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ec o.m ont – Volum e 10, Num ber 1, Ja n u a r y 2 0 1 8
I SSN 2 0 7 3 -1 0 6 X print v ersion – ISSN 2073-1558 online v ersion: http:/ / epub.oeaw.ac .a t / e co .mo n t
https:/ / dx.doi.org/ 10.1553/ ec o.mo n t-1 0 -1 s4 6
From natural forest to cultivated land: Lichen species diversity along land-use
gradients in Kanchenjunga, Eastern Nepal
Til Bikram Chongbang, Christine Keller, Michael Nobis, Christoph Scheidegger & Chitra Bahadur Baniya
Keywords: canopy openness, Canonical Correspondence Analysis, Ghunsa valley, hemispherical photography, land-use
change, lichen diversity
Profile
A b s t r a ct
This study aimed to evaluate the effects of elevation, land use and canopy openness on species richness and composition of lichens in Ghunsa valley of Kanchenjunga Conservation Area, Eastern Nepal. At five elevational levels, from 2 200 m
to 3 800 m, transects were established in four land-use types – cultivated land,
meadows, exploited and natural forests. Detrended Correspondence Analysis (DCA)
and Canonical Correspondence Analysis techniques were used to explore the lichen
species distribution patterns. Generalized linear models were applied to analyse
the impact of elevation and canopy openness on lichen species richness. Canopy
openness was measured by hemispherical photography. A total of 229 species
belonging to 71 genera were recorded. The length of the first DCA axis of 8.01 SD
units indicated a complete species turnover and high beta diversity along the elevation gradient. Exploited forests with lower canopy openness supported higher lichen
diversity than open meadows and cultivated areas. Significant differences in lichen
species richness were found for different land-use types, along the elevation gradient, and with varying canopy openness. A gradual increase of lichen species richness
from cultivated land to forests was observed. We concluded that substrate types that
depend on land-use types as well as canopy openness significantly affect the distribution of lichen communities.
Introduction
Lichen diversity along elevational gradients has
been analysed intensively in recent years (Bruun et
al. 2006; Grytnes et al. 2006; Pinokiyo et al. 2008;
Cobanoglu & Sevgi 2009; Baniya et al. 2010; Rai et
al. 2011; Baniya et al. 2012) as well as lichen diversity along land-use gradients (Bergamini et al. 2005;
Motiejûnaitë & Faùtynowicz 2005; Stofer et al. 2006;
Wolseley et al. 2006; Giordani et al. 2010). Similarly,
some recent studies are concerned with the influence
of canopy openness on species richness, diversity and
distribution of lichens (Li et al. 2011; Marmor et al.
2012; Li et al. 2013b, 2013a). However, effects of landuse related canopy openness on species richness and
composition of lichens have rarely been studied.
Land-use change determines vegetation cover,
species composition and distribution patterns of
plant communities (Tasser & Tappeiner 2002) and,
consequently, the variation in key characteristics of
host tree species, like their density, age and diameter,
which all influence the composition and distribution
of epiphytic lichen communities (Löbel et al. 2006;
Mežaka et al. 2008; Cobanoglu & Sevgi 2009; Li et al.
2011; Mežaka et al. 2012; Odor et al. 2013). Land-use
changes, habitat loss and degradation often decline lichen populations (Scheidegger & Werth 2009). Compared with other factors, changing light and moisture
Protected area
Kanchenjunga
Conservation Area
Mountain range
Himalaya
Country
Nepal
conditions are often the dominant factors to explain
differences in lichen diversity and abundance (Li et
al. 2013a).
Nepal is a mountainous country in the central Himalayas with an area of 147 181 km2. It is situated between China in the north and India in the east, south
and west. The elevation ranges from 60 m above sea
level in Terai to 8 848 m at Mt Everest, the highest
peak in the world (Chaudhary 1998).
In Nepal, lichens are found in all climatic zones.
However, floristic and ecological studies on lichens are
largely missing. The latest physiographic data of Nepal showed 29 % of the total land area covered by forests, 10 % by shrubs and degraded forests and 21 % by
cultivated land (MFSC 2009). Land-use and land-cover
change are substantial in Nepal; especially the forest
cover shows a drastic decline – even in protected areas.
For the Kanchenjunga Conservation Area (KCA), for
instance, Gautam and Watanabe (2004) found a decline in forest land cover by 14.9 % and grazing land
cover by 77.9 % between 1979 and 1992. This was the
result of an increase in cultivated land by 4.9 % and
shrubland by 19.7 %. KCA is a community-managed
protected area established in 1997 and handed over to
the KCA Management Council by the government of
Nepal in 2006. The shifting cultivation is a common
traditional farming system practiced in this protected
area by the local ethnic groups as their traditional oc-
Cho ng ba ng e t a l .
Nepal
0 45 90
180
270
360
km
Afghanistan
China
Nepal
Bhutan
Pakistan
India
Bangladesh
protected areas
0
rivers
5
10
20
km
Kanchenjunga conservation area
transect points
Figure 1 – Map of the study area showing the locations of the study sites.
cupation and livelihood. It also falls within the Sacred
Himalayan Landscape being developed by WWF Nepal (Aryal et al. 2010).
The main objective in the present study, therefore, is
to evaluate the effects of different land-use types, canopy openness on species richness and composition of
lichens along the elevational gradient in KCA, Eastern
Nepal. We hypothesized that (a) lichen diversity generally decreases from forests to open land and (b) highest
lichen diversity is reached in forests under intermediate
canopy openness.
Materials and methods
Study area
This study was carried out in Ghunsa of Eastern
Nepal between 2 200 m and 3 800 m (Figure 1). Ghunsa lies towards the north-eastern part of Nepal in
the KCA. KCA is located between 27º 24’–27º 57’ N
latitudes and 87º 39’–88º 12’ E longitudes, close to the
boarders of China in the North and India in the East.
KCA covers an area of 2 035 km2 between the Middle
Mountains and the high Himalayas, with an elevational
range from 1 200 m (Thiwa Khola) to Mt Kanchenjunga (8 586 m), the third-highest peak in the world. The
area includes three river valleys: Simbua, Ghunsa, and
Tamur (Anonymous 2011). KCA has diverse climatic
zones, including subtropical monsoon at 1 200 m to
alpine forests (above 4 000 m), where June to August
are the warmest months, with monthly maximums of
24.73°C to 24.81°C, and January is the coldest month,
with a maximum temperature of 13.8°C (Shrestha &
Ghimire 1996). KCA receives a good amount of monsoon rainfall from April / May to September / Octo-
ber, with a mean annual precipitation of 2 013 mm / yr
(Anonymous 2009).
Field methods and data collection
Land-use types were classified according to land
cover, disturbance frequency and intensity. At each elevational level, land-use gradients were stratified into
four land-use types (Scheidegger et al. 2010).
1. Natural forest: Forested area with very little or no
human disturbance. It includes mainly broad-leaved
trees and pine trees. This land-use type is often several hours walking distance away from human settlements.
2. Exploited forest: Disturbed and / or exploited forests used for extensive grazing and / or the collection of fodder and firewood, which are close to human settlements.
3. Meadow: Areas dominated by grasses and scattered
trees and shrubs. Grazed by domestic livestock like
sheep, goats, buffaloes, cows, yaks, and horses.
4. Cultivated land: Land extensively used for cultivation and including terraced fields. These arable
fields are often irrigated and fertilized.
Fieldwork was carried out in April 2012. Five elevation levels, from 2 200–3 800 m, with an interval of approximately 400 m were selected for the study. At each
level, the four land-use types were selected on both
sides of the Ghunsa river valley and two transects
of 2.5 m × 25 m each were studied at each land-use
type on both sides of the valley, which showed southeast and north-west facing aspects. A total of 72 of
80 planned transects were established, because not all
land-use types were found at each elevation level. The
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distance between two transects within the same landuse type was at least ten meters.
On each transect, elevation was recorded by Global Positioning System (Garmin, GPSmap60CSx) and
slope, and the direction of the slope was recorded by
a clinometer (Silva, Ranger). The growth form and substrate types were recorded. We considered the growth
forms crustose, foliose, fruticose and leprose, and the
substrate types corticolous (on bark), saxicolous (on
rock), muscicolous (on moss) and terricolous (on soil)
(Hale 1983). Hemispherical photographs were taken
using a digital camera (Coolpix995 Nikon) and fish-eye
lens (Fish-eye converter FC-E8 Nikon). The camera was
mounted at a height of 1.5 m above the ground on a
tripod and levelled with a bubble level.
with downweighting of rare species and found a gradient length 8.01 standard unit (SD) for the first axis.
This indicated the use of Canonical Correspondence
Analysis (CCA) (Lepš & Šmilauer 2003) and its implied
unimodal response model over a linear model like in
Redundancy Analysis (RDA) to analyse the relationships between species co-occurrence and environmental variables (i. e., elevation, land-use type and canopy
openness). All environmental variables were permuted
199 times during CCA to test for significant environmental variables. Direct correlations of environmental
predictors with CCA axes were also performed.
All statistical analyses were performed using the
vegan 2.4-0 package (Oksanen et al. 2016) under the
free statistical software environment R version 3.3.1
(R Core Team 2016).
Lichen identification and image analysis
Collected lichen specimens were examined at the
Laboratory of the Central Department of Botany,
Tribhuvan University, Kathmandu, Nepal, and at the
Swiss Federal Research Institute, WSL, Switzerland.
Identification of lichens was carried using the relevant keys and checklists (Awasthi 1991; Sharma 1995;
Awasthi 2007; Singh & Sinha 2010). Identified specimens were deposited at the Swiss Federal Research
Institute WSL, Switzerland.
Lichen species were categorized according to family,
growth forms, substrate type and photobiont types, i. e.
cyanobacteria or green algae, following the recent updated taxonomical classification (Lücking et al. 2016).
Data were organized in a relational database (MS Access). Hemispherical photographs were converted to
binary (black and white pixels) following the image
analysis manual described by Frazer et al. (1999). All
image analyses were performed using image-processing software, Gap Light Analyzer (GLA Version 2.0).
Statistical analysis
We calculated Pearson correlation coefficients between variables such as total lichen species richness,
growth forms, substrate types and photobiont types
(i. e. green algal and cyanobacterial lichen species richness) and canopy openness. TukeyHSD multiple comparison tests were used to test the effect of particular
land-use types on species richness of lichens. Generalized Linear Models (GLMs; McCullagh & Nelder 1989)
with quasi-poisson error distribution were performed
for modelling lichen richness. We build models with
linear only and linear and quadric predictor terms and
chose the final model parameterization according to
the significance of the quadratic term. Graphics were
made only for statistically significant models by using
GLM. GLMs were not built for species richness of
leprose, terricolous and muscicolous lichens because
of the scarcity of occurrence data.
Detrended Correspondence Analysis (DCA; Hill
& Gauch 1980) was used to determine the lengths of
the main gradient in species composition based on the
sample by species data matrix. We performed DCA
Results
A total of 518 lichen specimens were collected
from 72 transects, which included 229 lichen species
of 71 genera (Appendix 1). 95 species belonged to the
foliose growth form, 87 species were crustose, 44 species fruticose and 3 species were leprose. With regard
to the substrate preference, 157 species were corticolous, 55 saxicolous, 14 muscicolous and 3 terricolous
species. Green algal photobionts were associated with
205 lichen species, while the remaining 24 lichen species were associated with cyanobacteria. A TukeyHSD
test showed significant differences in lichen species
richness between cultivated and other land-use types
(p < 0.05) (Appendix 2a).
Species richness between land-use types
According to land-use types, 174 species were recorded from exploited forests with the highest number
of foliose lichens (77 species), followed by 172 species
on natural forests, dominated again by foliose lichens
(70 species). Likewise, the highest number of corticolous species (151 species) was recorded from natural
forests followed by exploited forests with 135 species.
Species richness and canopy openness
Total species richness showed a significant monotonic decline with canopy openness (Figure 2a). Such a
monotonic decline of species richness was also found
for specific growth forms, specific photobiont species
richness and species richness of corticolous of specific substrate types (Figure 2, Appendix 3). As an exception, a significant monotonic increase was found
for saxicolous species richness towards higher canopy
openness (Figures 2e, Appendix 3). An optimum of
total lichen richness was found at low canopy openness
with 20.1 species predicted at 10 % canopy openness,
with a gradual decline towards higher canopy openness
(Figure 2a). Similarly, species numbers of crustose and
fruticose lichens showed a decline towards higher
canopy openness, with a predicted species number of
6.4 and 5.2 species at 10 % canopy openness respec-
Cho ng ba ng e t a l .
Figure 2 – Relationship between lichen species richness and canopy openness. a–g: a) total species richness; b & c) species richness of specific grwoth forms;
d & e) species richness of specific substrate types and
f & g) specific photobiont species richness. The fitted
regression lines represent Model 1 (Appendix 3).
tively (Figures 2b & c). Regression analysis was not
performed for the leprose growth form because only
three species presented this feature. Regarding the four
substrate categories, corticolous lichen richness also
showed a gradual decline with increasing canopy openness, with an average of 19.8 species at 10 % canopy
openness (Figure 2d). In contrast, saxicolous lichen
richness had a positive trend with increasing canopy
openness with an average of 7.4 species at 85% canopy openness (Figure 2e). GLM was not performed
for muscicolous and terricolous species as their number was too low (14 and 3 species respectively). With
respect to photobiont type, both cyanolichens and
green algal lichens exhibited a significant decrease with
canopy openness, with an average of 2.9 species of cy-
anolichens, 17.2 species of green algal lichens at 10 %
canopy openness (Figures 2f & g) respectively.
Species richness along elevation
There is a significant correlation of the total lichen
species richness with the elevation and canopy openness (p ≤ 0.05). Total species richness of lichens and
species richness of specific growth forms, specific
substrate types and specific photobiont types, except
species richness of leprose, muscicolous, terricolous
lichens, showed a significant (p ≤ 0.05) monotonic increase with elevation (Figures 3a-g, Appendix 3). A
total richness of 21.9 species was predicted at 3 800 m
with a predicted species richness of 6.3 crustose, 10.3
foliose, 5.2 fruticose, 16.5 corticolous and 2.4 cyanoli-
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Figure 3 – Relationship between lichen species richness and elevation. a–g: a) total species richness; b–d)
species richness of specific grwoth forms; e) species
richness of specific substrate types and f & g) specific
photobiont species richness. The fitted regression lines
represent Model 1 (Appendix 3).
chens and 19.5 green algal lichen species at 3 800 m
(Figures 3a–g). The regression analysis results showing
the best selected model for each response variable is
shown in Appendix 3.
Species composition
The length of the first DCA axis was 8.01 SD units
(Table 2) that indicated a high beta diversity with almost complete species turnover between transects.
The first two DCA axes explained 12.3 % of the total
variance in the data matrix.
In CCA, the environmental variables elevation,
canopy openness and land-use explained 21 % of the
total species variation variance (Table 3). CCA axis I
was significantly correlated with elevation, while CCA
axis II was highly correlated with canopy openness
and land-use types (Figure 4, Appendix 2b). Along
the CCA axis I, the highest abundance of Aspicilia
contorta, Chaenotheca chrysocephala, Evernia mesomorpha,
Leptogium burnetiae, Umbilicaria indica var. indica and Usnea longissima showed more preference towards high
elevation, while species such as Cladonia scabriuscula,
Heterodermia comosa, Lecanora cenisia showed high preference towards low elevation. Likewise, along the
CCA axis II, species composition of Aspicilia caesiocinerea, Coccocarpia erythroxyli, Phaeophyscia ciliata, Umbilicaria badia, Xanthoria fallax showed higher abundance
towards higher canopy openness, while species like
Caloplaca farinosa, Hypogymnia vittata, Cladonia crispata
var. cetrariiformis, Usnea himalayana, Chaenotheca chryso-
Cho ng ba ng e t a l .
Table 1 – Environmental correlation coefficient matrix (Pearson correlation) among variables used during the study (p ≤ 0.05).
elv = elevation, cano = canopy openness, spn = total species number, cru = crustose species number, fol = foliose species number,
fru = fruticose species number, lep = leprose species number, cort = corticolous species, musc = muscicolous species, saxi = saxicolous
species, terr = terricolous species, blgrn = blue green algae and grnal = green algae
Elev
Canopy Spn
Cru
Fol
Fru
Lep
Cort
Musc Saxi
Terr
Blgrn Grnal
0.00
1.00
0.06
0.85
0.06
0.47
1.00
0.26
0.65
1.00
1.00
0.81
0.07
Canopy 0.29
0.00
1.00
1.00
1.00
1.00
1.00
0.06
1.00
0.09
1.00
0.28
1.00
Spn
0.00
0.05
0.00
0.00
0.00
0.00
1.00
0.00
0.06
1.00
1.00
0.00
0.00
Cru
0.02
0.05
0.00
0.00
0.00
0.00
1.00
0.00
0.78
1.00
1.00
0.00
0.00
Fol
0.00
0.18
0.00
0.00
0.00
0.00
1.00
0.00
0.61
1.00
1.00
0.00
0.00
Fru
0.01
0.05
0.00
0.00
0.00
0.00
1.00
0.00
0.00
1.00
1.00
0.00
0.00
Lep
0.41
0.64
0.21
0.10
0.47
0.50
0.00
1.00
1.00
1.00
1.00
1.00
1.00
Cort
0.01
0.00
0.00
0.00
0.00
0.00
0.33
0.00
0.61
1.00
1.00
0.00
0.00
Musc
0.02
0.29
0.00
0.02
0.01
0.00
0.67
0.01
0.00
1.00
1.00
0.37
0.07
Saxi
0.27
0.00
0.16
0.33
0.04
0.84
0.32
0.55
0.36
0.00
1.00
1.00
1.00
Terr
0.68
0.09
0.58
0.31
0.97
0.38
0.60
0.44
0.83
0.34
0.00
1.00
1.00
Blgrn
0.02
0.01
0.00
0.00
0.00
0.00
0.93
0.00
0.01
0.56
0.51
0.00
0.00
Grnal
0.00
0.09
0.00
0.00
0.00
0.00
0.14
0.00
0.00
0.09
0.61
0.00
0.00
Elev
Table 3 – DCA analysis summary of the study site.
Table 2 – DCA summary of the study site.
DCA Axis
I
Eigenvalues
0.63
II
0.50
III
0.40
IV
0.33
Axis lengths
8.01
6.30
4.28
3.77
Total inertia
9.16
Cumulative % vari6.87 12.32 16.71 20.30
ance of species data
cephala showed a higher abundance towards low canopy openness (Figure 4).
Similarly, species like Phaeophyscia endococcina, Lecanora polytropa, Umbilicaria badia, Parmotrema subarnoldii showed high abundance towards open habitats
and cultivated land, while species like Aspicilia cinerea,
Hypotrachyna scytophylla, Parmelina quercina, Heterodermia
obscurata, Stereocaulon paradoxum, Rhizoplaca chrysoleuca
showed high abundance in meadows. Likewise, Hypotrachyna cirrhata, Hypotrachyna nepalensis, Cetrelia cetrarioides, Parmotrema pseudonilgherrense, Usnea compressula
showed high abundance towards exploited forest and
species like Lobaria retigera, Cladonia crispata var. cetrariiformis, Hypogymnia vittata, Caloplaca farinosa, Nephromopsis ahtii showed high abundance towards natural forest
landscapes (Figure 4).
Inertia Proportion Rank
Total
9.15
1.00
Constrained
1.92
0.21
6
Unconstrained
7.23
0.79
29
Our study revealed a monotonic decrease in total lichen species richness with increasing canopy openness.
In the present study, low canopy openness of about
10 % still supported a high number of lichen species.
In the steep Himalayan mixed forests, canopy openness of 10 % and more is likely to provide sufficient
light into the forest stand and, in general, light limitation does not seem to be a major limiting factor for lichen species richness in the mountain forests of KCA.
Discussion
Lichen species richness and composition along
gradients of land use, canopy openness and
elevation
Our study indicated distinct effects of elevation,
land use and canopy openness on lichen species richness and composition. We found a considerable variation in lichen species richness among the four selected
land-use types, with decline of species richness from
forest to cultivated land. These findings are in accordance with other studies like Stofer et al. (2006), who
also observed decreasing lichen species richness from
natural forest landscape to open agricultural landscape
in a large-scale study covering several European biogeographic zones.
Figure 4 – Canonical Correspondence Analysis (CCA) of lichen composition constraint by elevation, canopy openness and
land-use types; C = Cultivated, M = Meadow, E = Exploited, and N = Natural. Arrow indicates the direction of increasing values and their length is proportional to the correlation
between the variable and the plot scores (not shown) on the two
ordination axes. Land-use types are shown as centroids. For full
names of species see Appendix 1.
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As trees are an important factor explaining lichen
species composition and richness (Mežaka et al. 2008;
Odor et al. 2013), meadows and natural forests seemed
to provide lichen-rich habitats because of a high diversity and abundance of trees. The exploited forest
type with varying disturbance intensity still maintained
a reasonable diversity of microhabitats for epiphytic
lichens, but some species that depend on semi-shaded
habitats and high moisture in natural forests, such as
corticolous lichens, are declining in exploited forests.
Pinokiyo et al. (2008) also found the maximum number of corticolous lichens in dense forest. In the present study, we found high saxicolous lichen richness
in meadows, because a high abundance of rocks and
boulders are exposed on meadows, where litter does
not continuously cover their surface. Exposed rock
surfaces can support more saxicolous lichens than in
closed forests, where slightly inclined rock surfaces are
often covered with litter. On cultivated lands, slightly
inclined rock surfaces are often disturbed by human
influence to remove them or to use them for various
activities related to farming. Frequent and intense
disturbance of rock surfaces in agricultural land is a
significant difference to European land-use gradients,
where Wolseley et al. (2006) recorded high saxicolous
richness in farmland including cultivated land.
The saxicolous species richness revealed a gradual
increase of species richness with increasing canopy
openness and reached an average of 7.4 species per
transect at 85 % openness, which corresponds to
meadows and open cultivated land. Rocks and boulders inside forest landscapes are primarily covered
by litter or mosses and also have a low exposition to
solar radiation. However, corticolous species richness showed a decline with increasing openness and
reached an average of 19.8 species at 10 % openness.
Because corticolous lichens in the studied land-use
gradients form a more species-rich species pool than
saxicolous species, the observed decline of saxicolous
species richness is overcompensated by a stronger increase of epiphytic lichens. As the lichen diversity is
related to tree diversity, density (Baniya et al. 1999; Li
et al. 2011; Li et al. 2013b, 2013a) and humidity (Pinokiyo et al. 2008), cultivated landscapes bear a limited
number of trees, shrubs and fewer rocks and boulders
as well as less humidity. The resulting lower epiphytic
lichen species richness cannot be compensated by an
increased density of boulders and bare rocks, which
are generally covered with lichen vegetation under an
open sky receiving direct solar radiation.
The lichen richness pattern is also closely related to
the management practices of the particular landscape,
e. g. the protected area. The traditional shifting cultivation practice common to this area (Aryal et al. 2010)
is significantly explained after finding of declining lichen richness pattern with open canopy. The shifting
cultivation practice opens up a landscape which seems
not to support lichen richness and its diversity pattern. Further, the shifting cultivation practice is also
common to Makalu-Barun areas of East Nepal. Thus,
future diversity of lichen seems in a difficult situation.
Conservation of lichen will automatically conserve the
landscape.
In addition to the differences between land-use
types, our study clearly indicated a distinct variation in
species richness along the elevational gradient studied.
We found a linear relationship with increasing elevation. Cobanoglu and Sevgi (2009) reported a similar
pattern for epiphytic lichens with elevations from
1 300 m to 1 900 m in Turkey. However, a majority
of former studies reported an unimodal relationship
(Bruun et al. 2006; Grytnes et al. 2006; Baniya et al.
2010, 2012). Unlike these studies, which generally covered long elevational gradients, our study was more
closely confined to a local scale, with an elevational
gradient covering temperate to subalpine forests, but
not reaching areas above the timberline. Therefore our
linear relationship can be interpreted as part of an unimodal relationship on larger scales.
Lichen species composition showed a strong species
turnover along CCA axis I (elevation) and CCA axis II
(land use-types). Natural and exploited forests supported diverse lichen vegetation which decreased towards
meadows and cultivated land. These results confirm
findings from European land-use gradients from forest
to agricultural land-use types (Stofer et al. 2006).
Conclusion
We conclude that besides elevation as a general
climate proxy, differences in land use, which directly
affect canopy openness, are the two main general factors of both lichen species richness and composition
in this area of the Himalayas in Nepal. Forests with
diverse habitats and relatively low canopy openness
harbour more lichen species than meadows and cultivated land. However, elevation and canopy openness
are not direct drivers. Canopy openness influences
light intensity and relative moisture on the forest floor
and tree trunks, which directly affect lichen diversity.
In addition, elevation serves as a general climate proxy
for temperature or precipitation, which more directly
influences both species richness and composition of
lichen communities. Highest species richness of lichens was reached at the highest altitudinal level of
our survey, indicating that the maximum total species
richness of lichens as well as the richness of most of
the studied species groups is at or above 3 800 m in
this part of the Himalayas.
Acknowledgement
We thank the Swiss National Science Foundation
(JRP IZ70Z0_131338 to CS, MN and K. K. Shrestha) for financial support. We are also grateful to all
members of the CDB-WSL Project, Prof. Dr. Krishna
Kumar Shrestha, Prof. Dr. Khadga Basnet, Dr. Jyoti
Pd. Gajurel, Mr. Shiva Devkota, Mr. Sanjeev Kumar
Cho ng ba ng e t a l .
Rai, Mr. Hem Bdr. Katuwal, Mr. Yam Aryal and Mr.
Rajesh Tamang for their kind support. We thank all local people of Ghunsa, KCA, and Utra Kumar Rai for
his help with collecting lichens in the field.
References
Anonymous 2009. Kanchenjunga conservation area.
Kanchenjunga conservation area management council.
Anonymous 2011. Kanchenjunga conservation area.
Kanchenjunga conservation area management council.
Aryal, K.P., E.E. Kerkhoffn, N. Maskey & R. Sherchan 2010. Shifting Cultivation in the Sacred Himalayan
Landscape: A case study in the Kanchenjunga Conservation
Area. WWF Nepal.
Awasthi, D.D. 1991. A key to the microlichens of
India, Nepal and Sri-Lanka. Bibliotheca Lichenologica 40:
1–137.
Awasthi, D.D. 2007. A compendium of the macrolichens
from India, Nepal and Sri-Lanka. India.
Baniya, C.B., G.P.S. Ghimire & B. Kattel 1999. Diversity of lichens in Nepal. Banko Jankari 9: 26–28.
Baniya, C.B., T. Solhøy, Y. Gauslaa & M.W. Palmer
2010. The elevation gradient of lichen species richness
in Nepal. The Lichenologist 42: 83–96.
Baniya, C.B., T. Solhøy, Y. Gauslaa & M.W. Palmer
2012. Richness and composition of vascular plants
and cryptogams along a high elevational gradient on
Buddha Mountain, Central Tibet. Folia Geobotanica 47:
135–151.
Bergamini, A., C. Scheidegger, S. Stofer, P. Carvalho, S. Davey, M. Dietrich, F. Dubs, E. Farkas, U.
Groner, & K. Kaerkkaeinen 2005. Performance of
macrolichens and lichen genera as indicators of lichen
species richness and composition. Conservation Biology
19: 1051–1062.
Bruun, H.H., J. Moen, R. Virtanen, J.A. Grytnes, L.
Oksanen & A. Angerbjörn 2006. Effects of altitude
and topography on species richness of vascular plants,
bryophytes and lichens in alpine communities. Journal
of Vegetation Science 17: 37–46.
Chaudhary, R.P. 1998. Biodiversity in Nepal: Status and
conservation: Bangkok, Thailand.
Cobanoglu, G. & O. Sevgi 2009. Analysis of the
distribution of epiphytic lichens on Cedrus libani in
Elmali Research Forest (Antalya, Turkey). Journal of
Environmental Biology 30: 205–212.
Frazer, G.W., C.D. Canham & K.P. Lertzman 1999.
Gap Light Analyzer (GLA), Version 2.0: Imaging software
to extract canopy structure and gap light transmission indices
from true-colour fisheye photographs, user’s manual and program
documentation. Simon Fraser University, Burnaby, British Columbia, and the Institute of Ecosystem Studies,
Millbrook, New York. (http://www.ecostudies.org)
Gautam, C.M. & T. Watanabe 2004. Reliability of
Land Use/Land Cover Assessment in Montane Nepal: A Case Study in the Kangchenjunga Conservation Area (KCA). Mountain Research and Development 24:
35–43.
Giordani, P., G. Incerti, G. Rizzi, F. Ginaldi, S. Viglione, I. Rellini, G. Brunialti, P. Malaspina & P. Modenesi 2010. Land use intensity drive the local variation of
lichen diversity in Mediterranean ecosystems sensitive
to desertification. Bibliotheca Lichenologica 105: 139–148.
Grytnes, J.A., E. Heegaard & P.G. Ihlen 2006. Species richness of vascular plants, bryophytes, and lichens along an altitudinal gradient in western Norway.
Acta oecologica 29: 241–246.
Hale, M.E. 1983. The biology of lichens. London.
Hill, M.O. & H.G. Gauch 1980. Detrended Corrrespondence Analysis: An Improved Ordination Technique. Vegatatio 42: 47–58.
Li, S., W.Y. Liu & D.W. Li 2013a. Bole epiphytic lichens as potential indicators of environmental change
in subtropical forest ecosystems in southwest China.
Ecological Indicators 29: 93–104.
Li, S., W.Y. Liu & D.W. Li 2013b. Epiphytic lichens
in subtropical forest ecosystems in southwest China:
Species diversity and implications for conservation.
Biological Conservation 159: 88–95.
Li, S., W. Liu, L. Wang, W. Ma & L. Song 2011.
Biomass, diversity and composition of epiphytic macrolichens in primary and secondary forests in the subtropical Ailao Mountains, SW China. Forest Ecology and
Management 261: 1760–1770.
Löbel, S., T. Snäll & H. Rydin 2006. Species richness patterns and metapopulation processes–evidence
from epiphyte communities in boreo nemoral forests.
Ecography 29: 169–182.
Lücking, R., B.P. Hodkinson & S.D. Leavitt 2016.
The 2016 classification of lichenized fungi in the Ascomycota and Basidiomycota-Approaching one thousand genera. The Bryologist 119: 361–416.
Marmor, L., T. Tõrra, L. Saag & T. Randlane 2012.
Species richness of epiphytic lichens in coniferous
forests: the effect of canopy openness. Annales Botanici
Fennici 49: 352–358.
McCullagh, P. & J.A. Nelder 1989. Generalized Linear
Models (2nd ed.). London.
Mežaka, A., G. Brūmelis & A. Piterāns 2008. The
distribution of epiphytic bryophyte and lichen species
in relation to phorophyte characters in Latvian natural
old-growth broad leaved forests. Folia Cryptogamica Estonica 44: 89–99.
Mežaka, A., G. Brūmelis & A. Piterāns 2012. Tree
and stand-scale factors affecting richness and composition of epiphytic bryophytes and lichens in deciduous woodland key habitats. Biodiversity and Conservation
21: 3221–3241.
MFSC 2009. Nepal Fourth National Report to the Convention of Biological Diversity. Singha Durbar, Kathmandu, Nepal: Ministry of Forests and Soil Conservation,
Government of Nepal.
Motiejûnaitë, J. & W. Faùtynowicz 2005. Effect of
land-use on lichen diversity in the transboundary region of Lithuania and northeastern Poland. Ekologija
3: 34–43.
53
54
R esearch
Odor, P., L. Kiraly, F. Tinya, F. Bortignon & J. Nascimbene 2013. Patterns and drivers of species composition of epiphytic bryophytes and lichens in managed
temperate forests. Forest Ecology and Management 306:
256–265.
Oksanen, J., F.G. Blanchet, M. Friendly, R. Kindt,
P. Legendre, D. McGlinn, P.R. Minchin, R.B. O’Hara,
G.L. Simpson, P. Solymos, M. H.H. Stevens, E. Szoecs
& H. Wagner. 2016. vegan: Community Ecology Package. R package version 2.4-0. http://CRAN.R-project.
org/package=vegan
Pinokiyo, A., K.P. Singh & J.S. Singh 2008. Diversity and distribution of lichens in relation to altitude
within a protected biodiversity hot spot, north-east
India. The Lichenologist 40: 47–62.
R Core Team 2016. R: A language and environment
for statistical computing. R Foundation for Statistical
Computing, Vienna, Austria. URL http://www.Rproject.org/.
Rai, H., R. Khare, R.K. Gupta, & D.K. Upreti 2011.
Terricolous lichens as indicator of anthropogenic
disturbances in a high altitude grassland in Garhwal
(Western Himalaya), India. Botanica Orientalis: Journal of
Plant Science 8: 16-23.
Scheidegger, C., M.P. Nobis & K.K. Shrestha 2010.
Biodiversity and livelihood in land-use gradients in an
era of climate change-outline of a Nepal-Swiss research project. Botanica Orientalis: Journal of Plant Science
7: 7–17.
Scheidegger, C. & S. Werth 2009. Conservation
strategies for lichens: insights from population biology. Fungal Biology Reviews 23: 55–66.
Sharma, L.R. 1995. Enumerations of the Lichens of Nepal. Biodiversiy Profile Project. Tec. Pub. No.3, DNPWC, Kathmandu, Nepal.
Shrestha, K.K., & S.K. Ghimire 1996. Plant diversity
inventory of poposed Kanchenjunga Conservation Area (Ghunsa and Simbua Valleys). Kathmandu, Nepal: WWF
Nepal Progaram Report series 22.
Singh, K.P. & G.P. Sinha 2010. Indian lichens: an annotated checklist. Botanical Survey of India, Ministry of
Environment and forests.
Stofer, S., A. Bergamini, G. Aragon, P. Carvalho,
B.J. Coppins, S. Davey, M. Dietrich, E. Farkas, K.
Karkkainen, C. Keller, L. Lokos, S. Lommi, C. Maguas,
R. Mitchell, P. Pinho, V.J. Rico, A.M. Truscott, P.A.
Wolseley, A. Watt & C. Scheidegger 2006. Species richness of lichen functional groups in relation to land use
intensity. The Lichenologist 38: 331–353.
Tasser, E. & U. Tappeiner 2002. Impact of land use
changes on mountain vegetation. Applied vegetation science 5: 173–184.
Wolseley, P.A., S. Stofer, R. Mitchell, A.M. Truscott,
A. Vanbergen, J. Chimonides & C. Scheidegger 2006.
Variation of lichen communities with landuse in Aberdeenshire, UK. Lichenologist 38: 307–322.
Authors
Til Bikram Chongbang – corresponding author
holds an M.Sc. in Botany from Tribhuvan University,
Kathmandu, Nepal. Now a member of the biology
teaching staff at the National School of Sciences,
Lainchour, Kathmandu, he is particularly interested
in lichens and ethnobotany. E-mail: tbchongbang@
yahoo.com
Christine Keller 1
is an expert in lichen taxonomy and ecology. She is
a member of the scientific staff at the Swiss Federal
Institute for Forest, Snow and Landscape Research
WSL, Birmensdorf. E-mail: christine.keller@wsl.ch
Michael Nobis 1
is a botanist and macroecologist with a research
focus on trait-environment relationships, species distribution modelling, species migration and biological
invasions. He is a member of the scientific staff at
the Swiss Federal Institute for Forest, Snow and Landscape Research WSL. E-mail: michael.nobis@wsl.ch
Christoph Scheidegger 1
focuses in his research on the conservation
biology of lichens and plants, population genetics
and landscape ecology, and biodiversity assessment
and evaluation at the landscape level. He recently
coordinated a research project in Nepal. He is a senior
scientist and group leader at the Swiss Federal Institute
for Forest, Snow and Landscape Research WSL.
E-mail: christoph.scheidegger@wsl.ch
Chitra Bahadur Baniya
is Associate Professor of Ecology and Resource
Management at the Central Department of Botany,
Tribhuvan University, Kathmandu, Nepal. His major field of interest is quantitative ecology of plants
and lichens. His present teaching and research mainly
focus on open-source programs, such as R statistical
packages and science in philosophy. E-mail: cbbaniya@
gmail.com
1
Swiss Federal Institute for Forest, Snow and
Landscape Research WSL, Zürcherstrasse 111, CH8903 Birmensdorf.
Cho ng ba ng e t a l .
Appendices
Appendix 1 – List of lichens, their family, growth forms, substrate groups, photobiont partner and frequency of occurrence along
land use types in the study area. Cru – crustose, Fol – foliose, Fru – fruticose, Lep – leprose, Cort – corticolous, Musc – muscicolous,
Saxi – saxicolous, Terr – terricolous, BGA – blue green alga, GA – green alga, C – cultivated land, M – meadow, E – exploited
forest, F – natural forest.
S.N. Name of Lichen species
Short
form
Family
Growth
form
Substrate Photobiont Frequency Land use
group
partner
(Number) types
1
Amandinea punctata (Hoffm.) Coppins & Scheid.
Ama pun Caliciaceae
Cru
Cort
GA
2
F
2
Aspicilia caesiocinerea (Nyl.ex Malbr.) Arnold
Asp cae
Cru
Saxi
GA
12
C, E, M, F
Megasporaceae
3
Aspicilia cinerea (L.) Körb.
Asp cin
Cru
Saxi
GA
4
C, E, F, M
4
Aspicilia contorta (Hoffm.) Körb.
Asp con
Cru
Saxi
GA
2
E, F
5
Aspicilia griseocinerea Räsänen
Asp gri
6
Bacidia laurocerasi (Delise ex Duby) Zahlbr.
Bac lau
7
Bacidia rubella (Hoffm.) A. Massal.
Bac rub
8
Bryoria himalayensis (Motyka) Brodo & D. Hawksw.
Bry him
Cru
Cort
GA
1
C
Ramalinaceae
Cru
Cort
GA
1
F
Cru
Cort
GA
2
F, E
Parmeliaceae
Fru
Cort
GA
1
F
9
Bryoria lactinea (Nyl.) Brodo & D. Hawksw.
Bry lac
Fru
Cort
GA
1
E
10
Bryoria smithii (Du Rietz) Brodo & D. Hawksw.
Bry smi
Fru
Cort
GA
3
M, F, E
11
Bryoria tenuis (Dahl) Brodo & D. Hawksw.
Bry ten
12
Buellia aethalea (Ach.) Th. Fr.
Bue aet
Caliciaceae
Fru
Cort
GA
5
E, M, F
Cru
Saxi
GA
1
F
13
Buellia inornata Zahlbr.
Bue ino
Cru
Cort
GA
1
E
14
Buellia montana H. Magn.
Bue mon
Cru
Cort
GA
2
M
15
Calicium subquercinum Asah.
Cal sub
16
Caloplaca chlorina (Flot.) Sandst.
Cal chl
Teloschistaceae
Cru
Cort
GA
4
F, E
Cru
Cort
GA
1
M
17
Caloplaca citrina (Hoffm.) Th. Fr.
Cal cit
Cru
Cort
GA
1
F
18
Caloplaca encephalarti (Kremp.) Zahlbr.
Cal enc
Cru
Cort
GA
1
E
19
Caloplaca farinosa Poelt & Hinter.
Cal far
Cru
Cort
GA
3
F
20
Caloplaca holocarpa (Hoffm.) Wade
Cal hol
Cru
Cort
GA
1
F
21
Caloplaca holochracea (Nyl.) Zahlbr.
Cal hol
Cru
Saxi
GA
1
M
22
Caloplaca isabellina Poelt & Hinter.
Cal isa
Cru
Saxi
GA
5
M, C, E
23
Candelaria indica (Hue) Vain.
Can ind
24
Candelariella vitellina (Hoffm.) Müll. Arg.
Can vit
25
Candelariella xanthostigma (Pers. ex Ach.) Lettau
Can xan
26
Cetrelia braunsiana (Müll.) W. Culb. & C. Culb.
Cet bra
Candelariaceae
Parmeliaceae
Fol
Saxi
GA
3
C, E, M
Cru
Saxi
GA
1
M
Cru
Cort
GA
2
E, M
Fol
Cort
GA
4
E, F, M
27
Cetrelia cetrarioides (Delise) W. Culb. & C. Culb.
Cet cet
Fol
Cort
GA
10
E, F, M
28
Cetrelia olivetorum (Nyl.) W. Culb. & C. Culb.
Cet oli
Fol
Cort
GA
1
E
29
Cetrelia pseudolivetorum (Asahina) W. Culb. & C. Culb.
Cet pse
30
Chaenotheca chrysocephala (Ach.) Th. Fr.
Cha chr
Coniocybaceae
Fol
Cort
GA
2
E, F
Cru
Cort
GA
3
F
31
Chaenotheca phaeocephala (Turner) Th. Fr.
Cha pha
Cru
Cort
GA
1
E
32
Chaenotheca trichialis (Ach.) Hellb.
Cha tri
Cru
Cort
GA
1
F
33
Chrysothrix candelaris (L.) Laundon
Chr can
34
Chrysothrix chlorina (Ach.) Laundon
Chr chl
35
Chrysothrix xanthina (Vain.) Kalb
Chr xan
36
Cladoina chlorophaea (Flörke ex Sommerf.) Spreng.
Cla chl
Chrysotricaceae
Cladoniaceae
Lep
Saxi
GA
1
M
Lep
Cort
GA
1
E
Lep
Cort
GA
1
F
Fru
Musc
GA
1
F
37
Cladoina corniculata Ahti & Kashiw.
Cla cor
Fru
Musc
GA
1
C
38
Cladonia coccifera (L.) Willd.
Cla coc
Fru
Musc
GA
6
E, M
39
Cladonia coniocraea (Flörke) Spreng.
Cla con
Fru
Musc
GA
4
E, F
40
Cladonia corymbescens Nyl. ex Leight.
Cla cor
Fru
Musc
GA
4
M, E
E, F
41
Cladonia crispata var. cetrariiformis (Delise) Vain.
Cla cri
Fru
Cort
GA
3
42
Cladonia fimbriata (L.) Fr.
Cla fim
Fru
Musc
GA
1
E
43
Cladonia furcata (Huds.) Schrad.
Cla fur
Fru
Cort
GA
4
E, F, M
44
Cladonia macilenta Hoffm.
Cla mac
Fru
Terr
GA
1
M
45
Cladonia macroptera Räsänen
Cla mac
Fru
Saxi
GA
1
E
46
Cladonia ramulosa (With.) Laundon
Cla ram
Fru
Musc
GA
1
M
47
Cladonia scabriuscula (Delise) Nyl.
Cla sca
Fru
Saxi
GA
2
E, M
48
Cladonia stellaris (Opiz) Pouzar & Vězda
Cla ste
Fru
Terr
GA
1
E
49
Cladonia subconistea Asahina
Cla sub
Fru
Cort
GA
2
E
50
Cladonia subsquamosa Kremp.
Cla sub
Fru
Cort
GA
1
E
51
Cladonia subulata (L.) F.H. Wigg.
Cla sub
Fru
Cort
GA
1
M
52
Cladonia verticillata (Hoffm.) Schaer.
Cla ver
53
Coccocarpia erythroxyli (Spreng.) Swinsc. & Krog
Coc ery
Coccocarpiaceae
Fru
Musc
GA
1
C
Fol
Saxi
BGA
2
M, C
54
Collema subconveniens Nyl.
Col sub
Collemataceae
Fol
Cort
BGA
2
F
55
Dibaeis baeomyces (L. f.) Rambold & Hertel
Dib bae
Icmadophilaceae
Cru
Saxi
GA
1
E
56
Coenogonium luteum (Dicks.) Kalb & Lücking
Coe lut
Coenogoniaceae
Cru
Cort
GA
3
F, E
57
Diploschistes scruposus (Schreb.) Norman
Dip scr
Ghraphidaceae
Cru
Saxi
GA
1
M
Caliciaceae
58
Diplotomma alboatrum (Hoffm.) Flot.
Dip alb
59
Diplotomma himalayense S. Singh & D.D. Awasthi
Dip him
60
Diplotomma proximatum (Magn.) S. Singh & D.D. Awasthi
Dip pro
61
Erioderma meiocarpum Nyl.
Eri mei
Pannariaceae
Parmeliaceae
Cru
Cort
GA
1
E
Cru
Cort
GA
3
E, M
Cru
Cort
GA
3
E, F
Fol
Cort
BGA
3
F, M
62
Evernia mesomorpha Nyl.
Eve mes
Fru
Cort
GA
5
F, M, E
63
Hypotrachyna cirrhata (Fr.) Divakar, A. Crespo, Sipman, Elix
& Lumbsch
Hyp cir
Fol
Cort
GA
5
E, M, F
64
Hypotrachyna nepalensis (Taylor) Divakar, A. Crespo, Sipman,
Elix & Lumbsch
Hyp nep
Fol
Cort
GA
6
F, E, M
65
Flavoparmelia caperata (L.) Hale
Fla cap
Fol
Saxi
GA
1
M
55
56
R esearch
S.N. Name of Lichen species
Short
form
Family
Graphidaceae
66
Glyphis cicatricosa Ach.
Gly cic
67
Graphis nigroglauca Leight.
Gra nig
Growth
form
Substrate Photobiont Frequency Land use
group
partner
(Number) types
Cru
Cort
GA
1
F
Cru
Cort
GA
1
F
68
Graphis pyrrhocheiloides Zahlbr.
Gra pyr
Cru
Cort
GA
3
E, F
69
Graphis rimulosa (Mont.) Trevis.
Gra rim
Cru
Cort
GA
1
F
70
Graphis scripta (L.) Ach.
Gra scr
Cru
Cort
GA
4
E, F
71
Graphis sikkimensis Nagarkar & Patw.
Gra sik
Cru
Cort
GA
5
F, E
72
Graphis sorediosa Nagarkar & Patw.
Gra sor
Cru
Cort
GA
1
F
73
Haematomma puniceum (Sm. ex Ach.) Massal.
Hae pun
Haematommataceae Cru
Cort
GA
6
F, M, E
Physciaceae
74
Heterodermia angustiloba (Müll. Arg.) D.D. Awasthi
Het ang
75
Heterodermia boryi (Fée) Kr.P. Singh & S.R. Singh
Het bor
Fol
Cort
GA
3
E, F
Fol
Cort
GA
6
E, M, C, F
76
Heterodermia comosa (Eschw.) Follman & Redon
Het com
Fol
Cort
GA
2
M, E
77
Heterodermia diademata (Taylor) D.D. Awasthi
Het dia
Fol
Cort
GA
1
F
78
Heterodermia firmula (Nyl.) Trevis.
Het fir
Fol
Cort
GA
1
E
79
Heterodermia incana (Stirt.) D.D. Awasthi
Het inc
Fol
Cort
GA
1
E
80
Heterodermia obscurata (Nyl.) Trevis.
Het obs
Fol
Saxi
GA
3
M, E
81
Heterodermia pellucida (D.D. Awasthi) D.D. Awasthi
Het pel
Fol
Cort
GA
1
E
82
Heterodermia pseudospeciosa (Kurok.) W. Culb.
Het pse
Fol
Cort
GA
1
E
83
Heterodermia rubescens (Räsänen) D.D. Awasthi
Het rub
Fol
Cort
GA
2
E, F
84
Heterodermia speciosa (Wulf.) Trevis.
Het spe
Fol
Cort
GA
3
E
85
Heterodermia togashii (Kurok.) D.D. Awasthi
Het tog
Fol
Cort
GA
6
E, M, F
86
Heterodermia tremulans (Müll. Arg.) W. Culb.
Het tre
Fol
Cort
GA
1
M
87
Heterodermia verrucifera (Kurok.) W.A. Weber
Het ver
Fol
Cort
GA
1
F
88
Hypogymnia hypotrypa (Nyl.) Rass.
Hyp hyp
89
Hypogymnia vittata (Ach.) Gasil.
Hyp vit
Parmeliaceae
Fol
Cort
GA
4
F, E
Fol
Cort
GA
2
F, E
90
Hypotrachyna crenata (Kurok.) Hale
Hyp cre
Fol
Saxi
GA
1
E
91
Hypotrachyna exsecta (Taylor) Hale
Hyp exs
Fol
Cort
GA
1
E
92
Hypotrachyna infirma (kurok.) Hale
Hyp inf
Fol
Cort
GA
1
F
93
Hypotrachyna majoris (Vain.) Hale
Hyp maj
Fol
Cort
GA
1
M
94
Hypotrachyna revoluta (Flörke) Hale
Hyp rev
Fol
Cort
GA
1
M
95
Hypotrachyna scytophylla (Kurok.) Hale
Hyp scy
Fol
Saxi
GA
4
M, C, E
96
Hypotrachyna sinuosa (Sm.) Hale
Hyp sin
Fol
Saxi
GA
4
C, M, E
97
Hypotrachyna sublaevigata (Nyl.) Hale
Hyp sub
Fol
Cort
GA
1
C
98
Lasallia freyana D.D. Awasthi
Las fre
Umbilicariaceae
Fol
Saxi
GA
1
M
99
Lecanora frustulosa (Dicks.) Ach.
Lec fru
Lecanoraceae
Cru
Saxi
GA
1
C
100 Lecanora albella (Pers.) Ach.
Lec alb
Cru
Cort
GA
3
M, E
101 Lecanora allophana (Ach.) Nyl.
Lec all
Cru
Cort
GA
1
E
102 Lecanora campestris (Schaer.) Hue
Lec cam
Cru
Saxi
GA
2
E, F
103 Lecanora cenisia Ach.
Lec cen
Cru
Saxi
GA
4
M, F, C
104 Lecanora chlarotera Nyl.
Lec chl
Cru
Cort
GA
8
F, E ,M
105 Lecanora intricata (Ach.) Ach.
Lec int
Cru
Saxi
GA
1
M
C, M
106 Lecanora polytropa (Ehrh.) Rabenh.
Lec pol
Cru
Saxi
GA
2
107 Lecanora rugosella Zahlbr.
Lec rug
Cru
Cort
GA
4
F, E
108 Lecanora saligna (Schrad.) Zahlbr.
Lec sal
Cru
Cort
GA
1
F
109 Lecanora strobilina Ach.
Lec str
Cru
Cort
GA
1
M
110 Lecanora varia (Hoffm.) Ach.
Lec var
Cru
Cort
GA
1
M
111 Lecidea betulicola (Kullh.) H. Magn.
Lec bet
112 Lecidea erythrophaea Flörke ex Sommerf
Lec ery
Lecideaceae
Cru
Cort
GA
1
F
Cru
Cort
GA
1
F
113 Lecidea fuscoatra (L.) Ach.
Lec fus
Cru
Saxi
GA
1
M
114 Lecidea vorticosa (Flörke) Körb.
Lec vor
Cru
Saxi
GA
1
M
115 Lecidella elaeochroma (Ach.) M. Choisy
Lec ela
Lecanoraceae
Cru
Cort
GA
1
M
116 Lepraria crassissima (Hue) Lettau
Lep cra
Stereocaulaceae
Cru
Saxi
GA
1
M
117 Lepraria ecorticata (J.R. Laundon) Kukwa
Lep eco
Cru
Saxi
GA
1
M
118 Lepraria membranacea (Dicks.) Vain.
Lep mem
Cru
Cort
GA
2
M, E
119 Leptogium askotense D.D. Awasthi
Lep ask
120 Leptogium burnetiae Dodge
Lep bur
Collemataceae
Fol
Cort
BGA
1
E
Fol
Cort
BGA
3
F, E, M
121 Leptogium chloromelum (Sw.) Nyl.
Lep chl
Fol
Cort
BGA
1
F
122 Leptogium cyanescens (Rabenh.) Körb.
Lep cya
Fol
Cort
BGA
1
F
123 Leptogium pedicellatum P.M. Jørg.
Lep ped
Fol
Cort
BGA
7
E, F, M
124 Leptogium saturninum (Dicks.) Nyl.
Lep sat
Fol
Cort
BGA
1
M
125 Lethariella cladonioides (Nyl.) krog
Let cla
Parmeliaceae
Fru
Cort
GA
1
M
126 Lobaria isidiosa (Müll. Arg.) Vain.
Lob isi
Lobariaceae
Fol
Cort
BGA
1
F
127 Lobaria pindarensis Räsänen
Lob pin
Fol
Cort
BGA
3
F
128 Lobaria retigera (Bory) Trev.
Lob ret
Fol
Cort
BGA
5
E, F
129 Melanelia panniformis (Nyl.) Essl.
Mel pan
130 Melanelia tominii (Oxner) Essl.
Mel tom
Parmeliaceae
Fol
Cort
GA
1
M
Fol
Saxi
GA
2
M
131 Menegazzia terebrata (Hoffm.) A. Massal.
Men ter
Fol
Cort
GA
6
E, F
132 Mycobilimbia hunana (Zahlbr.) D.D. Awasthi
Myc hum Lecideaceae
Cru
Terr
GA
1
C
133 Mycoblastus affinis (Schaer.) T. Schauer
Myc aff
Tephromelatacae
Cru
Cort
GA
2
F
134 Myelochroa subaurulenta (Nyl.) Elix & Hale
Mye sub
Parmeliaceae
Fol
Cort
GA
1
F
135 Nephroma isidiosum (Nyl.) Gyeln.
Nep isi
Nephromataceae
136 Nephroma nakaoi Asahina
Nep nak
Fol
Musc
BGA
1
M
Fol
Cort
BGA
4
F, E, M
137 Nephromopsis nephromoides (Nyl.) Ahti & Randl.
Nep nep
Parmeliaceae
Fol
Cort
GA
1
E
138 Ochrolechia androgyna (Hoffm.) Arnold
Och and
Ochrolechiaceae
Cru
Saxi
GA
1
F
139 Ochrolechia parellula (Müll. Arg.) Zahlbr.
Och par
Cru
Saxi
GA
1
F
140 Ochrolechia rosella (Müll. Arg.) Vers.
Och ros
Cru
Cort
GA
8
E, F, M
Cho ng ba ng e t a l .
S.N. Name of Lichen species
Short
form
Family
141 Parmotrema thomsonii (Stirt.) A. Crespo, Divakar & Elix
Par tho
Parmeliaceae
142 Parmelia squarrosa Hale
Par squ
Growth
form
Substrate Photobiont Frequency Land use
group
partner
(Number) types
Fol
Cort
GA
1
E
Fol
Cort
GA
2
M
143 Parmeliella cinerata Zahlbr.) P.M. Jørg.
Par cin
Fol
Cort
BGA
1
E
144 Parmelina quercina (Willd.) Hale
Par que
Fol
Cort
GA
4
F, C, E, M
145 Parmotrema cetratum (Ach.) Hale
Par cet
Fol
Cort
GA
2
E, F
146 Parmotrema latissimum (Fée) Hale
Par lat
Fol
Cort
GA
1
F
147 Parmotrema nilgherrense (Nyl.) Hale
Par nil
Fol
Cort
GA
5
E, F, M
148 Parmotrema praesorediosum (Nyl.) Hale
Par pra
Fol
Cort
GA
1
M
149 Parmotrema pseudocrinitum (Abbayes) Hale
Par pse
Fol
Cort
GA
1
M
150 Parmotrema pseudonilgherrense (Asahina) Hale
Par pse
Fol
Cort
GA
8
E, M, F
151 Parmotrema reticulatum (Taylor) M. Choisy
Par ret
Fol
Saxi
GA
2
C, E
152 Parmotrema saccatilobum (Taylor) Hale
Par sac
Fol
Cort
GA
1
E
153 Parmotrema sancti-angelii (Lynge) Hale
Par san
Fol
Saxi
GA
1
M
154 Parmotrema subarnoldii (Abbayes) Hale
Par sub
Fol
Saxi
GA
2
C, M
155 Parmotrema tinctorum (Despr. ex Nyl.) Hale
Par tin
Fol
Cort
GA
1
E
156 Parmotrema ultralucens (Krog) Hale
Par ult
Fol
Saxi
GA
1
M
157 Peltigera didactyla (With.) J.R. Laundon
Pel did
158 Peltigera dolichorrhiza (Nyl.) Nyl.
Pel dol
Peltigeraceae
Fol
Musc
BGA
1
E
Fol
Cort
BGA
3
F, E
159 Peltigera dolichospora (D.A. Lu) Vitik.
Pel dol
Fol
Cort
BGA
2
E, F
160 Peltigera malacea (Ach.) Funck
Pel mal
Fol
Musc
BGA
1
F
161 Peltigera membranacea (Ach.) Nyl.
Pel mem
Fol
Musc
BGA
2
F
162 Peltigera polydactylon (Neck.) Hoffm.
Pel pol
Fol
Musc
BGA
3
E, F
163 Peltigera praetextata (Flörke) Zopf
Pel pra
164 Pertusaria albescens (Huds.) M. Choisy & Wern.
Per alb
Pertusariaceae
Fol
Musc
BGA
1
C
Cru
Cort
GA
1
E
165 Pertusaria amara (Ach.) Nyl.
Per ama
Cru
Cort
GA
1
F
166 Pertusaria amarescens Nyl.
Per ama
Cru
Saxi
GA
2
M, E
167 Pertusaria commutata Müll. Arg.
Per com
Cru
Saxi
GA
1
E
168 Pertusaria composita Zahlbr.
Per com
Cru
Cort
GA
1
M
169 Pertusaria hemisphaerica (Flörke) Erichsen
Per hem
Cru
Cort
GA
1
F
170 Pertusaria krogiae A.W. Archer, Elix, Eb. Fisch., Killmann &
Sérus
Per kro
Cru
Cort
GA
1
E
171 Pertusaria lactea (L.) Arnold
Per lac
Cru
Cort
GA
1
F
172 Pertusaria ophthalmiza (Nyl) Nyl.
Per oph
Cru
Cort
GA
1
F
173 Pertusaria pertusa (Weigel) Tuck.
Per per
Cru
Cort
GA
2
E, F
174 Pertusaria psoromica A.W. Archer & Elix
Per pso
Cru
Cort
GA
2
M, F
175 Pertusaria umbricola A.W.Archer & Elix
Per umb
Cru
Cort
GA
2
E
176 Pertusaria velata (Turner) Nyl.
Per vel
Cru
Cort
GA
1
F
177 Pertusaria xanthoplaca Müll. Arg.
Per xan
178 Phaeophyscia ciliata (Hoffm.) Moberg
Pha cil
Physciaceae
Cru
Cort
GA
1
E
Fol
Cort
GA
4
C, E
179 Phaeophyscia endococcina (Körb.) Moberg
Pha end
Fol
Saxi
GA
2
M, C
180 Phaeophyscia hispidula (Ach.) Moberg
Pha his
Fol
Cort
GA
1
M
181 Phaeophyscia hispidula var. exornatula (Zahlbr.) Moberg
Pha his
Fol
Cort
GA
2
F
182 Phaeophyscia primaria (Poelt) Trass
Pha pri
Fol
Saxi
GA
1
C
183 Phaeographis extrusa (Stirt.) Zahlbr.
Phe ext
Graphidaceae
Cru
Cort
GA
1
F
184 Phlyctis argena (Ach.) Flot.
Phl arg
Phlyctidaceae
Cru
Cort
GA
1
F
185 Physcia caesia (Hoffm.) Fürnr.
Phy cae
Physciaceae
186 Physcia dilatata Nyl.
Phy dil
Fol
Saxi
GA
3
M, C
Fol
Cort
GA
1
E
187 Physcia semipinnata (Gmelin) Moberg
Phy sem
Fol
Cort
GA
1
E
188 Physcia tenella (Scop.) DC.
Phy ten
Fol
Saxi
GA
1
M
E, F, M
189 Platismatia erosa W. Culb. & C. Culb.
Pla ero
Parmeliaceae
Fol
Cort
GA
3
190 Polychidium stipitatum Vězda & W.A. Weber
Pol sti
Massalongiaceae
Fru
Cort
BGA
1
F
191 Porina chlorotica (Ach.) Müll.Arg.
Por chl
Porinaceae
Cru
Saxi
GA
1
F
192 Porpidia albocoerulescens (Wulfen) Hertel & Knoph
Por alb
Lecideaceae
Cru
Saxi
GA
1
F
193 Pyxine berteriana (Fée) Imsh.
Pyx ber
Caliciaceae
Fol
Cort
GA
2
F, E
194 Ramalina conduplicans Vain.
Ram con
Ramalinaceae
195 Ramalina hossei Vain.
Ram hos
Fru
Cort
GA
8
F, E, M
Fru
Cort
GA
4
E, F, M
196 Ramalina roesleri (Hochst) Hue
Ram roe
Fru
Cort
GA
3
M, F
197 Ramalina sinensis Jatta
Ram sin
Fru
Cort
GA
1
F
198 Rhizocarpon badioatrum (Flörke ex Spreng.) Th. Fr.
Rhi bad
199 Rhizocarpon obscuratum (Ach.) A. Massal.
Rhi obs
Rhizocarpaceae
Cru
Saxi
GA
2
E, M
Cru
Saxi
GA
1
M
200 Rhizoplaca chrysoleuca (Sm.) Zopf
Rhi chr
Lecanoraceae
Fol
Saxi
GA
2
M
201 Rinodina efflorescens Malme
Rin eff
Physciaceae
Cru
Cort
GA
1
M
202 Rinodina instrusa (Krempelh. in Nyl.) Mamle
Rin ins
Cru
Cort
GA
2
M, E
203 Rinodina lecideina H. Mayrhofer & Poelt
Rin lec
Cru
Saxi
GA
1
C
204 Rinodina sophodes (Ach.) A. Massal.
Rin spo
205 Sclerophora amabilis (Tibell) Tibell
Scl ama
Coniocybaceae
206 Stereocaulon paradoxum I.M. Lamb
Ste par
Stereocaulaceae
207 Stereocaulon piluliferum Th.Fr.
Ste pil
208 Sticta nylanderiana Zahlbr.
Sti nyl
209 Sticta praetextata (Räsänen) D.D. Awasthi
Sti pra
210 Sticta weigelii (Ach.) Vain.
Sti wei
211 Sulcaria sulcata (Lév.) Bystr. ex Brodo & D. Hawksw.
Sul sul
Lobariaceae
Parmeliaceae
Cru
Saxi
GA
1
M
Fru
Cort
GA
1
F
Fru
Saxi
GA
7
M, E, C
Fru
Saxi
GA
2
C, E
Fru
Cort
GA
2
F
Fru
Cort
GA
2
F, E
Fru
Cort
BGA
1
F
Fru
Cort
GA
2
F, E
212 Nephromopsis ahtii (Randl. & Saag) Randl. & Saag
Nep aht
Fol
Cort
GA
7
E, F
213 Nephromopsis laureri (Kremp.) Kurok.
Nep lau
Fol
Cort
GA
5
F, E, M
57
58
R esearch
S.N. Name of Lichen species
Short
form
Family
Growth
form
Substrate Photobiont Frequency Land use
group
partner
(Number) types
214 Umbilicaria badia Frey
Umb bad Umbilicariaceae
Fol
Saxi
GA
4
C, E,
215 Umbilicaria indica var. indica Frey
Umb ind
Fol
Saxi
GA
8
F, M, E
216 Umbilicaria vellea (L.) Ach. em. Frey
Umb vel
217 Usnea bailey (Stirt.) Zahlbr.
Usn bai
Parmeliaceae
Fol
Saxi
GA
3
M, F
Fru
Cort
GA
1
E
218 Usnea cirrosa Motyka
Usn cir
Fru
Cort
GA
8
E, F, M
219 Usnea compressa Taylor
Usn com
Fru
Cort
GA
5
F, M, E
220 Usnea cornuta Körb.
Usn cor
Fru
Cort
GA
4
E, M, F
221 Usnea himalayana Bab.
Usn him
Fru
Cort
GA
2
E, F
222 Usnea longissima Ach.
Usn lon
Fru
Cort
GA
3
F, E
223 Usnea pygmoidea (Asahina) Y. Ohmura
Usn pyg
Fru
Cort
GA
1
M
224 Usnea sp1 Dill. ex Adans.
Usn sp1
Fru
Cort
GA
3
M, E
225 Usnea sp2 Dill. ex Adans.
Usn sp2
Fru
Cort
GA
2
F, M
226 Verrucaria nigrescens Pers.
Ver nig
Verrucariaceae
Cru
Saxi
GA
1
C
227 Xanthoparmelia tinctina (Maheu & A. Gillet) Hale
Xan tin
Parmeliaceae
Fol
Cort
GA
1
M
228 Xanthoria fallax (Hepp) Arnold
Xan fal
Teloschistaceae
229 Xanthoria parietina (L.) Th. Fr.
Xan par
Appendix 2 – TukeyHSD test for multiple comparisons of
mean species richness of lichens between land-use types and b.
Biplot CCA scores.
a) TukeyHSD test for multiple comparisons of mean species
richness of lichens between land-use types.
Variables
Difference Lower
Upper p adjusted
Fol
Cort
GA
2
M, C
Fol
Cort
GA
1
F
b) Pearson correlations between environmental variables and
CCA axes.
Variables
CCA1
CCA2
0.964 −0.184
Elevation
Exploited forest
−0.005 −0.178
Natural forest
−0.187 −0.573
Exploited-Cultivated
14.46
3.93
24.99
0.00
Meadow
0.161
0.355
Natural-Cultivated
12.33
2.05
22.60
0.01
Canopy openness
0.193
0.755
9.90
−0.62
20.43
0.07
Natural-Exploited
−2.13 −12.09
7.82
0.94
Meadow-Exploited
−4.56 −14.77
5.66
0.63
Meadow-Natural
−2.42 −12.38
7.53
0.91
Meadow-Cultivated
Appendix 3 – Regression analysis results modelled for lichen species richness, growth forms, substrate types and photobiont types as
response variables and canopy openness and elevation as predictor variables. The Quasi-Poisson family error fitted in GLM (Generalized Linear Model). p-values refer to linear (linear model) or quadratic (linear & quadratic model) coefficient. p-value codes: 0 ‘***’
0.001 ‘**’ 0.01 ‘*’ 0.05 ‘.’ 0.1. ns (non-significant) for p>|0.1| which means marginal significance.
Predictor variables Response variables
Canopy openness
Total lichen richness
Crustose species richness
Fruticose species richness
Corticolous species richness
Saxicolous species richness
Cyanolichen species richness
Green algal lichen species
richness
Elevation
Total species richness
Crustose species richness
Model
Degrees of Residual
freedom
deviance
Deviance
explained
ΔD2
p(>|t value|)
Intercept
35
222.23
0
Linear
34
199.19
23.04 0.104
***
*
Linear & quadratic
33
192.35
6.85 0.134
ns
***
Intercept
35
65.36
0
Linear
34
58.56
6.80 0.104
.
Linear & quadratic
33
57.96
0.60 0.113
ns
***
Intercept
35
111.10
0
Linear
34
99.39
11.71 0.105
Linear & quadratic
33
90.94
8.45
*
0.18
0.09
***
Intercept
35
289.78
0
Linear
34
218.15
71.63 0.247
**
Linear & quadratic
33
204.48
13.67 0.294
ns
0
***
Intercept
35
104.63
Linear
34
79.87
24.76
0.23
**
Linear & quadratic
33
75.72
4.15
0.27
ns
Intercept
35
68.57
0
0.37
Linear
34
55.61
12.96 0.189
*
Linear & quadratic
33
54.72
0.89 0.202
ns
***
Intercept
35
189.80
0
Linear
34
174.93
14.87 0.078
.
Linear & quadratic
33
168.55
6.38 0.112
ns
0
***
Intercept
35
222.23
Linear
34
163.35
53.88
0.26
**
Linear & quadratic
33
160.42
2.93
0.28
ns
0
***
Intercept
35
65.36
Linear
34
56.24
9.12
0.14
*
Linear & quadratic
33
55.97
0.27
0.14
ns
Cho ng ba ng e t a l .
Predictor variables Response variables
Elevation
Foliose species richness
Fruticose species richness
Corticolous species richness
Cyanolichen species richness
Green algal lichen species
richness
Model
Degrees of Residual
freedom
deviance
Deviance
explained
ΔD2
p(>|t value|)
Intercept
35
137.8
Linear
34
104.81
32.99
0.24
0
***
**
Linear & quadratic
33
102.2
2.61
0.26
ns
***
Intercept
35
111.10
0
Linear
34
92.02
19.08 0.172
*
Linear & quadratic
33
91.65
0.37 0.152
ns
***
Intercept
35
289.78
0
Linear
34
235.08
54.7 0.189
**
Linear & quadratic
33
227.09
7.99 0.216
ns
0.23
Intercept
35
68.57
0
Linear
34
59.13
9.44 0.138
*
Linear & quadratic
33
58.38
0.75 0.149
ns
Intercept
35
189.80
0
Linear
34
140.67
49.13 0.259
***
**
Linear & quadratic
33
138.19
2.48 0.272
ns
Appendix 4 – Representative hemispherical photographs chosen from the analysed images characterizing transects in Ghunsa Valley, Kanchenjunga. (1 = 2 000 m, 2 = 2 600 m, 3 = 3 000 m, 4 = 3 400 m and 5 = 3 800 m; E = eastern slope, W = western slope;
c = cultivated land, e = exploited forest, m = meadows and f = natural forest).
1Ecb
1Emb
1Eea
1Efb
openness = 70.5 %
openness = 62.4 %
openness = 55.75 %
openness = 9.98 %
1Wcb
1Wmb
1Web
1Wfb2
openness = 84.76 %
openness = 82.81 %
openness = 37.05 %
openness = 38.46 %
2Ecb1
2Efb
2Wcb
2Wmb
openness = 62.35 %
openness = 31.48 %
openness = 65.46 %
openness = 58.33 %
2Web
2Wfb
3Emc
3Eeb
openness = 20.42 %
openness = 16.48 %
openness = 55.92 %
openness = 24.23 %
59
60
R esearch
3Efb
3Wcb
3Wmb
3Web
openness = 14.06 %
openness = 69.39 %
openness = 61.24 %
openness = 36.53 %
3Wfb
4Ecb
4Emb
4Eeb
openness = 17.19 %
openness = 55.98 %
openness = 7.16 %
openness = 50.71 %
4Efb
4Wcb
4Wmb
4Wee
openness = 21.51 %
openness = 61.36 %
openness = 61.04 %
openness = 18.34 %
4Wfb1
5Emb
5Eeb
5Efb
openness = 17.51 %
openness = 47.72 %
openness = 21.05 %
openness = 19.78 %
5Wcb
5Wmb
5Wec
5Wfc
openness = 47.85 %
openness = 62.88 %
openness = 35.16 %
openness = 31.24 %