diversity and distribution of terricolous lichens as indicator of habitat heterogeneity and grazing...

8/2/2019 Diversity and distribution of terricolous lichens as indicator of habitat heterogeneity and grazing induced trampling i

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O R I G I N A L P A P E R

Diversity and distribution of terricolous lichens

as indicator of habitat heterogeneity and grazing inducedtrampling in a temperate-alpine shrub and meadow

Himanshu Rai D. K. Upreti Rajan K. Gupta

Received: 16 August 2010 / Accepted: 6 October 2011/ Published online: 20 October 2011 Springer Science+Business Media B.V. 2011

Abstract Lichens are among the most sensitive biomonitors of ecosystem health and

human induced disturbances. Terricolous lichens of ChoptaTungnath (Garhwal, western

Himalaya, India) were analysed for their ability to indicate habitat variability and distur-

bances induced by livestock grazing. Terricolous lichens were sampled from 12 sites,

distributed across the three macrohabitats between 2,700 and 4,001 m, using 50 9 10 cm

narrow frequency grids having five 10 9 10 cm sampling units. The terricolous lichen

community of the area constituted, 20 species belonging to 10 genera, five families andfour growth forms. Altitude and relative humidity were the major habitat factors found

influencing the terricolous lichen community of the landscape. Fruticose and compound

soil lichen growth forms were found indicative of habitat disturbance largely caused by

grazing induced trampling. Terricolous lichen diversity of the area was delimited by

grazing pressure at mid-altitudes (3,0003,400 m) and by decreasing soil cover at higher

altitudes ([3,400 m).

Keywords Bioindicators Disturbances Garhwal Himalaya Grazing Trampling

Abbreviations

m Meter

cm Centimeter

H. Rai D. K. Upreti (&)Lichenology Laboratory, Plant Biodiversity and Conservation Biology Division, National BotanicalResearch Institute (CSIR), Rana Pratap Marg, Lucknow 226 001, Indiae-mail: [email protected]

H. Rai R. K. GuptaDepartment of Botany, Pt. L. M. S. Govt. P.G. College, Rishikesh, Dehradun,Uttarakhand 249201, India

123

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Introduction

Soils inhabited by terricolous lichens are good indicators of ecosystem stability and con-

tinuity (Will-Wolf et al. 2002). Requirements of substratum and environmental stability

make terricolous lichens highly sensitive indicators of various environmental perturbations(Eldridge and Tozer 1997; Grabherr 1982; Scutari et al. 2004; Lalley and Viles 2005;

Motiejunaite and Fautynowicz 2005; Lalley et al. 2006; St. Clair et al. 2007).

Although considerable attention has been accorded to documenting the lichens of India

(Upreti 1998; Awasthi 2007; Singh and Sinha 2010), investigations of their community

ecology have only recently begun, and those so far undertaken have not explored the

potential of terricolous lichens as a indicator group (Negi 2000; Negi and Upreti 2000;

Pinokiyo et al. 2008).

Terricolous lichens, constitutes about 9% of total lichen species recorded from India and

their distribution ranges from temperate (1,5003,000 m) to alpine habitats (C3,000 m).

Most important for these are alpine pastures in the Himalayas locally known as Bugyals.The native nomadic communities bring their livestock to the Bugyals in summer which

exerts both grazing and trampling pressures to the habitats. Though there are sporadic few

scattered studies on grazing pressures in temperate and alpine habitats of the region, they

lack any reference of lichens (Nautiyal et al. 2004).

Terricolous lichens, by virtue of their direct occurrence on soil, competition with other

ground vegetation and sensitivity to anthropogenic pressures are suitable indicators for

alpine regions (St. Clair et al. 2007). Here we report on a study of the distribution of

terricolous lichens in a temperatealpine habitat of Garhwal Himalayas, in order to assess

their potential as bioindicators of landscape heterogeneity and grazing induced trampling.

Materials and methods

Study area

ChoptaTungnath lies between 302803900302905100N and 79120900791302100E in the

Garhwal Himalayas of Rudraprayag district of Uttarakhand, and is a constituent habitat of

the Kedarnath wildlife sanctuary (Fig. 1). The area is famous for the 1,000 year-old holy

shrine of Tungnath, situated at 3,680 m on the ridge dividing the waters of the riversMandakini and Alaknanda (Fig. 1); the Tungnath temple is the highest Hindu shrine

dedicated to Lord Shiva.

The shrine and associated alpine grassland lies about 2 km below the Chandrashila Peak

(4,001 m). The road to Chopta is just below this ridge and provides the shortest bridle

approach path to Tungnath from Chopta, of about 4 km. The landscape of the area is

typically mountainous, with moderate to steep slopes (3060) and elevation rising from

2,700 to 4,001 m. The topography comprises ridges and exposed rocks with patches of flat

alpine grassland. The surface geology is of crystalline and metamorphic weathered bedrock

with sedimentary deposits formed during the Paleozoic. Soils are coarse-textured loam orsandy loam at lower altitudes, sandy at higher altitudes, and are well drained and acidic

(Sundriyal 1992). The climate is characterised by severe frosts, blizzards and hailstorms

for most of the year. Precipitation occurs as snow, hail, heavy rain and showers over the

year. Snowfall is from November to April, and snowmelt occurs during April and May

providing an abundance of soil water prior to the monsoon period. Maximum rainfall is in

JulyAugust (Nautiyal et al. 2001).

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The maximum monthly temperatures in the area varies from 1937C in the months of

MayOctober, while minimum temperatures as low as -15C are recorded during

DecemberFebruary (Negi 2000). The vascular plant vegetation is broadly temperate, with

mixed oak and coniferous forest, through subalpine forest, to alpine scrub or grassland

along an altitudinal gradient (Gadgil and Meher-Homji 1990).

Landuse by the local human population is mainly semipastoral agriculture based on

livestock grazing, agriculture, and the collection of fodder and fuel from Oak-Rhodo-

dendron woodlands. Open grasslands (Bugyals) at 2,9003,400 m are extensively used as

pasturelands by native nomads (Bhotiyas). In addition, land around the Tungnath shrine

and at Chopta is occupied during the pilgrimage season. Livestock grazing peaks during

MayOctober. About 4,0005,000 sheep/goat, 5070 buffalo, and 3040 horses and mules,

reach the alpine pastures of the area and cause heavy grazing pressure (Nautiyal et al.

2004). This grazing pressure peaks at mid-altitudes (3,000 to B3,400 m), where the

landscape is dominated by open alpine grass lands with gentler slopes than present at lower

altitudes (2,700 to B3,000 m). Terricolous lichens constitute 1.2% of total lichen biota ofthe area (Negi 2000). The most favoured microhabitats for soil lichens in the area are soil

patches amongst rocks, rock crevices, and the ground.

Field methods and data recording

Sampling of terricolous lichens was carried out in September 2008, along the bridle

approach path from Chopta to Chandrashila summit and in the intervening alpine pastures.

A stratified random sampling method was employed (Greig-Smith 1983; Krebs 1989). The

study area was stratified into three macrohabitat types based on the dominant vegetation

along the elevation gradient (Table 1).Twelve sites were selected from across the three macrohabitats (Table 1; Fig. 2). As the

study area is subjected to frequent trampling by grazing livestock, site selection was

dependent upon the availability of lichen rich patches in the landscape. Three land use

types were recognized in the 12 sampling sites (Table 1). Among these three land use types

(i.e. human settlement, pasture, and grassland), pasture predominated (Table 1).

Fig. 1 Location map of the study area

Biodivers Conserv (2012) 21:97113 99

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A frequency grid of 10 9 50 cm, divided into five sampling units of 10 9 10 cm was

used for lichen sampling (Scheidegger et al. 2002). The quadrat frame of the sampling unit

(10 9 10 cm) was marked off at 1 cm intervals to assist in estimating soil cover (Beymer

and Klopatek 1992). Three such grids were laid at each of the sampling sites; i.e. 15

10 9 10 cm sampling units were laid at each sampling site. Data on macroscale envi-

ronmental variables (temperature, elevation, and relative humidity) were taken for mac-

rohabitats/sites. Elevation was measured with a hand-held GPS unit (Garmin GPS 76S),

and relative humidity and temperature were recorded using a hygrometer (Huger-3123,

85 mm) and a maximum/minimum thermometer, respectively.

The lichen samples collected were examined and identified in the Lichenology Labo-

ratory of the National Botanical Research Institute, Lucknow, Uttar Pradesh, India. Lichens

were identified to species using a stereomicroscope, light microscope (morpho-anatomi-

cally), and chemically with the help of spot tests, UV light and standardized thin-layer

Table 1 Stratification, land use pattern and grazing pressure in sites of the three macrohabitats of ChoptaTungnath landscape

Macrohabitat Altituderange (m)

Geomorphic features Characteristicvegetation

Sites

Abbr. Landuse

GP

Oak-Rhododendron

mixed forest(ORMF)

2,7003,000 Temperate topographywith steep increase inaltitude; forest floorcovered with litter;rich in soil cover; flatpastureland less thanof mid-altitudes

Dominant vegetation ofQuercus

semecarpifolia

forming the uppervegetational stratafollowed by

Rhododendron spp.forming the lowerstory of vegetational

strata

C1 Hs 0

C2 Hs 1

C3 Ps 0

Rhododendron

rich middlealtitudesubalpineforest (RMSF)

3,0003,400 Subalpine topographywith increasing rockcover; moderate risein slope; less litter onfloor; adequate soilcover and flat plainsfor flourished growthof grasslandvegetation resultingin highestconcentration of flat

pasturelands

Dominant vegetation ofRhododendron

arboreum andRhododendron

campanulatum, alongwith isolated standsof Quercussemecarpifolia andfew Abies pindrowand Taxus baccatatrees

T1 Ps 4

T2 Ps 5

T3 Ps 8

T4 Ps 7

T5 Hs 3

Higher altitudealpinegrassland(HAGL)

3,4004,001 Open alpine grasslandwith massive exposedboulders; land-slidedominant area withvery scarce soil cover

Dominant vegetation ofherb species of

Anemone, Potentilla,Aster, Geranium,Meconopsis, Primulaand Polemonium, andscattered pockets ofshrubs of

Rhododendron,Anthopogon andJuniperus species

Cd1 Ps 2

Cd2 Gl 1

Cd3 Gl 0

Cd4 Gl 0

Hs Human settlements, Ps pastures, Gl grassland, GP grazing pressure (number of livestock heard)


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chromatography (Elix and Ernst-Russel 1993; Orange et al. 2001). Keys and monographsemployed included those of Ahti (2000), Awasthi (2007), McCune and Rosentreter (2007),

Rosentreter et al. (2007), and Saag et al. (2009). Representative specimens are deposited at

the lichen herbarium (LWG), National Botanical Research Institute (NBRI), Lucknow,

Uttar Pradesh, India.

Soil chemistry was recorded with reference to soil pH and soil bulk density. Soil pH was

measured using a digital pH meter (Hanna pHep HI98107) calibrated at pH 7 and 4, whereas

soil bulk density was measured by the core method (Arshad et al. 1996). Soil cover and data

regarding lichen species richness at all collection sites, their growth form diversity and associ-

ations with other ground vegetation, were carefully recorded. Grazing pressure was measured as

the number of livestock heard encountered at each site of sampling. A majority of heardcomprised 3035 individuals, and only two at lower altitudes (2,7003,000 m) were comprised

of 60 and 65 individuals, the lower limit of 3035 individuals was considered as one heard.

Data analysis

Terricolous lichen assemblage was quantitatively analysed for frequency, with reference to

species richness (number of species), growth form diversity, and associated plants in each

site (Curtis and McIntosh 1950; Pinokiyo et al. 2008). An indirect gradient ordination

method, principal component analysis (PCA), was used to summarise the compositional

differences between the sites (Gauch 1982; ter Braak and Prentice 1988; Pinokiyo et al.

2008). Pearsons correlation coefficients were calculated to compare explanatory variables

(i.e. altitude, temperature, relative humidity, soil cover, and grazing pressure) and response

variables (PCA axis score, soil pH and soil bulk density, associations with plants, species

richness and growth forms (Pinokiyo et al. 2008). Terricolous lichen groups were sought

through hierarchical cluster analysis (Ludwig and Reynolds 1988; Jongman et al. 1995)

Fig. 2 Terrain map of the study area showing the location of sampling sites (refer Table 1 forabbreviations)


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using Bray-Curtis distances and unweighted pair-group moving average (UPGMA) on

three criteria: distributions in all 12 sites, growth form type, and association with plants

(Scutari et al. 2004). The absolute constancy of each species in the study was calculated as

the number of sites in which the given species was present (Mueller-Domdois and

Ellenberg 1974; Scutari et al. 2004). Except PCA and cluster analysis, which were per-formed using multivar option in PAST 1.92 (Hammer et al. 2001; Hall 2005), all other

statistical analysis were made using SPSS (PASW) 17.

Results

Average community structure and patterns

The terricolous lichen assemblage recorded from the 12 sites in the ChoptaTungnath

landscape consisted of 20 species belonging to 10 genera and five families (Table 2).Soil lichens were found in 57 of the 180 sampling units laid (Table 2). A gradual

decrease in number of sampling units containing soil lichens was observed with increasing

altitude (Table 2). Cladoniaceae and Stereocaulaceae were the dominant families, fol-

lowed by Parmeliaceae, Physciaceae, and Ramalinaceae (Table 2). Four lichen growth

forms, leprose, foliose, fruticose, and combined (squamules as the primary thallus bearing

erect fruticose podetia) were encountered (Table 2). Soil lichens were preferentially

associated with vegetation (mosses, grasses, and other herbaceous flowering plants); only

one species (Everniastrum cirrhatum) was found growing without any association. There

were substantial differences in species constitution and abundance between the sites(Table 2). The majority of sampling units consisted of a single lichen species per unit.

Though there was no significant linear relationship between altitude and soil lichen

species diversity, polynomial fits to the data suggest that terricolous lichen diversity

decreases at mid-altitude (3,0003,400 m) and the distribution of data points along the

altitudinal gradients indicates that there were more terricolous lichen species at lower

altitude (2,7003,000 m), than the mid-and higher ([3,400 m) altitudes (Fig. 3).

Terricolous lichen species showed restricted distributions, and only three species

(Cladonia coccifera, C. pyxidata, and Stereocaulon foliolosum) occurred in five sites,

whereas 12 species (60%) were confined to a single site (Table 2). Cladonia scabriuscula

was recorded only from one site, in the high altitude macrohabitat (HAGL) (Table 2).Physicochemical characteristics of the landscape varied along the altitudinal gradient;

there was a gradual decrease occurred in the ambient temperature (maximum and mini-

mum) and relative humidity (%) (Fig. 4).

Among the various soil parameters assessed, soil cover decreased with increasing

altitude (Fig. 5), whereas soil pH and soil bulk density increased along the altitudinal

gradient from Chopta to Chandrashila (Fig. 6).

Principal community determinants

The dominant and co-dominant terricolous lichens differed within each site of the three

macrohabitats indicating distinct lichen assemblages in different macrohabitats (Table 2),

which was also confirmed by PCA (Fig. 7).

The PCA analysis required 13 components (axis) to account for 100% variation in the data

set. The first three axis of PCA explained 77% of variance, and each axis explained 38, 24 and

15%, respectively. The first two axes explained 63% of variance in the study. PCA axis 1 was


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Table2

Terricolous

lichenspeciesrecordedinthesamplingsitesofthreemacrohabitatsinChoptaTungnathlandscape

Species

Family

Morphological

group

ORMF

RMSF

HAGL

Sites

C1

C2

T1

T2

T3

T4

T5

Cd1

Cd2

Cd3

Cd4

Bryoriaconfusa

Parmeliaceae

Fr

X

X

Cetreliaolivetorum

Parmeliaceae

Fo

X

Cladoniacartilaginea

Cladoniaceae

Cd

X

X

Cladoniacoccifera

Cladoniaceae

Cd

D2(26.6

7)

D2(26.67

)

X

D2(13.3

3)

Cladoniafurcata

Cladoniaceae

Fr

X

X

Cladoniapyxidata

Cladoniaceae

Cd

D1(26.6

7)

D3(13.3

3)

D1(33.33

)

D2(13.3

3)

X

Cladoniascabriuscula

Cladoniaceae

Cd

X

Cladoniasubulata

Cladoniaceae

Cd

D3(13.3

3)

Everniastrumcirrhatum

Parmeliaceae

Fo

X

X

Heterodermiahypocaesi

a

Physciaceae

Fo

D2(20)

X

Heterodermiaobscurata

Physciaceae

Fo

X

Leprariacaeseoalba

Stereocaulaceae

Lp

X

Leprarianeglecta

Stereocaulaceae

Lp

D3(13.3

3)

X

D3(13.3

3)

Melaneliastygia

Parmeliaceae

Fo

X

Physconiagrisea

Physciaceae

Fo

X

Ramalinahossei

Ramalinaceae

Fr

X

X

Stereocaulonalpinum

Stereocaulaceae

Fr

X

X

X

Stereocaulonfoliolosum

var

strictum

Stereocaulaceae

Fr

D3(13.3

3)

D3(13.3

3)

X

D1(20)

X

Stereocaulonmassartian

um

Stereocaulaceae

Fr

X

Stereocaulonpomiferum

Stereocaulaceae

Fr

X

Samplingunitswithsoil

lichens

11

3

8

2

4

9

5

1

1

6

3

4


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Table2

continued

Species

Family

Morphological

group

ORMF

RMSF

HAGL

Sites

C1

C2

T1

T2

T3

T4

T5

Cd1

Cd2

Cd3

Cd4

Speciesrichness

(noofspecies)

9

2

5

4

1

2

5

1

1

3

3

5

XPresentinsite,

D1do

minantspecieswithinmacrohabitat,D2primaryco-dominantspecieswithin

macrohabitat,D3secondaryco-domin

antspecieswithinmacrohabitat,Lpleprose,

Fofoliose,

Frfruticose,

Cdcompound,

ORMFOak-R

hododendronmixed

forest,

RMSFRhododendronrichmid

dlealtitudesubalpineforest,

HAGLhigheraltitudeAlpinegrassland;valuesinparenthesesare

frequencyvalueofspec

iesforeachsite


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significantly correlated with combined growth forms and associative growth with grasses,

indicating their primary influence in differentiating soil lichen communities (Table 3). PCA

axis 2 was significantly correlated to altitude, relative humidity (%), species richness, growth

forms (leprose and fruticose), association with other plants (mosses, grasses, and other her-

baceous flowering plants), soil bulk density and grazing pressureindicating a secondary

influence in differentiating the lichen communities (Table 3).

Fig. 3 Soil lichen species richness (number of species) of all sites, arranged according to macrohabitats(see Table 1 for abbreviations). The dashed line is a best-fit trend polynomial

Fig. 4 Environmental attributes of macrohabitats (temperature and relative humidity), values are inmean standard error (see Table 1 for abbreviations)


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The altitude, temperature and relative humidity were significantly correlated with soil

parameters (bulk density and soil cover), diversity of growth forms, and grazing pressure

(Table 3). While there was no significant correlation between soil pH and ecological and

morphological elements of the soil lichen community, the soils bulk density and soil cover

Fig. 5 Percentage soil cover estimates in all 12 sites, arranged according to macrohabitats (refer Table 1for abbreviations). Values are in mean standard error (see Table 1 for abbreviations)

Fig. 6 Physico-chemical parameters estimated all 12 sites, arranged according to macrohabitats (seeTable 1 for abbreviations). Values are in mean standard error (see Table 1 for abbreviations)


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were significantly correlated with total lichen diversity and other ecological and mor-

phological elements of the terricolous lichen community of the study area (Table 3).

Lichen groups

Hierarchical cluster analysis resulted in three major clusters (IIII) which were subse-

quently divided into two, five and four groups, respectively (Table 4; Fig. 8).

Major cluster I, made up of two soil lichenized species (i.e. S. massartianum and

C. scabriuscula), in single site and associated with mosses was separated in two groups (A and

B) on the basis of different growth forms (fruticose and combined) types (Table 4; Fig. 8).

Major cluster II, constituted by nine lichenized species ( Bryoria confusa, Cladonia carti-laginea, C. coccifera, C. furcata, C. pyxidata, C. subulata, Heterodermia hypocaesia, Lep-

raria neglecta and S. foliolosum), was further divided into five groups (CG) primarily on the

basis of increasing constancy of lichenized species, increasing diversity in associations with

plants and dominance of combined growth form along the hierarchy (Table 4; Fig. 8). Major

cluster III, with nine species (E. cirrhatum, Ramalina hossei, S. alpinum, S. pomiferum,

Cetralia olivetorum, L. caeseoalba, H. obscurata, Physconia grisea, and Melanelia stygia),

was further divided into four groups (HK) on the basis of decreasing constancy, selectivity

for associations with mosses, and dominance of foliose growth forms (Table 4; Fig. 8).

Discussion

High altitude ecosystems and their vegetation are usually considered fragile and sensitive

to human impact (Grabherr 1982). Although soil is the major substratum available for

lichen colonization in the ChoptaTungnath landscape, it harbors fewer lichen species than

Fig. 7 PCA ordination plot of 12 soil lichen study site data in three macrohabitats of ChoptaTungnathlandscape, values in parentheses are PC axis scores for axis 1 and 2, respectively (see Table 1 forabbreviations)


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Table 3 Pearsons correlation coefficients between PCA axes and selected variables (significant correla-tions are tagged)

PC1 PC2 Alt MXT MNT RH SC Gz

Alt 0.210 0.638a

MXT -0.231 -0.295 -0.698a

MNT -0.229 -0.325 -0.727b 0.999b

RH -0.019 -0.689a -0.768b 0.218 0.264

SC -0.179 -0.210 -0.681a 0.908b 0.906b 0.193

Gz -0.314 0.581a 0.379 0.197 0.156 -0.791b 0.260

pH 0.075 0.248 0.270 0.077 0.063 -0.280 0.113 0.268

BD 0.055 0.667a 0.889b -0.726b -0.745b -0.578a -0.753b 0.196

SPN -0.225 -0.805b -0.501 0.286 0.305 0.459 0.205 -0.325

Lp-

0.276-

0.741

b-

0.585

a

0.498 0.507 0.304 0.486-

0.069Fo 0.103 -0.527 -0.251 0.186 0.204 0.403 0.071 -0.450

Cd -0.928b 0.168 -0.043 0.240 0.232 -0.122 0.176 0.466

Fr -0.148 -0.713b -0.679a 0.244 0.270 0.593a 0.256 -0.469

Ms -0.529 -0.659a -0.569 0.419 0.439 0.524 0.370 -0.186

Gr -0.697a 0.638a 0.315 0.076 0.050 -0.527 0.069 0.749b

Ag -0.451 -0.635a -0.690a 0.623a 0.634a 0.380 0.583a 0.076

pH BD SPN Lp Fo Cd Fr Ms Gr

Alt

MXT

MNT

RH

SC

Gz

PH

BD 0.090

SPN 0.105 -0.651a

Lp 0.036 -0.783b 0.868b

Fo 0.197 -0.372 0.665a 0.369

Cd -0.033 0.047 0.082 0.143 -0.084

Fr 0.000 -0.621a 0.721b 0.654a 0.246 -0.176

Ms 0.266 -0.591a 0.808b 0.751b 0.482 0.388 0.639a

Gr 0.175 0.351 -0.289 -0.209 -0.350 0.835b -0.447 -0.036

Ag 0.052 -0.793b 0.732b 0.875b 0.170 0.322 0.634a 0.818b -0.016

Alt Altitude, MXT maximum temperature, MNT minimum temperature, RH relative humidity(%), pH soilpH, BD soil bulk density, SCsoil cover (%), SPNnumber of species of soil lichens, Lp leprose growth form,Fo foliose growth form, Cd compound growth form, Fr fruticose growth forms, Ms soil lichen associations

with mosses, Gr soil lichen associations with grasses, Ag soil lichen associations with other angiospermicvegetation, Gz grazing pressure (number of live stock heard at each sampling site)a Correlation is significant at the 0.05 level (2-tailed)b Correlation is significant at the 0.01 level (2-tailed)


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the rock and wood releves. This disparity can be attributed to the instability of the ter-

restrial niche due to livestock grazing as the majority of the landscape is used as pasture

and experiences high trampling pressure which affects some key soil properties. The steepincrease in pH at mid-altitudes (3,0003,400 m) can be attributed to loss of humus by

trampling due to livestock grazing (Grabherr 1982). Trampling also increased soil bulk

density, which negatively influenced soil cover, total soil lichen diversity, and diversity of

particular growth forms (leprose, fruticose and compound). Further, the soil cover decrease

along the increasing altitudinal gradient was an additional factor constraining soil lichens

in the study area. Therefore species richness of soil lichens in the study area was deter-

mined both by grazing pressure and decrease in soil cover (Ahti 1959; Klein 1987; Helle

and Aspi 1983; Joly et al. 2009).

The correlation of ordination data (PCA axis scores) with the soil lichen communitys

morphological elements (growth forms) indicated that compound and fruticose were the

main defining growth forms for the soil lichen community of the area. The negative

significant relationship of these growth forms with PCA scores suggests their sensitivity to

grazing induced trampling (Grabherr 1982; Wolseley 1995). Further (as suggested by

PCA) the strong associative correlation of soil lichens to the ground vegetation, especially

mosses, can be attributed to their ability to tolerate longer periods of desiccation along with

Table 4 Constancy and frequency (%) of growth forms and associations of soil-lichens in all the 12 sites, inChoptaTungnath landscape

Lichengroup

Species Constancy Growth forms Associations

I A Stereocaulon massartianum 1 Fr (0.56) Ms (0.56)

B Cladonia scabriuscula 1 Cd (0.56) Ms (0.56)

II C Cladonia subulata 1 Cd (0.56) Ms (0.56)

D Heterodermia hypocaesia 1 Fo (0.56) Ms (0.56)

Bryoria confusa 1 Fr (0.56) Ms (0.56)

E Cladonia cartilaginea 2 Cd (1.11) Ms (0.56), Ag (0.56)

Cladonia furcata 1 Fr (0.56) Ms (0.56)

F Cladonia coccifera 5 Cd (6.11) Gr(6.11), Ag (0.56)

Cladonia pyxidata 5 Cd (7.22) Ms (6.11), Gr (2.22), Ag (2.78)

G Lepraria neglecta 3 Lp (3.33) Ms (3.33), Gr (1.11), Ag (2.78)

Stereocaulon foliolosum 5 Fr (5.00) Ms (5), Ag (0.56)

III H Everniastrum cirrhatum 2 Fo (1.11) Na (2.78)

I Ramalina hossei 2 Fr (1.11) Ms (1.11)

Stereocaulon alpinum 3 Fo (0.56) Ms (1.11)

J Stereocaulon pomiferum 1 Fr (0.56) Ms (0.56)

Cetrelia olivetorum 1 Fo (0.56) Ms (0.56)

Lepraria caeseoalba 1 Lp (0.56) Ms (0.56)

K Heterodermia obscurata 1 Fo (0.56) Ms (0.56)

Physconia grisea 1 Fo (0.56) Ms (0.56)

Melanelia stygia 1 Fo (0.56) Ms (0.56)

Lp Leprose, Fo foliose, Fr fruticose, Cd compound (squamules as primary thallus bearing erect fruticosebody as secondary thallus), Ms associative growth with mosses, Gr associative growth with grasses, Agassociative growth with other angiospermic herbaceous vegetation, Na non associative growth. Species arearranged according to groups resulting from cluster analysis. Values in parentheses are overall frequencyvalues for all sites studied


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the soil lichens (Sheard 1968; Rosentreter and Belnap 2003). Among the habitat factors,

the significant correlation of altitude and relative humidity with PCA axis scores empha-

sized their role in modulating the lichen community in the terrestrial domain of the area.

Hierarchical cluster analysis also fell in line with PCA where the three major clusters

were formed on the basis of dominance of compound and fruticose growth forms in the

first two clusters (I and II), and dominance of foliose growth forms in the last one (III). The

first two clusters contained soil lichen species, mainly distributed in highly grazed mac-

rohabitats of the landscape which further legitimize their position as indicators of distur-bances in the area (Grabherr 1982).

Though some studies on lichens of the region mention terricolous lichens but their role

as indicator of habitat variability and grazing disturbances is first time reported (Negi and

Gadgil 1996; Upreti and Negi 1998). A lower turnout of lichenized species on terrestrial

domains in this study is due to perturbation these regions are encountering in recent

decade. These high altitude grasslands are subjected to variety of pressures i.e. land-use

change, invasive species, elevated temperature and CO2 which are changing the very

skeleton the vegetational dynamics of these habitats. As terricolous lichens contribute in

nitrogen dynamics, surface soil stabilization, and significant source of carbon input inecosystem (Evans et al. 2003), studies on change in their communities in Himalayan

ecosystems can give a fare picture of the perturbations these regions are facing. In long

term gradual changes in cover and species composition of terricolous lichens, due to

elevated temperature, CO2 and UV radiation can be an efficient estimate of effect of global

climate change in Himalaya (Evans et al. 2003; Rai et al. 2010).

Fig. 8 Groups of terricolous lichens resulting from hierarchical cluster analysis based on ecological (sitedistribution and association with other ground vegetation) and morphological characters (growth form type)


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Conclusion

The temperate-alpine meadows of Garhwal Himalaya are representative of western

Himalayan alpine shrub and meadows communities. These are delimited by moister

eastern Himalayan alpine shrub and meadows in the east, and to the north grade into thedrier Central Tibetan Plateau alpine steppe. These habitats harbour most of the open

grasslands which are subject to high grazing pressures. Our investigation reveals the major

factors influencing the soil lichen community of the ChoptaTungnath meadows. The

terricolous lichens proved to be appropriate bioindicators of habitat heterogeneity, and

grazing pressure.

We consider that these findings can be confidently extrapolated for conservation and

management practices of these high altitude Himalayan pastures. Management practices of

the area should therefore focus on reduction of grazing pressure; rehabilitation of highly

disturbed area could also be considered but these are only likely to be effective in the long-

term due to slow rates of colonization by lichens. A controlled utilization and sustainableuse of the terrestrial resources must be encouraged, in order to decrease further deterio-

ration of these already fragile biomes. Though the finding presented here are from a

relatively small area, the study suggests that region-specific monitoring of terricolous

lichen communities can be utilised in the development of conservation and management

practices.

Acknowledgments We are grateful to Director, National Botanical Research Institute, Lucknow andUniversity Grant Commission, India, for providing necessary laboratory facilities and financial assistance.This research work is dedicated to Late Dr. D.D. Awasthi (28 Sept. 192221 Aug. 2011) pioneer Lichen-

ologist, whose work laid foundation for systematic lichenological research in Indian subcontinent.

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