The Lichenologist 42(1): 83–96 (2010

)
doi:10.1017/S0024282909008627

© British Lichen Society, 2009

The elevation gradient of lichen species richness in Nepal
Chitra Bahadur BANIYA, Torstein SOLHØY, Yngvar GAUSLAA
and Michael W. PALMER
Abstract: This study of elevation gradients of lichen species richness in Nepal aimed to compare
distribution patterns of different life-forms, substratum affinities, photobiont types, and Nepalese
endemism. Distribution patterns of lichens were compared with elevational patterns shown by a wide
range of taxonomic groups of plants along the Nepalese Himalayan elevational gradient between
200–7400m. We used published data on the elevation records of 525 Nepalese lichen species to
interpolate presence between the maximum and minimum recorded elevations, thereby giving
estimates of lichen species richness at each 100-m elevational band. The observed patterns were
compared with previously published patterns for other taxonomic groups. The total number of lichens
as well as the number of endemic species (55 spp.) showed humped relationships with elevation. Their
highest richness was observed between 3100–3400 and 4000–4100m, respectively. Almost 33% of the
total lichens and 53% of the endemic species occurred above the treeline (>4300m). Non-endemic
richness had the same response as the total richness. All growth forms showed a unimodal relationship
of richness with elevation, with crustose lichens having a peak at higher elevations (4100–4200m) than
fruticose and foliose lichens. Algal and cyanobacterial lichen richness, as well as corticolous lichen
richness, all exhibited unimodal patterns, whereas saxicolous and terricolous lichen richness exhibited
slightly bimodal relationships with elevation. The highest lichen richness at mid altitudes concurred
with the highest diversity of ecological niches in terms of spatial heterogeneity in rainfall, temperature,
cloud formation, as well as high phorophyte abundance and diversity implying large variation in bark
roughness, moisture retention capacity, and pH. The slightly bimodal distributions of saxicolous and
terricolous lichens were depressed at the elevational maximum of corticolous lichens.
Key words: altitude, endemism, Himalaya

Introduction
The Himalayas are the highest mountain
range in the world and show a rich diversity
of eco-climatic zones. In such a landscape,
elevational gradients are of particular interest
in applied as well as in theoretical ecology
(Wolf 1993). The pattern of changes in
species-richness with elevation characterizes
C. B. Baniya (corresponding author) and T. Solhøy:
Department of Biology, University of Bergen, Allégaten
41, P.O. Box 7803, N-5020 Bergen, Norway. Email:
chitra.baniya@bio.uib.no
Current address: Central Department of Botany,
Tribhuvan University Kirtipur, Kathmandu, Nepal.
Y. Gauslaa: Department of Ecology and Natural
Resource Management, Norwegian University of Life
Sciences, P.O. Box 5003, NO-1432 Ås, Norway.
M. W. Palmer: Department of Botany, Oklahoma
State University, 104 LSE Stillwater OK 74078
USA 405-744-7717.

the vegetation in a simple but powerful way.
For example, Yoda (1967) found gradually
decreasing tree species richness with increasing altitude. Later, Hunter & Yonzon (1993)
reported a similar pattern for Nepalese birds
and mammals. A number of studies have
produced macro-scale elevational gradients
for the whole of Nepal, using interpolation
methods on data from secondary sources.
Several groups of organisms have been
studied and exhibit maximum richness at
group-specific intermediate elevations, referred to as a unimodal, or humped, elevation pattern. For example, Himalayan
vascular plants showed their highest total
richness at altitudes between 1500 to 2500m
and their highest endemic richness at 4000m
(Grytnes & Vetaas 2002; Vetaas & Grytnes
2002), while fern richness peaked at 2000m
(Bhattarai et al. 2004). However, orchids

84

THE LICHENOLOGIST

showed their maximum richness at 1600m
whereas the highest orchid endemic richness
was bimodal with two separate peaks at 1200
and 3300m (Acharya 2008). Liverworts and
mosses had their maximum richness at 2800
and 2500m, respectively, and the peak for
endemic liverwort richness occurred at
3300m (Grau et al. 2007).
Relatively few studies on elevational richness patterns for lichens have been undertaken (Kessler 2000; Bruun et al. 2006;
Grytnes et al. 2006) and among terrestrial
photosynthetic organisms they are the only
major group that has not yet been investigated in this respect in Nepal. They are
among the most successful organisms in extreme environments such as cold arctic and
alpine environments where few other plants
can grow (e.g. Schroeter et al. 1994; Kappen
et al. 1996). Lichens also show a high diversity as eipiphytes and thus may benefit from
high diversity of trees and shrubs with
species-specific bark chemistry, texture and
stability (Kessler 2000; Bruun et al. 2006;
Grytnes et al. 2006). This paper describes
and interprets the Himalayan elevational distribution of lichen species richness in Nepal.
Material and Methods
Study area
Nepal is a mountainous country in the central
Himalayas between 26°22# to 30°27# N and 80°04# to
88°12# E with an area of 147 181 km2. The elevation
ranges from 60m above sea level to 8848m (Mt. Everest).
The country is surrounded by the Indo-Gangetic Plain
in the east, west and south, and the Tibetan Plateau
towards the north. Parallel mountain ranges run southeast and north-west dissecting the country into northsouth running river valleys that are associated with fertile
plains, inner valleys, lakes, and the outer flood plains.
The Indian monsoon beginning in May and ending in
September is the main source of precipitation (more
than 80%). Monsoon rain forms after condensation of
water vapour arising from the Bay of Bengal, and consequently there is a spatial gradient of monsoon rainfall
decreasing from east to west. Precipitation also occurs in
winter formed from central Asian water vapour and
diminishes eastwards.
Physiographically, Nepal has been divided into five
zones (Hagen 1969; Upreti 1999). 1) A 30–40km wide
southernmost plain is called Terai between 60 and 300m.
It is the most fertile, densely populated zone with a tropical climate (between 22–27°C during winter and above

Vol. 42

35°C during the warmest summer months); it is mainly
cultivated but has still some tropical forests. 2) The
Siwalik located from 700 to 1500m in the foothills of
the Mahabharat Range. 3) The Mahabharat Range lies
between 1500 and 3000m; it belongs to the Midlands or
lesser Himalaya and includes the principal inner valleys
such as Kathmandu and Pokhara. The climate is mild
throughout the year and receives the highest annual rainfall (above 4000mm) south of Annapurna, central Nepal.
It includes the lower cloud zone at 2000 to 2500m (Miehe
1989, 1990). 4) The Greater-Himalaya extends above
2700 m. Here the climate varies from dry summers to cold
winters with snow, and subzero temperatures on the
mountain tops. This zone includes the uppermost limit of
cloud-zone forest in Nepal at 4000m (Miehe 1989, 1990).
5) Trans-Himalaya lies towards the northern side of the
Greater-Himalaya with elevations above 3000m. It is in
the rain-shadow zone with an average annual rainfall
between 200 and 400mm.
Vegetation is one of the prime determinants of lichen
diversity and distribution (Awasthi 2007). In Nepal,
vegetation has been studied and classified into 8 elevational forest zones (Yoda 1967; Stainton 1972, 2001;
Dobremez & Jest 1969; Dobremez 1976; Miehe 1982,
1989). 1) Tropical forest between 60 and 1000 m
(Stainton 1972) dominated by Shorea robusta Gaertn.,
Dalbergia sissoo Roxb., Adina cordifolia (Willd. ex Roxb.)
Benth. & Hook. f. ex Brandis, Terminalia spp., Lagerstroemia spp., Michelia champaca L., and Bombax ceiba L.
2) Subtropical forest occurring between 1000 to 2000m
with Pinus roxburghii Sarg. as the dominant species on
south-facing slopes in the Midlands. 3) Lower temperate broad-leaved forest at 1700–2400m in the east
and 2000–2700m in the west. Dominant trees include
Alnus nitida (Spach.) Endel., Castanopsis tribuloides A.
DC., Castanopsis hystrix A. DC., Lithocarpus pachyphylla
(Kurz.) Rehder., and Quercus spp. Moister slopes between 1700–2200m are dominated by Cinnamomum
spp. and represent the lower temperate mixed broadleaved forest. Upper temperate broad-leaved forests are
represented by various types. Drier southern slopes at
2200–3000m are dominated by Schima wallichii (DC.)
Korth., Castanopsis indica (Roxb.) Miq., and Quercus
semecarpifolia J. E. Smith. Moist, north and west
slopes between 2500–3500m from central to east
Nepal are dominated by Acer spp., Rhododendron spp.,
Aesculus spp. and Juglans spp. 4)Temperate coniferous
forests between 2000 and 3700m dominated by Pinus
wallichiana A. B. Jack. Other conifers include Cedrus
deodara (Roxb.) G. Don, Picea smithiana (Wall.) Boiss.,
Juniperus indica Bertol., Larix himalaica W. C. Cheng &
L. K. Fu, Larix griffithiana Carriere, Cupressus torulosa D.
Don, and Tsuga dumosa (D. Don) Eichler. 5) Sub-alpine
forests occurring between 3000 and 4100m (Stainton
1972) dominated by Betula utilis D. Don, Abies spectabilis
(D. Don) Mirb., and Rhododendron spp. This forest
includes the climatic treeline at 4100–4300m. 6) Open,
low-alpine shrub communities including Caragana spp.,
Lonicera spp., Rosa spp., Sophora spp., Rhododendron
anthopogon D. Don, R. lepidotum Wall. ex G. Don,
Ephedra gerardiana Wall. ex C. A. Meyer, and Hippophae
tibetana Schldl. 7) Mid-alpine zone with dwarf-shrubs

2010

Elevation gradients of lichens in Nepal—Baniya et al.

but largely dominated by herb communities. 8) Nival
zone begins above the mid-alpine zone with more
lichens and fewer mosses and angiosperms. Within the
nival zone, there are vascular plants of open wind exposed areas as well as those tolerant of snow-lie.
Data sources
The main source of data for this study is the Lichens of
Nepal (Sharma 1995). However, many species reported
in Sharma (1995) lack elevation data and since then
many new species have been described. Other relevant
and easily accessible literature as well as the recent keys
for both microlichens and macrolichens of Nepal,
India and Sri Lanka by Awasthi (1991; 2007) have also
been checked. In addition, the following literature on
Nepalese lichens (Lamb 1966; Poelt 1966a, b, 1974;
Bystrek 1969; Poelt & Reddi 1969; Abbayes 1974; Jahns
& Seelen 1974; Kurokawa 1974; Mitchell 1974;
Schmidt 1974; Veˇzda & Poelt 1975; Hellmich & Poelt
1977; Awasthi 1986; Vitikainen 1986; Awasthi &
Mathur 1987; Miehe 1990; Awasthi 1991, 2007;
Esslinger & Poelt 1991; Poelt & Hinteregger 1993;
Sharma 1995; Baniya 1996; Jørgensen 2001) have provided information on the elevation ranges of Nepalese
lichens. These sources have provided elevational distribution data for 525 species, including 55 Nepalese
endemics (Appendix 1). Among the 525 species, 172,
including 29 endemics, are reported from above the
treeline (> 4300m). In addition, data about life-forms
(crustose, foliose and fruticose), photobiont types
(cyanobacteria and green algae) and five substratum
types (tree bark, wood, mosses, rocks and soil) were
obtained. Tripartite lichens were treated as cyanobacterial. There were 50 species of lichens for which elevational data or other important information are either
incomplete or lacking; these species were excluded from
the analysis. Lignicolous and muscicolous substrata had
been reported for a few species (5 and 17, respectively)
and were thus also excluded during the analyses involving substrata. Lichen nomenclature follows Awasthi
(1991; 2007).
The altitudinal range of lichens in Nepal, 200–
7400m, was divided into 73 bands of 100m each and a
complete data matrix for all species was assembled.
Presence of a species indicates that the species occurs in,
or has been collected in the past, from that elevation
band and absence means either that the species does not
occur or has previously not been collected from that
elevation. A species is assumed to be present in all
possible 100m bands between its upper and lower elevation limits. For example, a lichen that has elevational
occurrences between 210 to 451m in the literature falls
between the 200- and 500-m bands. A list of the lichens
included in this study together with their elevation
ranges is provided in Appendix 1.
Species richness is an estimate of the total number of
lichen species occurring in each 100m elevation band.
This is a macro-scale study (gamma diversity, sensu
Whittaker 1972) that covers the entire elevational range
of Nepal. Fifty-six taxa, varieties, forms or subspecies,
were treated as species (Appendix 1).

85

Data analysis
Patterns related to total lichen species richness and
their sub-sets (life forms, algal partners, and substrata)
as responses and their elevations as a predictor variable were extracted by using a cubic smooth spline (s)
within the framework of Generalized Additive Models
(GAM, Hastie & Tibshirani 1990; Heegaard 2004) with
a default of c. 4 degrees of freedom. Response variables
are counts; thus, the variance changes with the mean and
negative predictions are meaningless. We found overdispersion in our data. We thus applied a Quasi-poisson
family error distribution with a logarithmic link function
(Crawley 2006). We confirmed our assumption of
normal distribution of error after the Q-Q diagnostic
plots plotted against residuals. The change in deviance
follows the F-distribution. We used R 2.7.0 (R Development Core Team 2008) to analyse our data and
smoothers were fitted with library GAM (Hastie &
Tibshirani 1990). GAM was used because it is a
non-parametric approach that does not make a priori
assumptions about the species-elevational relationship.
One of the main purposes of this study is to contrast
observed lichen richness patterns with other studies.
Biogeographic data have biases and constraints, such as
a focus on specific taxa and/or a restriction to easily
accessible landscapes. However, these problems are
shared among data sets with various taxonomic groups
(Grytnes & Vetaas 2002; Vetaas & Grytnes 2002;
Bhattarai et al. 2004; Grau et al. 2007). Thus, a comparison of empirical patterns among available studies is
possible.

Results
The 525 lichen species recorded (Appendix
1) represented 40 families and 121 genera.
Among these lichens, 35·4% were crustose,
46·3% were foliose and 18·3% were fruticose. A total of 12% had a cyanobacterial
photobiont and 88% had a green algal photobiont. Lichens endemic to Nepal (n=55) represented 10% of the total Nepalese lichen
flora. There were 172 species reported from
above the treeline (R 4300m) and 29 of
these were endemics. Carbonea vorticosa in
Nepalese Himalaya at 7400m was the
world’s highest reported lichen. Heterodermia
pseudospeciosa represented one of the lichens
occurring at the lowest elevation ranges in
Nepal (150–2100m).
The total lichen species richness showed
a unimodal relationship with elevation. The
maximum modelled total richness occurred
at 3100–3400m (Fig. 1A), whereas the
observed maximum richness (144 species)

86

THE LICHENOLOGIST

Vol. 42

F. 1. Relationship between elevation and lichen
species richness in Nepal. A, total lichen species richness; B, total endemic lichen species richness. The
fitted regression lines represent statistically significant
(P % 0·001) smooth spline (s) after using GAM with
approximately 4 degrees of freedom.

was at approximately 4000m. Endemic
lichens also had a unimodal relationship with
elevation, but their maximum modelled
richness occurred at 3900–4400 m (Fig. 1B)
approximately 800m higher than the peak
for total species. Non-endemic richness
matched closely the elevational patterns of
the total species richness (data not shown).
Species richness of all three growth forms
of lichens had unimodal responses to elevation (Fig. 2A–C). Among the three morphological types, crustose lichens peaked at the
highest altitude (4100–4200m) while foliose
lichens had their maximum richness at
consistently lower altitudes (2400–2500m).
Fruticose lichens peaked at an intermediate
elevation (3200m). With respect to photobiont type, cyanolichens exhibited maximum
richness at lower altitudes (2900–3000m;
Fig. 3A) than green algal lichens (3300–
3500m; Fig. 3B).
Among the five substratum categories
studied, corticolous lichens had a clear unimodal relationship to elevation, with the
highest species richness occurring between
2500 and 2700m (Fig. 4A) in the lower part
of the temperate forests. Saxicolous and terricolous lichens had slightly bimodal patterns

F. 2. Relationship between elevation and lichen
species life-form richness in Nepal. A, foliose lichen
species richness; B, fruticose lichen species richness; C,
crustose lichen species richness. The fitted regression
lines represent statistically significant (P % 0·001)
smooth spline (s) after using GAM with approximately 4
degrees of freedom.

with a prominent hump at 3900–4200m
(Fig. 4B & C).
The regression analysis results showing
the best selected model for each response
variable are recorded in Appendix 2.

Discussion
Total lichen species richness in Nepal varies
strongly with elevation (Figs 1A & B) in line
with previous findings for vascular plants
(Grytnes & Vetaas 2002; Vetaas & Grytnes
2002; Bhattarai & Vetaas 2003; Bhattarai
et al. 2004) including orchids (Acharya
2008), and for bryophytes (Grau et al. 2007).
In all these studies, as well as studies dealing
with lichens from other countries (Wolseley
& Aguirre-Hudson 1997; Negi 2000, 2003;
Wolf & Alejandro 2003; Pinokiyo et al.

2010

Elevation gradients of lichens in Nepal—Baniya et al.

87

F. 3. The relationship between elevation and lichen
richness for lichens with different photobionts. A, cyanolichen species richness; B, green algal lichen species
richness. The fitted regression lines represent statistically significant (P % 0·001) smooth splines (s) after
using GAM with approximately 4 degrees of freedom.

2008), species richness tends to peak at
intermediate altitudes. The various major
taxonomic groups exhibit maximum richness
at different altitudes: vascular plants (1500 to
2500m; orchids 1600m, ferns 2000m), liverworts 1800m, mosses 2500m and as high as
3100 to 3400m for lichens. Thus, total richness for lichens occurs at higher elevations
than for any of the other groups studied.
The total lichen species richness in the
Himalayas peaked in the upper cloud-forest
zone (following the vegetation classification
of Hagen 1969; as modified by Miehe 1982;
1989). This is the zone of the highest rainfall
on southernmost slopes (> 4000mm), and
the lowest rainfall (< 300mm) on northernmost slopes where the Himalayan föhn
causes local drying (Miehe 1989). The zone
with maximum lichen richness represents
the temperate zone with extremely large
local variations in water availability and the
accompanying gradients in vegetation cover.
According to Bhattarai et al. (2004), the high
annual rainfall (> 4000mm) and cool summer temperatures (14–17°C) in this zone are
likely to favour forests.The large number of
temperate broad-leaved and coniferous trees

F. 4. The relationship between elevation and lichens
preferring specific substrata. A, corticolous lichen
species richness; B, saxicolous lichen species richness;
C, terricolous lichen species richness. The fitted regression lines represent statistically significant (P % 0·001)
smooth splines (s) after using GAM with approximately
4 degrees of freedom.

with bark differing in roughness, moisture
retention capacity and pH present a wide
variety of habitats for lichens. The slightly
bimodal peaks for saxicolous (Fig. 4B) and
terricolous lichens (Fig. 4C) tend to show the
lower peak in species richness occurring at
the same altitude at which corticolous species
reach their maximum richness (Fig. 4A).
These patterns may be determined by habitat availability and specificity. Corticolous
lichens apparently peak at elevations with
a high abundance of forests, implying a
reduced occurrence of natural and well-lit
terricolous and saxicolous habitats.
Fruticose lichens have their maximum
richness at a substantially higher altitude
than the foliose lichens (Figs 2A & B). A
similar relationship, but on a very different
scale, can be seen within forest canopies

88

THE LICHENOLOGIST

where the biomass of fruticose lichens increases and that of foliose lichens decreases
with height above the ground (Goward 1998;
Campbell & Coxson 2001; Gauslaa et al.
2008). In well-lit and open high altitude forests and the upper canopies of trees, fruticose
lichens may have the advantage in being able
to utilise light from all directions, whereas
many foliose and flat lichens maximize the
harvest of more or less unidirectional light in
shady positions in dense forests and on low
canopy branches (Gauslaa et al. 2009). The
foliose lichen richness peaks in the same zone
as the highest richness for mosses (Grau et al.
2007), taxa that are also relatively shade
tolerant. Furthermore, finely dissected fruticose lichens on twigs lack thick boundary
layers. Thus, they are more closely coupled
to ambient air than flat foliose lichens and
absorb water vapour more readily from the
air as discussed by Jonsson et al. (2008). By
contrast, flat lichens and bryophyte carpets
are more dependent on rainwater.
The elevation range between 4000 to
4300m is the sub-alpine zone of Nepal
(Stainton 1972). This humid upper-cloud
zone (Miehe 1989) is densely colonized by
epiphytic mosses reflecting the high humidity
(Grau et al. 2007). This humidity may also
facilitate high lichen richness associated with
open canopies with more light facilitating
lichen growth (Gauslaa et al. 2007).
A high degree of isolation creates endemism (Cox & Moore 2000), and isolation
might increase with elevation. High mountain peaks with an exceptionally cold and
harsh climate act as islands (Grau et al.
2007), and phytogeographical isolation
can lead to localized speciation events. The
occurrence of 29 endemic species (17%) out
of the 172 total species above the treeline
(R 4300 m) emphasises the link between
elevation and endemism. The Nepalese
lichen endemism rate is higher than that for
mosses (3 out of 480 spp. i.e., < 1%) and
liverworts (33 out of 368 spp. i.e., 9%), but
slightly less than that for vascular plants (303
out of 1957 spp. i.e., 16%). Above 4500m
both total and endemic lichen diversity (Figs
1A & B) decline until 6100m, which is the
highest reported elevation for lichens en-

Vol. 42

demic to Nepal. Cold winds leading to rapid
freezing and drying may reduce competition
from surviving vascular plant species (Grime
1977). The harsh climate might lead to various specializations and thus facilitate formation of endemic species (Aptroot & Bungartz
2007).
Land area might impose a unimodal richness pattern. In general, the larger the area
sampled, the greater will be the number
of species encountered (Rosenzweig 1995;
Qian et al. 2007). A decreasing trend in land
area in each 500m band occurs with increasing elevation (Vetaas & Grytnes 2002) as
estimated from the Digitized Map by the
International Centre for Integrated Mountain Development (ICIMOD, Kathmandu).
This is not always a common pattern (Körner
2007). If we consider species-area relationships, more species should be expected towards lower elevations than we observed.
However, lichen diversity can be greatly
limited by lack of long ecological continuity
in forests (Rose 1976) and lichens may be
less prevalent at low altitudes because of
dense forest canopies and intensive land-use.
Furthermore, as lichen species richness also
tends to be universally greater in cool or cold
climates (Mattick 1953), we believe that the
strong reduction in lichen richness at low
elevations is real and not a sampling artefact.
In conclusion, lichens and various subgroups of lichens exhibit unimodal patterns
similar to those found in other major taxonomic groups, but the highest total lichen
richness peaks at higher altitudes than in
any other group. The maximum lichen richness occurred at the altitude with the highest
diversity of ecological niches in terms of
spatial heterogeneity, rainfall, temperature,
cloud formation, as well as high phorophyte
abundance and diversity implying large variation in bark roughness, moisture retention
capacity, and pH.
We are grateful to the Norwegian State Education Loan
Fund (Lånekassen) for providing funding and the
Central Department of Botany, Tribhuvan University,
Nepal, for providing study leave for CBB. Thanks are
due to John Birks for his valuable comments, Walter
Obermayer and Per Magnus Jørgensen for provision of
literature on lichens and to Govind Ghimire and Bijaya

2010

Elevation gradients of lichens in Nepal—Baniya et al.

Kattel for their feedback. Thanks are also due to Louise
Olley, Teuvo Ahti and Tor Tönsberg for their help in
revising the species list and to the Senior Editor, Peter
Crittenden and two anonymous referees for their useful
comments and suggestions.
R
Abbayes, H. D. (1974) Cladonia du Nepal. In Khumbu
Himal 6 (W. Hellmich & J. Poelt, eds): 111–116.
München: Universitätsverlag Wagner Ges.
M. B. H. Innsbruck.
Acharya, K. P. (2008) Orchid species richness along a
Himalayan elevation gradient. M. Sc. thesis, Bergen
University.
Aptroot, A. & Bungartz, F. (2007) The lichen genus
Ramalina on the Galapagos. Lichenologist 39: 519–
542.
Awasthi, G. (1986) Lichen genus Usnea in India. Journal
of the Hattori Botanical Laboratory 61: 333–421.
Awasthi, D. D. (1991) A key to the microlichens of
India, Nepal and Sri Lanka. Bibliotheca Lichenologica 40: 1–337.
Awasthi, D. D. (2007) A Compendium of the Macrolichens
from India, Nepal and Sri Lanka. Dehra Dun: Bishen
Singh & Mahendra Pal Singh.
Awasthi, D. D. & Mathur, R. (1987) Species of the
lichen genera Bacidia, Badimia, Fellhanera and
Mycobilimbia from India. Proceedings of Indian Academy of Sciences, Bangalore. 97: 481–503.
Baniya, C. B. (1996) Floristic composition of lichens of
Sikles (Kaski) and Shivapuri (Kathmandu) and
their ecology. M. Sc. thesis, Tribhuvan University,
Kathmandu, Nepal.
Bhattarai, K. R. & Vetaas, O. R. (2003) Variation in
plant species richness of different life forms along a
subtropical elevation gradient in the Himalayas,
east Nepal. Global Ecology and Biogeography 12:
327–340.
Bhattarai, K. R., Vetaas, O. R. & Grytnes, J. A. (2004)
Fern species richness along a central Himalayan
elevational gradient, Nepal. Journal of Biogeography
31: 389–400.
Bruun, H. H., Moen, J., Virtanen, R., Grytnes, J. A.,
Oksanen, L. & Angerbjörn, A. (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.
Bystrek, J. L. (1969) Die Gattung Alectoria. In Khumbu
Himal 6 (W. Hellmich & J. Poelt, eds): 17–24.
München: Universitätsverlag Wagner Ges.
M. B. H. Innsbruck.
Campbell, J. & Coxson, D. S. (2001) Canopy microclimate and arboreal lichen loading in subalpine
spruce-fir forest. Canadian Journal of Botany 79:
537–555.
Cox, C. B. & Moore, P. D. (2000) Biogeography: an
Ecological and Evolutionary Approach. London:
Blackwells.
Crawley, M. J. (2006) Statistics: an Introduction using R.
London: John Wiley & Sons, Ltd.

89

Dobremez, J. F. (1976) Le Nepal: Écologie et Biogeographie. Paris: Centre National de la Recherche
Scientifique.
Dobremez, J. F. & Jest, C. (1969) Carte Écologique de la
Region Annapurna Dhaulagiri. Paris: Centre
National de la Recherche Scientifque.
Esslinger, T. L. & Poelt, J. (1991) Parmelia masonii,
a new lichen species (Ascomycota) from the
Himalayas. Bryologist 94: 203–206.
Gauslaa, Y., Palmqvist, K., Solhaug, K. A., Holien, H.,
Hilmo, O., Nybakken, L., Myhre, L. C. & Ohlson,
M. (2007) Growth of epiphytic old forest
lichens across climatic and successional gradients.
Canadian Journal of Forest Research 37: 1832–1845.
Gauslaa, Y., Lie, M. & Ohlson, M. (2008) Epiphytic
lichen biomass in a boreal Norway spruce forest.
Lichenologist 40: 257–266.
Gauslaa, Y., Palmqvist, K., Solhaug, K. A., Hilmo, O.,
Holien, H., Nybakken, L. & Ohlson, M. (2009) Size
dependent growth in two old-growth associated
macrolichen species. New Phytologist 18: 683–692.
Goward, T. (1998) Observations on the ecology of
the lichen genus Bryoria in high elevation conifer
forests. Canadian Field Naturalist 112: 496–501.
Grau, O., Grytnes, J.-A. & Birks, H. J. B. (2007) A
comparison of altitudinal species richness patterns
of bryophytes with other plant groups in Nepal,
Central Himalaya. Journal of Biogeography 34:
1907–1915.
Grime, J. P. (1977) Evidence for the existence of three
primary strategies in plants and its relevance to
ecological and evolutionary theory. American
Naturalist 11: 1169–1194.
Grytnes, J. A., Heegaard, E. & Ihlen, P. G. (2006)
Species richness of vascular plants, bryophytes, and
lichens along an altitudinal gradient in western
Norway. Acta Oecologica 29: 241–246.
Grytnes, J. A. & Vetaas, O. R. (2002) Species richness
and altitude: a comparison between null models
and interpolated plant species richness along the
Himalayan altitudinal gradient, Nepal. American
Naturalist 159: 294–304.
Hagen, T. (1969) Report on the Geological Survey of
Nepal: Preliminary Reconnaissance. Zurich: Orell
Füssli Arts Graphiques.
Hastie, T. J. & Tibshirani, R. J. (1990) Generalised
Additive Models. London: Chapman & Hall.
Heegaard, E. (2004) Trends in aquatic macrophyte
species turnover in Northern Ireland – which factors
determine the spatial distribution of local species
turnover? Global Ecology and Biogeography 13:
397–408.
Hellmich, W. & Poelt, J. (eds) (1977). Khumbu Himal 6.
München: Universitätsverlag Wagner Ges.
M. B. H. Innsbruck.
Hunter, M. L. & Yonzon, P. (1993) Altitudinal distributions of birds, mammals, people, forests, and
parks in Nepal. Conservation Biology 7: 420–423.
Jahns, H. M. & Seelen, E. J. R. (1974) Baeomyces-Funde
aus dem Himalaya. In Khumbu Himal 6 (W.
Hellmich, & J. Poelt, eds): 101–108. München:
Universitätsverlag Wagner Ges. M. B. H. Innsbruck.

90

THE LICHENOLOGIST

Jonsson, A. V., Moen, J. & Palmqvist, K. (2008) Predicting lichen hydration using biophysical models.
Oecologia 156: 259–273.
Jørgensen, P. M. (2001) The lichen genus Erioderma
(Pannariaceae) in China and Japan. Annales
Botanici Fennici 38: 259–264.
Kappen, L., Schroeter, B., Scheidegger, C.,
Sommerkorn, M. & Hestmark, G. (1996) Cold
resistance and metabolic activity of lichens below
0°C. In Life Sciences: Space and Mars Recent Results
18: 119–128. Oxford: Pergamon Press Ltd.
Kessler, M. (2000) Altitudinal zonation of Andean
cryptogam communities. Journal of Biogeography
27: 275–282.
Körner, C. (2007) The use of ‘altitude’ in ecological
research. Trends in Ecology and Evolution 22:
569–574.
Kurokawa, S. (1974) Anaptychia of the NepalHimalaya. In Khumbu Himal 6 (W. Hellmich, & J.
Poelt, eds): 109–110. München: Universitätsverlag
Wagner Ges. M. B. H. Innsbruck.
Kuusinen, M. (1996) Epiphyte flora and diversity on
basal trunks of six old-growth forest tree species in
southern and middle boreal Finland. Lichenologist
28: 443–463.
Lamb, L. M. (1966) Die Gattung Stereocaulon. In
Khumbu Himal 1 (W. Hellmich, ed): 349–352.
München: Universitätsverlag Wagner Ges.
M. B. H. Innsbruck.
Mattick, F. (1953) Lichenologisches Notizen. Berichte
der Deutschen Botanischen Gesellschaft 66: 263–276.
Miehe, G. (1982) Vegetationsgeographische Untersuchungen im Dhaulgiri und Annapurna Himalaya,
Dissertationes Botanicæ. Journal Cramer, Vaduz 66:
66.
Miehe, G. (1989) Vegetation patterns in Mount Everest
as influenced by monsoon and föhn. Vegetatio 79:
21–32.
Miehe, G. (1990) Langtang himal-flora und vegetation als
klimazeiger und-zeugen im Himalaya. Ph. D. thesis,
Universität Göttingen.
Mitchell, M. (1974) Die Gattung Leptogium sect. Mallotium im Himalaya. In Khumbu Himal 6 (W.
Hellmich, & J. Poelt, eds): 121–126. München:
Universitätsverlag Wagner Ges. M. B. H.
Innsbruck.
Negi, H. (2000) On the patterns of abundance and
diversity of macrolichens of Chopta-Tunganath in
the Garhwal Himalaya. Journal of Biosciences 25:
367–378.
Negi, H. (2003) Lichens: a valuable bioresource for
environmental monitoring and sustainable development. Resonance 8: 51–58.
Pant, G. & Awasthi, D. D. (1989a) Caliciales from India
and Nepal. Biovigyanam 15: 3–27.
Pant, G. & Awasthi, D. D. (1989b) Lichen genus Catillaria s. lat. in India. Proceedings of Indian Academy of
Sciences (Plant Science). 99: 369–384.
Pinokiyo, A., Singh, K. P. & Singh, J. S. (2008) Diversity and distribution of lichens in relation to altitude
within a protected biodiversity hotspot, north-east
India. Lichenologist 40: 47–62.

Vol. 42

Poelt, J. (1966a) Die Gattung Ochrolechia. In Khumbu
Himal 1 (W. Hellmich, ed): 251–261. München:
Universitätsverlag Wagner Ges. M. B. H.
Innsbruck.
Poelt, J. (1966b) Die Lobaten Arten der Sammelgattung
Lecanora, Lecanoraceae. In Khumbu Himal 3 (W.
Hellmich, ed): 187–202. München: Universitätsverlag Wagner Ges. M. B. H. Innsbruck.
Poelt, J. (1974) Die Gattungen Physcia, Physciopsis und
Physconia. In Khumbu Himal 6 (W. Hellmich, & J.
Poelt, eds): 57–99. München: Universitätsverlag
Wagner Ges. M. B. H. Innsbruck.
Poelt, J. & Hinteregger, E. (1993) Beiträge zur Kenntnis
der Flechtenflora des Himalaya. Bibliotheca Lichenologica 50: 1–247.
Poelt, J. & Reddi, B. V. (1969) Candelaria und Candelariella. In Khumbu Himal 6 (W. Hellmich, & J.
Poelt, eds): 1–16. München: Universitätsverlag
Wagner Ges. M. B. H. Innsbruck.
Qian, H., Fridley, J. D. & Palmer, M. W. (2007) The
latitudinal gradient of species-area relationships
for vascular plants of North America. American
Naturalist 170: 690–701.
R Development Core Team (2008) R: A language and
environment for statistical computing version 2.7.0.
Rose, F. (1976) Lichenological indicators of age and
environmental continuity in woodlands. In
Lichenology: Progress and Problems (D. H. Brown,
D. L Hawksworth & R. H. Bailey, eds): 279–307.
London: Academic Press.
Rosenzweig, M. L. M. L. (1995) Species Diversity in
Space and Time. Cambridge: Cambridge University
Press.
Schmidt, A. (1974) Chaenotheca und Coniocybe. In
Khumbu Himal 6 (W. Hellmich, & J. Poelt, eds):
133–143. München: Universitätsverlag Wagner
Ges. M. B. H. Innsbruck.
Schroeter, B., Green, T. G. A., Kappen, L. & Seppelt,
R. D. (1994) Carbon dioxide exchange rate at subzero temperatures. Field measurements on Umbilicaria aprina in Antarctica. Cryptogamic Botany 4:
233–241.
Sharma, L. R. (1995). Enumeration of Lichens of
Nepal. Euroconsult, Biodiversity Profiles Project,
Publication No. 3. Nepal Government, 1–109.
Stainton, J. D. A. (1972) Forests of Nepal. London: John
Murray Publisher, Ltd.
Stainton, A. (2001) Flowers of the Himalaya: a Supplement. Oxford: Oxford University Press.
Upreti, B. N. (1999) An overview of the stratigraphy and
tectonics of the Nepal Himalaya. Journal of Asian
Earth Sciences 17: 577–606.
Vetaas, O. R. & Grytnes, J. A. (2002) Distribution of
vascular plant species richness and endemic richness along the Himalayan elevation gradient in
Nepal. Global Ecology and Biogeography 11:
291–301.
Veˇzda, A. & Poelt, J. (1975) Die Gattungen Dimerella
und Pachyphiale. In Khumbu Himal 6 (W. Hellmich,
& J. Poelt, eds): 127–132. München: Universitätsverlag Wagner Ges. M. B. H. Innsbruck.

2010

Elevation gradients of lichens in Nepal—Baniya et al.

Vitikainen, O. (1986) Peltigera dolichospora, a new
Himalayan-western Chinese lichen. Lichenologist
18: 387–390.
Whittaker, R. H. (1972) Evolution and measurement of
species diversity. Taxon 21: 213–251.
Wolf, J. H. D. (1993) Diversity patterns and biomass of
epiphytic bryophytes and lichens along an altitudinal gradient in the Northern Andes. Annals of the
Missouri Botanical Garden 80: 928–960.
Wolf, J. H. D. & Alejandro, F.-S. (2003) Patterns in
species richness and distribution of vascular epiphytes in Chiapas, Mexico. Journal of Biogeography
30: 1689–1707.

91

Wolseley, P. A. & Aguirre-Hudson, B. (1997) The ecology and distribution of lichens in tropical deciduous
and evergreen forests of northern Thailand. Journal
of Biogeography 24: 327–343.
Yoda, K. (1967) A preliminary survey of the forest
vegetation of eastern Nepal II. General description,
structure and floristic composition of sample plots
chosen from different vegetation zones. Journal of
the College of Arts and Sciences, Chiba University
National Science Series 5: 99–140.

Accepted for publication 28 March 2009

Appendix 1. Elevation ranges of lichen
species of Nepal.
Name of lichen species

Altitudinal
range (m)

Alectoria ochroleuca
Allocetraria flavonigrescens*
A. globulans
A. oakesiana
A. sinensis
A. stracheyi
Anthracothecium himalayense
A. leucostomum
Arctocetraria nigricascens
Arctoparmelia subcentrifuga
Aspicilia cinerea
Awasthia melanotricha*
Bacidia millegrana
B. nigrofusca
B. personata
B. rubella
B. spadicea
Baeomyces pachypus
B. roseus
Bryonora castanea var. castanea
B. castanea var. euryspora
B. curvescens
B. pulvinar var. microspora*
B. pulvinar var. pulvinar*
B. pulvinar*
B. reducta
B. rhypariza var. cyanotropha
B. rhypariza var. lamaina
B. rhypariza var. rhypariza
B. selenospora*
B. stipitata*
B. yeti
Bryoria bicolor
B. confusa

4600–5100
4600–4800
3700–4700
3900–4200
4830
3000–5000
1650
2400–2600
3800
4600
4300
4350–4510
1800–3200
3100
300–1800
300
2600
2000–4200
2000–2500
3750
4420
3500–5000
4720–5070
4830–5100
4720–5390
3200
3750–3900
5100–5200
4000–4100
3800–4400
3000–5080
4190–5200
3000–5050
3450

Appendix 1. Continued
Name of lichen species

Altitudinal
range (m)

B. furcellata
B. himalayana
B. implexa
B. lactinea
B. lanestris
B. levis
B. nadvornikiana
B. nepalensis*
B. nitidula
B. perspinosa
B. poeltii
B. smithii
B. variabilis
Buellia geophila
B. granularis
B. inornata
B. papillata
Bulbothrix isidiza
B. meizospora
B. setschwanensis
Calicium abietinum
C. lenticulare
C. viride
Caloplaca arnoldii
C. aureosora
C. borealis var. borealis
C. borealis var. oligosperma
C. castellana
C. cerina var. cerina
C. cerina var. chloroleuca
C. cerinopsis*
C. cirrochroa
C. cirrochroopsis*
C. citrina
C. cupreobrunnea*

3000
3900–4000
3700–4000
3300–3450
3800–3900
3450
3800–3900
3900–4000
3900
3800–4500
4400
3000–4511
3800–4000
4200
2100
700–900
4650
900–2180
2100–2300
1260–2300
3000–3300
2250
3000–3300
3500–4000
2500–3000
3650
4000
4000–5540
3200
4950–5050
1800–3200
3900–4500
1480
3750
2700

92

THE LICHENOLOGIST
Appendix 1. Continued

Vol. 42

Appendix 1. Continued

Name of lichen species

Altitudinal
range (m)

Name of lichen species

C. cupulata*
C. epiphyta
C. epithallina
C. exsecuta var. aphanes*
C. farinosa*
C. grimmiae
C. holocarpa
C. holochracea
C. insularis
C. isabellina*
C. leptocheila
C. lithophila
C. lobulascens*
C. lypera*
C. maura*
C. obliterans
C. ochroplaca
C. phoenicopta*
C. praeruptorum*
C. procerispora*
C. rinodinopsis
C. sancta*
C. saxicola var. chamaeleon
C. saxicola var. saxicola
C. saxifragarum
C. tetraspora
C. ulcerata*
C. variabilis
Calvitimela aglaea
C. armeniaca
Candelaria crawfordii
C. sphaerobola
Candelariella aurella
C. coralliza
C. grimmiae
C. himalayana
C. nepalensis
C. sorediosa
C. vitellina var. glacialis
C. vitellina var. vitellina
Canomaculina subsumpta
C. subtinctoria
Canoparmelia aptata
C. ecaperata
C. eruptens
Carbonea vorticosa
Catapyrenium cinereum
C. daedaleum
Catillaria leptocheiloides
Catolechia wahlenbergii
Cetraria ambigua
C. nepalensis*
Cetrelia braunsiana

3200–3800
4400–4500
4350–4450
2060
3000–3500
3500
3700–5000
1700–2300
4600–5540
4100–5200
5000
3200–4850
1500–1800
2000–3000
4950–5000
3200–3700
1800–3000
3200
3200–3650
4100–5000
2000–3000
4000–4340
3200–4000
2900–5000
4950–5000
4400–5000
3500–3750
2000–3500
5000
5000
1300–3000
3000–3900
1600–3500
3800–5000
4250–5000
3700–5100
5000–5400
3700–5200
4900–5400
1600–5540
1200–1500
1500
1500–2000
1300–2350
5000
5000–7400
4300–5080
3900–5080
2100
4500
3400–5390
4500
3150

C. cetrarioides
C. olivetorum
Cetreliopsis rhytidocarpa ssp. langtangi
Chaenotheca brunneola
C. chrysocephala
C. furfuracea
C. hispidula
C. phaeocephala
C. stemonea
Chrysothrix chlorina
Cladia aggregata
Cladonia amaurocraea
C. awasthiana
C. calyciformis
C. cariosa
C. carneola
C. cartilaginea
C. ceratophyllina
C. chlorophaea
C. ciliata
C. coccifera
C. corniculata
C. corymbescens
C. delavayi
C. fenestralis
C. fimbriata
C. fruticulosa
C. furcata
C. humilis
C. laii
C. luteoalba
C. macilenta
C. macroptera
C. mongolica
C. nitida
C. ochrochlora
C. pocillum
C. pyxidata
C. ramulosa
C. rangiferina
C. scabriuscula
C. singhii
C. squamosa
C. stellaris
C. stricta
C. subconistea
C. subulata
C. yunnana
Coccocarpia erythroxyli
C. palmicola
Collema nepalense*
C. poeltii*
C. pulcellum

Altitudinal
range (m)
2000–3000
2850
2880
3000–3556†
2700–3879†
3000
†2947–3250
3000–3869†
3000
3100–3200
3900–4000
4530–5230
950–5100
1500–2700
3100
3900–4100
1100–2000
3000–4600
2000–5350
4480
2000–4000
1650–2300
1500–5100
2200–5230
3900–5100
3900–4250
1500–5100
1700–3900
1200–4250
2700–4600
3200–3660
1500–3800
2700–3900
1600–4500
3900
1500–3500
2000–3600
4000–4100
2000–3366
3400–4500
2845–3000
750–2700
1800–4000
4400–4500
3300–4000
2500–3500
2200–2700
3300–4000
3150
1800
3900–4000
3900–4000
1350–1500

2010

Elevation gradients of lichens in Nepal—Baniya et al.
Appendix 1. Continued

Name of lichen species

C. rugosum
C. subconveniens
C. substipitatum
Dermatocarpon miniatum var. miniatum
D. vellereum
Dibaeis sorediata
Dimelaena oreina
Dimerella lutea
Diploicia canescens
Diploschistes muscorum
D. muscorum subsp. bartlettii
D. muscorum subsp. muscorum
D. nepalensis
D. scruposus
Dirinaria aegialita
D. applanata
D. consimilis
Erioderma meiocarpum
Eumitria pectinata
Evernia mesomorpha
Everniastrum cirrhatum
E. nepalense
E. rhizodendroideum
Flavocetraria cucullata
F. nivalis
Flavocetrariella leucostigma
F. melaloma
Flavoparmelia caperata
Flavopunctelia flaventior
Fuscopannaria poeltii*
F. praetermissa
Glyphis cicatricosa
Graphis scripta
G. subglauconigra
Haematomma puniceum
Heterodermia angustiloba
H. awasthii
H. boryi
H. comosa
H. dactyliza
H. dactyliza f. serpens
H. dendritica
H. diademata
H. dissecta var. dissecta
H. firmula
H. flabellata
H. himalayensis
H. incana
H. isidiophora
H. japonica
H. obscurata
H. pellucida
H. propagulifera

93

Appendix 1. Continued

Altitudinal
range (m)

Name of lichen species

Altitudinal
range (m)

1800
1260
3000–4000
3200
2000–3600
2500
3200–4884
3000–4000
4650
900–2160
900–2100
1500–2160
900
2000–3000
1800
600
600
3000–3050
400–2000
3962–4572
1900–3300
1410–3600
2490–3800
3600–4500
3800–5300
3500–4500
3900–4200
2250–2743
2250–2800
3500
†3805–4000
1600
1600–3366
2000–3200
1800
1800–2300
1900
1500–3000
1500–2500
2100
3800–5150
1800
410–3807†
1500–3000
1200–2200
3100
1500–2000
1800–2100
2100–2250
3000–4000
1400–3000
2500–3992
1500–3000

H. pseudospeciosa
H. punctifera
H. rubescens
H. speciosa
H. togashii
H. tremulans
Hyperphyscia granulata
H. minor
Hypogymnia delavayi
H. hypotrypa
H. vittata
Hypotrachyna adducta
H. crenata
H. exsecta
H. flexilis
H. imbricatula
H. infirma
H. koyaensis
H. majoris
H. neodissecta
H. osseoalba
H. revoluta
H. rhabdiformis
H. scytophylla
H. sinuosa
H. sublaevigata
Immersaria athroocarpa
Ingvariella bisporus
Ioplaca pindarensis
Lasallia freyana
L. pertusa
Lecanora adolfii
L. amorpha*
L. chlarotera
L. chondroderma
L. demissa
L. emodi*
L. formosa
L. garovaglii
L. hellmichiana*
L. himalayae*
L. kirra*
L. lesleyana*
L. meridionalis
L. muralis var. dubyi
L. muralis var. muralis
L. phaeodrophthalma
L. rubina var. australis
L. rugosella
L. sherparum*
L. somervellii
L. sulphurea
L. terestiuscula

150–2100
2250
1800–2800
1650–2100
3000–4000
1500
1200
1200–1400
1800–4080
3600–4050
2800–4200
2100
2500–2800
1800–2300
2160
1400–2600
1000–2100
1500–2100
1650
2250–2550
1100–2200
2000–2250
2500–2800
2250–3250
3500
2250
3900–4340
3750
2900–5200
3800–3900
3800–5050
5540
5000
200
3800–5400
2700–4000
5540
4850–5860
3000–5050
1800–3500
3900–4850
4400–4500
5029–5334
1600–3366
4930
3500–4340
4100–4500
3300–4250
2732–3200
3300–4600
3700–5500
4300–5639
4500–5200

94

THE LICHENOLOGIST
Appendix 1. Continued

Vol. 42

Appendix 1. Continued

Name of lichen species

Altitudinal
range (m)

Name of lichen species

Altitudinal
range (m)

L. tschomolongmae*
Lecidea advena
L. auriculata
L. bella*
L. brachyspora
L. bucculenta
L. diducens
L. epiiodiza
L. fuscoatra var. indecora
L. haerjedalica var. gyrodisca
L. himalaica
L. khumbuensis
L. lactea
L. leptoboloides
L. molybdochroa
L. poeltii
L. secernens
L. silacea
L. steineri
L. tessellata
Lecidella carpathica
L. dimelaenophila
L. stigmatea
Leprocaulon arbuscula
Leproplaca chrysodeta
Leptogium asiaticum
L. askotense
L. azureum
L. brebissonii
L. burnetiae var. burnetiae
L. cochleatum
L. delavayi
L. delavayi f. fuliginosulum
L. isidiosellum
L. javanicum
L. pedicellatum
L. phyllocarpum
L. resupinas
L. saturninum
L. trichophorum
Lethariella cladonioides
Letrouitia domingensis
Lobaria discolor
L. isidiosa
L. kurokawae
L. pindarensis
L. pseudopulmonaria
L. retigera
L. subretigera
Lobothallia alphoplaca
L. praeradiosa
Melanelia tominii
Melanohalea poeltii*

3000–5000
5000–5540
5000–5200
4950–5000
4300–5900
5000–5540
4720
4850
4340
5150–5200
5150–5200
5540
5460
4340–5200
3900–4000
4500–5000
5000
5000
5150–5200
5080–5639
3900–4000
3800–3900
4520–5200
1900–3100
3300–4850
3900–4100
1500–1800
1410
1800–3200
1500–2250
1800
3000–4100
3900–4000
1500
1410–1800
1500–3000
2000–3100
1600
1500–2100
1450–1500
4724
240
3150
2800–3962
1800–3400
2700–4000
2550–4050
1600–3650
2800
4400
4000–4500
3200–3600
4500–4600

Menegazzia terebrata
Mycobilimbia hunana
Myelochroa aurulenta
M. entotheiochroa
M. subaurulenta
Nephroma helveticum var. helveticum
N. nakaoi
Nephromopsis ahtii
N. isidioidea
N. nephromoides
N. pallescens
N. stracheyi
Ochrolechia bryophaga
O. glacialis
O. margarita
O. rosella f. sorediascens
O. subviridis
O. trochophora
Pachyphiale himalayensis
Parmelaria subthomsonii
P. thomsonii
Parmelia adaugescens
P. erumpens
P. latissima var. marmariza
P. masonii*
P. meiophora
P. omphalodes
P. ricasolioides
P. squarrosa
P. submutata
P. sulcata
Parmelina tiliacea
Parmelinella simplicior
P. wallichiana
Parmelinopsis expallida
P. minarum
Parmotrema austrosinense
P. cooperi
P. hababianum
P. maclayanum
P. melanothrix
P. mellissii
P. nilgherrensis
P. praesorediosum
P. pseudonilgherrense
P. pseudotinctorum
P. rampoddense
P. ravum
P. reticulatum
P. sancti–angelii
P. stuppeum
P. tinctorum
P. ultralucens

1300–4000
1650
1410–2250
2000–2200
1950
2160–2800
2800
4090–4530
2900–3100
2400–3600
2400–3048
2700–3300
3900–4000
5000–5200
5000
3500
2900
3000–4000
3900–4000
1800–3200
1800–3366
3300–3600
1900–2200
2900
3000–6100
1800–3350
3500–4500
2700
3000–3800
2800
3000–3366
2100
2000–2900
1800–2400
1800
1500
750–3000
1500–2100
1500–2000
1500
2400–3200
1500
1080–2800
1410–1800
1800–3530
2900–3366
900–2800
1500–2500
1500–2500
1410–2100
1800
240–2250
1500

2010

Elevation gradients of lichens in Nepal—Baniya et al.
Appendix 1. Continued

95

Appendix 1. Continued

Name of lichen species

Altitudinal
range (m)

Name of lichen species

Altitudinal
range (m)

P. yodae*
Peltigera canina
P. dolichorrhiza f. dolichorrhiza
P. dolichospora*
P. elisabethae
P. malacea
P. membranacea
P. polydactylon
P. pruinosa
P. rufescens
P. scabrosa
Pertusaria hemisphaerica
Phaeographina pyrrhochroa
Phaeophyscia endococcina
P. endococcina var. khumbuensis
P. endococcinoides var. megalospora
P. hispidula var. exornatula
P. hispidula var. hispidula
P. lygaea*
P. primaria*
P. pyrrhophora*
P. sciastra
Phlyctella indica
Physcia aipolia
P. caesia
P. clementei
P. dilatata
P. dubia
P. phaea
P. stellaris ssp. intestiniformis*
P. tribacia
P. tribacioides
Physciella nepalensis*
Physconia distorta
P. enteroxantha
P. grisea
P. muscigena
Physma byrsaenum
Placidium squamulosum
Pleopsidium chlorophanum*
Porpidia aerolotera
P. crustulata
P. elegantior
P. hydrophila
P. macrocarpa
Protoparmelia badia var. badia
P. effigurans*
Punctelia borreri
P. rudecta
P. subrudecta
Pyrenula cayennensis
P. immersa
Pyxine coccifera

2300
3150–3300
1800–2250
3000–4100
1350
2700–3300
1900–2400
1950–2920
1800
2100
3800
2400–3366
2100
3800–3900
3900
3900–4000
365
3800–3900
4500–4600
3800–3900
1420
3900–5200
1800
1800–2200
3900–5000
3500–4000
1600–5000
5000
4340–5000
4250–4340
3800–4600
1400
1400
3000–4000
2740
1400–4200
4250–5540
1350
4400–4500
3500–4000
5150–5200
2800–5000
4700
3900–4000
2950–4000
3750–4000
3750
1600–3200
1500–3200
1500–2500
3000–3200
2000
750

P. meissnerina
P. philippina
P. sorediata
Ramalina conduplicans
R. farinacea
R. flabelliformis*
R. hossei var. hossei
R. sinensis
R. subfarinacea
Rhizocarpon geographicum
Rhizoplaca chrysoleuca var. chrysoleuca
R. melanophthalma var. obscura
R. peltata
Sagema potentillae
Sclerophora coniophaea
Solorina bispora
Stereocaulon claviceps
S. foliolosum var. foliolosum
S. foliolosum var. botryophorum
S. foliolosum var. strictum
S. glareosum
S. himalayense
S. myriocarpum
S. paradoxum
S. piluliferum
S. pomiferum
S. sasaki var. sasaki
Sticta henryana
S. nylanderiana
S. platyphylloides
S. praetextata
S. weigelii var. weigelii
Sulcaria sulcata
S. virens
Tephromela aglaea
T. armeniaca
T. glacialis*
T. siphulodes var. siphulodes*
Thamnolia vermicularis
Thelenella luridella
Tremolecia atrata
Tuckneraria laureri
Tylophoron moderatum
Umbilicaria badia
U. cinereorufescens
U. decussata var. decussata
U. decussata var. rhizinata*
U. indica var. indica
U. indica var. nana*
U. krascheninnikovii
U. leiocarpa
U. nenella
U. nepalensis

300
900–1800
1410–1800
2133
2700
3200
1200–2500
2700–4267
1610
4500
2100–5150
4270
3660
4000
3000
3600–4200
4000
3300–4000
3600–4600
2400–3600
4200–4400
2500–5400
3900–5303
2000–3366
2160–5150
2500–4700
3600–4200
3657–3962
1800–3600
1900–3150
2100–3500
800–2250
3150–3300
3000–3600
5000–5200
5000–5200
4500
4830
3869–5455
1410
5000–5860
3200–4900
1800
2550
4250–5200
5100–5500
4950–5000
1800–3150
4200–4300
3700–5100
5150
5100
3600–5100

96

THE LICHENOLOGIST
Appendix 1. Continued

Vol. 42

Appendix 1. Continued

Name of lichen species

Altitudinal
range (m)

Name of lichen species

Altitudinal
range (m)

U. thamnodes
U. trabeculata
U. vellea
U. yunnana
Usnea aciculifera
U. baileyi
U. compressa
U. dendritica
U. galbinifera var. subfibrillosa*
U. himalayana
U. longissima
U. montisfuji
U. nepalensis
U. norkettii*
U. pectinata
U. pseudomontisfuji
U. robusta

3600–5050
5000–5100
3600–5050
2550
1200–2250
1500–2400
2400–2700
1200–3000
2200
1800–3300
2700–3750
2200–3900
3800–4000
3000–3500
3962
3200
2500–3000

U. rubicunda
U. splendeus
U. thomsonii
Xanthoparmelia coreana
X. dentata
Xanthoparmelia isidiosa
X. mexicana
X. nepalensis*
Xanthoria borealis
X. elegans
X. fallax
X. fulva
X. sorediata
X. ulophyllodes var. ulophyllodes

2700
1800–3300
1800–3150
3400–3850
4480–4572
2800–3200
3070–3700
3900
3900–4000
3750–6000
3200–3400
2700–3200
3200–5000
3200–4850†

* species endemic to Nepal
†Louise Olley’s unpublished personal observation

Appendix 2. The elevational gradient of lichen species richness regression analysis
results modelled after different species richness as response variables and their
elevation as predictor variable. The Quasi-poisson family of error fitted in the
GAM model after the cubic smooth spline (s) with approximately 4 degrees of
freedom. (P%0·05)
Response variables

Null df

Res. df

Total species
Endemic species
Crustose species
Foliose species
Fruticose species
Cyanolichens
Green algal lichens
Corticolous
Saxicolous
Terricolous

72
72
72
72
72
72
72
72
72
72

68
68
68
68
68
68
68
68
68
68

D2
0·945
0·915
0·914
0·941
0·961
0·944
0·941
0·970
0·908
0·895

Deviance
3598·1
362·3
1008
1764·9
1252·2
632·2
3047·7
2374·3
1075·7
422·3

Df = degree of freedom, Res. = residual, D2 = regression coefficient of determination

F

Pr (>F)

304·5
205
181·4
288·4
560·9
338·4
280·9
588
190·2
192·8

<0·001
<0·001
<0·001
<0·001
<0·001
<0·001
<0·001
<0·001
<0·001
<0·001