Transcript

Forest Ecology and Management, 29 (1989) 151-163 151 Elsevier Science Publishers B.V., Amsterdam - - Printed in The Netherlands

Patterns of Litter Nutr ient Concentrat ion in Some P lantat ion Ecosys tems

S.C. SHARMA and P.K. PANDE

Forest Ecology Branch, Forest Research Institute, New Forest, Dehra Dun 248006 (India)

(Accepted 2 November 1988)

ABSTRACT

Sharma, S.C. and Pande, P.K., 1989. Patterns of litter nutrient concentration in some plantation ecosystems. For. Ecol. Manage., 29: 151-163.

Patterns of litter nutrient concentrations in plantations of Shorea robusta (sal), Tectona gran- dis (teak), Pinus roxburghii (pine) and Eucalyptus sp. (eucalypts) were studied. Concentration of Ca and N were greater than K, Mg and P, in all fractions of litter, irrespective of species. The nutrient concentrations, in general, were related to the tissue longevity and the species life-forms. The nutrient-concentration changes in the leaf-tissue were negatively correlated with the mag- nitude of the leaf-fall: in the case of sal for all the nutrients, in teak and pine for N and P only, and in eucalypts for none of the nutrients. These changes were due to the back-translocations of nutrients from the leaves prior to leaf-fall. The variation in the concentration of leaf-litter nu- trients is not a species' attribute but depends upon the combined effect of soil nutrient status, growth of the stand and tree growth formations.

INTRODUCTION

Of geochemical, biogeochemical and biochemical mineral-flow pathways in terrestrial ecosystems, the third-named assumes importance in redistribution and conservation of nutrients within the standing crop and in determining quantity of nutrients in litter-fall. Metabolically active leaves continue to drive nutrients up to a maximum till maturity; thereafter, in certain circumstances, depending upon the plant growth-form and stature associated with age, not- withstanding the site characteristics, nutrient contents often decline to a min- imum as a result of senescence-caused retranslocations (cf. Stachurski and Zimka, 1975; Charley and Richards, 1983). The changing litter nutrient con- centrations decisively affect plant nutrition and within-stand nutrient cycling. In recent years, therefore, much emphasis has been placed on studies of nu- trient retrieval from senescing leaves (Chapin and Kedrowski, 1983; Vitousek, 1984).

Litter-fall seasonality and litter accretion in plantations of sal (Shorea ro-

0378-1127/89/$03.50 © 1989 Elsevier Science Publishers B.V.

152 S.C. SHARMA AND P.K. PANDE

busta Gaertn. ), teak (Tectona grandis L.), pine (Pinus roxburghii Sarg. ) and Eucalyptus spp. have been reported in a previous communication (Pande and Sharma, 1986). This paper presents a comparative account of litter nutrient concentrations and their seasonal variations in different tree plantations, with the aim of ascertaining the role of these variations on the plantations' nutrient cycling.

THE SITE

Plantations of sal, teak, pine and eucalypts were located in the demonstra- tion area of the Forest Research Institute and Colleges, Dehra Dun, India. This zone, at Long 77 ° 52' 12" E, Lat. 30 ° 20'40" N and altitude 640 m a.s.l., has been categorised as Siwalik Sal Zone of semievergreen vegetation, with associated species such as Anogeissus latifolia, Buchanania lanzan, Woodfordia [ruticosa, Indigofera pulchella, Eulaliopsis binata and Terminalia tomentosa in dry areas, and Lagerstroemia parviflora, Eheritia laevis and Mallotus philippensis in moist areas (Champion and Seth, 1968).

Tree species Mallotus philippensis, Syzygium cumini, Cinnamomum cam- phora and Flacourtia ramontchi were common in teak as well as in pine plan- tations. In addition, Michelia champaca was also present in pine plantations. In sal plantation, Urena lobata, Lantana camara, Clerodendrum viscosum, Murraya koenigii, Litsea glutinosa were characterized as undergrowth. Jasmi- num multiflorum was common in both sal and teak plantations. In eucalypts, Achyranthes aspera was the most dominant herbaceous species, Lantana ca- mara as dominant shrub. Other relevant details of the plantations and soil characteristics are given in Tables 1 and 2, respectively.

The climatic data was obtained from Forest Influence Branch of the Insti- tute. Mean air temperature was 20 °C over the 1960-1984 period, daily average

TABLE 1

Characteristics of the plantations

Plantation Year of Area Average Density Annual Total Litter k 2 plantation (ha) Dbh (no. ha- 1) litterfall biomass %1

(cm) (kgha -1) (kgha -1)

Sal 1926 0.32 26.9 700 11271 390 0163 2.89 1.97 Teak 1926 0.30 28.8 410 6950 129 5804 5.36 1.21 Eucalypts 1972 1.13 9.1 1700 7070 57 9133 12 .21 1.05 Pine 1926 0.50 38.1 385 9676 141 5505 6.84 1.31

1Percentage of total above-ground biomass. 2Decomposition constant. The values of k are calculated using l/x~, where l is litter production; and xsB, litter accumulation (Olson, 1963 ). 3Estimated from derived equations of Negi (1984); 4Kaul et al. (1979); and ~Kaul et al. (1981.).

LITTER NUTRIENT PATTERNS IN PLANTATIONS

TABLE 2

Soil characteristics of different plantations

153

Plantation Soil Soil Carbon + + Nitrogen + + depth pH + (%) (%) (cm)

C : N Texture

Sal 0-30 6.3-6.6 3.6 0.24 15.12 Loam Teak 0-30 6.8-7.4 4.3 0.27 15.81 Clay loam Eucalypts 0-30 6.0-6.6 3.7 0.23 16.35 Clay loam Pine 0-30 6.4-6.6 3.5 0.22 15.95 Silty clay loam

+Annual range for 12 months. + +Average value of 12 months. Source: Pande (1986).

ranged from 13.3°C (January) to 27.8°C (June). The mean annual rainfall for the period in question was 2042 mm, the bulk of which occurred during June-September.

M E T H O D S AND MATERIALS

Three permanent quadrats of 5 X 5-m size were randomly placed in each plantation. All the quadrats were initially cleared and swept of any deposited debris. A total of 9 X 12 samples in each plantation was considered for the es- timation of annual litter production. Monthly estimation of litter-fall was made by collecting the litter from these quadrats, and then sorting it into leaves, twigs, fruits wherever possible, and 'miscellaneous', consisting of litter of other than the main species and other unidentified organic matter. From each quad- rat, triplicate samples of each litter fraction were collected and brought to the laboratory for determining the oven-dry weight (80°C). Phosphorus was de- termined by ammonium-molybdate-blue method (Vogal, 1961), K and Ca flame-photometrically, and Mg by atomic absorption spectrophotometry. Ni- trogen was determined by the Kjeldhal method (Piper, 1942 ). All the results are expressed on the oven-dry-weight basis.

RESULTS

Litter and leaf-faU patterns The monthly leaf and litter-fall (kg ha -1) as total of leaves, twigs, repro-

ductive parts and other miscellaneous components are shown in Fig. 1. All the plantations showed unimodal leaf-fall patterns, except eucalypts wherein it was bimodal. The peak leaf-fall period in sal was during the months of March- April, and in teak and pine during April-May. In eucalypts, two peaks were observed, first during the months of October-November and the other in April-

1 5 4 S.C. S H A R M A AND P.K. PANDE

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Fig. 1. Variations in monthly litter-fall magnitude.

May. Table 1 shows the total annual acquisition of litter-fall and litter-fall as percent of the total above-ground biomass, along with the other characteristics of the plantations. Although total annual litter input was greatest in sal, yet as percentage of above-ground biomass it was least among all the plantations.

Litter nutrient concentrations Monthly variations in nutrient concentration and average annual status of

nutrients in leaf and twig litter, along with their percent coefficients of vari- ance, are shown in Fig. 2 and Table 3, respectively. Percent concentrations of Ca and N were higher than K, Mg and P, in all the litter fractions. Leaf litter showed higher concentrations of nutrients than twig litter. In general, litter nutrient concentrations were greater in the deciduous species sal and teak than in the evergreen pines and eucalypts. Considerable monthly variations in the concentration of leaf-litter nutrients were observed in all species. In all the plantations, starting from the initial months (August-September), N-concen- tration increased to a maximum during the 4th and 5th months, thereafter declined to a minimum value during the 8th to 10th months, and again rose till the final month. Potassium, in the cases of teak and eucalypts, showed higher

LITTER NUTRIENT PATTERNS IN PLANTATIONS 155

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Fig. 2. Nutrient concentration changes in leaf and twig litter.

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LITTER NUTRIENT PATTERNS IN PLANTATIONS 157

2 x l ~ T Nitrogen = TEAK • • • • : PINE ~ l) • • • SAL

-- X X EUCALYPTS

3 x I

2x I ~ Phosphorus

• •

i:~d 2 X TEAK

P o t a s s i u m

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• SAL

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I x l , , , , , , IO' " . . . . . . . ,'O 2 . . . . . ,'0 3 . . . .

LEAF-FALL ( kg ho -I) Fig. 3. Relationship between nutrient concentration and magnitude of leaf-fall.

values during the initial months, decreased in the following months and again increased towards the final months. However, in pine, a decrease in K concen- tration was noticed from initial to 5th month, followed by an increase and decrease during 7th and 10th months, respectively. Calcium concentration, on

158 s.c. SHARMA AND P.K. PANDE

TABLE4

Correlation coefficients (r) between monthly leaf-fall and nutrient concentration along with their regression equations

Species Leaf-fall (kg ha -1) vs. nutrients (%)

N P K Mg

Sal - 0.797** - 0.846** - 0.798** - 0.737** log y = 0.3002 logy= -0.7453 logy= -0.0536 log y = 0.5166 -0.1137 log x -0.1935 log x -0.1429 log x -0.09868 og x

Teak - 0.583* - 0.783** - 0.533 .... - 0.491 .... log y-- 0.3299 log y = - 0.3427 -0.1481 log x -0.3513 log x

Eucalypts -0.417 ns -0.166 .... -0.091 .... -0.073 ....

Pine - 0.910"* -0.631" - 0.165 .... - 0.122 .... log y = 0.3285 log y = - 0.9833 - 0.2063 log x - 0.0351 log x

Pooled data -0.569** -0.576** -0.310" -0.341" of the log y = 0.2714 log y = 0.850 logy= -0.7158 logy= -0.6617 four -0.1271 log x -0.1469 log x -0.0799 log x -0.05344 log x species

n.s. = not significant; *= significant at 5%; **= significant at 1% probability level.

t h e o t h e r h a n d , s h o w e d v e r y l i t t le v a r i a t i o n in a n y o f t h e species . M a g n e s i u m c o n c e n t r a t i o n s in sal a n d t e a k s h o w e d a r ise f r o m in i t ia l to 5 t h m o n t h , d e c l i n e d t h e r e a f t e r , a n d s h o w e d l i t t le v a r i a t i o n s in t h e fo l lowing m o n t h s . N o c o n s i s t e n t p a t t e r n for t h i s n u t r i e n t cou ld be n o t i c e d for p i n e o r euca lyp t s .

A l t h o u g h t h e tw ig l i t t e r s h o w e d c o n s i d e r a b l e m o n t h l y v a r i a t i o n s in n u t r i e n t c o n c e n t r a t i o n s , t h e p a t t e r n wa s n o t c o n s i s t e n t in e i t h e r o f t h e species (Fig. 2, T a b l e 3 ).

Leaf-litter nutrient concentration vs. leaf-faU (kg ha- 1) L e a f - l i t t e r n u t r i e n t c o n c e n t r a t i o n s were n e g a t i v e l y c o r r e l a t e d w i t h t h e m a g -

n i t u d e o f leaf- fa l l (Fig. 3, T a b l e 4) . Sal s h o w e d s i gn i f i c an t nega t ive r e l a t i on - sh ip for N, P , K a n d Mg, t e a k a n d p i n e o n l y for N a n d P. H o w e v e r , in t h e case o f euca lyp t s , no s i gn i f i c a n t c o r r e l a t i o n c o u l d be e s t ab l i shed . P o o l e d d a t a for i nd iv idua l n u t r i e n t s o f all t h e spec ies a lso s h o w e d s i gn i f i c an t c o r r e l a t i o n s w i t h t h e m a g n i t u d e o f leaf-fal l .

DISCUSSION

T h e re su l t s o n t h e m a g n i t u d e o f l i t t e r - fa l l in d i f f e r e n t p l a n t a t i o n s revea l a d e c r e a s i n g t r e n d w i t h i n c r e a s i n g b i o m a s s , an o b s e r v a t i o n also m a d e b y o t h e r

LITTER NUTRIENT PATTERNS IN PLANTATIONS 159

workers (Ebermayer, 1876; Miller, 1984). As such, litter-fall magnitude is ex- pected to increase rapidly till canopy closure, thereafter parallel net primary productivity, and during the advanced phase of growth reduce to a value rep- resenting just net primary production of the stand itself (Miller, 1984). Thus, eucalypts, being a younger stand, registered a proportionately higher magni- tude of litter-fall than the advanced-aged plantations of sal, teak and pine (Table 1 ). The seasonal leaf-fall peak of March-April shown by all the species may be related to the drier climate prevailing in the area of plantations during that period. However, the peak leaf-fall period of October-November shown only by eucalypts may be regarded as genetically determined (Pande and Sharma, 1986).

Mean leaf-litter nutrient concentrations except nitrogen were almost the same in all species (Table 3). The higher P concentration in pine than the other plantations can be attributed to enhanced uptake and mineralization in the rhizosphere of conifers (Fisher and Stone, 1969). Distinct differences in the concentration of N between evergreens and deciduous species (Table 3, Fig. 2), are related to their respective growth forms (Chapin, 1980; Miller, 1984). The greater leaf-litter nutrient status in pine and eucalypts compared to results reported elsewhere (Table 5) were probably due to the enriched soil nutrient status of plantations. Similar results for various tropical plantations have also been reported by Seth et al. (1963) and Srivastava et al. (1972) for Pinus roxburghii, Das and Ramakrishnan (1985) for P. kesiya, and George (1977) and Negi (1984) for Eucalyptus hybrid (Table 5 ). The concentrations of Ca and N were more than the other nutrients, irrespective of species (Table 3). High concentrations of Ca and N in the litter of tropical trees have also been reported by Vitousek (1984).

As compared to leaf litter, the low nutrient concentrations in twig litter may be attributed to non-photosynthetic:photosynthetic-tissue ratios and tissue longevity. Low nutrient concentrations in perennial tissues have also been re- ported by Gosz et al. (1972) and Atiwill et al. (1978).

In general, monthly variations of N, P and K were more than Ca and Mg (Table 3). These variations could be associated with leaching or withdrawal prior to leaf-fall. The highly significant negative correlation of N and P with monthly magnitude of leaf-fall for pooled data (Table 4), also suggest their retrieval back to the living tissues, as N and P are generally the minor constit- uents of canopy washing (Morton, 1977). However, variations in K concen- tration may also be accounted forby leaching (Tuckey, 1970; Gosz et al., 1973 ). Calcium, being a structural element of the tissues, showed little variation.

Invariably, the variations in the nutrient concentration in leaf litter of de- ciduous plantations (sal and teak ) were more than evergreen, except N in pine (Table 3 ) and were also negatively correlated with the magnitude of leaf-fall (Fig. 3, Table 4). Indeed, in deciduous species, the immediate demand of the resumption of entire foliage mass every year necessitates the recovery of

160 s.c. SHARMA AND P.K. PANDE

TABLE5

Nutrient concentrations in leaf-litter at different localities of the world

Species/community Nutrient concentration (%)

N P K Ca Mg

Authority

Gymnosperms (U.S.A.) Angiosperms (U.S.A.) Tropical forests (Guatemala) Eucalyptus camaldulensis (swamp forest, Australia) *Eucalyptus hybrid 1.11 (India) *Eucalyptus hybrid 1.16 (India) * Pinus patula 0.64 {Tanzania) *P. caribea 0.45 (Nigeria) P. kesiya (India)

0.58-1.25 0.04-0.10 0.22-0.39 0.55-2.16 0.14-0.32

0.31-1.01 0.09-0.28 0.40-1.18 0.99-3.84 0.22-0.77

1.61-1.88 0.05-0.07 0.15-0.29 0.66-2.13 0.37-0.87

0.91-1.09 0.05-0.09 0.21-0.29 1.67-1.98 0.23-0.25

0.06 0.54 1.24 0.16

0.07 0.56 1.27 0.15

0.03 0.14 0.81 0.24

0.04 0.22 0.60 0.17

1.35-1.45 0.07-0.08 0.83-0.86 0.26-0.29 0.12-0.15

*P. roxburghii (India) P. roxburghii (India) *Shorea robusta (India) S. robusta (India)

*S. robusta (India) *Tectona grandis (India) T. grandis (India)

T. grandis (India) S. robusta (India) P. roxburghii (India) Eucalyptus spp. ( India )

1.05 0.19 0.56 0.66 0.13

0.58-1.37 0.10-0.19 0.25-0.61 0.77-1.50 0.08-0.19

0.92 0.19 0.38 1.54 0.20

0.71-1.57 0.11-0.18 0.23-0.58 1.25-1.56 0.11-0.28

1.15 0.18 0.34 1.35 0.15 0.98 0.21 0.37 2.46 0.11

0.67-1.33 0.12-0.20 0.28-0.60 1.34-2.34 0.09-0.25

0.67-1.53 0.03-0.13 0.40-0.90 0.80-1.60 0.13-0.26 0.81-1.53 0.03-0.11 0.20-0.60 0.60-1.30 0.13-0.26

0.44-1.12 0.08-0.10 0.40-0.90 0.60-0.80 0.09-0.18

0.64-1.14 0.05-0.09 0.40-0.90 1.30-1.60 0.13-0.20

Lutz and Chandler (1946) Lutz and Chandler (1946) Ewel (1976)

Briggs and Maher (1983)

Negi (1984)

George (1977)

Lundgren (1978)

Egunjobi and Onweluzo (1979) Das and Ramakrishnan (1985) Seth et al. (1963)

Srivastava et al. (1972) Seth et al. {1963)

Srivastava et al. (1972) Negi (1984) Seth et al. (1963)

Srivastava et al. {1972)

Present study

*Mean values.

LITTER NUTRIENT PATTERNS IN PLANTATIONS 161

nutrients through retranslocation from the dying foliage, because acquisition of nutrients through active absorption is costlier than for carbon (Bloom et al., 1986). In eucalypts, nutrient concentrations, although not significantly correlated with monthly magnitude of leaf-fall, showed variations in their monthly concentrations (Tables 3 and 4). These variations may be attributed to, firstly, the age-differential population of canopy foliage, as production of new foliage is also at the expense of old leaves (Alvim, 1964), and secondly, to premature leaf-fall caused by competing population of foliage for food and/or nutrients. Thus, mobile nutrients are liable to be retranslocated to the devel- oping tissues during this period.

The mode and magnitude of leaf-fall and its nutrient concentrations are among the most important features of nutrient cycling in a plantation ecosys- tem. The variation in the concentration of leaf-litter nutrients is not a species' attribute but depends upon the combined effect of soil nutrient status, growth of the stand and tree growth formations.

Study of litter nutrient status and differential pattern of nutrients in leaf- fall is of importance, especially in conservation of nutrients in the manage- ment of short-rotation forestry on a perpetual basis, where nutrient loss through leaf-fall may exceed the nutrient demand, as in evergreens, which are less de- manding on nutrients compared to deciduous trees. Further, the bimodal pat- tern of leaf-litter deposition, high turnover rate; the higher amounts of nu- trient return to the forest floor in eucalypts compared to sinusoidal pattern of deciduous species, all will result in asynchronous nutrient-related ecosystem feedback links. The importance of these variations is considerable in well- drained soils, where leaching loss of nutrients during wet periods is possible and likely to remove significant amounts of community nutrients (Rogers and Westman, 1977; Lamb, 1985), thus leaving the soil impoverished after every rotation of plantations.

REFERENCES

Alvim, P., 1964. Tree growth periodicity in tropical climates. In: M.H. Zimmermann (Editor), The Formation of Woods in Forest Trees. Academic, London, pp. 479-496.

Attiwill, P.M., Guthrie, H.B. and Leuning, R., 1978. Nutrient cycling in Eucalyptus obliqua (L'Herit) forest. I. Litter production and nutrient return. Aust. J. Bot., 26:79-91.

Bloom, A.J., Chapin, F.S. III and Mooney, H.A., 1985. Resource limitation in plants - An eco- nomic analogy. Annu. Rev. Ecol. Syst., 16: 363-392.

Briggs, S.V. and Maher, M.T., 1983. Litter fall and leaf decomposition in a river red-gum (Euca- lyptus camaldulensis) swamp. Aust. J. Bot., 31: 307-316.

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