Ecological Research (2000) 15, 435447

Leaf litter decomposition of canopy trees, bamboo and moss

in a montane moist evergreen broad-leaved forest on Ailao
Mountain, Yunnan, south-west China
Wenyao Liu,1,2* John E. D. Fox2 and Zaifu Xu1
1
Kunming Division, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences,
Kunming 650223, China and 2School of Environmental Biology, Curtin University of Technology,
GPO Box U1987, Perth WA 6845, Australia

Litter decomposition and nutrient release of selected dominant synusiae in an old-growth, evergreen,
broad-leaved mossy forest on Ailao Mountain, Yunnan, south-west China, were studied over a 22-
month period. The species studied were three dominant tall tree species, Lithocarpus xylocarpus
Markg., Lithocarpus chintungensis Hsu et Qian and Castanopsis wattii A. Camus; one dominant under-
story species (the bamboo Sinarundinaria nitida Nakai); and a mixture of dominant mosses (includ-
ing Homaliodendron scalpellifolium Fleisch, Symphyodon perrottetti Mont., Herberta longifolissa Steph. and
Bazzania albicans Horik.). Fast initial litter decomposition was followed by lower rates. Decompo-
sition rates of canopy species and bamboo leaf litter appear to be controlled by the initial concen-
tration of lignin, nitrogen (N) and phosphorus (P) more than by morphological features of the leaves.
The decay rate of moss litter was less correlated with nutrient composition and lignin concentra-
tion in initial mass. The order of decomposition rates was Castanopsis wattii > L. xylocarpus > L. chin-
tungensis > bamboo > moss. The decomposition rate constants (k) of the leaf litter for the canopy species
L. xylocarpus, L. chintungensis and Castanopsis wattii were 0.62, 0.50 and 0.64, respectively, and 0.40
and 0.22 for bamboo and moss, respectively. Turnover time (1/k) for the three canopy species was
1.61 years, 2.0 years and 1.55 years, respectively, and 2.50 years and 4.55 years for bamboo and
moss, respectively. The N and P concentration in the decomposing leaf litter increased in the first
6 months and then decreased over the remaining period. There was a relatively rapid initial loss of
potassium (K), followed by a slight increase. Each of calcium (Ca) and magnesium (Mg) decreased
with time whereas iron (Fe) and manganese (Mn) increased with time to some extent. Nutrient
release from decomposing leaf litter was in the order of K > Mg > Ca > N > P > Mn > Fe, except for
bamboo (Sinarundinaria nitida) K > Ca > P > N > Mg > Mn > Fe.

Key words: decomposition; litter; montane moist evergreen broad-leaved forest; nutrient dynam-
ics; SW China.

INTRODUCTION plants (Waring & Schlesinger 1985). The rate of

Litter decomposition and nutrient mineralization
presents an important source of nutrients for
primary production in terrestrial ecosystems (Swift
et al. 1979; Vitousek 1984; Berg et al. 1993;
Kchy & Wilson 1997). It is estimated that nutri-
ents released by litter decomposition make up
7090% of the total nutrient requirement of
*Author to whom correspondence should be
addressed. Email: rliuw@alpha2.curtin.edu.au
Received 1 March 2000.
Accepted 17 July 2000.
decomposition is an important determinant of the
biomass and productivity of terrestrial forest
ecosystems. Litter decomposition and mineraliza-
tion are affected by physico-chemical factors
(temperature, moisture and pH), litter quality
and decomposer organisms (Swift et al. 1979;
Anderson & Swift 1983; Songwe et al. 1995).
Decomposition rate varies considerably between
forest types and within sites (Anderson & Swift
1983; Swift & Anderson 1989; Lisanework &
Michelsen 1994). There is more organic matter in
the soil and a greater standing crop of litter in
forests at higher altitudes on tropical and sub-
tropical mountains (Grubb 1971; Edwards 1977;
436 W. Liu et al.
Tanner 1981). In Yunnan, south-west China, the
old-growth, evergreen, broad-leaved forest is rep-
resented by trees of the genera Lithocarpus and Cas-
tanopsis. This association is distributed mainly on
tropical and subtropical mountains with an eleva-
tion of 20002600 m, over some 2648 km2 in
Yunnan province (Bureau of Environment Conser-
vation of Yunnan 1998). Due to the abundance of
rainfall, high elevation and high humidity, this
forest type has been named: middle mountain
humid evergreen broad-leaved forest in Yunnan
( Jin 1983; Wu 1987). Ailao Mountain National
Nature Reserve (NNR), one of the key national
parks of China, includes a total area of 504 km2 of
this type of forest. Although there have been
several studies comparing structure and species
composition of primary and secondary forest in the
area (Young & Wang 1989; Young et al. 1992),
information on litter decomposition and nutrient
dynamics in the old-growth forest is limited. Ailao
Mountain occurs in the transition area between the
southern and northern subtropical forest forma-
tions ( Jin 1983; Wu 1987; Qiu et al. 1998).
Decomposition and mineralization in this transi-
tion zone have been little investigated.
Due to the particular environments of richer
rainfall, high humidity and elevation, there is a
well-developed bamboo understory and an abun-
dant moss epiphyte population in the old-growth
forest. These components are likely to have differ-
ent nutrient cycling characteristics. In the present
study, we examined decomposition rates and nutri-
ent dynamics of dominant canopy species, an
understory species and epiphytic moss in old-
growth forest on Ailao Mountain NNR, south-
west China. The objectives were to characterize
variation in amount and quality of leaf litter, decay
rates and release of nutrients through litter decom-
position to the soil within this particular environ-
ment and forest where nutrient cycling had not
been studied before.
METHODS
Study site
The study was conducted in the Xujiaba region
(2432N, 10101E), in a protected section of
Ailao Mountain NNR, Yunnan, south-west China.
This encompasses the largest tract of old-growth,
evergreen, broad-leaved forest in China (Qiu et al.
1998). The study site is within a 5100 ha pristine
area of undulating hills and crests, between 2400
and 2600 m altitude, and close to the Forest
Ecosystem Research Station (FERS) of Kunming
Institute of Ecology, Chinese Academy of Sciences.
The Institute has established permanent vegeta-
tion study plots and accumulated basic soil data.
There is a meteorological station and the Institute
operates a simply equipped field laboratory at
Xujiaba FERS.
The mean annual temperature is 11.3C; the
warmest month ( July) is 16.4C and the coldest
month ( January) is 5.4C. The accumulated tem-
perature 10C is 3049.5C. The mean annual
rainfall is 1931 mm, of which 85% is concentrated
in the rainy season from May to October. The mean
annual humidity is 86%, with humidity >90%
from June to December and 6070% from January
to May. The annual evaporation is 1485.5 mm,
which is less than the rainfall (Fig. 1). The rela-
tively warm climatic regime enables the evergreen
broad-leaved forest of the Xujiaba region to exist
with some of the features of subtropical forests
(Qiu et al. 1998).
The upper canopy of the forest is 1825 m high
and contains a number of tree species including
Castanopsis wattii A. Camus, Hartia sinensis Dunn,
Lithocarpus xylocarpus Markg., Lithocarpus chintun-
gensis Hsu et Qian, Machilus viridis Hand.-Mazz.
Manglietia insignis Blume and Schima noronhae
Rainfall and evaporation (mm)
400 20
Temperature (c)
300 15
200 10
100 5
0 0
Jan. Feb. Mar. Apr. May Jun. Jul. Aug.Sept.Oct. Nov. Dec.
(Months)
Fig. 1. Mean monthly rainfall, evaporation and tem-
perature (19911999) at Xujiaba on Ailao Mountain,
altitude 2450 m. Data from the Forest Ecosystem
Research Station, Kunming Insititute of Ecology,
Chinese Academy of Sciences. (), Rainfall (mm);
( ), Evaporation (mm); , Temperature.

Reinw. ex Blume. A variable second story of small
trees or tall shrubs 312 m tall includes: Camellia
forrestii Coh. Stuart, Eriobotrya bengalensis Hook. f.,
Eurya obliquifolia, Litsea elongata Benth. et Hook.
f., Rhododendron leptothrium I. B. Balf, Vaccinium
duclouxii Hand.-Mazz. and Symplocos species. A
bamboo, Sinarundinaria nitida Nakai, 1 4 m tall,
locally dominates the understory to 70% cover
(Qiu et al. 1998). The forest is a mossy forest ( Jin
1983). It contains abundant epiphytes (moss, ferns
and lichens) with much moss cover on tree boles
and branches, exposed rock and dead wood. The
ubiquitous litter layer is generally 37 cm deep.
Soils developed under the forest are yellow-brown
in colour, acidic in reaction (pH 4.2 4.9) and are
high in organic matter and nitrogen content (Deng
et al. 1983; Qiu et al. 1998). Table 1 shows the
chemical properties of the surface soil.
Decomposition
The litterbag method was used in the study. This
method has been recognized as a very useful proce-
dure for understanding decomposition and nutrient
dynamics of litter in forest ecosystems (Swift et al.
1979; Berg et al. 1993), although the decay rate
measured by this method may be underestimated
because of the restriction of macrofauna entry into
the litterbags (Ewel 1976; Sundarapandian &
Swamy 1999). Decomposition and nutrient release
were examined using freshly fallen foliage of five
leaf types. Three of these were canopy dominants:
Castanopsis wattii, Lithocarpus xylocarpus and Litho-
carpus chintungensis, that comprise 62% of total
aboveground biomass 47% of total leaf biomass and
45% of the total leaf litter of this forest. The fourth
leaf type was the understory bamboo Sinarundinaria
nitida that accounts for 99% of total shrub biomass
(Xie et al. 1996). The final leaf type was mixed moss
Table 1 Chemical characteristics of soil (010 cm) of the old-growth evergreen broad-leaved forest in Xujiaba
region of Ailao Mountain
pH Organic matter Total N
(H2O) (%) (%) C/N
4.37 7.48 0.52 14.4
(0.41) (0.26) (0.11) (1.68)
Values are mean, n = 3 with SD in parentheses. The data are summarized from Deng et al. (1983) and Qiu et al. (1998).
Leaf litter decomposition in montane forest 437
growing on tree stems. You (1983) identified the
four main moss species as Homaliodendron scalpelli-
folium Fleisch, Symphyodon perrottetti Mont., Herberta
longifolissa Steph. and Bazzania albicans Horik.
The 4 ha (200 200 m) research plot used in
this study was located near the Xujiaba FERS.
Fifteen 1 1 m litter traps made of four pieces of
wood 15 cm deep with a fine 2 mm nylon mesh
nailed to the base, 1 m above the ground, were
located randomly in a subplot (100 100 m)
within the research plot. Litter was collected at
monthly intervals from the traps for 9 years
(19911999). The mean annual litterfall was
7.12 t ha-1 with a bimodal seasonal pattern in lit-
terfall: the first litterfall peak occurred in April
May and the second peak in OctoberNovember.
Leaf litter of the three canopy species and the
bamboo used for the decomposition was collected
from the traps on 30 October 1997. Simultane-
ously, moss was collected randomly from the bark
of more than 20 trees at 11.5 m above the ground,
when the moss was relatively dry, from sampling
areas of 20 20 cm. All collected materials were
dried to constant weight in an oven at 60C for
48 h. Samples of 5 g (weighed to the nearest
0.01 g) for dried leaves and mixed mosses (or 3 g
for the lower specific gravity bamboo leaves) were
placed in 2 mm mesh nylon bags of 15 20 cm,
50 bags for each litter type. These were then sewn
with thread. Three samples of each litter type were
set aside for determination of initial oven dry
weight and initial leaf nutrient concentrations. On
28 November 1997, 10 litterbags of each type
were placed in each of five, 10 10 m blocks,
pegged out on the ground in the research plot.
Blocks were placed approximately 100 m apart.
One litterbag of each type was retrieved randomly
from each of the five blocks at intervals: one every
3 months for 12 months and thereafter one every
P2O5 K2O CaO MgO Al2O3 Fe2O3
(%) (%) (%) (%) (%) (%)
1.35 1.12 0.36 0.74 15.39 8.76
(0.005) (0.04) (0.02) (0.03) (2.38) (1.12)
438 W. Liu et al.

2 months for 10 months (i.e. five bags of each leaf
type were removed at each retrieval). Bags were
cleaned to remove extraneous materials, arthropods
and sand. Litter was dried at 80C to constant mass
and weight. From the mean dryweight the decom-
position rate constant (k) was calculated for each
species using the exponential relationship X1/X0 =
ekt, where X0 is initial dryweight and X1 is the dry-
weight remaining at collection time (t) in years
(Olson, 1963). Decomposition rate constants were
determined to compare decomposition rates
among the five litter types.
Chemical analysis
Initial samples and residual samples for six collec-
tion times over a 22-month period were milled to
yield 3 g of finely ground samples for chemical
analysis. Three samples taken from the mixed
whole litter of five litterbags were analyzed for
each leaf type at each retrieval. Total N was deter-
mined by the micro-Kjeldahl method after digest-
ing 0.5 g subsamples in K2Cr2O7H2SO4 (Institute
of Soil Academia Sinica 1978). Samples for P, K,
Ca, Mg, Mn and Fe were wet digested in a mixture
of HNO3, H2SO4 and HClO4 solution. Concentra-
tion of P was obtained colorimetrically by the
chloro-molybdophosphonic blue color method
(Institute of Soil Academia Sinica 1978). Potas-
sium, Ca, Mg, Mn and Fe were analyzed by an
atomic absorption spectrophotometer (AAS-932;
GBC Scientific Equipment, Melbourne, Australia).
Lignin concentrations were determined gravimet-
rically after acid digestion (72% H2SO4) and cor-
rection for ash content following the Van Soest
method (Cotrufo et al. 1999).
Statistical analysis
The initial chemical composition and the weight
loss of the decomposing litter were analyzed
statistically using one-way analysis of variance fol-
lowed by the multiple comparison Fishers test at
P < 0.05 to detect whether significant differences
occurred between the five leaf types. All statistical
analyses were performed using StatViewTM(Abacus
Concepts 1986).
RESULTS
Leaf litter quality
Initial foliar nutrient levels differed among leaf
types and in several cases differences were signifi-
cant (Table 2). Canopy trees had similar levels of
foliar nitrogen, but moss had most nitrogen. Litho-
carpus species had most carbon and highest C : N
ratio. Lithocarpus chintungensis had most lignin.
Castanopsis wattii had most foliar P, least lignin and
lowest Lignin : P and N : P ratios. Bamboo foliage

Table 2 Initial chemical composition of five leaf types in the old-growth forest in Ailao Mountain

Lignin N P C
Species (%) (%) (%) (%) Lignin/N Lignin/P C:N N:P

Lithocarpus 42.29 b 1.25 ab 0.091 b 48.64 b 34.05 b 466.37 c 39.13 a 13.76 c

xylocarpus (1.99) (0.10) (0.008) (0.91) (3.12) (33.34) (2.59) (1.34)
Lithocarpus 47.86 a 1.29 a 0.074 c 50.62 a 37.31 ab 647.84 b 39.48 a 17.37 b
chintungensis (1.35) (0.09) (0.005) (1.00) (1.90) (28.41) (2.28) (0.44)
Castanopsis 33.60 c 1.30 a 0.101 a 42.23 d 25.98 c 333.95 e 32.67 b 12.70 c
wattii (1.18) (0.06) (0.006) (1.11) (1.22) (17.79) (0.83) (0.94)
Sinarundinaria 42.55 b 1.08 b 0.054 d 36.31 e 39.51 a 789.92 a 33.77 b 20.96 a
nitida (1.84) (0.14) (0.006) (0.95) (3.28) (35.26) (3.31) (0.40)
Moss 34.16 c 1.39 a 0.083 bc 43.38 c 24.71 c 410.24 d 32.12 b 16.61 b
(0.98) (0.09) (0.004) (0.78) (1.63) (17.39) (1.56) (0.52)
F 82.140 4.452 39.007 140.415 24.785 186.599 8.489 40.440
P 0.0001 0.0347 0.0001 0.0001 0.0001 0.0001 0.0056 0.0001
Values are mean, n = 3 with SD in parentheses. Same letters indicate samples which are not significantly different (P < 0.05)
within a column, using Fishers test.
 Leaf litter decomposition in montane forest 439

had least nitrogen, phosphorus and carbon with
highest lignin : N, lignin : P and N : P ratios. Moss
(with most N) had lowest lignin : N and C : N
ratios.
Litter decomposition
Changes in mean dryweight of litter of all leaf
types during the decomposition period were
recorded (Table 3). Differences between bags at
each sampling occasion for all leaf types tended to
increase with time. In general, there were lower
values of the coefficient of variation taking five leaf
period decreased to 0.0440.057 g g-1 for canopy
species, 0.039 g g-1 for bamboo and 0.029 g g-1
for moss.
Castanopsis wattii leaf litter decomposed faster
than other species (Table 3; Fig. 2). The lowest
decomposition was found in mixed mosses. After
22 months, 49% of moss mass remained while
C. wattii had only 24%. Although bamboo (S.
100

Weight remaining (%)

types and all sampling times, ranging from 0.01 80
to 0.18. Higher coefficient of variation between
bags was found at month 18 of exposure when the 60
litter weight increased significantly. Castanopsis
wattii had the highest coefficient of variation.
Loss of dryweight followed the same trend for all 40

leaf types (Fig. 2). Decomposition was more rapid

in the first 12 months, in which the mean monthly 20
decomposition rate was 0.0530.074 g g-1 for the 0 3 6 9 12 15 19 21 24
three canopy species, 0.048 g g-1 for bamboo, and Time (months)
0.045 g g-1 for mixed moss. Between 14 and 18
Fig. 2. Dry weight remaining as a percentage of the
months all leaf types increased in weight. In the initial mass in decomposing litter of five leaf types in
subsequent 4 months the rates of decomposition of the old-growth forest on Ailao Mountain. (), Litho-
all leaf types decreased rapidly. The mean monthly carpus xylocarpus; (), Lithocarpus chintungensis; (), Cas-
decomposition rate within the 1422-month tanopsis wattii; (), Sinarundinaria nitida; ( ) , Moss.

Table 3 Mean dryweight (g) SD (n = 5) of litter of five leaf types decomposed in the old-growth forest in Ailao
Mountain

Time Lithocarpus Lithocarpus Castanopsis Sinarundinaria

(months) xylocarpus chintungensis wattii nitida1 Moss F P

0 5.00 5.00 5.00 5.00 5.00

3 4.26 0.06 a 4.16 0.11 ab 3.92 0.11 c 4.07 0.30 abc 4.00 0.16 bc 2.843 0.059
6 3.44 0.11 b 3.81 0.08 a 3.30 0.11 b 3.93 0.21 a 4.00 0.16 a 21.870 0.0001
9 2.76 0.13 b 3.38 0.19 a 2.74 0.11 b 3.57 0.35 a 3.66 0.24 a 20.496 0.0001
12 1.91 0.13 d 2.48 0.18 c 1.92 0.24 d 2.90 0.15 b 3.38 0.28 a 43.101 0.0001
14 1.70 0.28 c 2.28 0.27 b 1.73 0.05 c 2.42 0.29 b 2.99 0.20 a 23.788 0.0001
16 2.06 0.19 c 2.56 0.29 b 2.22 0.28 bc 3.20 0.32 a 3.48 0.33 a 26.480 0.0001
18 2.38 0.42 b 2.64 0.15 b 2.58 0.46 ab 2.80 0.42 ab 3.04 0.16 a 2.755 0.0644
20 1.76 0.48 c 2.37 0.30 b 1.57 0.14 c 2.22 0.11 b 2.86 0.28 a 35.865 0.0001
22 1.44 0.11 c 1.72 0.18 c 1.21 0.18 d 1.93 0.15 b 2.46 0.17 a 51.372 0.0001
Figures in parentheses are proportion of the initial weight remaining at different interval. Same letters indicate samples which
are not significantly different (P < 0.05) within a row, using the Fisher test.
1
In the case of S. nitida, the original dryweight was 3.00 g. Values in this table have been converted in proportion, as if the
original weight was 5.00 g. Actual mean weight of decomposing litter was in order 2.44, 2.36, 2.14, 1.74, 1.45, 1.92, 1.68,
1.33 and 1.16 g.
440 W. Liu et al.
nitida) has thin and less rigid leaves (density
0.017 0.001 g cm-2), its decomposition rate was
lower than the canopy species with thicker
and tough leaves (densities of foliage of 0.030
0.002 g cm-20.035 0.002 g cm-2).
With Olsons (1963) model, C. wattii leaf litter
had the highest decomposition rate constant (k),
with a turnover time (1/k) of 1.6 years and the time
required for 95% loss being 4.5 years. Lithocarpus
xylocarpus, L. chintungensis and bamboo followed in
order. The mixed moss material decayed more
slowly, requiring 9.4 years for 95% decomposition
(Table 4).
Concentration of nutrients in
decomposing litter
Concentrations of N, Mn and Fe in the residual
tissue increased steadily over the 22 months. There
was a slighter increase of N and Mn for tree foliage
compared with bamboo and moss (Fig. 3). The P
concentration in all species also increased in the
first 12 months followed by a gradual decrease
later.
Although initial K levels differed among leaf
types, the changes in concentration of this element
were similar for all five leaf types. Major reductions
in K occurred in the first 6 months, followed by
little change subsequently. The Ca concentration
of bamboo was half that of other leaf types, except
for C. watti. Despite some fluctuation, the Ca levels
were fairly constant. Changes in Mg concentration
were variable, but increased levels occurred at
1214 months, and as with Ca decreased to 12
months and then increased to 16 months, except
for bamboo.
Table 4 Coefficients of regressions used to estimate decomposition rate constants (k), turnover time (1/k) and time
required for 95% loss for five leaf types
Regression coefficient
Species X0 (%) k (year-1)
Lithocarpus xylocarpus 91.8356 0.6211
Lithocarpus chintungensis 95.2019 0.4964
Castanopsis wattii 91.5513 0.6447
Sinarundinaria nitida 91.5513 0.4513
Moss 93.5223 0.3118
The regression was of the form Xt = X0e-kt, where Xt is the percentage of the mass remaining at time t (years), X0 is the initial
mass in percent dryweight, and k is the decomposition rate constant. **P < 0.01.
Absolute amount of nutrient in
decomposing litter
The observed change in concentrations of most
nutrient elements did not correspond to net
immobilization of nutrients. Change in the
absolute amount of nutrient during decomposition
is a function of loss in mass and of change in rela-
tive nutrient concentration in the remaining
litter. Change in the absolute amount of each
nutrient is expressed as a percentage of the initial
amount that is remaining (Fig. 4). Canopy species
showed initial N immobilization and then later
release, while bamboo and moss had a net release
of N over 22 months. By 22 months the net release
levels of N from L. xylocarpus, L. chintungensis, C.
wattii, S. nitida and moss were 43, 47, 60, 30 and
39% of the initial amount, respectively. With the
exception of C. wattii and L. xylocarpus, P in
decomposing litter increased in the first 12
months, especially for moss, reaching 152% at 6
months and 130% at 12 months, of the initial
level. By 22 months the net release levels of P from
the five litter types were 57, 62, 72, 44 and 40%,
respectively, of the initial amount. More than 60%
of K was released from all species within 6 months,
and >75% by 12 months with little subsequent
loss. By 22 months the net release levels of K from
the five litter types were 81, 82, 91, 85 and 77%,
respectively, of the initial amount. During the 22-
month period there was a net release of Ca from all
the litter types with rapid initial loss, followed by
a more gradual release. Both Ca and Mg increased
from 12 to 14 months, after initial declines. At 22
months the net release levels of Ca were 73, 69,
83, 63 and 53% respectively, and the net release
Turnover time Time (years) required
r2 (1/k) (year-1) for 95% loss
0.8526** 1.61 4.68
0.8864** 2.01 5.94
0.8237** 1.55 4.51
0.8754** 2.22 6.44
0.8913** 3.21 9.39
 Leaf litter decomposition in montane forest 441

(a) (b)
2.5 0.2

0.15
2

0.1

1.5
0.05

1 0

(c) (d)
0.8 1.2

Nutrient concentration (%)

0.6 0.9

0.4 0.6

0.2 0.3

0 0
Nutrient concentration (%)

(e) (f)
0.3 0.3

0.2 0.2

0.1 0.1

0 0
0 3 6 9 12 15 18 21 24
(g) Time (months)
1.5

1.2

0.9

0.6

0.3

0
0 3 6 9 12 15 18 21 24
Time (months)

Fig. 3. Change in nutrient concentration in decomposing litter of five leaf types in the old-growth forest on Ailao
Mountain. (a) Nitrogen; (b) phosphorus; (c) potassium; (d) calcium; (e) magnesium; (f ) manganese; (g) iron.(),
Lithocarpus xylocarpus; (), Lithocarpus chintungensis; (), Castanopsis wattii; (), Sinarundinaria nitida; ( ) , Moss.
442 W. Liu et al.

(a) (b)
120 160

100
120
80

60 80

40
40
20

0 0

(c) (d)
100
100

Nutrient remaining (%)

80 80

60 60

40 40
Nutrient remaining (%)

20 20

0 0
(e) (f)
120 200

100
150
80

60
100
40

20 50

0
0
(g)
2500 0 3 6 9 12 15 18 21 24
Time (months)
2000

1500

1000

500

0
0 3 6 9 12 15 18 21 24
Time (months)

Fig. 4. Change in absolute amount of nutrient in decomposing litter of five leaf types in the old growth forest on
Ailao Mountain. (a) Nitrogen; (b) phosphorus; (c) potassium; (d) calcium; (e) magnesium; (f) manganese; (g) iron.
(), Lithocarpus xylocarpus; (), Lithocarpus chintungensis; (), Castanopsis wattii; (), Sinarundinaria nitida; ( ) , Moss.

levels of Mg were 61, 53, 81 and 46% for canopy
species and moss respectively, however, Mg in
bamboo litter increased slightly. In canopy species,
Mn had an initial release and then had small
increases at 18 months, while Mn in bamboo and
moss material was greater, reaching 118% and
171% of the initial amount, respectively. Iron was
an accumulative element in this experiment: some
2641425% of the initial amount was accumu-
lated in all leaf types over the 22-month period.
For the 22-month period, the aggregate rate of
nutrient release for all leaf types followed the
sequence of K > Ca > Mg > P > N > Mn > Fe, except
for bamboo (S. nitida) where the corresponding
order was K > Ca > P > N > Mg > Mn > Fe.
DISCUSSION
Litter decomposition of all leaf types was initially
rapid, followed subsequently by a slower rate.
Faster initial rates may reflect the decay of easily
decomposable tissues and readily mineralizable
elements (Swift et al. 1979; Swift & Anderson
1989; Kumar & Deepu 1992; Songwe et al. 1995;
Sundarapandian & Swamy 1999). Residual tissues
decompose less readily. Relatively slower decay
rates in later stages may be attributable to the
accumulation of more resistant or stable constitu-
ents in the residual litter mass (Sundarapandian &
Swamy 1999). Leaching is the main factor influ-
encing weight loss of litter during the initial
stages (Swift et al. 1979; Blair 1988; Berg et al.
1993; Songwe et al. 1995). Pardo et al. (1997)
reported that leaching and decomposition were
halted during summer dryness. Higher relative
loss of mass during the rainy season than during
the dry season is a feature of tropical forest
(Sundarapandian & Swamy 1999). The timing of
placement of the litterbags is an important factor
that affects decomposition rates in regions where
distinct dry and wet seasons alternate. In Xujiaba,
the rainy season occurs from June to October and
the dry season from November to May. Lithocarpus
species litterfalls were highest in AprilMay and
OctoberNovember. The decomposition of leaf
litter experiment commenced early in the dry
season. During the study period, samples taken at
6 and 18 months coincided with the change from
dry to wet weather. Loss of the dryweight of the
Leaf litter decomposition in montane forest 443
decomposing litter was higher in the rainy period
than in the dry period, although some rainfall
occurred early in the dry season, which resulted in
higher decomposition rates. During the height of
the dry season (between 3 and 6 months, and
between 14 and 18 months, or MarchMay 1998
and 1999) lower decomposition rates of litter
occurred for all leaf types, presumably because
with scarce moisture, less leaching can occur at
those times.
Decomposition rates between leaf types differed
significantly. Varying rates of decomposition have
been related to initial nutrient composition and
lignin concentration in leaves (Singh & Gupta
1977; Melillo et al. 1982; Upadhyay 1993;
Lisanework & Michelsen 1994; Couteaux et al.
1995; Bubb et al. 1998; Murphy et al. 1998;
Cotrufo et al. 1999). Other authors have attributed
variability to morphological features of foliage
(Edwards 1977; Songwe et al. 1995). Wedderburn
and Carter (1999) emphasized that N and lignin
concentration and the lignin : N ratio in leaf litter
appear to be likely controls of the rate of decay. Of
the three canopy species in the present study, C.
wattii and L. xylocarpus had lower lignin and higher
N and P concentrations in initial leaves and these
decomposed faster than that of L. chintungensis,
which had higher lignin concentration and lower
N and P concentration. It has also been shown
that the proportion of lignin, N and P in litter
affects decomposition rate (Melillo et al. 1982;
Lisanework & Michelsen 1994; Couteaux et al.
1995). Phosphorus deficiency, higher N : P and
lignin : P ratios in decomposing leaf litter are asso-
ciated with lower decomposition rates (Lisanework
& Michelsen 1994; Gallardo & Merino 1999).
Bamboo leaves decomposed more slowly than
canopy species leaves. Bamboo leaves are thinner,
less rigid and of lower density than leaves of
canopy species, but they are distinguished by
higher lignin : N, lignin : P and N : P ratios,
although lignin in bamboo leaves is not lowest
among the tested litter species. These results
suggest that decomposition rates of canopy species
and bamboo in the same environmental conditions
may be controlled more by nutrient composition
and lignin concentration than by their morpho-
logical features.
In the case of moss, the decomposition rate
seems less related to initial nutrient concentration.
444 W. Liu et al.
Daubenmire and Prussco (1963) found poor
intraspecific correlation between N content and
decomposition rates. Lower decomposition rates in
species with higher N and P concentrations also
occur in tropical forest (Sundarapandian & Swamy
1999). Moss had higher N and P concentrations in
initial material. After an initial rapid weight loss,
moss subsequently decomposed more slowly. The
lower rate of decomposition of moss may be asso-
ciated with its higher content of repellent tissues
(Nakatsubo et al. 1997). That bryophytes are effec-
tive in uptake and slow release of nutrients in
humid mossy forest is reported elsewhere (Li &
Vitt 1997; Nakatsubo 1997; Nakatsubo et al.
1997). The higher initial nutrient concentration of
mosses could be attributed to the presence of many
microbes (fungi, acarids, arthropods, etc.) in moss
tissues and to atmospheric deposition captured by
such species (Coxson & Nadkarni 1995). These
loosely attached nutrients would be readily leached
in the first stage, leaving more resistant structural
materials that may decay more slowly. The resist-
ance to decomposition and the low potential to
immobilize nitrogen for moss litter is in agree-
ment with results reported elsewhere (Coxson &
Nadkarni 1995; Hobbie 1996; Nakatsubo et al.
1997).
The dryweight of the decomposing litter
increased 820% within 1218 months of the
start of the experiment. Apparent increases of the
bagged leaf litter weight over some months were
observed in five species of tropical forest in
Cameroon (Songwe et al. 1995). This could be due
to the entry of organisms or other contaminants
excluded by the mesh size used (Swift et al. 1979;
Songwe et al. 1995). At 19 months, the dryweight
of the decomposing litter decreased. This was the
time of onset of the summer wet season and the
increased moisture may have washed the debris.
The decomposition rates of the canopy species in
the present study were high, compared to values
reported for temperate forests. Decomposition con-
stants of some temperate hardwood species ranged
from 0.08 to 0.47 (Melillo et al. 1982), and for pine
species from 0.307 to 0.46 (Das & Ramakrishnan
1985). Songwe et al. (1995) reported that mean
decomposition constants varied from 1.60 to 4.18
for five species with different leaf morphological
characteristics in tropical forest. These gave times
required for 50% decay of 1.8 to 4.6 months and
turnover time (1/k) of 2.2 to 7.7 months. In the
present study, the time required for 95% decay
ranged from 4.5 to 9.4 years, higher levels than
other reported ranges. Thus our decomposition
rates for canopy species in the middle mountain
humid, evergreen, broad-leaved forest lie in the
upper part of the range for temperate forest and in
the lower part of the range recorded for montane
tropical forest. This is consistent with our under-
standing of the status of these forests as being in a
transitional or intermediate category in relation to
environmental conditions.
The N and P concentrations in litter of all
species increased in the first 12 months, followed
by a decrease corresponding with the subsequent
loss in mass. The absolute amounts of these two
elements in the decomposing litter varied with the
different decay rates between species, as elsewhere
(Palm & Sanchez 1990; Lisanework & Michelsen
1994; Songwe et al. 1995). High concentrations of
N and P in decomposing litter are affected by
atmospheric deposition, leaching, insect frass and
fungal accumulation (Swift et al. 1979; Lisanework
& Michelsen 1994). The N, and especially P,
concentration in decomposing residues results in
an increase in P-deficient substrates (Alexander
1976). The high accumulation of P in decompos-
ing litter in the initial stage has also been observed
in other forests (Upadhyay & Singh 1989; Palm &
Sanchez 1990; Lisanework & Michesen 1994).
Songwe et al. (1995) noted that the high initial
build up of N and P in decomposing litter might
reflect the degree of limitation of these elements
in litter and soil. At Xujiaba, P concentration in
the soil is relatively low. The fresh leaf litter of
canopy species and of bamboo have a higher N : P
ratio. In decomposing litter, this ratio decreased
initially and then subsequently increased with
decreased decay rate. It is not unusual to find a
high N : P ratio in fresh litter in soils where P is
limited, affecting litter decomposition (Lugo &
Murphy 1986; Lisanework & Michelsen 1994).
This suggests P limitation in the soil substrates, a
hypothesis supported by the low available P found
in all soils (Michelsen et al. 1993). The high
ratio of N : P indicated that P was limiting the
litter decomposition of the natural forest in
the Ethiopian highland in the beginning and
at the end (Lisanework & Michelsen 1994). Vogt
et al. (1986) noted that the ideal N : P ratio for

decomposers was 10 and that higher ratios limited
litter decomposition. In the present study, P
seemed to limit the decomposition of the canopy
species, bamboo and moss litter both at early and
at later stages.
ACKNOWLEDGEMENTS
We thank the Chinese Academy of Sciences and
The Foundation for Natural Sciences of Yunnan
Province, China, for financial assistance through a
research grant. The Australian Department of
Employment, Education, Training and Youth
Affairs (DEETYA), is thanked for provision of an
Overseas Postgraduate Research Scholarship to L.
Wenyao to undertake a PhD degree at the School
of Environmental Biology, Curtin University of
Technology. The Management Authority of the
Ailao Mountain Natural Reserve is thanked for
granting permission to undertake the research
reported. Thanks are also due to Messrs Yang
Gun-ping, Li Shou-chang and Li Da-wen for
assistance in fieldwork. Ms Fu Yun and Ms Xie Ke-
jing are particularly thanked for their assistance
with chemical analysis of soil samples.
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