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HUSSAIN et al. 167 Tropical Ecology 49(2): 167-181, 2008 ISSN 0564-3295 © International Society for Tropical Ecology www.tropecol.com Species composition and community structure of forest stands in Kumaon Himalaya, Uttarakhand, India M. SHAH HUSSAIN* # , AISHA SULTANA # , JAMAL A. KHAN & AFIFULLAH KHAN Department of Wildlife Sciences, Aligarh Muslim University, Aligarh 202 002, India Abstract: The paper describes species composition and community structure of 23 forest stands in Kumaon Himalaya (28° 43' 55" to 30° 30' 12" N and 78° 44' 30" to 80° 45' E), between altitudes 1500-3000 m. A total of 902 plots was sampled following plot sampling method. Density and diversity measures were calculated for different vegetation layers of each stand. TWINSPAN identified 19 tree communities and 17 ground vegetation communities. The distribution of tree species on DCA axis 1 indicated influence of altitudinal gradient while the second axis of DCA indicated canopy cover and shrub diversity. The first component of PCA represented open to close canopy forest, while the second reflected increase in shrub density and diversity. Quercus floribunda-Rhododendron arboreum group had maximum tree density and Abies pindrow-Betula utilis the minimum. Myrsine africana was the dominant shrub species. Maximum tree species diversity and richness were recorded for Daphiadhura site whereas the Vinaiyak site expressed maximum shrub diversity. Pinus wallichiana, Betula utilis, Tsuga demosa, etc. were found to be rare tree species. Resumen: El artículo describe la composición de especies y la estructura de la comunidad de 23 rodales de bosque en Kumaon Himalaya (28° 43' 55" a 30° 30' 12" N, y 78° 44' 30" a 80° 45' E), ubicadas entre las altitudes de 500-3000 m. En total se muestrearon 902 parcelas, utilizando el método de muestreo con área. Se calcularon medidas de densidad y diversidad para las diferentes capas de vegetación de cada rodal. El análisis TWINSPAN identificó 19 comunidades arbóreas y 17 comunidades del suelo del bosque. La distribución de las especies arbóreas en el eje 1 del DCA indicó una influencia del gradiente altitudinal, mientras que el segundo eje del DCA indicó la cobertura del dosel y la diversidad de arbustos. El primer componente de un PCA representó bosque de dosel abierto a cerrado, mientras que el segundo reflejó el incremento en la densidad de arbustos y su diversidad. El grupo Quercus floribunda-Rhododendron arboreum tuvo la máxima densidad de árboles, y el de Abies pindrow-Betula utilis, la mínima. Myrsine africana fue la especie de arbusto dominante. Los valores máximos de diversidad y riqueza de especies arbóreas fueron registrados en el sitio Daphiadhura, mientras que en el sitio Vinaiyak se expresó la máxima diversidad de arbustos. Se encontró que Pinus wallichiana, Betula utilis y Tsuga demosa, entre otras, fueron especies raras. Resumo: O artigo descreve a composição e a estrutura de comunidade de 23 parcelas florestais em Kumaon Himalaia (28° 43' 55" to 30° 30' 12" N e 78° 44' 30" to 80° 45' E), entre as altitudes de 1500-3000 m. Um total de 902 parcelas foram amostradas seguindo o método de amostragem de quadrados. As medidas da densidade e diversidade foram calculadas para diferentes andares de vegetação em cada parcela. A análise TWINSPAN identificou 19 * Corresponding Author; e-mail: [email protected], [email protected] # Present Address: Centre for Environmental Management of Degraded Ecosystems, School of Environmental Studies, University of Delhi, Delhi 110007, India

Transcript of Species composition and community structure of forest ... Hussain.pdf · HUSSAIN et al. 169...

HUSSAIN et al. 167

Tropical Ecology 49(2): 167-181, 2008 ISSN 0564-3295

© International Society for Tropical Ecology

www.tropecol.com

Species composition and community structure of forest stands in

Kumaon Himalaya, Uttarakhand, India

M. SHAH HUSSAIN*#, AISHA SULTANA#, JAMAL A. KHAN & AFIFULLAH KHAN

Department of Wildlife Sciences, Aligarh Muslim University, Aligarh 202 002, India

Abstract: The paper describes species composition and community structure of 23 forest

stands in Kumaon Himalaya (28° 43' 55" to 30° 30' 12" N and 78° 44' 30" to 80° 45' E), between

altitudes 1500-3000 m. A total of 902 plots was sampled following plot sampling method.

Density and diversity measures were calculated for different vegetation layers of each stand.

TWINSPAN identified 19 tree communities and 17 ground vegetation communities. The

distribution of tree species on DCA axis 1 indicated influence of altitudinal gradient while the

second axis of DCA indicated canopy cover and shrub diversity. The first component of PCA

represented open to close canopy forest, while the second reflected increase in shrub density

and diversity. Quercus floribunda-Rhododendron arboreum group had maximum tree density

and Abies pindrow-Betula utilis the minimum. Myrsine africana was the dominant shrub

species. Maximum tree species diversity and richness were recorded for Daphiadhura site

whereas the Vinaiyak site expressed maximum shrub diversity. Pinus wallichiana, Betula

utilis, Tsuga demosa, etc. were found to be rare tree species.

Resumen: El artículo describe la composición de especies y la estructura de la

comunidad de 23 rodales de bosque en Kumaon Himalaya (28° 43' 55" a 30° 30' 12" N, y 78°

44' 30" a 80° 45' E), ubicadas entre las altitudes de 500-3000 m. En total se muestrearon 902

parcelas, utilizando el método de muestreo con área. Se calcularon medidas de densidad y

diversidad para las diferentes capas de vegetación de cada rodal. El análisis TWINSPAN

identificó 19 comunidades arbóreas y 17 comunidades del suelo del bosque. La distribución

de las especies arbóreas en el eje 1 del DCA indicó una influencia del gradiente altitudinal,

mientras que el segundo eje del DCA indicó la cobertura del dosel y la diversidad de

arbustos. El primer componente de un PCA representó bosque de dosel abierto a cerrado,

mientras que el segundo reflejó el incremento en la densidad de arbustos y su diversidad. El

grupo Quercus floribunda-Rhododendron arboreum tuvo la máxima densidad de árboles, y el

de Abies pindrow-Betula utilis, la mínima. Myrsine africana fue la especie de arbusto

dominante. Los valores máximos de diversidad y riqueza de especies arbóreas fueron

registrados en el sitio Daphiadhura, mientras que en el sitio Vinaiyak se expresó la máxima

diversidad de arbustos. Se encontró que Pinus wallichiana, Betula utilis y Tsuga demosa,

entre otras, fueron especies raras.

Resumo: O artigo descreve a composição e a estrutura de comunidade de 23 parcelas

florestais em Kumaon Himalaia (28° 43' 55" to 30° 30' 12" N e 78° 44' 30" to 80° 45' E), entre as

altitudes de 1500-3000 m. Um total de 902 parcelas foram amostradas seguindo o método de

amostragem de quadrados. As medidas da densidade e diversidade foram calculadas para

diferentes andares de vegetação em cada parcela. A análise TWINSPAN identificou 19

* Corresponding Author; e-mail: [email protected], [email protected] #Present Address: Centre for Environmental Management of Degraded Ecosystems, School of Environmental Studies,

University of Delhi, Delhi 110007, India

168 FOREST STRUCTURE IN KUMAON HIMALAYA

comunidades arbóreas e 17 comunidades de vegetação rasteira. A distribuição das espécies

arbóreas no eixo DCA indicam a covertura pelo copado e a diversidade dos arbustos. A primeira

componente do ACP representou florestas com copado aberto a fechado, enquanto a segunda

reflectiu o aumento na densidade e diversidade dos arbustos. O grupo Quercus floribunda-

Rhododendron arboreum apresentou a densidade arbórea máxima e o Abies pindrow-Betula

utilis o mínimo. A Myrsine africana foi a espécie arbustiva dominante. A diversidade arbórea

máxima e riqueza foi registada para a estação Daphiadhura enquanto a estação Vinaiyak

expressou a diversidade arbustiva máxima. Encontrou-se que a Pinus wallichiana, Betula

utilis, Tsuga demosa, etc. eram espécies arbóreas raras.

Key words: Forest conservation, Himalaya, Himalayan forests, India, ordination,

rare species, vegetation community.

Introduction

Himalaya, the youngest mountain system of

the world, constitutes an important link between

the vegetation of the southern peninsular India

on the one hand, the eastern Malaysian, the

north-eastern Sino-Japanese and the northern

Tibetan areas on the other (Puri et al. 1983).

Several studies have described the vegetation of

Kumaon (Dhar et al. 1997; Rikhari et al. 1989a;

Singh et al. 1984; Singh & Singh 1984; Singh &

Singh 1987; Singh et al. 1987; Tiwari & Singh

1985; Upreti et al. 1985) and Garhwal Himalaya

(Anthwal et al. 2006; Kumar & Bhatt 2006;

Nautiyal et al. 2004). Some of the studies

described altitudinal variation in vegetation

(Adhikari et al. 1992; Saxena et al. 1985) and

reported that vegetation types differ with change

in altitude. However, Puri et al. (1983) observed

that geology and soils may exercise a far greater

influence on the distribution of vegetation types

than the altitude or climate.

Some pioneering contributions on

phytosociology (Ralhan et al. 1982; Saxena &

Singh 1982) and population structure (Saxena et

al. 1985; Singh et al. 1987) of certain central

Himalayan forest types have already been

reported. The main objectives of this paper are: (i)

to describe structural attributes (density, species

diversity and richness) of tree, shrub and ground

layers; (ii) to identify predominating communities

of trees and ground vegetation, and plant species

of special concern, and to further identify the

localities having rare plant communities needing

protection.

Materials and methods

Study area

The study was conducted in 23 forest stands

including Binsar Wildlife Sanctuary (BWS) and

Askot Wildlife Sanctuary (AWS), in five districts;

Almora, Bageshwer, Champawat, Naini Tal and

Pithoragarh of Kumaon Himalaya (28° 43' 55" to

30° 30' 12" N latitude and 78° 44' 30" to 80° 45' E

longitude) covering an area of 21,032 km² in the

Uttarakhand state of India (Fig. 1, Table 1).

Kumaon Himalaya is altitudinally divisible into

subtropical (300 to 1500 m), temperate (1500 to

3500 m) and alpine (>3500 m) zones (Saxena et al.

1985). Annual rainfall peaks at about 1200 m

altitude (4100 mm) and gradually declines to 670

mm at 2700 m (Saxena et al. 1985).

Sampling procedures

Overall 902 sampling plots were laid in the 23

forest stands (details in Table 1). Vegetation was

sampled along existing forest trails which passed

through all the major habitat types to permit

sampling of different habitats in equal proportion

at each site. In each stand, sampling plots were 50

m apart, and each was laid at a distance of 10 m

from the edge of the trail on either side to avoid

sampling of the relatively disturbed vegetation.

Number of plots sampled in different stands is

given in Table 1.

At each sampling plot, a 10 m radius circular

plot (Dombois & Ellenberg 1974) was established.

Trees > 4 m height were considered mature trees.

Species and their individuals were recorded for the

HUSSAIN et al. 169

estimation of density, species diversity and species

richness. Shrub layer was quantified in 3 m radius

circular plots whereas ground vegetation (herbs

and grasses) was estimated in 0.5 m x 0.5 m

quadrats at four places within the 10 m radius

sampling plot. Tree cover was measured at 5 m

distance from the sampling point in four different

directions using gridded mirror of 25.4 x 25.4 cm

(10 x 10 inch) dimension, divided into 25 equal

grids. Grids covered with > 50% foliage were

counted and expressed in terms of percent tree

cover (Dombois & Ellenberg 1974).

Fig 1. Location of 23 surveyed forest stands in Kumaon Himalaya.

170 FOREST STRUCTURE IN KUMAON HIMALAYA

Density and diversity estimates

Density of trees, shrubs and ground vegetation

was calculated following Greig - Smith (1983). The

diversity values for each layer (tree, shrub and

ground vegetation) were calculated using Shannon

- Wiener’s diversity index following Magurran

(1988). The species richness was calculated using

Margelef’s species richness index (Magurran

1988). One-way ANOVA (Zar 1984) was used to

detect significant differences in density, diversity

and richness for the vegetation layers in all the

surveyed forest stands.

Classification and ordination of species and

sites

The vegetation was classified on the basis of

tree species and dominant ground vegetation

(shrub and herb species) using TWINSPAN (Two-

way indicator species analysis) computer program

(Hill 1979 a). Same data matrix was used for

ordination of tree species as well as sites (forest

stands) through Detrended Correspondence

Analysis (DCA) in computer program DECORANA

(Hill 1979 b). The vegetation attributes (density,

diversity and richness) of the 23 sites along with

altitude, number of stumps, lopped trees and cattle

dung were subjected to Principal Component

Analysis (PCA).

Stepwise multiple regression was used to

obtain correlation between the DCA axes, and

measured environmental and vegetation

attributes. All data matrices were standardized

following Zar (1984) to achieve normality and

reduce heteroscadesticity.

Rarity index

A rarity index was generated to identify rare

tree species of Kumaon. For this purpose two

parameters were taken into account-

a) Qualitative proportion of each tree species in

Kumaon (PQA)

Table 1. Location details and distribution of sampling plots in different forest stands of Kumaon

Himalaya. RF = Reserve forest, POF = Privately owned forest, WLS = Wildlife sanctuary, CF =

Community forest, Alt. = Altitude, m = meter, asl = above sea level, BWS = Binsar Wildlife Sanctuary.

Forest stand District Status Sampling

Plots

Alt. Range

(m asl.)

Coordinates

Kilbery Naini Tal RF 40 2085- 2240 29°25'24.3"N 79°2.6'24.3"E

Vinaiyak Naini Tal RF 40 2130- 2290 29°27'45.4"N 79°24'31.8"E

Kunjakharak Naini Tal RF 45 2040- 2430 29°39'N 79°18'58.1"E

Maheshkhan Naini Tal RF 40 1820- 2090 29°24'16.2"N 79°33'50.6"E

Gager Naini Tal RF 40 1860- 2220 29°25'11.4"N 79°30'31.9"E

Mukteshwer Naini Tal RF 49 1800- 2260 29°28'34.1"N 79°38'28.1"E

Jilling Naini Tal POF 20 1860- 2010 29°22'1.6"N 79°37'E

BWS Bageshwer WLS 75 1990- 2260 29°42'3.2"N 79°45'E

Pandavkholi Almora CF 40 2460- 2590 29°48'19.5"N 79°27'E

Sitlakhet Almora RF 15 1880- 1980 29°42'3.2"N 79°45'E

Jageshwer Almora RF 26 2060- 2200 29°39'3.2"N 79°50'52.5"E

Gasi Bageshwer RF 40 2140- 2370 30°04'48.4"N 80°E

Dhakuri Bageshwer RF 55 2470- 2825 30°13'19.5"N 79°55'26.3"E

Wachham Bageshwer RF 50 2410- 2935 30°07'25"N 79°54'37.5"E

Sunderdunga Bageshwer RF 36 2560- 2780 30°13'30.3"N 79°54'18.5"E

Pindari Bageshwer CF 39 2200- 2960 30°11'11.3"N 79°59'30"E

Daphiadhura Pithoragarh WLS 40 2020- 2440 29°54'N 80°20'E

Majtham Pithoragarh WLS 40 1595- 2250 29°53'N 80°22'E

Gandhura Pithoragarh WLS 50 1710- 2045 29°51'40"N 80°14'16.9"E

Sobala Pithoragarh WLS 40 2190- 2650 30°04'16.2"N 80°34'15"E

Duku Pithoragarh WLS 48 1930- 2530 29°56.3'N 80°30'E

Munsiary Pithoragarh RF 25 2655- 2770 30°05'3.2"N 80°14'41.3"E

Mechh Champawat CF 10 1810- 1830 29°16'16.2"N 80°12'18.8"E

HUSSAIN et al. 171

(PQA) = Number of stands the concerned tree

species was encountered/total number of stands (23)

b) Quantitative proportion of each tree species in

Kumaon (PQI)

(PQI) = Number of individuals of each tree

species / number of individuals of all tree species

where the concerned species was encountered

Rarity index for each tree species = PQA + PQI

Fig. 2. TWINSPAN classification of (a) 63 species into 19 groups based

on the tree species data of Kumaon Himalaya, and (b) classification of 23

surveyed forest stands into 8 groups.

(a)

(b)

172 FOREST STRUCTURE IN KUMAON HIMALAYA

Results

Tree species classification

Nineteen broad communities have been

recognised in Kumaon. Overall, 63 tree, 56 shrub,

90 herb and 21 grass species were recorded in the

23 forest stands. A total of five homogenous groups

of tree species, in relation to the environmental

variables, was identified through TWINSPAN

analysis (Fig. 2). The left arm of the first dichotomy

contained 25 species, which was further divided into

two groups. First negative group consisted of four

species which were characteristic of site 8 (for site

code see Fig 1). Second positive group also consisted

of two homogenous groups. The first group

contained Quercus semecarpifolia as a dominant

species at sites 12, 13 and 17. The second group

consisted of Abies pindrow and Taxus baccata as

dominant species and represented mixed coniferous

habitat. Further subdivisions did not provide any

additional ecological information. The right arm of

the first dichotomy had 38 tree species, which were

further divided into two groups. First negative

group contained 21 species. These tree species were

mainly encountered at sites 2, 4, 5, 7, 18, 19 and 23.

At site 2, Q. leucotricophora, Pinus wallichiana and

Cedrus deodara were the dominant tree species

while at site 23 the dominant species was Q.

lanuginosa (Table 2).

Tree species ordination

DCA ordination successfully handled the

variation in tree species communities from low to

high altitude. All the sites and tree species showed

meaningful distribution on axis 1 and axis 2 of

DCA (Fig. 3 a). The first axis (eigen value = 0.389)

represents an altitude gradient (low to high). Sites

14, 15, 16, 20, 21, and 22 occupied extreme end of

the first axis and represented TWINSPAN group 3

(A. pindrow, T. baccata, Betula utilis, Tsuga

Table 2. Communities and their characteristic tree species with their codes (in parentheses used in

DECORANA computer program) in Kumaon based on TWINSPAN classification. Comm. = Community.

Groups Comm. Forest stand Tree species

1 1 BWS Acer caesium (20), Aesculus indica (23), Swida sp.(42),

Betula alnoides (44)

2 2 Daphiadura Gasi, Quercus semecarpifolia (4), Toona serrata (14),

Dodecademia grandiflora (37),

Dhakuri Symplocos sp. (50)

3 3, 4, 5, 6 Pindari, Sobala, Duku

Wachham, Sunderdunga,

Munsiary

Abies pindrow (15), Jugulans regia (34), Prunus cerasoides

(63), Betula utilis (28) Rhododendron barbatum (7), Taxus

baccata (16), Tsuga demosa (35), Pyrus vestita (52),

Zanthoxylum armatum (58), Ficus palmata (46), Morus

serrata (47), Symplocos chinensis (51), Prunus cornuta (60),

Debregeasia hypoleuca (59), Acer cappadocicum (53),

Fraxinus sp. (48), Dendroephthoe falcate (43)

4 7, 8, 9, 10, 11, 12, 13 Vinaiyak, Maheshkhan,

Gager, Jilling, Majtham,

Gandhura, Mechh

Cedrus deodara (17), Cupressus torulosa (25), Cassia fistula

(62), Quercus lanuginose (5), Engelhardia spicata (36),

Ficus auriculata (41), Daphnephyllum himalense (38),

Quercus glauca (3), Pinus roxburghii (19), Myrica esculenta

(24), Maytenus rufa (54), Benthamidia capitata (39),

Phoenix humilis (61), Castanopsis tribuloides (40), Quercus

leucotricophora (1), Pyrus pashia (13), Pinus wallichiana

(18), Euonymus tingens (10), Swida oblonga (27),

Macaranga pustulata (33), Picea smithiana (45)

5 14, 15, 16, 17, 18, 19 Mukteshwer, Kilbery,

Pandavkholi, Sitlakhet,

Jageshwer, Kunjakharak

Litsea umbrosa (30), Populus ciliate (49), Persea duthiei (9),

Rhododendron arboreum (6), Alnus nepalensis (21),

Viburnum mullaha (22), Ilex dipyrena (26), Stranvissia

nausea (57), Meliosma dillenaeafolia (29), Rhamnus

triqueter (56), Quercus floribunda (2), Fraxinus micrantha

(32), Symplocos theifolia (12), Lindera pulcherrima (31),

Lyonia ovalifolia (8), Euonymus pendulus (11), Viburnum

coriacieum (55)

HUSSAIN et al. 173

demosa etc.), while the low altitude sites 4, 10, 18

and 23 represented TWINSPAN groups 4 & 5 (P.

roxburghii, Q. leucotricophora, Pyrus pashia, Q.

floribunda).

The second axis (eigen value = 0.254) appeared

to reflect the canopy cover from open to close. The

species associated with open canopy were Q.

glauca, Q. lanuginosa, Cassia fistula,

Zanthoxylum armatum, Q. semecarpifolia and the

representative sites were 13, 14, 18, 19 and 23,

while close canopy areas were Gasi, Sobala, Duku

and Pandavkholi and the associated tree species

were Q. semecarpifolia, Toona serrata, Symplocos

theifolia (Fig. 3 b). These interpretations are

largely confirmed by the results of PCA. The PC 1

explained 44.26% variance and represented a

gradient of open to close canopy forest with tree

density and diversity in increasing order. PC 2

explained 18.26% of variance and reflected an

increasing pattern in shrub density and diversity.

The distribution of species and sites was same on

the two axes of PCA as on DCA axes (Fig. 3 c).

Fig 3. (a) DCA ordination of tree species of Kumaon Himalaya on two axes extracted by DECORANA

computer program. Numbers refer to different species, (b) DCA ordination of 23 surveyed forest stands on

two axes. DCA axis 1 is related to altitude and slope. DCA axis 2 is related to canopy cover. Numbers

refer to different sites, and (c) PCA ordination of surveyed forest stands on two extracted components. PC

1 is related to canopy cover and diversity of trees. PC 2 is related to the abundance of grasses and shrubs.

Numbers refer to various sites.

(a) (b)

(c)

174 FOREST STRUCTURE IN KUMAON HIMALAYA

DCA axis 1 was positively correlated with

altitude (53.5% variance); slope explained

additional variation; thus 62.1% cumulative

variance was accounted for by the model (Table 3).

Axis 2 appeared to reflect shrub characters and

canopy cover gradient, as it was positively

correlated with these two gradients (Table 3).

Shrub and herb species classification

A total of 17 broad communities

comprising 52 shrub and herb species was

recognized and five major homogenous groups

were identified (Fig. 4 a). The left arm of the first

dichotomy contained 37 species, which was further

divided into 19 and 18 species. The three major

Table 3. Multiple regression analysis of Axis 1 and Axis 2 of DCA with vegetation attributes.

Combination of Variables Relationship R²

Axis 1 Altitude + 0.535**

Altitude & Slope + 0.655**

Axis 2 Shrub diversity + 0.32*

Shrub diversity, Grass richness + 0.544**

Shrub diversity, Grass richness, Canopy cover + 0.635**

Shrub diversity, Grass richness, Canopy cover, Shrub density + 0.714**

* p < 0.05, ** p<0.001

Fig. 4. TWINSPAN classification of (a) 52 species into 17 groups based on the

ground vegetation data of Kumaon Himalaya, and (b) classification of 23 sites

into 9 groups of ground vegetation data.

(a)

(b)

HUSSAIN et al. 175

communities formed in the left armed dichotomy

were Desmodium gangeticum, Pyracantha

crenulata and Rubus biflorus. These communities

were found at different sites (Fig. 4 b & Table 4)).

The right arm of dichotomy comprised 15 species,

which were further divided into one and 14 species

(Fig. 4 a). Five homogenous groups were combined

to form two communities.

Species composition

Tree density (ha-1) was significantly different

at all sites (F22, 879 = 14.13 , p = 0.00). It was high

at site 12 (995 ha-1) and site 5 (915 ha-1) compared

to the rest of the forest stands. Tree species

diversity and richness also varied significantly

between the sites (F22, 879 = 15.47, p = 0.00 and

F22, 879 = 13.61, p = 0.00, respectively) (Table 5).

Maximum tree diversity and richness were found

at site 17 (1.53 and 1.76 respectively) in Askot

Wildlife Sanctuary. Shrub density (ha-1) was also

significantly different among the forest stands

(F22, 879 = 24.04, p = 0.00) and it was highest at site

6 (28158 ha-1) while lowest at site 23 (6852 ha-1).

Shrub diversity (F22, 879 = 16.21, p = 0.00) and

richness (F22, 879 = 16.37, p = 0.00) also differed

significantly between the sites (Table 5). Diversity

was highest at site 2 while richness was high at

site 6 (1.48).

Although significant differences in tree density

did not occur at group level, the maximum tree

density was recorded for Q. floribunda -

Rhododendron arboreum group and the minimum

for A. pindrow - B. utilis group. Shrub density was

also not significantly different among groups but it

was maximum for Q. floribunda - R. arboreum and

minimum for Q. semecarpifolia - T. serrata group

(Table 6).

Taking all sites together the tree layer was

dominated by Q. floribunda (181 trees ha-1) followed

by R. arboreum (175 ha-1), Q. lanuginosa (167 ha-1)

and A. pindrow (151 ha-1) (Table 7). Myrsine

africana (6521 plants ha-1) was the dominant shrub

species in the Kumaon followed by Nerium sp. (3671

ha-1) and Athyrium sp. (3055 ha-1) (Table 7).

Species of special conservation concern

Using the individuals of tree species sampled

in all the 23 forest stands, the generated rarity

index value ranged from 0.03 to 0.40. The tree

species having rarity index value 0.03-0.20 were

considered rare. B. utilis (0.03), Tsuga demosa

(0.06), Q. glauca (0.06), P. wallichiana (0.09),

Taxus baccata (0.12), Cupressus torulosa (0.15),

Picea smithiana (0.16), A. pindrow (0.16) and C.

Table 4. Vegetation communities and their characteristic ground species in Kumaon Himalaya based

on TWINSPAN classification. BWS = Binsar Wildlife Sanctuary.

Groups Comm. Forest stand Tree species

1 1, 2, 3 Pandavkholi, Jageshwer,

Gasi

Desmodium elegans, Gaultheria nummulanoides, Mahonia

sp., Valeriana wallichii, Origanum vulgare, Boehmenia

rugulosa, Polygonum recumbens

2 4,5,6,7 Dhaphiadhura, Majtham,

Gandhura, Sobala, Duku,

Dhakuri

Cotoneaster acuminata, Asparagus racemosus, Adiantum

venustum, Pyracantha crenulata, Pteris biaurita, Myrsine

africana, Rubus peniculata, Desmodium gangeticum

3 8, 9, 10, 11, 12 Kilbery, Vinaiyak, Gager,

Kunjakharak, Jilling,

Maheshkhan, Sitlakhet,

Mukteshwer, Munsiyari

Wikstroemia canescens, Nerium sp., Daphne papyracea,

Athyrium sp, Rubus biflorous, Berberis aristata,

Boeninghausienia albiflora, Polystichum sp, Thalictrium

foliolosum, Pteridium sp., Indigofera heterantha, Randia

tetrasperma, Arundinella nepalensis, Hypericum

oblongifolium, Rubus ellipticus, Rhamnus virgatus,

Bistorta amplexicaulis, Hedychium spicatum

4 13, 14 BWS, Mechh Cratagus sp., Urtica dioca, Arisaema flavam, Geranium

wallichianum, Argemone maxicana

5 15, 16, 17 Wachham, Sunderdunga,

Pindari

Euphorbia prolifera, Skimmia laureola, Thamnocalamus

spathiflorus, Berginia legulata, Leptodermis kumaonensis,

Circium wallichii, Deutzia staminea, Aechmanthera

gossypina, Trachelospermum lucidum, Polystichum

squarossum

176 FOREST STRUCTURE IN KUMAON HIMALAYA

HUSSAIN et al. 177

deodara (0.17) were found to be rare tree species in

Kumaon. Except for Q. glauca, most of the above-

mentioned rare tree species were found in Pindari

region. P. wallichiana, C. torulosa, P. smithiana

and B. utilis were also found in Vinaiyak reserve

forest.

Table 6. Tree species density (TDEN ha-1, S.E.) along with tree species diversity (TDIV), tree species

richness (TRIC), shrub density (SDEN ha-1, S.E.), shrub diversity (SDIV) and shrub richness (SRIC) in

five homogenous groups of 19 tree communities of Kumaon Himalaya based on TWINSPAN classification

(S.E. refers to standard error).

Homogenous group TDEN± S.E. SDEN± S.E TDIV TRIC SDIV SRIC

Aesculus indica- Betula alnoides 744 ± 416 14402 ±10490 1.3 1.4 1.1 1.1

Quercus semecarpifolia- Toona serrata 704 ± 203 6915 ± 431 1.2 1.3 0.9 0.8

Abies pindrow- Betula utilis 493 ± 37 10263 ± 799 1.0 1.0 0.8 0.7

Quercus leucotricophora- Pyrus pashia 667 ± 68 14580 ± 2719 1.3 1.3 1.1 1.1

Quercus floribunda-Rhodendron arboreum 714 ± 70 16835 ± 2863 1.3 1.3 1.3 1.2 Table 7. Mean values of density (ha-1) of major tree and shrub species of surveyed forest stands of

Kumaon Himalaya (S.E. refers to standard error).

Tree species Density S.E. Shrub species Density S.E.

Quercus leucotricophora 139 93 Argemone maxicana 2064 1918

Quercus floribunda 181 102 Arundinella nepalensis 2954 1400

Quercus glauca 59 47 Athyrium sp. 3055 1595

Quercus semecarpifolia 108 46 Berberis aristata 985 319

Quercus lanuginosa 167 71 Euphorbia prolifera 1862 1232

Rhododendron arboreum 175 75 Cratagus sp. 1492 281

Lyonia ovalifolia 114 55 Daphne papyracea 1809 1190

Persia duthiei 110 93 Desmodium gangeticum 2021 1252

Euonymus tingens 70 38 Indigofera heterantha 1667 959

Symplocos theifolia 130 107 Mahonia sp. 554 205

Pyrus pashia 50 18 Myrsine africana 6521 4760

Toona serrata 52 24 Nerium sp. 3671 2656

Abies pindrow 151 97 Polystichum sp. 1527 631

Taxus baccata 151 97 Pteridium sp. 2190 1687

Cedrus deodara 110 77 Pteris sp. 3047 1391

Pinus wallichiana 92 60 Pyracantha crenulata 813 357

Pinus roxburghii 81 44 Rubus biflorus 1618 759

Acer caesium 76 35 Rubus ellipticus 1955 136

Alnus nepalensis 56 27 Urtica dioca 5017 7525

Viburnum mullaha 82 41

Aesculus indica 47 19

Myrica esculenta 86 33

Cupressus torulosa 112 37

Ilex dipyrena 70 37

Swida oblonga 68 28

Betula utilis 85 37

Litsea umbrosa 92 53

Lindera pulcherrima 118 65

Jugulans regia 70 22

Tsuga demosa 94 25

Swida sp. 110 91

178 FOREST STRUCTURE IN KUMAON HIMALAYA

Discussion

Various phytosociological techniques are

employed to study forest communities and their

relationships (Noy-Meir & Austin 1970; Whittaker

& Gauch 1973). We applied TWINSPAN, DCA and

PCA ordination techniques to analyse the

distribution of forest vegetation in Kumaon

Himalaya with satisfactory results. A somewhat

similar approach was used by Adhikari et al.

(1992) and Rihkari et al. (1989b) but their study

was confined to a small area of Kumaon.

The polythetic divisive classification divides

sites into groups on the basis of all the species

information. In our study this division was made

on the basis of species composition for the entire

sites. As suggested by Margules (1986),

representativeness should be used as the first

stage in selecting nature reserves. By classifying

sites into groups with different species

composition, one can ensure that all the major

groups are represented in the selection.

Tree species density was significantly different

among the present sites. It was substantially high

at Gasi and Gager. In this study, tree density at

Gager was higher than that estimated by Rikhari

et al. (1989 a). The diversity index values for all

the sites in the present study were higher than

what has been reported by Rikhari et al. (1989a),

Saxena & Singh (1982) and Singh & Singh (1984)

for forests in different localities of Kumaon

Himalaya. P. roxburghii and Q. semecarpifolia are

typical west Himalayan elements and are poorly

represented in Nepal and further east (Ohsawa et

al. 1986). Two other Oak forests (Q.

leucotricophora and Q. floribunda) are widely

distributed in the west with higher concentration

in the central Himalaya (Singh & Singh 1986).

Both these forests were distributed in most of the

surveyed sites except at higher elevation. A.

pindrow, T. baccata and B. utilis form sub-alpine

forest throughout the Himalaya (Dhar et al. 1997).

A. pindrow community dominated in some of the

stands at AWS, Pindari, Wachham, Sunderdunga

and Munsiary. Tree density and diversity and

shrub density and diversity were low in A. pindrow

– B. utilis group as diversity and richness decrease

at higher elevations (Rawal & Pangtey 1994; Singh

et al. 1994). Similar results are reported from

other areas also (Brithers & Spingarn 1992; Knops

et al. 1995). Q. semecarpifolia forest was

represented at Daphiadhura, Gasi and Dhakuri.

Tree diversity range (0.72 - 1.53) was similar as

recorded by Dhar et al. (1997) for Q. semecarpifolia

forest in AWS (1.41) but shrub diversity range (0.6

- 1.1) reported by them was lower than what has

been reported here (1.36).

In our study, Q. leucotricophora forest

represented the elevation range 1800 - 2300 m

(1200 - 2300 m by Singh & Singh 1986; 1700 - 2100

m by Singh et al. 1994), while Q. semecarpifolia

forest was present between 2200 - 3000 m altitude

range (2400 - 3600 m by Singh & Singh 1986; 2366

- 3000 m by Singh et al. 1994). The Shannon-

Wiener (H') diversity values were also similar to

those reported by others (Singh et al. 1994). These

values were also similar to those reported for

temperate communities in adjacent Nepal

Himalaya (Ohsawa et al. 1975) and elsewhere

(Monk 1967).

As reported by Dhar et al. (1997), > 50%

species of this region are non-native species. The

area has received plant elements from adjoining

regions of tropical Asia (Indo-China and Indo-

Malaya, Mani 1974) and Indo-Gangetic plains

(Spate 1957). The distribution of non-native

species is known from the Himalaya (Maheswari

1962). The change in native flora because of non-

native species could lead to long-term change in

ecosystem processes (Ramkrishnan & Vitousek

1989).

Some species such as Alnus nepalensis is fast

growing. Ohsawa (1991) considered it a ‘habitat

pioneer’ species, as it can occupy the newly formed

habitats. Similarly, the expanding P. roxburghii

poses serious threat to native Oak (Q.

leucotricophora and Q. floribunda) in whole of the

Kumaon, as it has been reported earlier also

(Singh & Singh 1987). The ecological nature of P.

roxburghii does not allow other broad-leaf species

to replace it, and P. roxburghii will continue to

hold a site indefinitely once it occupies it. (Singh et

al. 1984). All Oak species are facing severe threats

because of the demand for fodder and fire-wood.

This leads to reduction in seed production (Saxena

& Singh 1984). Other valuable tree species such as

A. pindrow, T. baccata, T. demosa and C. deodara

are felled because of their timber value. A.

pindrow community was mainly represented in

Pindari but A. pindrow and C. deodara had a good

population size also in Vinaiyak reserve forest.

Protection of this community is necessary.

HUSSAIN et al. 179

Numerical methods make no claim to being

objective, as the very choice of method is a

subjective decision (Birks 1987). However, the

main advantage of using numerical methods in

evaluating representativeness was that they

summarized information about the range in

variation in species composition found in whole of

the Kumaon Himalaya in an effective and

meaningful way. All plant species of special

conservation concern were found in Pindari and

Vinaiyak reserve forest. Both of these forests are

facing severe threats (Hussain et al. 2000) so

immediate action is required for conserving

important floral communities of these areas.

Conclusions

Overall 63 tree, 56 shrub, 90 herb and 21 grass

species were recorded in the 23 forest stands of

Kumaon Himalaya. The distribution of the tree

communities in these forest stands was governed

mainly by the gradients of altitude, slope and

canopy cover.

Some tree species such as B. utilis, Tsuga

demosa, Taxus baccata, C. deodara were recorded

rare. These species and their communities should

be protected in whole of Kumaon Himalaya. All

the plant communities as well as their associated

biodiversity are, in general, threatened and in

order to protect the whole range of biodiversity,

these plant communities need to be conserved.

Acknowledgements

Authors are thankful to the Ministry of

Environment & Forests, Govt. of India, for funding

the present study. We are grateful to Chief

Wildlife Wardens (Uttar Pradesh) Mr. Ashok

Kumar Singh and Dr. R.L. Singh for giving

permission to carry out fieldwork in Kumaon

Himalaya. We also thank Prof. Wazahat Hussain,

Department of Botany, AMU, Aligarh, for

reviewing the manuscript and providing valuable

suggestions. We are thankful to reviewers for

giving valuable suggestions. Thanks are also due

to Dr. S.S. Samant, G. B. Pant Institute of

Himalayan Environment and Development,

Almora (Uttarakhand) for identifying plant

species. We also thank Ms. Huma Waseem for

proof reading of the manuscript.

References

Adhikari, B.S., M. Joshi, H.C. Rikhari & Y.S. Rawat.

1992. Cluster Analysis (Dendrogram) of high

altitude (2150-2500 m) forest vegetation around

Pindari glacier in Kumaun Himalaya. Journal of

Environmental Biology 13: 101-105.

Anthwal, A., R.C. Sharma & A. Sharma. 2006. Sacred

groves: Traditional way of conserving plant

diversity in Garhwal Himalaya, Uttaranchal. The

Journal of American Sciences 2: 35-38.

Birks, H.T.B. 1987. Recent methodological developments

in quantitative descriptive biogeography. Annales

Zoologici Fennici 24: 165-178.

Brithers, T. & A. Spingarn. 1992. Forest fragmentation

and alien plant invasion of central Indiana old

growth forest. Conservation Biology 6: 91-100.

Dhar, U., R.S. Rawal & S.S. Samant. 1997. Structural

diversity and representativeness of forest vegetation

in a protected area of Kumaun Himalaya, India:

Implication for conservation. Biodiversity and

Conservation 6: 1045-1062.

Dombois, M.D. & H. Ellenberg. 1974. Aims and Methods

of Vegetation Ecology. John-Wiley & Sons, New York.

Greig-Smith, P. 1983. Quantitative Plant Ecology.

Blackwell, Scientific Publications, Oxford UK 359S.

Hill, M.O. 1979 a. TWINSPAN-a FORTRAN program for

arranging multivariate data in an ordered two- way

table by classification of the individuals and

attributes. Ecology and Systematics. Cornell

University, Ithaca, New York, USA.

Hill, M.O. 1979 b. DECORANA-a FORTRAN program

for detrended correspondence analysis and

reciprocal averaging. Ecology and Systematics.

Cornell University, Ithaca, New York, USA.

Hussain, M.S., A. Sultana & J. A. Khan. 2000. A Study

of Threats to Biodiversity Conservation of Middle

Altitude Oak Forest in Kumaon Himalaya. Final

Technical Report MoEF, Department of Wildlife

Sciences, A.M.U., Aligarh.

Knops, J.M.H., J.R. Grifin & A.C. Royalty. 1995.

Introduced and native plants of the Hestings

reservation central coastal California a comparison.

Biological Conservation 71: 115-123.

Kumar, M. & V. Bhatt. 2006. Plant biodiversity and

conservation of forests in foot hills of Garhwal

Himalaya. Lyonia 11: 43-59.

Magurran, A.E. 1988. Ecological Diversity and its

Measurement. University Press, Cambridge.

Maheswari, J.K. 1962. Studies on naturalized flora of

India pp. 156-170. In: P. Maheshwari, S. John &

I.K. Vasil (eds.) Proceedings of a Summer School of

Botany, Darjeeling. Calcutta, Sree Saraswati Press.

180 FOREST STRUCTURE IN KUMAON HIMALAYA

Mani, M.S. 1974. Biogeography of the Himalayas. pp.

664-681. In: M.S. Mani (ed.) Ecology and

Biogeography in India. The Hague.

Margules, C.R. 1986. Conservation evaluation in

practice. pp. 289-314. In: M.B. Usher (ed.) Wildlife

Conservation Evaluation. Chapman & Hall,

London.

Monk, C.D. 1967. Tree species diversity in the eastern

deciduous forest with particular reference to North-

Central Florida. American Naturalist 101: 173-187.

Nautiyal, M.C., B.P. Nautiyal & V. Prakash. 2004.

Effect of grazing and climatic changes on Alpine

vegetation of Tungnath, Garhwal Himalaya, India.

The Environmentalist 24: 125-134.

Noy-Meir, I. & M.P. Austin. 1970. Principal component

ordination and simulated vegetation data. Ecology

51: 551-552.

Ohsawa, M. 1991. Montane evergreen broad-leaved

forests of the Bhutan. pp. 89-156. In: M. Ohsawa

(ed.) Life Zone Ecology of Bhutan Himalaya II.

Japan: Chiba University Press.

Ohsawa, M., P.R. Shakya & M. Numata. 1975. Forest

vegetation of the Arun Valley, east Nepal. pp. 99-

143. In: M. Numata (ed.) Mountaineering of Mt.

Makalu and Scientific Studies in Eastern Nepal. 11,

1971 Japan: Chiba University Press.

Ohsawa, M., P.R. Shakya & M. Numata. 1986.

Distribution and succession of west Himalayan

forest types in the eastern part of the Nepal

Himalaya. Mountain Research and Development 6:

143-157.

Puri, G.S., V.M. Meher-Homji, R.K. Gupta & S. Puri.

1983. Phytogeographical ecology. pp. 115-210. In:

Forest Ecology. 2nd edn. Oxford & IBH Publishing

Company.

Ralhan, P.K., A.K. Saxena & J.S. Singh. 1982. Analysis

of forest vegetation at and around Naini Tal in

Kumaun Himalaya. Proceedings of Indian National

Science Academy B 48: 121-137.

Ramakrishnan, P.S. & P.M. Vitousek. 1989. Ecosystem

level processes and the consequences of biological

invasion. pp. 281-300. In: J.A. Drake, M.A. Mooney,

F. di Castri, R.H. Groves, F.J. Kruger, M. Rejmanek

& M.Williamson (eds.) Biological Invasions: A

Global Perspective. SCOPE 37: John Wiley and

Sons.

Rawal, R.S. & Y.S.P. Pangtey. 1994. High altitude forest

in a part of Kumaun, central Himalaya. Proceedings

of Indian National Science Academy B60: 557-564.

Rikhari, H.C., P.K. Ralhan & S.P. Singh. 1989 a.

Phytosociology and population structure of chir-pine

forests in Kumaun Himalaya. Annals of Biology 5:

129-140.

Rikhari, H.C.; R. Chandra & S.P. Singh. 1989b. Pattern

of species distribution and community characters

along a moisture gradient within an Oak zone of

Kumaon Himalaya. Proceedings of Indian National

Science Academy B55: 431-438.

Saxena, A.K. & J.S. Singh. 1982. A phytosoiological

analysis of woody plant species in forest

communities of a part of Kumaun Himalaya.

Vegetatio 50: 3-22.

Saxena, A.K. & J.S. Singh. 1984. Tree population

structure of certain Himalayan forests and

implications concerning the future composition.

Vegetatio 58: 61-69.

Saxena, A.K., S.P. Singh & J.S. Singh. 1985. Population

structure of forests of Kumaun Himalaya.

implication for management. Journal of

Environmental Management 19: 307-324.

Saxena, A.K., T. Pandey & J.S. Singh. 1985. Altitudinal

variation in the vegetation of Kumaon Himalayas.

pp. 43-66. In: D.N. Rao. K.J. Ahmed. M. Yunus &

S.N. Singh (eds.) Perspectives in Environmental

Botany. Print House, Lucknow.

Singh, J.S. & S.P. Singh. 1984. An Integrated Ecological

Study of Eastern Kumaun Himalaya with Emphasis

on Natural Resources. Vol. 1-3, Final Report

(HCS/DST/187/76). Kumaun University, Naini Tal.

Singh, J.S. & S.P. Singh. 1987. Structure and

functioning of central Himalayan chirpine forest

ecosystem. Current Science 56: 383-391.

Singh, J.S., Y.S. Rawat & S.P. Chaturvedi. 1984.

Replacement of Oak forest with Pine in the

Himalaya affects the nitrogen cycle. Nature 311: 54-

56.

Singh, R.S., P.K. Rahlan & S.P. Singh. 1987.

Phytosociological and population structure of mixed

Oak conifer forest in a part of Kumaun Himalaya.

Environmental Ecology 5: 475-487.

Singh, S.P. & J.S. Singh. 1986. Structure and function of

the central Himalayan Oak forest. Proceedings of

Indian National Science Academy 96: 159-189.

Singh, S.P., B.S. Adhikari & D.B. Zobel. 1994. Biomass

productivity, leaf longevity and forest structure in

the central Himalaya. Ecological Monograph 64:

401-421.

Spate, O.H.K. 1957. India and Pakistan: A General and

Regional Geography. London: Mathuen & Co.

Tiwari, J.C. & S.P. Singh. 1985. Analysis of woody

vegetation in mixed Oak forest of Kumaun

Himalaya. Proceedings of Indian National Science

Academy 51: 332-347.

Upreti, N., J.C. Tewari & S.P. Singh. 1985. The Oak

forests of Kumaun Himalaya (India). Composition,

diversity and regeneration: Mountain Research and

Development 5: 163-174.

HUSSAIN et al. 181

Whittaker, R.H. & H.G. Jr. Gauch. 1973. Evaluation of

ordination techniques. Ordination and

Classification of Communities. Part V 11: 287-321.

Zar, J.H. 1984. Biostatistical Analysis. Prentice-Hall,

Englewood Cliffs, NJ.