Download vegetation dynamics in the western himalayas, diversity indices and

Sci., Tech. and Dev., 31 (3): 232-243, 2012
VEGETATION DYNAMICS IN THE WESTERN
HIMALAYAS, DIVERSITY INDICES AND CLIMATE
CHANGE
SHUJAUL M. KHAN1*, SUE PAGE2, HABIB AHMAD3, HAMAYUN SHAHEEN4
AND DAVID HARPER5
1
Department of Botany, Hazara University, Mansehra, Pakistan.
Department of Geography, University of Leicester, UK
3
Department of Genetics, Hazara University, Mansehra, Pakistan.
4
Department of Botany, Azad Kashmir University, Muzaffarabad, Pakistan.
5
Department of Biology, University of Leicester, UK
2
Abstract
Vegetation provides the first tropic trophic level in mountain ecosystems and hence
requires proper documentation and quantification in relation to abiotic environmental variables
both at individual and aggregate levels. The complex and dynamic Himalayas with their
varying climate and topography exhibit diverse vegetation that provides a range of ecosystem
services. The biodiversity of these mountains is also under the influence of diverse human
cultures and land uses. The present paper is not only first of its kind but also quite unique
because of the use of modern statistical techniques for the quantification of Diversity Indices of
plant species and communities. The vegetation was sampled in three categories, i.e., trees,
shrubs and herbs, as follows: a height of ≥ 5m were classified in the tree layer, shrubs were all
woody species of height 1m and 5m and, finally, the herb layer comprised all herbaceous
species less than 1m in height. The presence/absence of all vascular plants was recorded on preprepared data sheets (1, 0 data). For the tree layer, the diameter of trees at breast height was
measured using diameter tape. Coverage of herbaceous vegetation was visually estimated
according to Daubenmire and Braun Blanquet methods. It gives overall abundance of vascular
plants on one hand and composition of these species on the other. Data was analysed in
Canonical Community Coordination Package (CANOCO) to measure diversity indices of plant
communities and habitat types. Results for five plant communities/habitat types indicated that
plant biodiversity decreased along the altitude. Shannon Diversity Index values range between
3.3 and 4. N2 index and Index of Sample Variance were also designed. All of these Diversity
Indices showed the highest values for the communities/habitats of north facing slopes at middle
altitudes. Higher plant diversity at these slopes and altitudes can be associated to the period of
snow cover which is longer and a relatively denser tree cover as compared to the southern
slopes and hence the soil has high moisture which supports high biodiversity in return. Global
warming causes desertification in number of fragile mountain ecosystem around the globe.
These findings suggest that species diversity decreases along the measured ecological gradient
under the influence of deforestation coupled with global climatic change.
Keywords: Biodiversity, Diversity Index, Climate, Mountain Ecosystem, Western Himalayas,
Vegetation.
Introduction
Mountains are the most remarkable land
forms on earth surface with prominent vegetation
zones based mainly on altitudinal and climatic
variations. Variations in aspects also enhance
habitat heterogeneity and bring microenvironmental variation in vegetation pattern
(Clapham, 1973; Khan et al., 2011b). In northwestern Pakistan, three of the world’s highest
mountain ranges, i.e., the Himalayan, Hindu Kush
*Author for correspondence E-mail: [email protected]
VEGETATION DYNAMICS IN THE WESTERN HIMALAYAS, DIVERSITY INDICES AND CLIMATE CHANGE
and Karakoram, come together, ensuring high
floral diversity and phytogeographic interest.
Plant biodiversity survives at the edge of life at
these high mountains where climatic changes are
more visible and species extinction very rapid.
Studies on mountainous vegetation around the
globe show that ecological amplitude of alpine
species shifted to even higher elevations over the
recent decades. Species preferred temperatures in
higher altitudinal zones. Therefore, many alpine
species are under the risk of extinction due to
their requirements for germination and
reproduction (Grabherr et al., 1994; Holzinger et
al., 2008). Unlike the eastern Himalayas, where
monsoon-driven vegetation predominates under
higher rainfall and humidity (Chawla et al., 2008;
Dutta and Agrawal, 2005; Anthwal et al., 2010;
Behera et al., 2005; Roy and Behera, 2005), the
vegetation in the western Himalayas in general
(Chawla et al., 2008; Kukshal et al., 2009;
Shaheen et al., 2011; Ahmad et al., 2009; Dickoré
and Nüsser, 2000; Shaheen et al., 2012) and in the
Naran Valley (Khan et al., 2011b) in particular
have closer affinities with that of the Hindukush
mountains, which have a drier and cooler climate
(Ali and Qaiser, 2009; Wazir et al., 2008;
Noroozi et al., 2008). A recent study showed that
with the passage of time, homogeneity takes place
in the vegetation of a region due to continuous
dominance of certain resistant and vigorous
species. This phenomenon is further enhanced by
selective utilisation of species by human
intervention (Del Moral et al., 2010). Similarly,
the tree-line vegetation and indicator species shift
upwards due to climate change.
Mountainous vegetation has manifold
functions, not only within the system where it
exists regionally in the lowland ecosystem by
regulating floods and flow in streams and
globally in combating the climate change and
greenhouse
effects.
Regionally,
shrubby
vegetation of high altitude regulates avalanche
movements and protects soil but in majority of
places, it is threatened due to human and climatic
influences (Hester and Brooker, 2007). Plant
biodiversity is necessary for regulation of overall
system in the mountain, e.g., the Himalayas is the
birth place for 10 largest rivers in the Asia and a
big and important carbon sink. Ecological
changes in the Himalayas affect global climate by
bringing changes in temperature and precipitation
233
patterns of the world. Himalayan Vegetation is
diverse and range from tropical evergreen species
in the south east to thorn steppe and alpine
species in the north western parts (Behera and
Kushwaha, 2007). Melting of its snow in a
regular fashion is related to its vegetation cover.
Irregular loss of its ice might have dangerous rise
in world sea-levels (Xu et al., 2009). These
mountains are extremely sensitive to global
climatic change. Such hazardous glimpses have
already been observed in the form of flood in
Pakistan, India, China and Thailand in the last
three years.
The Naran Valley is located at the far west of
the Western Himalayas on the border with the
Hindu Kush range which lie to the west and near
to the Karakorum Range to the north (Figure 1).
Due to this transitional location the valley host
representative vegetation types from all three
mountain ranges. Monsoon winds are main
source of precipitation and also a primary force of
controlling erosion and climatology over millions
years of time and thus modify its climate,
topography and vegetation of Himalayas but in
the western Himalaya, especially in Naran Valley,
high mountains situated at the opening of the
valley act as barriers to the incoming summer
monsoon from the south and limit its saturation
into upper northern parts. Thus, summers remain
cool and relatively dry and make most of the
valley as a dry temperate-type of habitat. There
are clear seasonal variations. Total average
annual precipitation is low at only 900-1000mm
but there is heavy snowfall in winter which may
occur any time from November to April (average
annual snowfall 3m). The range reflects a sharp
increase in depth of snow with increasing altitude.
There is a distinct wet season in January-April
whilst the driest months of the year are JuneNovember. As far as temperature of the area is
concerned, most of the year, it remains below
10°C. December, January, February and March
are the coldest months of the year in which
temperature remains around the freezing point or
even below most of the times. June to August is
the main season for growth, with average daytime
temperatures in the range 15-20°C. Geologically,
the valley is located where the Eurasian and
Indian tectonic plates meet and where the arid
climate of the western Eurasian mountains gives
way to the moister monsoon climate of the Sino-
234
Japanese region (Qaiser and Abid, 2005;
Takhtadzhian and Cronquist, 1986; Kuhle, 2007).
The valley thus occupies a unique transitional
position in the region. In remote mountainous
valleys like the Naran, the complexity of
ecosystems, their inaccessibility and the cost and
time factors make it extremely difficult to observe
each and every aspect of vegetation features.
Perhaps those were the reasons that in spite of its
high phytogeographic importance, there have
been no previous quantitative studies of the
vegetation in this valley. The present study is not
only the first of its kind but also quite unique
because of the use of modern statistical
techniques for the quantification of plant species
and
communities
along
geo-climatic
environmental gradients that has limited
comparator studies in this region (Wazir et al.,
2008; Saima et al., 2009; Dasti et al., 2007). The
Naran Valley, which is floristically located in the
Western Himalayan province of the IranoTuranian region, forms a botanical transitional
zone between the moist temperate (from the
South East) and dry temperate (from the North
West) vegetation zones of the Hindu Kush and
Himalayan mountain ranges, respectively. A
SHUJAUL M. KHAN ET AL.
phyto-climatic gradient of vegetation based on
life forms further emphasizes the nature of the
area and makes apparent the transitional position
of the region, though predominated by alpine
species (Khan et al., 2011b & c).
The quantitative approaches to vegetation
description and analyses deployed in this study
not only fill methodological deficiencies (Khan,
2012) and gaps in the literature, i.e., evaluation of
diversity indices but also provide a firm basis for
extending this approach to the adjacent mountain
systems that are in need of up to date vegetation
mapping (Fosaa, 2004; Mucina, 1997). In
addition, the present study documents and
provides suggestions for the conservation of
mountain plant biodiversity under a scenario of
continuous human exploitation and climate
change. It is necessary to maintain ecosystem
services in general and food security in particular,
not only within mountain system but also for the
people and ecosystems of the lowlands that
depend on those mountains (Rasul, 2010;
Manandhar and Rasul, 2009; Sharma et al., 2010;
Khan et al., 2011a).
Fig. 1. Physiographic map showing the elevation zones and position of the Naran Valley, western
Himalayas (study area) in relation to Karakorum and Hindu Kush mountains
52
SHUJAUL M. KHAN ET AL.
Methodology
In total of 432 quadrats in 144 replicates, the
vegetation was sampled in three layers i.e., trees,
shrubs and herbs. Quadrats sizes used in this
study were 10x5 (for trees layer), 5x2 (for shrubs
layer) and 1x0.5 (for herbs layer). Trees had a
height of ≥ 5m, shrubs were all woody species of
height in the range 1-5 m and the herb layer
comprised all herbaceous species less than 1m in
height. The presence of all vascular plants was
record on pre-prepared data sheets (1, 0 data). For
the tree layer, the diameter of trees at breast
height was measured using diameter tape
(Figure 2). This enabled an assessment of tree
cover values in the quadrats. For the shrub and
herb layers, abundance/cover values were visually
estimated according to a regulated BraunBlanquet scale later on modified by Daubenmire
(Table 1) (Braun-Blanquet et al., 1932;
Daubenmire, 1968). Absolute and relative values
of density, cover and frequency of each vascular
plant species at each station were calculated using
phytosociological formulae to finally calculate
Important Value Index (McIntosh, 1978).
Fig. 2. Measuring diameter of tree species through diameter tape in a 10x5m quadrat in the field at
Lalazar, Naran.
Cover Class
1
2
3
4
5
6
Table 1. Braun Blanquet covers classes modified by Daubenmire.
Range of Percent Cover
Midpoint
0-5%
2.5%
5-25%
15.0%
25-50%
37.5%
50-75%
62.5%
75-95%
85%
95-100%
97.5%
52
In the past, it was a very common technique
to classify plant communities based on Important
Value Index (IVI) of plants species. In this sort of
characterization
and
classification
in
phytosociology, the species with the highest
Importance Values IV are considered as dominant
plant species and communities are attributed with
them. Even today, it is a very common method of
classifying plant communities. The present day
revolution of computer based technology in every
field of science has modified the older techniques.
The IVs were calculated by adding the values of
relative density, cover and frequency and then
dividing by 3. In our study, we use IV data
mainly for ordination purposes.
Data analysis
Ecologists use number of diversity indices for
measuring the plant species diversity in different
habitat types and plant communities. CANOCO
version 4.5 (Ter Braak 1988, Ter Braak 1989, Ter
Braak and Smilauer 2002) was used to analyse
IVI data to assess diversity indices. The main
objective of these statistical packages is to
formulate the study more precisely. As
phytosociology is concerned with vegetation
itself, the sites in which it occurs and the
environmental variables related with those sites
hence need to be examined in a statistical
framework (Kent and Coker, 2002; 1995,
Lambert and Dale, 1964). Plant biologists often
need to test hypotheses regarding the effects of
investigational factors on whole groups of species
(Khan et al., 2011b; Shaheen et al., 2011;
Anderson et al., 2006). CANODRAW is a utility
of CANOCO and was used to generate data
attribute
plots
(graphic
forms)
of
indicator/characteristic species. It also gives the
opportunity to utilise index of number of species,
Shannon Weiner index, index of species richness,
evenness, sample variance, etc. Diversity indices
were calculated at the community through data
attribute plot under the DCA function of
CANODRAW.
Results
Floristic composition of the Naran Valley
A total of 198 plant species belonging to 150
genera and 68 families were recorded at the 432
replicates of relives/quadrats. The division into
major taxonomical groups (Table 2) indicates that
SHUJAUL M. KHAN ET AL.
dicotyledonous angiosperms are the most
abundant. Five plant communities were identified
and discussed in our first paper from this project
(Khan et al., 2011b). There is a clear altitudinal
zonations, i.e., (i) the lower altitude (2450-3250
m) dominated by temperate vegetation and (ii)
the higher altitude (3250-4100 m) dominated by
subalpine and alpine vegetation. The most
abundant plant family was Asteraceae
(Compositae) with seventeen species and a 25%
share of all species. Rosaceae represented by
fourteen species (with a 21% share) was the
second most species rich family in the study area.
Lamiaceae,
Ranunculaceae,
Poaceae
and
Polygonaceae were represented by 13, 12, 11 and
10 plant species respectively. The remaining
families all had less than 10 species each.
Importance Values of the top 15 abundant species
are given in Table 3.
Table 2. Taxonomic divisions of recorded plant species.
No. of
No. of
Taxonomic distribution
Families
Species
Dicot
53
161
Monocot
9
23
Subtotal of Angiosperms
62
184
Gymnosperms
3
8
SUBTOTAL OF
65
192
SPERMATOPHYTA
Pteridophyta (Ferns and allies)
3
6
TOTAL NO. OF PLANT
68
198
SPECIES
Table 3.
S.No.
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
Important Value Index (IVI) of the top 15
species reported from the region.
Importance
Name of the Species
Value (IV)
Abies pindrow
584
Betula utilis
582
Sambucus weghtiana
446
Juniperus communis
338
Pinus wallichiana
323
Salix flabillaris
160
Rosa webbiana
137
Artemesia absinthium
135
Rhododendron hypenanthum
116
Fragaria nubicola
112
Berberis pseudoumbellata
111
Thymus linearis
109
Iris hookeriana
104
Cedrus deodara
103
Viola canescens
96
Based upon plant growth habit, 12 trees, 20
shrubs and 166 herbs shared 6, 10 and 84%,
VEGETATION DYNAMICS IN THE WESTERN HIMALAYAS, DIVERSITY INDICES AND CLIMATE CHANGE
respectively, in the plant diversity of the
Fig. 3.
237
vegetation of the Naran Valley (Figure 3).
Percent share of various habit forms of vegetation of the Naran Valley.
Species richness along environmental
gradients
Analyses show that the main influencing
environmental variables are altitude, aspect (slope
direction) and soil depth and that both species
richness and diversity vary along these gradients.
Analysis of the elevation gradient showed that
species richness was higher at lower altitudes and
lower at higher elevations. Species β-diversity
gradually decreased along the altitudinal gradient
as soil depth and temperature decreased.
Similarly species richness on slopes with a
northern aspect was slightly higher than those
with a southern aspect (Fig. 4).
Fig. 4. Average β-diversity of plant species along the altitudinal gradient.
52
Diversity indices using DCA analysis
Data attribute plots using Deterended
Correspondence Analysis (DCA) were used to
calculate diversity indices. The first sort of
diversity index based on species richness was the
calculation of species number per community
(Figure 5). The highest number of plant species
was reported in community 2 at middle altitude
northern aspect habitats followed by lower
altitude valley bottom habitats. The high diversity
of these habitat types can be attributed to high
soil depth with high moisture retaining capacity
and relatively high temperature at those
SHUJAUL M. KHAN ET AL.
somewhat lower altitudes. The lowest number of
species scored in the index was for the
community number 5 at peak elevations above
the tree line habitats.
The Shannon Diversity index values range
between 3.3 and 4. Being constituted of both
Northern and Southern aspect stations,
community 1 has the highest value of 3.98
amongst all the groups whilst community 5 has
the lowest value due to narrow ecological
amplitude (Figure 6). High values of index are
due to the inclusion of considerable number of
stations in a single community.
Fig. 5. Index of species number (alpha diversity) amongst all community types through DCA data
attribute plots along the gradients.
Fig. 6. Shannon Weiner diversity index at community level through DCA data attribute plots show the
gradient of the community.
52
The N2 index is the reciprocal of Sampson
diversity index and is available in the
CANODRAW utility of CANOCO. This index
showed the highest value for community 2
followed by community 1 (Figure 7).
Index of sample variance showed the highest
variance at community 2 amongst the groups. As
SHUJAUL M. KHAN ET AL.
the altitudinal pattern (Figure 4) showed that
species richness is optimum at the middle altitude
especially on north facing slopes. This index
reconfirmed the phenomenon of species richness
phenomenon through sample variance index
(Figure 8).
Fig. 7. N2 diversity index at community level through DCA data attribute plots along the gradient of the
community.
Fig. 8. Index of Sample Variance amongst all community types through DCA data attribute
plots along the gradient.
The pattern of plant communities in the
valley is largely determined by aspect and
altitude. Four plant communities can be separated
on the basis of these two variables, whilst one
community is established under the combined
effect of lower altitude and greater depth of soil
(the third most important variable). Relatively
higher summer temperatures and soil moisture are
the co-variables associated with the low elevation
and deeper soil. Predominantly, community 1
reflects the latitudinal gradient of vegetation from
moist temperate to dry temperate along the valley
on either side of the River Kunhar at lower
altitudes. Species diversity and richness are
240
optimal at the middle altitudes (2800-3400m), in
contrast to the lower altitudes (2400-2800m)
where direct anthropogenic activities have had
their greatest impact.
Discussion
Diversity amongst plant communities/habitat
types
Short summer, deep snow, low temperature,
intense solar radiation and cold speedy winds in
the valley in general and higher altitude and
special slope orientation in particular result xeric
condition for plant growth and hence the 𝛽diversity is gradually decreasing both along the
altitudinal and latitudinal gradients of the valley.
Though drawing a sharp line in any natural
ecosystem of mountains is difficult as the rapid
micro climatic and edaphic variations overlap
each other due to number of driving agencies and
historical perspectives but based on some
indicator species, vegetation zonation and
association can be established. Both at habitat and
community level, the plant species richness and
diversity is strongly influenced by elevation,
aspect and soil depth which in turn are associated
with precipitation and soil moisture. Various
diversity indices including number of species,
Shannon Weiner, sample variance and N2 show
maximum values for the middle altitude northern
aspect plant community (Com. 2) followed by the
subalpine (Com. 4) and middle altitude southern
aspect plant communities (Com. 3). Similar
patterns of diversity across altitudinal gradients
have been observed in other studies in the
Himalayan regions (Kharkwal et al., 2005;
Tanner et al., 1998; Vázquez and Givnish, 1998).
Implications of the present project for future
studies
Observing biodiversity is a lifelong process
and needs inputs from various disciplines from
the natural as well as the social sciences. For
better ecosystem management and protection,
plant biodiversity should be studied at species,
community and ecosystem levels. The
mountainous valleys need more botanical
exploration for the complete description of their
plant taxa, abundance and conservation status.
There are also extensive gaps in information on
ecosystem services in the Himalayan region.
Most of the vegetation studies have been carried
out solitarily either based on scientific approaches
SHUJAUL M. KHAN ET AL.
or public perception for making inventories. We
also believe and propose that such approach must
be statistically sound and communicable to
conservationists, planners, politicians and policy
makers. Furthermore, we suggest that the
identification of indicator species for specific
habitats will assist in monitoring and assessment
of the biological diversity on the one hand and the
effects of climatic change on the other. In
addition, a broad scale deterioration of the natural
ecosystems due to agriculture, expansion of
roads, increases in population and deforestation
cause enormous losses to the natural vegetation.
In spite of IUCN recommendations, there is very
limited and small scale documentation on Red
List Categories for plant species in the Himalayas
as well as in Pakistan more generally (Ali, 2008;
Pant and Samant, 2006; Chettri et al., 2008).
Being a member state of the CBD, Pakistan
should give top priority to this issue as addressed
by Khan (2012) and Khan et al., (2011b & c).
The three mountain ranges i.e., the
Himalayas, Hindu Kush and Karakorum, provide
essential ecosystem services to millions of people
across 10 countries of the world (Dong et al.,
2010; Khan et al., 2007). These services include
provisioning, regulating, supporting and cultural
services. In the present study, the main focus was
on vegetation mapping and provisioning services
in one of the Himalayan valley. Most of the
provisioning services considered in the present
study can further be evaluated at molecular and
biochemical levels in the future. Beyond the
direct role of plant biodiversity in the
socioeconomics of the mountain people, it is
indispensable for the people living in the plains
and hence this mountain valley can be studied for
the evaluation of the other kinds of services,
including those of regulatory nature, that it
provides on a broader level, e.g., flood control,
erosion control, irrigation water and hydropower
development. These mountains also provide
Supporting Services, e.g., soil formation,
biogeochemical and nutrient cycling, etc. These
mountain systems host characteristic biodiversity
on the one hand and a long established and
unique cultural diversity on the other.
Preservation of their indigenous knowledge and
its utilization in environmental management can
also be potential topics for future studies (Fig. 9).
VEGETATION DYNAMICS IN THE WESTERN HIMALAYAS, DIVERSITY INDICES AND CLIMATE CHANGE
241
Fig. 9. Sketch showing the potential topics for future to study different aspects of biodiversity of the
western Himalayas
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