, if the disturbance were to continue. Thus forests under
continuous and persistent anthropogenic pressure may come to be dominated by species with
abiotic mode of dispersal. Such changes in community structure in turn may have negative effect on populations of pollinator and seed dispersers thus leading to lower levels of biodiversity.