Biomass and net primary productivity of Bhabar ...

Forest Ecology and Management 176 (2003) 217±235

Biomass and net primary productivity of Bhabar Shisham forests in central Himalaya, India Neelu Lodhiyala, L.S. Lodhiyalb,* a

Department of Botany, D.S.B. Campus, Kumaun University, Nainital 263002, Uttaranchal, India Department of Forestry, D.S.B. Campus, Kumaun University, Nainital 263002, Uttaranchal, India

b

Received 4 October 2001; received in revised form 20 February 2002; accepted 10 May 2002

Abstract This paper illustrates biomass (dry weight per unit area) and net primary productivity (NPP) in 5-, 10- and 15 years old Shisham (Dalbergia sissoo Roxb.) forests planted after clear cutting of Sal (Shorea robusta) mixed broad-leaved tree species in Bhabar (a nutrient poor and low water table site) adjacent to foothills in Kumaun of central Himalaya. The linear regression equations for all the aboveground and belowground components of trees and shrubs were developed for each forest. The forest ¯oor biomass, litter input and understorey vegetation were also determined from each forest. The tree density was 625 trees ha 1 for each forest. The basal area, biomass, forest ¯oor litter mass, tree litter fall and NPP of trees increased with increase in forest age, whereas the herb biomass and NPP signi®cantly P < 0:01 decreased with increasing forest age. The total vegetation biomass and NPP ranged from 52.5 (5 years) to 118.1 t ha 1 (15 years) and 11.4 (5 years) to 14.8 t ha 1 per year (15 years old), respectively. The biomass accumulation ratio (BAR) for different tree components increased with increase in forest age. The BAR ratio ranged from 5.2 in the 5 years old to 8.9 in 15 years old forest. # 2002 Elsevier Science B.V. All rights reserved. Keywords: Dalbergia sissoo; Biomass; Net primary productivity; Forest ¯oor litter; Litter fall; Turnover time; Biomass accumulation ratio; Bhabar belt; Central Himalaya

1. Introduction Shisham (Dalbergia sissoo Roxb.) is a medium to large sized gregarious and deciduous tree having thick rough greyish brown bark. This species occurs throughout the sub-Himalayan tract and outer Himalayan valleys from Indus to Assam usually up to 900 m but sometimes ascending to 1500 m. Shisham descends the river valleys and plains. This species generally occurs naturally in ravine, freshly degraded *

Corresponding author. Present address: M.P. Niwas, Stoneleigh Compound, Tallital, Nainital 263002, Uttaranchal, India. Tel.: 91-5942-36754

sites. The tree species is self-sown by seeds and root suckers and can be propagated by arti®cial methods for large area plantations. According to Champion and Seth (1968), shisham is a primary seral type species always found in association with Khair (Acacia catechu Willd.) trees. In central Himalaya, it is most typically found on alluvial ground along the bed of streams, rivers and mostly in freshly exposed sandy soils. This species prefers porous soil with adequate moisture. It shows marked preference to soil composed of sand, pebbles and boulders in riverbeds. According to Troup (1921), shisham is indigenous only to the sub-Himalayan and Bhabar areas and has been introduced elsewhere by

0378-1127/02/$ ± see front matter # 2002 Elsevier Science B.V. All rights reserved. PII: S 0 3 7 8 - 1 1 2 7 ( 0 2 ) 0 0 2 6 7 - 0

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N. Lodhiyal, L.S. Lodhiyal / Forest Ecology and Management 176 (2003) 217±235

man. Trees shed its leaves from October to February and new lea®ng occurs in the months of April. The tree is a strong light demander right from the seedling stage. It has a good coppicing power in the age 5±15 years. Forest biomass is de®ned as the total amount of dry weight of forest trees per unit area of land at any time. However, net primary productivity (NPP) is the rate of dry matter accumulation in a given period. The quantity of tree biomass per unit area of land constitutes the primary data needed to understand the ¯ow of materials and water through forest ecosystem (Swank and Schreuder, 1974). Lieth and Whittaker (1975) pointed out that if forest biomass is to be measured and analysed in its proper context as a part of production, this gives an over all picture of ecosystem functioning. According to Lodhiyal and Lodhiyal (1997), the rising demand of energy from renewable sources has generated new ideas and turn attention to woody biomass production system. Plantation of indigenous tree species in short rotation cycle are going to play a vital role in this regard. An interesting example is D. sissoo, has shown promising growth and productivity, and can be raise well under agroforestry system and harvested in short period in irrigated condition. Shisham produces wood with numerous marketing options and can be propagated by seeds and vegetative parts. Shisham forest provides daily needs such as fuel wood, small timber for agricultural implements and fodder to the poor people residing in nearby forest areas. The increasing trend of plantations of an indigenous tree species are widely gaining popularity due to their higher biomass (dry matter) accumulation per unit area, better nutrient conservation ef®ciency and suitability in nutrient poor sites. The increasing pressure of growing population in the country as well as adverse ecological changes in the natural forest of D. sissoo has warned the scientists to know more about their sustainability. Therefore, present investigation aims at providing the detailed information on the structure and functioning of Shisham forests growing in Bhabar belt (a plain area with nutrient poor and low water table site) adjoins in Kumaon of Indian central Himalaya. Some studies have been made over a long time regarding the dry matter production of leguminous indigenous forest in India (Pacholi, 1997) but no study has been carried out in this region. This paper describes the results of dry matter production and

dynamics on related age changes (from 5-, 10- and 15 years old) of shisham forests raised in 4 m 4 m spacing, the plant to plant and row to row distances were 4 m each way, respectively. We compared various productivity aspects of relatively middle age of Bhabar Shisham forests with those of middle aged Tarai Shisham forests, exotic plantations and natural forests of the region and elsewhere. In this context, we have followed an ecological approach of management by considering the ¯ow of dry matter including litter inputs both in terms of quantity and time regime. The major objectives of this study were: (i) to estimate the biomass and NPP pattern in three Shisham forests (i.e. 5-, 10- and 15 years old) in Bhabar of central Himalaya, and (ii) to assess whether these Bhabar Shisham forests do better than the central Himalayan natural forests and exotic plantations of the region and elsewhere. Keeping in mind the above objectives, we approached through an ecological graph based on the dry matter estimates of Shisham forests for their best management growing in the central Himalayan Bhabar region of India. 2. Material and methods 2.1. Description of study sites The study sites were located between 288430 and 298370 N latitude and 798200 and 798230 E longitude at an altitude of 300 m in Bhabar belt (a nutrient poor and low water table site) in the district Nainital (between 50 and 60 km from Kumaun University, Nainital) of the Indo-Gangetic plains in south of the outer Shiwalik range of central Himalaya, India. The climate of study area is sub-tropical and monsoon type. There are three seasons in a year, viz., winter (November±February), summer (April±midJune) and rainy season (mid-June±mid-September). The months of October and March are transitional periods and are known as autumn and spring, respectively. The average monthly rainfall ranged from 0.5 mm in February (minimum) to 109.9 mm in July (maximum) (Fig. 1). The average minimum and maximum temperature was 4.5 8C (January) to 25.4 8C (July) and 17.3 8C (December) to 37.0 8C (May), respectively (Fig. 1).


219

Fig. 1. Ombrothermic diagram based on 2 years (1996±1997) meteorological data of the study area taken by Meteorological Department, GB Pant University of Agriculture and Technology, Pantnagar, Udham Singh Nagar, Uttaranchal, India.

The slope of Bhabar area is above 2.5 m per kilometre. The soil is coarse textured and nutrient poor with low water table. The soil of Bhabar is sandy loam with boulders. However, the low water table of Bhabar is due to coarse soil with low water holding capacity, high porosity and low nutrient content. Therefore, considered it contributes low production than Tarai (a nutrient rich and high water table site), an adjacent area of the central Himalaya, India. 2.2. Soils Thirty six soil samples (nine samples in each subplot, three samples from each diameter at breast height (dbh) class) were collected in September 1996, January 1997 and May 1997 from each forest a using soil corer placed randomly at three depths 0±10, 10±20 and 20±30 cm. Soil texture, bulk density, moisture content, water holding capacity, and porosity was determined according to Misra (1968), Lodhiyal (2000) and Lodhiyal et al. (2002). The soil pH extract was determined using a digital pH meter, the nitrogen concentration using a micro-Kjeldahl technique (Peach and Tracy,

1956), phosphorus by spectrophotometer and potassium by ¯ame photometer (Jackson, 1958). 2.3. Biomass 2.3.1. Sampling design The total forest area planted with Shisham trees was 37 ha. Of this area, 5-, 10- and 15 years old forest shared 10, 12 and 15 ha, respectively. The 5-, 10- and 15 years old Shisham forests were raised in the year 1991, 1986 and 1981, respectively, after clear cutting of indigenous species: A. catechu Willd., Adina cordifolia(Roxb.) Ridsdale, Albizzia lebbeck Benth., Bauhunia variegata Linn., Bombax ceiba Linn., Butea monosperma (Lamk) Taub., Mallotus philippensis (Lam.) Muell.-Arg., Shorea robusta Gaertn. f., Syzygium cumini (Linn.) Skeels, Toona ciliata M. Roem. and Trewia nudi¯ora Linn. in the area. The tree density in each Shisham forest was 625 trees ha 1, respectively, in 5-, 10- and 15 years old forest because of similar spacing (with plant to plant and row to row distances of 4 m 4 m in each way). A 1 ha plot was sampled in each forest, which

220


includes four replicate sub-plots of 50 m 50 m (0.25 ha). To ®nd out the accurate results, 100 trees in each sub-plot (total 400 trees (64%) in each forest) were measured. The height and diameter (dbh, 1.37 m) of trees were measured by Ravi's Multimeter and Tree Callipers, respectively, in each forest. After the vegetation analysis of Shisham forest, all the measured trees of each forest were divided into three diameter classes: 9.0±13.0, 13.0±16.0, 16.0±19.0 cm in 5 years old forest; 13.0±17.0, 17.0±21.0, 21.0± 25.0 cm in 10 years old forest; and 19.0±22.0, 22.0±25.0, 25.0 28.0 cm in 15 years old forest. For estimating density, the shrub and herb individuals were sampled in 50 quadrats of 2 m 2 m and 1 m 1 m, respectively. To estimate the biomass of Shisham forest, the selective harvest technique was adopted (Ovington, 1962; Newbould, 1967; Lodhiyal and Lodhiyal, 1997; Lodhiyal et al., 2002). Total 12 trees in each forest (three trees in each sub-plot, four from each dbh class) were harvested. A regression equation was developed for each tree component for the estimation of biomass. Harvested trees were cut into 1±2 m logs. Aboveground (bole wood, bole bark, branch, twig, leaf, reproductive parts, i.e. in¯orescence, ¯owers and fruits) and belowground (stump root, lateral roots and ®ne roots) components were assessed. The roots (stump root and lateral roots) were excavated to 2 m3 volume of soil for each harvested tree in each forest. The ®ne roots (roots 5 mm diameter associated with mycorrhizae) were sampled by digging out of three randomly distributed 25 cm 25 cm 25 cm block of soil around harvested tree in each dbh class of each sub-plot (total of 12 from each forest). The fresh weight of all components was determined in the ®eld using a heavy weight spring balance and pan balance. Samples of approximately 500 g (fresh weight) of each component from each forest were brought separately to the laboratory and oven-dried at 60 8C to constant weight. Using the fresh/dry weight factor, the dry weight of tree was estimated. Regression equations were developed for each tree components. The data were subjected to the regression in the form Y a bX, where Y is the dry weight of a component (kg), and X is the dbh aboveground level (cm, per tree). The mean diameter value for each diameter class was used in the regression equation of the different components to obtain an estimate of

mean biomass. This value was then multiplied by tree density in that diameter class. The total biomass of trees for each forest was calculated by summing biomass of each diameter class. Ten individuals of each shrub species differing in height and diameter were harvested and the roots were recovered to depth of 50 cm. The harvested material was separated into foliage, stem and roots. A regression equation was developed for each component to estimate the shrub biomass. Component wise average biomass of each shrub species (i.e. woody perennials) multiplied by the respective density per hectare in each forest. Individual wise biomass of the shrub species summed up to get a total shrub biomass for each forest. For herbaceous biomass, herbs were harvested at their peak (in rainy season; September 1997) from 10, 50 cm 50 cm, quadrats. The harvested material was divided into aboveground and belowground components. Fresh and dry weight was determined for each shrub and herb component. The total vegetation biomass was obtained by summing biomass values of trees, shrubs and herbs for each forest. 2.4. NPP After the selection of four permanent sub-plots (total area: 1 ha) in each age forest, 100 trees in each sub-plot were marked with white paint (just below breast height, 1.37 m from the base) in September 1996 to assess diameter and height increments at annual intervals. The height and diameter of marked trees were measured again in September 1997. The mean diameter and height increments for each diameter class were then calculated. The NPP of different tree components (bole wood, bole bark, branch, twig, leaf and reproductive parts in aboveground, and stump root, lateral roots and ®ne roots in belowground parts) was calculated using the allometric equations as followed by Lodhiyal, L.S., Singh, R.P., Singh, S.P., 1995; Lodhiyal, N., Lodhiyal, L.S., Pangety, Y.P.S., 2002 and Lodhiyal and Lodhiyal (1997). The net increase in biomass DB B2 B1 yielded the annual biomass accumulation. The sum of the DB values for the different components yielded net biomass accretion in the trees. Values for litter fall in the forests were added to the respective components (leaf and twigs). Considerable mortality of ®ne roots


occurs in actively growing trees. However, a basic knowledge of the control of ®ne root production requires a better understanding of soil ecosystems (Nadelhoffer and Raich, 1992). The biomass of ®ne roots could not be estimated repeatedly during the course of study so we followed the methods of Kalela (1954), Orlov (1968) and Ogino (1977). We assumed that the ®ne root mortality was equivalent to one-®fth of leaf litter fall; but present estimates of ®ne root production may be a gross underestimation (Harris et al., 1980; Vogt et al., 1982, 1986; Fogel, 1983; Lodhiyal, L.S., Singh, R.P., Singh, S.P., 1995; Lodhiyal, N., Lodhiyal, L.S., Pangety, Y.P.S., 2002; Lodhiyal and Lodhiyal, 1997; Lodhiyal, N., 2000), and may differ from the real value as Nadelhoffer and Raich (1992) found no correlation between ®ne root production and leaf litter fall for a large data set. However, it must be pointed out that all methods, including those based on related biomass estimations of ®ne roots, are subject to uncertainties and possible biases (Lauenroth et al., 1986; Sala et al., 1988). The net production value of each component was summed across diameter classes to give total net production of trees in each forest. Twenty individuals of each dominant shrub species were measured for estimation. The four common dominant shrubs were (Lantana camara var. aculeata (Linn.) Moldeweke, Murraya koenigii (Linn.) Spreng., Clerodendrum viscosum Venten. and Pogostemon benghalense(Burm.f.) Kuntze) differing in height and diameter were marked with white paint and their basal diameter was measured in September, 1996. After 1 year the marked individuals were remeasured. Increases in biomass of these shrubs were calculated using the regression equations, and to this value was added the foliage biomass (assuming 100% turnover of leaves in 1 year) to obtain net annual production. The average production of a species when multiplied by the density yielded the total production of that species. Summing net production values for all species gave total shrub production for a site. The biomass (aboveground and belowground) of herbs at all sites was determined during their peak growth period in September 1996. This value was assumed equal to net herb production. The sum of net production values of trees, shrubs and herbs yielded the total annual NPP of vegetation in each Bhabar Shisham forest.

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2.5. Litter fall Litter fall was studied for 1-year period from July 1996 to June 1997. The litter input was measured by randomly placing 20 litter traps (®ve litter traps for each dbh class in each sub-plot) on the forest ¯oor of each Bhabar Shisham forest. Each trap was 50 cm 50 cm with 15 cm high wooden sides ®tted with a nylon net bottom. Litter was collected at monthly intervals on each sampling date from July 1996 to June 1997 and brought to the laboratory in polyethylene bags. Samples were then sorted into leaf, wood, reproductive parts and other components. The wood component comprised bark and twigs. Samples of separated components were cleaned and weighed after oven drying at 60 8C to constant weight. After weighing, the litter components were ground and retained for nutrient analysis. 2.6. Forest ¯oor biomass Forest ¯oor litter biomass data were collected using 15 quadrats (of size 1 m 1 m) placed randomly in each forest once in each season, i.e. rainy, winter and summer. In each quadrat, all the components were categorised into: (a) fresh leaf litter; (b) partially and more decomposed litter; (c) wood litter includes twigs, bark and branches; (d) miscellaneous litter (consisting in¯orescences, ¯owers and fruits, litter parts of shrubs); and (e) herbaceous litter (living and dead) following Rawat and Singh (1988), Lodhiyal, L.S., 1990; Lodhiyal, N., 2000; Lodhiyal, L.S., Singh, R.P., Singh, S.P., 1995; Lodhiyal, N., Lodhiyal, L.S., Pangety, Y.P.S., 2002 and Lodhiyal and Lodhiyal (1997). In each quadrat, all the herbaceous standing shoots (live and dead) were harvested at ground level (Line, 1959; Green, 1959; Lodhiyal, L.S., 1990; Lodhiyal, N., 2000; Lodhiyal, L.S., Singh, R.P., Singh, S.P., 1995; Lodhiyal, N., Lodhiyal, L.S., Pangety, Y.P.S., 2002; Lodhiyal and Lodhiyal, 1997). Forest ¯oor material was collected carefully, avoid soil contamination. Collected material was taken to laboratory in polyethylene bags, cleaned of soil particles, and weighed after oven drying at 60 8C for constant weight. 2.7. Turnover of litter The turnover rate (K) of litter was calculated by following Jenny et al. (1949), Olson (1963), Lodhiyal

222


and Lodhiyal (1997), Lodhiyal (2000) and Lodhiyal et al. (2002): K

A AF

where A is the annual increment of litter (i.e. annual litter fall) and F the biomass of the litter at steady state. Turnover time (t, years) is the reciprocal of the turnover rate and is expressed as t 1=K. In the present study, F was the standing crop of partially and more decomposed litter during the winter season and A was annual tree litter fall plus shrub litter (equal to foliage biomass) plus herb litter (equal to peak aboveground herb biomass). 3. Results 3.1. Forest structure and soil characteristics Table 1 summarises forest structure and physicochemical properties of soil studied in three Shisham forests. The basal area increased signi®cantly P < 0:01 with age of forest, however, the soil parameters such as sand, silt, clay, porosity, soil moisture per cent (in rainy and winter seasons), soil pH and soil

nutrient (NPK) concentrations decreased; however, other soil parameters increased with increasing age of forest (Table 1). 3.2. Biomass The regression coef®cient for all the aboveground and belowground components of trees is given in Table 2; however, the allometric equations were developed for estimating biomass of the different shrub components based on the basal diameter, are given in Table 3. The selection of independent variable, diameter at breast height, was occasioned by the ease and accuracy in these making measurements. It is evident from r2-values in Table 2, the relations of biomass with dbh were found to be quite satisfactory. The calculation of biomass through X2h method (h being height) has not been used because the r2 the values from such equations did not indicate much improvement over those obtained with dbh(X). Therefore, the regression model Y a bX was used for forest biomass computations. The average height of trees was 6.8, 10.8 and 14.8 m while the shrub height was 1.8, 2.3 and 2.4 m in 5-, 10and 15 years old forest, respectively. The annual diameter increment in trees was 2.3±3.1 cm while in

Table 1 Forest structure and soil characteristics in Bhabar Shisham forest (D. sissoo Roxb.) at different ages in central Himalaya Parameters

Altitude (m) Forest area (ha) Tree density (ha 1) Basal area (m2 ha 1) Bulk density of soil (g cm 3) Soil texture of forest (%, in 0±30 cm) Sand Silt Clay Porosity (%, in 0±30 cm) Soil moisture (%, in 0±30 cm) Water holding capacity (%, in 0±30 cm) Soil pH extract(in 0±30 cm) Soil nutrient concentration (%, in 0±30 cm) N P K

Age of Shisham forest (years) 5

10

15

300 10 625 9.2 1.0

300 12 625 19.0 1.03

300 15 625 27.7 1.06

37.2 33.3 29.5 61 16.6 85.5 6.5

36.1 31.3 32.5 59 15.4 86.0 6.3

35.2 29.3 35.5 59 15.0 86.4 6.2

0.155 0.69 0.012 0.38 0.046 0.65

0.149 0.62 0.011 0.50 0.043 0.57

0.144 0.52 0.011 0.41 0.040 0.69

N. Lodhiyal, L.S. Lodhiyal / Forest Ecology and Management 176 (2003) 217±235 Table 2 The relationship between the biomass and tree components (Y, kg per tree) and diameter at breast height (X cm) for Bhabar Shisham (D. sissoo Roxb.) forests (the equation used was Y a bX)a Component

Age of Shisham forests (years) 5

Bole wood

10

15

a b r2

3.0374 2.7698 0.972

11.0369 3.6005 0.935

22.0780 2.8541 0.953

Bole bark

a b r2

0.6509 0.5936 0.972

2.2528 0.7346 0.935

4.2900 0.5538 0.953

Branchb

a b r2

0.5876 0.5369 0.972

2.2074 0.7201 0.935

4.7404 0.6164 0.953

Twigc

a b r2

0.2604 0.2374 0.972

0.9702 0.3168 0.935

1.9716 0.2583 0.953

Leaf

a b r2

0.3337 0.3052 0.972

1.1037 0.3600 0.935

1.7329 0.2238 0.953

Reproductive parts

a b r2

0.0744 0.0678 0.972

0.3088 0.1008 0.934

0.4256 0.0922 0.955

Stump rootd

a b r2

0.6128 0.6231 0.972

2.1194 0.6914 0.935

3.4393 0.4626 0.955

Lateral rootse

a b r2

0.3962 0.3617 0.972

1.4529 0.4750 0.935

2.8214 0.3849 0.958

Fine rootsf

a b r2

0.1538 0.1413 0.972

0.6120 0.2012 0.934

1.3691 0.1649 0.948

a

All equations signi®cant at P < 0:001. Shoots of larger dimension without leaves. c Current shoots bearing leaves. d Main root of tree includes about 30 cm basal part of bole. e Lateral branches of main root with a diameter >5 mm. f Roots with a diameter 5 mm associated with mycorrhizae. b

shrubs it was 0.18±0.30 cm, respectively, in 5±15 years old Shisham forests. The total vegetation biomass increased with increase in forest age from about 52.5 in the 5 years old to 118.1 t ha 1 in the 15 years old forest. Of this, tree layer shared maximum about 84±93% (Table 4). The total tree biomass (dry weight) also increased with age from 43:9 0:98 in the 5 years old to

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Table 3 Allometric relationship between biomass of shrub components (Y, kg per tree) and basal diameter (D, cm) for four shrub species under Shisham (D. sissoo Roxb.) forests in Bhabar belt of central Himalayaa Species

Component Intercept (a)

Slope (b)

r

r2

5 years old forest L. camara Stem Foliage Roots M. koenigii Stem Foliage Roots C. viscosum Stem Foliage Roots P. benghalense Stem Foliage Roots

1.2928 1.3097 1.1902 1.3190 1.3903 1.1760 1.2514 1.3348 1.1053 1.3793 1.3748 1.2078

8.0568 2.2851 4.8251 7.8571 2.0422 4.4768 7.4286 1.8695 5.2631 5.8033 1.7255 4.2408

0.989 0.983 0.975 0.997 0.993 0.979 0.983 0.958 0.929 0.989 0.997 0.990

0.978 0.966 0.951 0.994 0.986 0.960 0.966 0.918 0.865 0.978 0.995 0.981


1.2541 1.4460 1.3522 1.2332 1.1728 1.1090 1.2673 1.2036 0.8017 1.3602 1.3407 0.9089

6.3347 1.5576 3.1159 6.2038 2.0163 3.6662 7.6988 2.7265 7.3843 6.7181 2.1662 6.0994

0.979 0.996 0.987 0.990 0.993 0.989 0.968 0.984 0.987 0.986 0.978 0.967

0.959 0.992 0.974 0.980 0.986 0.979 0.938 0.968 0.974 0.974 0.956 0.935


1.6415 1.4610 1.4610 1.0385 1.1142 1.1081 1.1062 1.0353 0.9778 1.2619 1.2328 1.2128

3.6484 1.1457 2.8511 4.5665 1.8202 3.0192 7.7200 3.5724 4.2409 6.5200 2.1775 3.5109

0.976 0.994 0.988 0.995 0.996 0.996 0.969 0.981 0.927 0.975 0.983 0.993

0.953 0.987 0.977 0.990 0.993 0.992 0.963 0.963 0.859* 0.952 0.967 0.986

a *

All the values are signi®cant at P < 0:001. Signi®cant at P > 0:05.

109:6 1:46 in the 15 years old forest. Of which, aboveground accounted for 80±82% (Table 4). The contribution of bole, branch and reproductive parts to total tree increased while leaf and root components

224


Table 4 Biomass (t ha 1) in tree, shrub and herb layers of Bhabar Shisham forest at different ages in central Himalaya Vegetation


Components

10

15

Tree layer 43.9 0.98 Percentage allocation in Bolea 59.6 13.7 Branchb Leaf 5.4 Reproductive part 1.2 Coarse rootsc 17.5 Fine roots 2.5

74.4 1.08

109.6 1.46

Shrub layer 6.2 1.34 Percentage of allocation in 67.2 Abovegroundd Belowground 32.8

7.3 1.42

60.2 14.4 5.0 1.4 16.2 2.8

66.5 33.5

60.6 15.7 4.0 1.5 15.2 3.0 6.5 1.48 68.0 32.0

Herb layer 2.4 1.84 Percentage of allocation in Aboveground 75.8 Belowground 24.2

75.0 25.0

74.0 26.0

Total vegetation

83.9

118.1

52.5

2.2 1.96

Table 5 Forest ¯oor litter biomass (t ha 1, average across seasons) and turnover of litter of Bhabar Shisham forests at different ages in central Himalaya

2.0 2.01

a

Bole wood bole bark, which accounted for 10.0±10.5% of the values. b Branch twig, which accounted for 4.2±4.6% of the values. c Stump root (main root) lateral roots (lateral branches of main root), which accounted for 6.4±6.9% of the values. d Stem foliage, which accounted for 14.5±18.8% of the values.

decreased with age of forest. The proportions of biomass stored in roots (stump, lateral and ®ne roots) were found to be 18±20% (Table 4). Analysis of variance showed signi®cant P < 0:01 variation in total tree biomass (dry weight) and its components among the different age of forests. 3.3. Forest ¯oor biomass The forest ¯oor litter mass increased with age of Shisham forest. The seasonal mean total forest ¯oor biomass (dry weight including herbaceous litter) approximately doubled with age, the values being 2.9 in the 5 years old to 5.4 t ha 1 in the 15 years old forest (Table 5). However, the herbaceous biomass (dry weight) both live and dead showed a reverse trend. The turnover time increased slightly with age, it was 1.26 years in the 5 years old to 1.33 years in the 15 years old forest (Table 5).

Age of Shisham forests (years) 5 1

Forest floor litter (t ha )

10

15

2.9 3.15 4.8 3.82 5.4 3.96

Percentage allocation in Fresh leaf litter 8.4 Partially and more 25.1 decomposed litter Wood litter 6.4 Miscellaneous littera 30.0 Herbaceous litterb 9.5 Turnover rate 0.79 kg per year) Turnover time (t, yr) 1.26

9.5 28.7

9.9 29.6

11.4 32.7 29.1 0.76

19.2 38.0 22.5 0.75

1.31

1.33

a

This includes the reproductive parts of trees and shrubs. This includes the herbaceous dead herbaceous live, which accounted for 3.5±6.2% of the values. b

3.4. Litter fall The total annual litter fall ranged from 2.3 to 4.4 t ha 1 per year and increased signi®cantly P < 0:01 with forest age. Of these values, leaf litter accounted for about 67±68% and reproductive parts for 2±4% (Table 6). 3.5. NPP The NPP of vegetation ranged between 11.4 t ha 1 per year in the 5 years old and 14.8 t ha 1 per year in Table 6 Annual litter production (t ha 1 per year) of Shisham (D. sissoo Roxb.) forests in Bhabar belt of central Himalaya Litter components


10

Leaf litter Wood littera Reproductive litterb Other litterc

1.54 0.09 0.04 0.61

Total

2.28 (100)

a

(67.5) (3.9) (1.8) (26.8)

2.46 0.30 0.09 0.76

15 (68.1) (8.3) (2.5) (21.0)

3.61 (100)

This includes barks, twigs and branches. This includes the in¯orescence, pods and fruits. c This includes the leaf litter of shrubs. b

2.96 0.38 0.16 0.87

(67.7) (8.7) (3.7) (19.9)

4.37 (100)

N. Lodhiyal, L.S. Lodhiyal / Forest Ecology and Management 176 (2003) 217±235 Table 7 NPP (t ha 1 per year) in trees, shrubs and herbs of Bhabar Shisham forests in central Himalaya Vegetation

showed about 31% increase with age from 5 years old to 15 years old forest (ratio being 19.0 and 24.9, respectively). However, the bole branch production to FSC ratio declined by 4% from 5- to 15 years old age. The proportional increase in the aboveground and belowground tree part was 50 and 32%, respectively (Table 7). A signi®cant P < 0:01 regression equation has been developed between age and biomass, age and NPP, biomass and NPP, FSC and NPP, litter fall and NPP, photo and non-photosynthetic, and root shoot components of Shisham trees (Table 8).


10

Tree layer 8.4 1.96 10.9 2.10 Percentage allocation in component Bolea 57.4 57.2 13.3 14.3 Branchb Leaf 5.3 4.8 Reproductive parts 1.1 1.3 Coarse rootsc 16.8 15.3 Fine roots 6.1 7.1 Shrub layer 0.6 0.89 Percentage of allocation in 66.6 Abovegroundd Belowground 33.3

0.7 0.99 65.2 34.8

12.2 2.18 60.6 13.7 3.5 1.5 13.2 7.5 0.6 1.06

3.6. Biomass accumulation ratio

67.2 32.8

75.0 25.0

74.0 26.0

Total vegetation

13.8

14.8

11.4

2.2 2.05

15

Herb layer 2.4 1.92 Percentage of allocation in Aboveground 75.8 Belowground 24.2

225

The biomass accumulation ratio (BAR, biomass/ NPP) has been used to characterise the production ef®ciency of Shisham forests. This expresses the quantity of biomass retained in per unit of net production of trees. The total BAR was increased with age of (5.2 in the 5 years old to 8.9 in the 15 years old) forest (Table 9).

2.0 2.24

a

Bole wood bark, which accounted for 8.7±10.1% of the values. b Branch twigs (current shoots bearing leaves which accounted for 4.0±4.9% of the values. c Stump, root (main root) lateral roots (lateral branches of main root, which accounted for 6.0±6.2% of the values. d Stem foliage, which accounted for 13.6±18.8% of the values.

3.7. Pattern of dry matter transfer A compartment graph of dry matter ¯ow in 5-, 10and 15 years old Shisham forests is given in Fig. 2A± C. The mean annual incident solar isolation is 5956 MJ h 1 per year. Of this, about 2978 MJ ha 1 per year (50% of the total isolation) is considered photosynthetically available. The biomass of vegetation was 52 103 kg ha 1 (5 years) and 118 103 kg ha 1 (15 years old). Trees accounts for 83%, shrubs for 12% and herbs for 5% in the 5 years old, and trees 92.8%, shrubs 5.5% and

the 15 years old forest (Table 7). Of this, tree layer NPP increased from 8.4 t ha 1 per year in 5 years old to 12.2 t ha 1 per year in 15 years old forest (Table 7). In comparison to this rise of about 90% the NPP/FSC (net primary productivity/foliar standing crop) ratio

Table 8 Regression constant and coef®cients relating to certain parameters (independent variables) of Bhabar Shisham forests at different ages in central Himalayaa Variables

Intercept (a)

Slope (b)

Correlation coefficient (r2)

Age (years) vs. biomass (t ha 1) Age (years) vs. NPP (t ha 1 per year) Biomass (t ha 1) vs. NPP (t ha 1 per year) FSC (t ha 1) vs. NPP (t ha 1 per year) Litter fall (t ha 1) vs. NPP (t ha 1 per year) Photo (t ha 1) vs. non-photosynthetic components (t ha 1) Root (t ha 1) vs. shoot (t ha 1)

10.247 6.660 4.872 6.839 4.872 0.144 8.736

6.574 0.384 2.1078 3.413 2.108 1.768 4.936

0.998 0.965 0.999 0.946 0.999 0.808 0.999

a

All values are signi®cant at P < 0:01.

226


Table 9 BAR (biomass/net production of tree) in different components of Bhabar Shisham forest at different ages in central Himalaya Components

Bole wood Bole bark Branch Twig Leaf Reproductive parts Stump root Lateral roots Fine roots Total


10

15

5.4 5.4 5.4 5.4 5.4 5.9 5.5 5.4 2.2 5.2

7.2 7.0 7.2 6.0 7.1 7.4 7.2 7.2 2.7 6.8

8.7 10.3 10.3 10.4 10.3 9.1 10.3 10.1 3.6 8.9

herbs 1.7% in the 15 years old forest of the total dry matter. In the tree layer ratio of aboveground photo: non-photosynthetic ratio ranged from 0.05 (15 years) to 0.07 (5 years) and that of root: shoot ratio 0.22 (15 years) to 0.25 (5 years old forest). NPP is 11 103 kg ha 1 per year in 5 years and 15 103 kg ha 1 per year in 15 years old forest. The share of tree, shrub and herb layers in the total net production of the vegetation is 73.2, 5.8 and 21.0% in the 5 years old forest, and 82.2, 4.3 and 13.5% in the 15 years old forest, respectively. In the tree layer, the largest proportion of net production is allocated to bole (57.4%) followed by branch (13.3%) in the 5 years old, and bole (60.6%) followed by branch (13.7%) in the 15 years old forest. The shrubs were accounted for 4.3 (15 years) to 5.8% (5 years) of the total vegetation dry matter. The proportioning of net production in shrubs follows in the order: stem (48.4± 53.0%) followed by roots (32.8±34.8%) and foliage (13.6±18.8%) (see Table 7). In herbs, the aboveground components command a majority of net production 74±76% in terms of dry matter, respectively, 15- and 5 years old forest. The restitution of biomass litter formation is 4169 in the 5 years old to 5559 kg ha 1 per year in the 15 years old forest. Of the total litter fall from tree layer, leaf litter constitutes 46.4% (5 years) and 53.3% (15 years old forest). The biomass restitution in term of dry matter equals 18.5% (5 years) to 24.3% (15 years) of the total annual production of trees and 13.5% (5 years) to 19.9% (15 years old forest) that of total vegetation. The mean standing crop of

litter on the forest ¯oor ranged from 2920 (5 years) to 5370 kg ha 1 per year (15 years), which is 36.2 (5 years) to 46.2% (15 years) of the tree biomass, and 26.2 (5 years) to 37.7% (15 years) of the total vegetation. Decomposition of litter at soil surface as indicated by the turnover rate, 3294 (5 years) to 4169 kg ha 1 per year (15 years). This amounts to 78.9 (5 years) to 74.9% (15 years) of the total litter fall. At the end of annual cycle 876 (5 years) to 1390 kg ha 1 per year (15 years) remains and is carried out over to next year. The herbaceous vegetation at all sites mostly annual, however, a few perennials were also present, therefore, in present study are assumed 100% root mortality. In 5 years old forest, the root mortality of frees, shrubs and herbs amounts to 309, 179 and 580 kg ha 1 per year, respectively. However, in 15 years old forest, trees amounts to 593, shrubs 245 and herbs 520 kg ha 1 per year. 4. Discussion The estimation of biomass is pre-requisite for the study of dry matter ¯ow and functioning of forests. Biomasses were estimated in three forests at the 5 years age intervals growing in Bhabar, a plain area in foothills having less nutrient and low water table in central Himalayan Mountain, India. Comparison with forests of almost similar age indicates that present estimates of biomass are conspicuously higher than the value 103 t ha 1 reported for 8 years. D. sissoo forest in plain area (a rich nutrient site with high water table) (Pacholi, 1997). However, it was lower than 186 t ha 1 reported for 24 years old D. sissoo forest (Sharma et al., 1988), 193 t ha 1 for Tectona grandis forest (Negi et al., 1995), 141 t ha 1 for 30 years old T. grandis plantation (Jha, 1995), 121 t ha 1 for Eucalyptus hybrid (Negi and Sharma, 1985), 175 t ha 1 for Populus deltoides (Kaul and Sharma, 1983), 200±710 t ha 1 reported for more than 100 years old S. robusta (Rana, 1985), 250 t ha 1 for Quercus leucotrichophora (Rana and Singh, 1990), 112±300 t ha 1 for Eucalyptus grandis (Tandon et al., 1988). The values of aboveground biomass 35±90 t ha 1 in the present study are comparable with those of Eucalyptus hybrid (36±39 t ha 1, Negi and Sharma, 1985),


227

Fig. 2. Compartment models for dry matter distribution in 5-, 10- and 15 years old Bhabar Shisham (D. sissoo Roxb.) forests (A±C). Rectangles represent compartments for standing crop of dry matter, arrows represent net ¯ux rate. The three circular enclosures represent the total annual values of usable solar radiation (MJ ha 1 per year), total net production (kg ha 1 per year) and total disappearance (kg ha 1 per year). Compartment values are in kg ha 1 and turnover rate in kg ha 1 per year.

228


Fig. 2. (Continued ).

P. deltoides (44 t ha 1, Raizada and Srivastava, 1989), P. deltoides (71.7 t ha 1, Kaul et al., 1983), Prosopis juli¯ora (16±92 t ha 1, Gurumurti et al., 1987), D. sissoo (41 t ha 1; Tewari, 1994), D. sissoo of 5 years age (82 t ha 1, Sharma et al., 1988).

The aboveground biomass values of Shisham forests are lower than the value 161 t ha 1 for D. sissoo (Sharma et al., 1988), 113.3 t ha 1 for Pinus roxburghii (Chaturvedi, 1983), 166.8 t ha 1 for Populus tremuloides (Alban et al., 1978), 282 t ha 1 for P.


229

Fig. 2. (Continued ).

tremuloides (Perla and Alban, 1979), 275 t ha 1 for 11 years old E. grandis (Tandon et al., 1988), 240 t ha 1 for 33 years old Alnus rubra (Zavitkovski and Stevens, 1972) and 561 t ha 1 for S. robusta forest (Rana, 1985). The aboveground biomass estimates are com-

pared with natural and exotic vegetation of the world (Table 10). The proportion of biomass stored in roots were found to be 18±20% of the total tree biomass, which are comparable than the values 19±21% reported for

230

Forest type

Location

Aspen±maple±birch S. robusta Oak forest Oak forest E. obliqua E. grandis

USA India India India Australia India

E. regnans E. saligna P. roxburghii Eucalyptus hybrid Eucalyptus hybrid

Australia Australia India India India

P. deltoides Marsh P. deltoides (I C) P. deltoides Marsh D. sissoo D. sissoo D. sissoo Tarai Shisham forests Bhabar Shisham forests

India India India India India India India India

Age

± >100 >100 >100 51 5 10 10 8 ± 10 2 3 5 14 10 9 3 5 24 5±15 5 10 15

Density

± ± ± ± 865 1650 689 1075 829 ± 1223 2000 2000 2000 ± ± ± 10000 ± 467 625 625 625 625

Aboveground biomass (t ha 1)

Percentage of allocation Bole

Branch

Twig

Foliage

Reproductive parts

95.4 70.2 301.5 263.2 298.2 97.6 275.1 319.0 129.8 113.0±283.0 21.9 3.2 18.5 54.1 44.5 105.4 151.6 41.4 82.0 16.2 41.8±103.1 35.1 60.3 89.8

79.5 43.9 51.0 51.7 91.2 83.8 92.0 82.1 80.9 76.1 62.1 75.2 77.7 80.8 69.9 74.4 71.9 38.6 40.6 65.5 64.4±66.1 59.6 60.2 60.6

17.6 51.0 44.4 40.5 6.5 6.8 3.9 13.0 13.4 19.4 15.4 5.6 7.0 6.6 13.7 13.0 16.0 29.2 34.9 14.2 7.8±8.4 9.5 10.0 11.1

± ± ± ± ± 4.6 2.0 ± 1.2 ± 8.8 1.9 1.8 1.6 ± ± ± 12.0 ± 2.6 3.6±4.2 4.2 4.4 4.6

2.9 5.1 4.6 7.8 2.3 4.6 2.0 4.8 4.4 4.5 13.7 17.2 14.5 10.2 7.4 12.6 12.1 ± 6.5 ± 4.2±6.4 5.4 5.0 4.0

± ± ± ± ± ±

References

± 0.08 ± ± ± ± 0.7 ± ± ± ± ± ± 0.8±1.2 1.2 1.4 1.5

Crow (1978) Singh (1979) Rawat (1983) Negi et al. (1983) Attiwill (1979) Tandon et al. (1988) Tandon et al. (1988) Frederick et al. (1985a) Frederick et al. (1985b) Chaturvedi (1983) Pandey et al. (1987) Bargali et al. (1992) Bargali et al. (1992) Bargali et al. (1992) Raizada and Srivastava (1989) Singh (1989) Singh (1989) Tewari (1994) Sharma et al. (1988) Sharma et al. (1988) Lodhiyal et al. (2002) Present study


Table 10 Comparisons of biomass distribution (%) in aboveground tree components of certain forests and plantations of the world


5±8 years old poplar and eucalypt plantations (Lodhiyal et al., 1995; Bargali et al., 1992), much less than the value of 43% reported for a eucalypt ecosystem (Westman and Rogers, 1977). The present values are near to 15±18% reported for poplar forest (Lodhiyal and Lodhiyal, 1997) and higher than the value 8±15% reported for temperate forests (Whittaker and Woodwell, 1971; Larsen et al., 1976), 10±12% eucalypt forest (Feller, 1980). The photo: non-photosynthetic ratio (aboveground 0.05 in the 15 years to 0.07 in the 5 years old) and root: shoot ratios (0.22 in the 15 years old to 0.25 in the 5 years old) are similar to that of natural forests of central Himalaya (Chaturvedi and Singh, 1987; Rawat and Singh, 1988), eucalypt plantations (Bargali et al., 1992), poplar plantations (Lodhiyal and Lodhiyal, 1997). The variation in the ratios may be related to the production ef®ciency of tree components with age (Lodhiyal and Lodhiyal, 1997). The herb biomass decreased with increase in forest age was due to shade of trees and allelopathic effects. According the Shiva and Bandopadhyay (1985) germination of undercanopy species decline because of chemicals releases from tree leaves. Thus, the combined effects of both allelopathic chemicals and shade of trees may be the real cause in decline of herbaceous dry matter production. The herbs contribute signi®cantly to the total production at 15 years (13.5%) and 5 years age (21.0%) and return of organic matters through litter about 37% in both 5 years and 15 years old, respectively, and thus contributed favourably to soil structure. However, herbs seem to be important competitor of the D. sissoo at 5 years age, a signi®cantly marked decline (21% compared to 13.5%) in herb production from 5- to 15 years old forest was accomplished by 47.6% rise in tree production (compared to the yearly rise 10.5± 23.5%) in subsequent years. In the 15 years age Shisham forest, tree production is 31.5% higher compared to 5 years age forest, this is also one another cause of decline in herb production. However, herbs contributed signi®cantly (in decreasing order 13.5± 21.0% in 5±15 years age) to total production and return of organic matter through litter fall (36.5± 37.4%) compensate the decline of herbs to keep the ecosystem much productive in relation to better soil structure compared to 7.7 and 20.7%, respectively, for herb and litter production in poplar plantations

231

(Lodhiyal and Lodhiyal, 1997). In 15 years old forest, the herbs contributed 10% signi®cantly to total production and return of organic matter through litter for 30% have compensated the loss of herb biomass in forest for better soil improvement. These values are higher than the values (8 and 21%) reported for fast growing poplar plantations (Lodhiyal and Lodhiyal, 1997). Our estimates of forest ¯oor biomass (2.9± 5.4 t ha 1) falls within the values 0.8±6.5 t ha 1 reported for Eucalyptus hybrid (Bargali et al., 1992) and lower than 8.4 t ha 1 for 4 years old poplar plantation (Lodhiyal and Lodhiyal, 1997) and close to the value 4.6±6.1 t ha 1 for low density poplar plantation (Lodhiyal et al., 1995). The turnover time was 1.26±1.33 years, which is similar to the values 1.2±1.3 years reported for Eucalyptus species (Richards and Charley, 1977; Bargali et al., 1992) and lower than 1.04±1.07 years reported for low and high density poplar plantations (Lodhiyal et al., 1995; Lodhiyal and Lodhiyal, 1997). However, present estimates are much lower than 3.5 years reported for Eucalyptus obliqua (Attiwill et al., 1978). Thus, the variability in the litter fall from year to year emphasises one of the problems of estimating litter decompositions rates from litter fall and the weight of accumulated litter on the forest ¯oor (Lamb, 1985). Our litter estimates (2.3±4.4 t ha 1 per year) fall within the range 2.0± 6.7 t ha 1 per year reported for poplar plantations (Lodhiyal and Lodhiyal, 1997) and much lower than 12.0 t ha 1 per year reported for litter fall value is higher than 1.3±2.6 t ha 1 per year for P. tremuloides (Crow, 1974; Van Cleave and Noonan, 1975) and somewhat close to the value 4.2 t ha 1 per year reported for Populus ``Tristis'' plantation (Zavitkovski, 1981). The NPP estimates (11.5±14.8 t ha 1 per year) are higher than 7.7 t ha 1 per year reported for 24.4 years old D. sissoo (Sharma et al., 1988) 7.0±10.0 t ha 1 per year for conifer forests (Rodin and Bazilevick, 1967), 10.0 t ha 1 per year for Gmelina arborea (Pacholi, 1997), 11 t ha 1 per year for Cryptomeria japonica (Tadaki et al., 1965) and 11.4 t ha 1 per year for tropical moist forest (Golley et al., 1975) present estimates are close to 14.6±15.7 t ha 1 per year reported for dry deciduous forests (Singh, 1974), but lower than 22.3 t ha 1 per year for D. sissoo (Pacholi, 1997), 19±24 t ha 1 per year for tropical rain forests (Warner, 1970; Bullock, 1981), 24± 32 t ha 1 per year for poplar plantations (Lodhiyal

232


Table 11 Comparison of total NPP with other forests and plantations of the world Forest type

Location

NPP (t ha

Tropical rain forest Dry deciduous forest Tropical moist forest Rianj dominated forest Sal old growth forest Chir-pine forest Conifer forests

Malaysia India ± India India India USSR

19.2 14.6±15.7 11.4 15.5 16.0±18.9 17.3 7.0±10.0

Pine forest G. arborea forest Cassia siamea forest C. japonica Tropical rain forest Camelia japonica Tropical rain forest Low density poplar plantation (8 years old) High density poplar plantation (4 years old)

USA India India Japan Thailand Japan Java India India

11.0 10.0 18.7 10.9 28.6 29.4 24.3 24.5 32.4

Eucalyptus plantation (8 years old) Sissoo plantation (24 years old) D. sissoo forest Tarai Shisham forests (5±15 years old) Bhabar Shisham forests (5-, 10- and 15 years old)

India India Bihar (India) Central Himalaya (India) Uttaranchal (India)

23.4 7.7 22.3 12.6±20.3 11.4±14.8

et al., 1995; Lodhiyal and Lodhiyal, 1997), 19 and 23 t ha 1 per year for Sal forest (Singh and Singh, 1987) and eucalypt plantation (Bargali et al., 1992). Some estimates of productivity reported for different forests in the world are given in Table 11. The biomass/FSC ratio increased (18.5 in 5 years old to 24.8 in 15 years old), however, NPP/FSC ratio declined from 3.5 in 5 years to 2.7 in the 15 years old with age of forest increase. Similarly the root:shoot and photo:non-photosynthesis ratios are also decreased with increase in forest age. This indicates that as the forest age increased the tree canopy size increases, consequently the sunlight penetration to the lower part of leaves decline, which results lower production ef®ciency of leaves. According to Lodhiyal and Lodhiyal (1997) lower production ef®ciency of leaves largely because of its shorter growing period, i.e. deciduous nature of trees with higher photosynthetic ef®ciency and nutrient rich with thin leaf characteristics. A signi®cant decline in production of herbs was reported with increase in forest age. Our estimation of NPP are lower compared to 18± 28 t ha 1 per year reported for central Himalaya

1

per year)

References Bullock (1981) Singh (1979) Golley et al. (1975) Rana et al. (1989) Singh and Singh (1989) Rana et al. (1989) Rodin and Bazilevick (1967) Whittaker (1966) Pacholi (1997) Pacholi (1997) Tadaki et al. (1965) Kira et al. (1967) Kan et al. (1965) Warner (1970) Lodhiyal et al. (1995) Lodhiyal and Lodhiyal (1997) Bargali et al. (1992) Sharma et al. (1988) Pacholi (1997) Lodhiyal et al. (2002) Present study

forests (Chaturvedi and Singh, 1987; Rawat and Singh, 1988; Rana et al., 1989; Singh and Singh, 1992) and 24±32 t ha 1 per year for poplar plantations (Lodhiyal et al., 1995; Lodhiyal and Lodhiyal, 1997). It can be argued that productivity of present Bhabar Shisham forests in age of 5±15 years is not much lower than the normal range of 20 t ha 1 per year at the its rotation age of 60 years. But these estimates of production can be increased by raising trees in closer with better inputs and also some extent to provide more time as these forest crop are growing on nutrient poor environment at younger age. However, our estimates are close to the stable temperate forest (12± 15 t ha 1 per year) and falls within the general range (10±20 t ha 1 per year) of world's forest estimates (Whittaker, 1975). The BAR increased with age of forest, which express the quantity of biomass retained per unit of net production. Our estimate of BAR (5.2±8.9) close to the value 4.9±7.7 for low-density poplar plantations (Lodhiyal et al., 1995) and 5.9 for eucalypt plantation (Bargali et al., 1992) and higher than (0.6±3.7) reported for high density young age poplar plantations


(Lodhiyal and Lodhiyal, 1997). Thus, these estimates of BAR suggests that higher dry matter can be produced with increase in age of Shisham forests, Therefore, BAR not only depends on the species and site condition but also the age of forest. It is concluded that present estimates of Shisham forests hold a promise of dry matter production at 5± 10 years age compared to fast growing poplar harvested at 4±8 years age and conventional forests of higher rotation age (50±100 years) of the region. The poor soil fertility and low water table are unfavourable for forest crop production but these present forests has maintained its productivity because of the following reasons: (i) leguminous characters of Shisham trees, (ii) higher nutrient conservation ef®ciency of trees, and (iii) run off materials brought from the Himalayan rivers. According to Lodhiyal and Lodhiyal (1997), the optimal size of FSC determines the better wood production. Therefore, it is necessary to enhance the relationship between NPP and FSC of present forests. Besides these conclusions, more observations are needed to depict the long-term effects of tree harvesting to maintain their productivity and soil fertility status in relation to sustainability of Shisham forest in the Bhabar zone of central Himalaya. Acknowledgements Authors are thankful to Professor Y.P.S. Pangtey, Department of Botany, Kumaun University, Nainital, India, for encouragement and suggestions throughout the study period, and Professor R.P. Singh, Head, Department of Forestry, Kumaun University, Nainital, India, for providing necessary facilities. References Alban, D.H., Perela, D.A., Schaegel, B.E., 1978. Biomass and nutrient distribution in aspen, pine and spruce stands on the same soil type in Minnesota. Can. J. For. Res. 8, 290±299. Attiwill, P.M., 1979. Nutrient cycling in an Eucalyptus obliqua (L' Herit) forest. III. Growth, biomass and net primary production. Aust. J. Bot. 27, 439±458. Attiwill, P.M., Guthrie, H.B., Leuning, R., 1978. Nutrient cycling in an Eucalyptus obliqua (L' Herit) forest. I. Litter production and nutrient return. Aust. J. Bot. 26, 76±91.

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