全 文 ：Journal of Forestry Research, 12(3):169-175 (2001) 169
Community features of Indocalamus wilsoni in the Shennongjia Na-
tional Nature Reserve, China
LI Zhao-hua, Manfred Denich
(Center for Development Researches (ZEF), Bonn University, D-53113 Bonn, Germany)
Abstract: Four vegetation types, namely coniferous and broadleaved mixed forest, secondary deciduous broadleaved forest,
open shrubs and meadow, with dwarf bamboo (Indocalamus wilsoni) are compared on their floristic composition, life form,
community structure, and survivorship of bamboos. Disturbance of conifer-logging two decades ago promoted the species di-
versity, but negatively influenced the survivorship of dwarf bamboo which is dominate on the forest floor. Open shrubs bear
more species diversity and more geophytes in its life-form spectrum than its background vegetation subalpine meadow, how-
ever, dwarf bamboo is growing much better in the open shrubs than in the meadow. It seems dwarf bamboo probably favor-
ites to occur in the vegetation at gentle slopes with a mediate canopy cover. Its mass flowering in the secondary deciduous
forest suggests that a simultaneous flowering and following dieback maybe irrupt in the recently coming years over the subal-
pine Shennongjia in China, which will be not only alter the floristic composition and community structure of the old bamboo
stands, but also influence the survivorship of this rare species.
Key words: Indocalamus wilsoni; Community structure; Survivorship
CLC number: S795.9.02 Document code: A Article ID: 1007-662X(2001)03-0169-07
Introduction 1
Dwarf bamboo, Indocalamus wilsoni (Rendle) C. S.
Chao et C. D. Chu (Syn. I. nubigenus (Keng f.) Yi ex H. R.
Zhao et Y. L. Yang), is a rare species which discretely en-
demics in the subalpine belt of the Shennongjia National
Nature Reserve (110°0305-110°3350 E, 31°2120-
31°3020 N) with an elevation ranged from 1 700 m up to
2 500 m (Zhu et al. 1994; Zheng 1993; Zhu et al. 1999). As
a monocarpic plant, it usually forms dense understorey,
with a height varied from 0.5 to 1.2 m, of forest and shrubs
or extends into the meadow to form bamboo mosaics. The
life history of many subalpine bamboos is characterized by
an unusual flowering habitat. They often flower to die si-
multaneously over a wide area after vegetative reproduc-
tion for several decades (Qin 1985; Yi 1988; Campbell
1987; Campbell 1991; Tian 1991; Wang et al. 1991; Taylor
et al. 1992; Makita 1992; Taylor & Qin Z. S. 1996; Zhou &
Huang 1996). This die back is an endogenous disturbance
that might promote tree seedling establishment and alter
community structure, composition or both. Meanwhile, the
existing community structure and composition should influ-
ence the bamboo regeneration thus may shift the vegeta-
tion in the old bamboo stands.
We noted in September 2000 that dwarf bamboo started
to mass flowering in a secondary broad-leaved forest and
Biography: LI Zhao-hua (1964-), male, Ph. D Candidate in the Ceter for
Development Researches (ZEF) of Bonn University, ZEF, Bonn Univer-
sity, Walter-Flex-Str. 3, D-53113 Bonn, Germany. E-mail:
ZLI9@hotmail.com
Received date: 2001-05-05
Responsible editor: Chai Ruihai
about 30%-50% of bamboo culms in the stand had already
withered away. To protect this rare species and observe its
regenerative process after flowering and the future suc-
cession of its associated vegetation, an ecological study is
immediately needed since there is no any ecological atten-
tion have been given to this bamboo species yet. This
study is attempted to demonstrate the present community
features of bamboo stands on the aspects of floristic com-
position, life form, cover structure, and survive situation of
this dwarf bamboo.
Study area
The Shennongjia National Nature Reserve, located at
the western terminus of the Daba mountain range on the
northern-west border of Hubei Province, comprises 70 467
hm2 of steep rugged mountains and elevations that range
from 480 m to 3 105.4 m. The extreme topographic relief
coupled with the elevation range provides habitats for an
incredible diverse flora and the climatic conditions that ex-
tend from northern subtropics through cold temperate (Zhu
et al, 1999; Ban et al. 1996). The zonal climate in Shen-
nogjia is a transitional type of the north subtropical and the
warm-temperate moist monsoon, controlled mainly by sub-
tropical circulation, but there is a complex climate gradient
that corresponds to elevation, however the dwarf bamboo
Indocalamus wilsoni occurs only from 1 700 m to 2 500 m
along the elevation. In this belt, with the increase of altitude,
the mean annual temperature varies from 7.4°C to 4.8°C
and annual precipitation increases from about 1 800 mm
up to 2 500 mm.
After a general vegetation survey, the ecological investi-
gation of dwarf bamboo was carried out in four selected
LI Zhao-hua et al. 170
sites: Small Qianjiaping (GPS location: 31°23336N,
110°24476E, Alt. 2040 m), Qianjiaping (31°23 345N,
110°24500E, Alt. 1990 m), White-Water Valley
(31°27704N, 110°09489E, Alt. 2300 m) and Tiger Ditch
(31°27648N 110°09528E Alt. 2450 m). These sites re-
spectively represented four main community types of dwarf
bamboo in the Shennongjia National Nature Reserve: a)
bamboo under coniferous and deciduous broad-leaved
mixed forest (Small Qianjiaping), b) bamboo under decidu-
ous broad-leaved forest (Qianjiaping), c) bamboo within
subalpine open shrubs (White-Water Valley) and d) bam-
boo in subalpine meadow (Tiger Ditch).
The Small Qianjiaping plot (Plot A as blow) was located
on a gently rolling slope of a mountain floor, and the Qian-
jiaping plot (Plot B) was located on the same mountain at a
concave slope (Table 1). Both plots face an alluvial basin
which was covered by hygrophytic grasses from the gen-
era of Carex, Junicus and Scirpus and scrubs such as
Salix carprea var. inica, Crataegus wilsoni, Lespedeza
Formosa and Prunus szechuanica. The original mountain
vegetation type on this region belongs to the coniferous
and broadleaf mixed forest (Ban 1995), but most of this
forest formation in this area had been destroyed due to
heavy logging of Pinus armandii in 1970s and early 1980s
before the establishment of the nature reserve. Plot A is an
easily accessible site where Dwarf bamboo is keeping in
the original mixed forest, and Plot B. represents the eco-
logical features of the bamboo in the disturbed community
of deciduous broadleaved forest. We set up Plot B also
because the bamboo here is starting to mass flower, and a
further observation will be carried on at this site.
Table.1. Environmental characteristics of the Indocalamus wilsoni stands
Plot Slope Aspect Alt. (m) Landform Soil type Veg. type
Small Qianjiaping (A) 15º S40ºE 2040 Floor slope Brown soil Forest
Qianjiaping (B) 35º S70ºE 1990 Concave slope Brown soil Forest
White-Water Valley(C) 25º S15ºW 2300 Ridge slope Dark-brown soil Shrubs
Tiger Ditch (D) 5º N60ºE 2450 Crest terrace Meadow soil Meadow
White-Water Valley plot (Plot C) and Tiger Ditch plot
(Plot D) situated at the south part of the reserve. Here tec-
tonic uplift and fluvial erosion have produced a deep
incised topography in the landform. Slopes are steep (30-
70o), but the crests and terraces of the mountain are rela-
tively plain. Where the vegetation is quite complex and
poorly understood. Most of steep slopes are covered by
secondary vegetation of deciduous broad-leaved forest
with arrow bamboo (Fargesia spp.) as the understorey,
while the original dominant conifer Abies fargesii remains
at the deep valley. Open shrubs and meadow are relatively
stable vegetations, which occupy the upper plateaus with
arrow bamboo clumps. Dwarf bamboo frequently occurs as
density understorey of the open shrub or as mosaic within
the meadow. White-Water plot (Plot C) and Tiger Ditch plot
(Plot D) show the ecological characteristics of the draft
bamboo in these two community types.
Methods
All field data were collected between September 2000
and April 2001. Four representative plots were selected
after an overall observation in the reserve. In each plots,
we set up a 20 m×20-m quadrat to quantify the upper sto-
rey. All living trees and high-shrubs taller than 2 m in height
were numbered, their heights, subaxle heights, diameters
at breast height (DBH), and crown projection were meas-
ured. In the 20 m×20-m quadrat, we also set up several
2m×2-m sub-quadrats for recording understorey and 1m
×1-m sub-quadrats for recording the draft bamboo Indo-
calamus wilsoni and herbs. Of each species in the under-
storey and herb layer the height, coverage, dominance,
and life form (using Raunkiaers system) were noted. A
particular attention was given to the bamboo on its com-
munity features.
Life form spectra were calculated based on the cover-
abundance rate for each species on a 5-points scale and
each is assigned a sociability index (Moore & Chapman,
1986), also derived from a 5-point scale: 1- Growing once
in a place, singly; 2- Grouped or tufted; 3- In troops, small
patches or cushions; 4- In small colonies, in extensive
patches or forming carpets; 5- In great crowds or pure
populations. The floristical similarity between plots based
on presence/absence data of species was calculated using
the Similarity Index of Sørensen (ISs): ISs=W/(A+B)´100%,
where W is the number of common species and A and B
the number of species in plot A and B respectively. To
compare the species diversities among the plots, Shannon
index and McIntoshs index (Kent, 1996) were calculated
for each plot.
For understanding the relationship between the canopy
cover and the survivorship of dwarf bamboo, a detailed
cover profile for each individual stand was presented, and
the living situation including height, density, cover of bam-
boo was analyzed.
Results
Floristical composition and diversity
There are 38 dominant species were checked out in our
four plots, which allocated in Plot A, B, C, and D are 15, 22,
24 and 18 respectively. The floristic similarity (Table 2)
shows that Plot C and D are very similar in species com-
position (ISs as high as 0.71), and Plot A and B are also
Journal of Forestry Research, 12(3):169-175 (2001) 171
similar in the floristic aspect (ISs = 0.65), while Plot A and
D are very different in the species composition (ISs = 0.24).
It suggests that the floristic feature has not changed dra-
matically when zonal mixed forest (Plot A) degraded to
deciduous broad-leaved forest (Plot B) in the past two
decades, and open shrubs (Plot C) has a tightly floristic
connection with its backgrounds vegetation of meadow.
Table 2. Sørensen similarity index of species in dwarf bam-
boo stands of Plot A, B, C and D
A B C D
A 0.35 0.54 0.76
B 0.65 0.57 0.65
C 0.46 0.43 0.29
D 0.24 0.35 0.71
Note: Down-left section shows the similarity (ISs) and upper-right
section shows the difference (D= 1 – ISs)
Although all stands are rich in species comparing with
the coniferous -- arrow bamboo (Fargesia spp) formation at
the same altitude (Li 1994), the difference of diversity be-
tween stands is quite clear. Both Shennons and McIn-
toshs indices shows that the species diversity of the open
shrubs (Plot C) is the highest, the meadow ranks (Plot D)
the second highest, and the secondary deciduous forest
(Plot B) and the original mixed forest (Plot A) rank as the
third and the forth respectively (Fig. 1 & 2). The diversity
indices indict only a slight difference between plot A and
Plot B, but their floristic compositions are very different. In
Plot B, when conifer Pinus armandii was cut down, the
sun-like species Betula albo-sinensis, Populus davisina,
Acer mono developed as the over-storey and Lindera ob-
tusiloba, Crataegus wilsoni, Cotoneaster acutifolius, Vibur-
num ichsngense developed as the sub-storey (Table 3).
Usually the logging gaps created an opportunity to the es-
tablishment of both conifers and broad-leaved trees (Taylor
1990; Taylor et al, 1996; Li 1996; Shotaro et al, 1998), but
in Plot B the dense understorey of bamboo I. wilsoni may
reduce seedling success of conifers on the ground since
we checked no seedlings of P. armandii in our quadrats.
There is no clear difference of the ground herbs between
Plot A and B, which are dominated by some shade-
tolerance species such as Smilacina japonica, Rubia cardi-
folia, Oxalis grtffithii, Paris quadrifolia and ferns (Dryopteris
sp. and Pteridium sp.).
The open shrubs (Plot C) are remarkably rich in species,
however its overstorey is usually composed by several
common species such as Malus huphensis, M. kansuensis,
Sorbus huphensis, and Crataegus wilsoni. We noted that S.
huphensis can reach as high as 20 m at low altitude al-
though it performances as a shrub with height only about
2-4 m in the area above 2200 m. There is no distinctive
boundary between open shrubs and subalpine meadow
(Plot D) in our study site since individual shrub clumps are
frequently distributed in the meadow and species composi-
tion of the understorey of open shrubs is nearly as same
as that of the meadow which is mainly composed by grass
species of Arundinella hirta, Deyeuxia henryi, Imperata
cylindrical, Carex spp, and herb species such as Polygo-
num suffultum, Geranium henryi, Paris quadrifolia, Allium
prattii, Smilacina japonica, as well as several fern species
0
0.2
0.4
0.6
0.8
1
1.2
1.4
1.6
Plot A Plot B Plot C Plot D
S
he
nn
on
d
iv
er
si
ty
in
de
x
Fig. 1. Shennon diversity index of species in I. wilsonii stands.
It shows Open shrubs (Plot C) > Meadow (D) > secondary decidu-
ous forest (B) > mixed forest (A)
0
2
4
6
8
10
12
Plot A Plot B Plot C Plot D
M
cl
nt
os
h
s
di
ve
rs
ity
in
de
x
Fig. 2. McIntosh’s diversity index of species in I. wilsonii
stands.
It shows Open shrubs (Plot C) > Meadow (D) > secondary decidu-
ous forest (B) = mixed forest (A)
Life form and cover structure
The life form spectra (Fig. 3) show a similarity between
mixed forest and deciduous broad-leaved forest. The
above-ground phytomass in Plot A and B are made up for
over 40% by phanerophytes, as against 20% in open
shrubs (Plot C) and 6% in meadow (Plot D). Nevertheless
the higher proportion of phanerophytes in Plot B than in
Plot A suggests that the logging disturbance has enhanced
the recruitment of trees and shrubs. In Plot C, phytomass
is mostly dominated by Geophytes (38%), which is clearly
different from its background vegetation of meadow (Plot
D). The latest plot is mostly dominated by hemicrypto-
phytes (37%).
LI Zhao-hua et al. 172
In subalpine Shennongjia, where a thick cover of snow
protects the rhizomes of dwarf bamboo and the bulbs of
bulb species such as Smilacina japonica, Paris quadrifolia,
and Allium prattii from the damage of frost, all of our plots
are considerably rich in cryptophytes which survive the
cold winter with buds or shoot apices below ground. Al-
though Fliervoet classified the giant bamboo (Phyllostachy
pubescens) as phanerophytes (Fliervoet et al. 1989), we
placed the dwarf bamboo in the context of geophytes
mainly because its new shoots emerge in the next growing
season only from the underground winter-buds of the rhi-
zomes. The dominance of geophytes on the open shrubs
also suggests that the shrub canopy positively influenced
the distribution of the geophytes and negatively on the
hemicrytophytes.
Fig. 3. Life form spectra of the stands of Indocalamus wil-
sonii based on the abundance-cover scales of each species.
Ph = Phanerophytes, Ch = Chamae-phytes, H = Hemicrypto-
phytes, G = Geophytes, Th = Therophytes
The vertical structure of mixed forest in Plot A consisted
three layers: canopy (higher than 8 m), bamboo and
scrubs (8-0.5 m), ground-herbs (lower than 0.5 m). Fig. 4A
shows the coverage distribution of each layer from the
main species. Conifer Pinus armandii covered about 28%
of the stand area, while three main broad-leaved tree spe-
cies Salix polyclona, Chaenomeles cathayensis and
Prunus szchuanica covered 31%, 26% and 14% respec-
tively. Considered the canopy overlap, we estimated that
the real canopy coverage is about 60%. The understorey is
absolutely dominated by I. wilsoni which covered over 90%
of forest floor in our quadrats. Under the dwarf bamboo
layer, the thin herbs covered only 20% of the ground.
Comparing with Plot A, the vertical layers in deciduous
broad-leaved forest of Plot B are not very clear (Fig. 4B),
but it can roughly be calculated as four layers: canopy,
shrubs, bamboo and ground herbs. Three most dominate
deciduous tree species Betula albo-sinensis, Populus
daviviana and Acer mono covered 38%, 20% and 16% of
the stand respectively with the height between 10 m and
20 m. The real canopy coverage is about 85%, which
means the canopy in Plot B is much closer than in Plot A.
The shrubs are usually high 2 to 6 m, where they formed a
sub-canopy with a real coverage of about 30%-45%. Bam-
boo layer covered 70-80% of forest floor, and among it 30-
50% are withered culms after flowering. We noted in the
gaps and forest margins, the bamboos are still alive quite
well. It suggests that the dense canopy maybe trigged the
flowering of Indocalamus, or at least, shorted its life cycle.
Since the dieback of Indocalamus, more herbs are invers-
ing in the bamboo stands so that 30%-40% ground was
covered by the herbs.
Like the mixed forest in Plot A, the open shrubs in Plot C
also contain three layers: shrub layer (3-8 m), bamboo
layer (0.4-0.8 m) and herbaceous layer (5-40 cm). Total
canopy cover of shrub layer is about 30%, which is mainly
formed by three common species of Chaenomeles cathay-
ensis, Sorbus huphensis and Malus kansuensis (Fig. 4C).
The understory layer covered about 90% of the stand floor,
in which Indocalamus contributed with 60% of the total
cover, while the other 40% came from perennial grasses
and ferns. The meadow is usually considered as a mono-
layered vegetation types, but in plot D, its canopy is formed
by three clearly separate strata (Fig. 4D): upper grass stra-
tum (0.6-1.8 m), bamboo stratum (0.3-0.6 m), and low her-
baceous stratum (5-30 cm). The upper stratum is domi-
nated by some typical meadow grasses in subalpine
Shennongjia: Arundinella hirta, Imperata cylindrical and
Trisetum henryi. The total cover of upper grasses is about
35% against the stand, and bamboo stratum and low her-
baceous stratum presented the coverage as 80% and 25%
respectively. In the bamboo layer I. wilsoni contributed 50%
of the canopy, and the ferns are poorly represented in this
stratum.
Discussion
In our study site of Shennongjia, the survivorship of
dwarf bamboo closely responded to the composition, struc-
ture, and canopy cover of its associated vegetation. Forest
canopy density has a strong influence on the growth of
bamboo. In the Plot A, where with a mature forest canopy
small-intermidediate-sized canopy gaps had taller bamboo
with greater coverage than beneath tree crowns of Plot B
(Table 3). This indicts the disturbances of conifer logging in
the bamboo stands probably promoted the recruitment of
trees and shrubs, which formed a denser canopy, and then
the closed canopy reduced the annual growth of new bam-
boo culms that sprout once every year from its under-
ground rhizome. During the past two decades the mean
height of bamboo layer in the forests has reduced from 80-
95 cm in Plot A to 40-65 cm in Plot B, and the coverage
also reduced from 90%-95% to 60%-80%.
On the other hand, bamboo grew much weaker in the
meadow without upper canopy than under open shrubs
with large gaps. The mean height of bamboo layer in the
0
5
10
15
20
25
30
35
40
45
50
Plot A Plot B Plot C Plot D
Li
fe
-f
or
m
s
pe
ct
ru
m
(
%
)
Ph Ch H G Th
Journal of Forestry Research, 12(3):169-175 (2001) 173
open shrubs lower than in both of the mixed forest and the
secondary deciduous broad-leaved forest may be ex-
plained by variation of altitude since Plot C is located about
400 m higher than that of Plot A and B. Thus our investiga-
tion suggests that Indocalamus very favorites in the vege-
tation occurred at gentle slopes with a mediate overstorey
canopy.
(m) A
14
12
11
10
9
8
6
3
2
1.5
1.0
0.9
0.8
0.7
0.6
0.5
0.4
0.3
0.2
0.1
0
0 10 20 30 40 50 60 70 80 90 100
Coverage (%)
Pinus armandii Salix polyclona Ground herbs
Malus huphensis Chaenomeles cathayensis
Prunus szechuanica Indocalamus wilsoni
(m) C
8
7
6
5
4
3
2
1.0
0.9
0.8
0.7
0.6
0.5
0.4
0.3
0.2
0.1
0
0 10 20 30 40 50 60 70 80 90 100
Coverage (%)
Chaenomeles cathayensis Herbs
Sorbus huphensis Grasses
Malus kansuensis Herbs
Indocalamus wilsoni Ferns
(m) B
20
19
18
17
16
15
14
13
12
11
10
8
6
5
2
1.0
0.8
0.6
0.5
0.4
0.2
0
0 10 20 30 40 50 60 70 80 90 100
Coverage (%)
Betula albo-sinensis Populus davidana
Pinus armandii Acer sp.
Other broad-leaved Trees Indocalamus wilsoni(Died)
Indocalamus wilsoni (living ) Draft shrubs Herbs
(m) D
2.0
1.8
1.6
1.4
1.2
1.0
0.9
0.8
0.7
0.6
0.5
0.4
0.3
0.2
0.1
0
0 10 20 30 40 50 60 70 80 90 100
Coverage (%)
Grasses Herbs
Indocalamus wilsoni Rubus sp
.
Fig. 4 (A-D). Coverage-profiles of four vegetation types with Indocalamus wilsoni
(A. Mixed forest, B. Deciduous broad-leaved forest, C. Open shrubs, and D. meadow)
LI Zhao-hua et al. 174
Table 3. Living performance of Indocalamus in different vege-
tation types
Plot Height
(cm)
Density
(culm/m²)
Coverage
(%)
A 80-95 88 90-95
B 40-65 104* 60-80
C 60-75 73 60-70
D 40-60 55 40-50
A--mixed forest, B--broad-leaved forest, C.--open shrubs,
D--meadow.
* including 30%-40% of died bamboos
The density of Indocalamus shows a special pattern in
Plot B due to the flowering in the past two years. In our
quadrats, the total density of bamboo is as higher as 104
culms/m2, but 30%-50% of the culms died back after flow-
ering in the past two years. Our investigation on its rhi-
zome system shows that the flowering death temperately
stimulated the vegetative regeneration of new shoots,
which is very different from the most dominant subalpine
bamboos from the genera of Fargesia in Shennongjia. The
underground part of Frgesia is a sympodial system and the
shoots can only be produced from its densely packed
pseudo-rhizomes, while Indocalamus bears an amphpodial
system which contains both spreading monopodia and
packed sympodia. When Fargesia flowering, there are no
new shoots are sprouted during the flowering period and
its underground system usually decays away in the follow-
ing years. When Indocalamus flowering, we noted its sym-
podium system ceased to generate new shoots, but its
spreading rhizome was continually to extend and produce
new shoots and new packed rhizomes. The extraordinary
increase of the recruitment of new culms may be initially
promoted by the gaps formed by flowered culms.
Although the vegetative growth of Indocalamus has not
ceased immediately corresponding to the mass flowering,
there is no evidence to indict that this mass flowering will
not spread to over all of its habitats in Shennongjia. In fact,
we checked out at the site that in some quadrats, over
90% of bamboo culms died after flowering and there was
no any new shoots occurred in the year of 2000 and 2001,
but we had not collected any bamboo seeds in the soil
neither discovered seedling on the stands. So that, a fur-
ther observation is needed to focus on whether its regen-
eration after flowering is established from seeds or com-
pensated by new culms from its living rhizomes.
When a mountain bamboo species flowers, it is usually
dieback over a large area and this moment we have no
practicable methods to control the mass flowering and
succeeding death (Yi 1994; Taylor et al. 1991; Akifumi.
1992; Huang 1998). However, our suggestion for the pro-
tection of this rare species is that I. wilsoni is very sensitive
to the changes of upper canopy. Protecting its associated
vegetation probably is one of the best ways to maintain its
survivorship in the reserve.
Acknowledgement
We would like to thank Mr. Z. Q. Zhu, vice director of the
Shennongjia reserve, Mr. B. Y. Zhao, director of the re-
search institute of the reserve for thorough support in the
field. Particular thanks goes to Mr. B. Liu in Hubei Univer-
sity, Mr. C. Y. Xu, Mr. S. Q. Wang in the reserve, and other
couples of anonymous local people who assisted our
fieldwork. The work is a part of bamboo research project
within ZEF (Center for Development Researches) of Bonn
University, financed by GTZ, Germany.
References
Akifumi, M. 1992. Surviveship of a monocarpic bamboo grass,
Sasa kurilensis, during the early regeneration process after
mass flowering [J]. Ecological Research, 7: 245-254
Ban, J.D., Qi, G.S. & Liu, L.D. 1995. Studies on the natural vege-
tation of Mt. Shennongjia in western Hubei [C]. In: Ban, J. D. et
al. (eds) Studies on the Vegetation of Western Hubei. Wuhan:
Publish House of the Central China University of Technology
and Science, 9-89 (in Chinese with English abstract)
Barbour, M.G., Burk, J.H. & Pitts, W.D. 1980. Terrestrial plant
ecology [M]. The Benjamin/Cummings. California: Publishing
Company,
Campbell, J.J.N. 1987. The history of Sino-Himalayan bamboo
flowering, droughts and sun-spots [J], Journal of Bamboo re-
search, 6, 1-15
Campbell, J. J. N. 1991. Sino-Himalayan bamboos: twards a syn-
thesis of western and eastern knowledge [J]. Journal of Ameri-
can Bamboo Society, 8(1&2), 12-18
Carter, J., Ackleh, A.S., Leonnard, B.P. & Wang, H. B. 1999. Giant
panda (Ailuropoda melanoleuca) population dynamics and
bamboo (subfamily Bambusoidieae) life history: a structured
population approach to examining carrying capacity when the
prey are semelparous [J]. Ecological Modelling, 123: 207-223
Fliervoet, L.M., Zhong, Z.C., Liu, Y.C. et al. 1989. Divisity and
aboveground structure of the understoreys of Phyllostachys pu-
bescens grove on Mount Jinyun, Sichuan, China [J]. Flora, 182,
203-219
Greig-Smith, P. 1983. Quantitative plant ecology (3 ed.) [M]. Ox-
ford: Alden Press.
Huang, S. T. 1998. A probe to bamboo flowerings cause and pre-
ventions [J]. Journal of Bamboo Research, 17(2), 53-55 (in Chi-
nese)
Jenzen, D.H. 1976. Why bamboo wait so long to flower [J]. Annual
Review of Ecology and Systematics, 7: 347-391
Li, B. & Ban, J.D. 1995. Studies on farges fir formation in Mt.
Shennongjia [C]. In: Studies on the Vegetation of Western Hubei
(ed Ban, J. D. et al), Publish House of the Central China Univer-
sity of Technology and Science, Wuhan, 159-168 (in Chinese
with English abstract)
Moor, P. D. & Chapman, S.B. 1986. Methods in plant ecology
(Second edition) [M]. Oxford: Blackwell Scientific Publications,
Shotaro, Y. & Eiichi S. 1998. The effects of sika-deer browsing on
the biomass and morphology of a dwarf bamboo, Sasa nip-
Journal of Forestry Research, 12(3):169-175 (2001) 175
ponica in Mt. Ohdaigahara [J]. Forest Ecology and Management,
103: 49-56
Taylor, A. H., Reid, D.G., Qin, Z.S. & Hu, J.C. 1991. Spatial pat-
terns and environmental associates of bamboo (Bashania fan-
giana Yi) after mass-flowering in southwestern China [J]. Bulle-
tin of the Torrey Botanical Club, 118(3): 247-254
Taylor, A.H. & Qin, Z.S. 1993. Structure and dynamics of bamboos
in the Wolong Natural Reserve, China [J]. American Journal of
Botany, 80(4): 375-384
Taylor, A.H., Qin, Z.S. & Liu, J. 1996. Structure and dynamics of
subalpine forests in the Wang Lang Natural Reserve, Sichuan,
China [J]. Vegetatio, 124: 25-38
Tewari, D.N. 1993. A monography on bamboo [J]. International
Book Distributors, Dehra Dun
Tian, X.H. 1991. The natural regeneration of mountain bamboos
after mass flowering in Qinling [J]. Journal of Bamboo Research,
10(2), 26-31 (in Chinese)
Wang, J.X., Ma, Z.G., Liu, C.X. & Gan, L.M. 1991. Preliminary
study on the growth and development rules of Fargesia denu-
data bamboo [J]. Journal of Bamboo Research, 10(3), 38-48 (in
Chinese)
Yi, T. P. 1997. Compendium of bamboos in Sichuan [M]. Beijing:
Forestry Publish House of China, (in Chinese)
Zhang, J.Y. 1985. Investigation on the flowering, fruizing, and
regeneration behaviour in Precalamus utilis Keng [J]. Journal of
Bamboo research, 4: 86-88, (in Chinese)
Zheng, Z., Zhao, Z. E., Fu, S. X. et al. 1984. Plants in Shennongjia
[M]. Wuhan: Peoples Publishing House of Hubei (in Chinese)
Zheng, Z. 1993. A preliminary study on the flora of vascular plants
in Shennongjia, China [J]. Journal of Wuhan Botanical Studies,
11(2): 137-148 (in Chinese with English abstract)
Zhou, S. X. & Huang J.Y. 1996. Study on the population density of
young Bashania fangiana after natural regeneration [J]. Journal
of Bamboo Research, 15(4), 9-18 (in Chinese)
Zhu, S.L., Ma, L.S. & Fu, M.Y. 1994. Compendium of Chinese
bamboo [M]. Beijing: Forestry Publishing House, (in Chinese)
Chinese abstracts i
《林业研究》（英文版）2001年第 12卷第 3期
中文摘要
(Chinese abstracts attached to Journal of Forestry
Research, Vol. 12, No.3 (2001))
火炬松体细胞胚胎发生和干化体细胞的氧化物酶活性/唐巍
（O-23, E. S. King Village, 2900 Ligon St., North
Carolina State University, Raleigh, NC 27607, USA）//
Journal of forestry Research, -2001,12(3):147-152.
培养于附加2，4-D、BA和KT的愈伤组织诱导培养基上
的火炬松成熟合子胚在培养 3-9周后形成白色、半透明、有
光泽的粘性愈伤组织。这类愈伤组织形成于成熟合子胚的子
叶，但当用 NAA或者 IBA代替愈伤组织诱导培养基中的 2，
4-D时，它的诱导频率明显降低。这种粘性愈伤组织在分化
培养基上形成体细胞胚。体细胞胚经过去 50μm ABA和
8.5%PEG600处理后成为耐干化胚。扫描电镜观察表明，萌发处
理36小时后，耐干化胚恢复到干化处理之前的状态且大小和
形态正常，而不耐干化胚不能恢复到干化处理之前的状态且
表面撕裂。过氧化物酶活性的分析结果表明，耐干化胚有更
高的过氧化物酶活性。耐干化胚的高过氧化物酶活性可能与
催化H2O2的分解和保护体细胞胚免受氧化的伤害有关。
关键词：火炬松；体细胞胚胎发生；耐干化胚；过氧化物酶
活性
不同 N源对兴安落叶松苗木生长，养分吸收及离子平衡的影
响/陈永亮（东北林业大学，哈尔滨 150040），韩士杰（中国
科学院沈阳应用生态研究所，沈阳 110016），苏宝玲（沈阳
大学，沈阳 110016）//Journal of Forestry Research, -2001,
12(3):153-156.
本文研究了不同 N源对落叶松苗木干物重、养分吸收及
离子平衡的影响。结果表明：施铵态 N的苗木干物重小于对
照处理；施硝铵态N的苗木生长与对照和施硝态N的苗木差
别不大，但较施铵态 N的苗木生长要好。铵态 N与硝铵态 N
处理苗木地上部的阳离子总浓度受 N水平的影响很小，而硝
态 N处理时则阳离子浓度增加；各处理对地上部阴离子浓度
的影响均不大。在根部除硫酸根与钙离子外，阳离子与阴离
子浓度均变化不大。硝态N处理的苗木具有较高的C-A值；
△(C-A)/△Norg值在硝态N处理的苗木中随N供应量的增加而
略有变化，但铵态N处理则明显下降；硝铵态N处理苗木的
△(C-A)/△Norg值界于铵态N处理与硝态N处理之间。
关键词：N源；生长；养分吸收；离子平衡
杨树冰核细菌溃疡病发病主导因素及防治技术/项存悌，董爱
荣，刘学峰（东北林业大学，哈尔滨 150040，中国），李淳
（黑龙江省森防站），原树忠（黑龙江省森林植物园），张景
华（黑龙江省阿城市林业局）//Journal of Forestry
Research, -2001, 12(3):157-164.
通过对杨树冰核细菌分离培养、活性测定和室内外接种
试验，研究了冰核细菌的种类、分布、数量、致冻性和致病
性，结果表明冰核活性（INA）细菌普遍存在于东北地区杨树
上，在春秋季节霜冻条件下造成杨树冻害，是杨树冰核活性
细菌溃疡病发生的主导因素。通过对不同杨树品种品评调查
和人工接种试验，选出了适合东北不同地区的优良杨树抗病
品种、品系。壮苗试验表明，将杨树插穗进行沙藏处理和地
膜覆盖育苗可避免杨树霜冻害和旱害，减轻冰核细菌对杨树
苗木的侵染，又能提高苗木的生长。利用强冰核活性细菌专
化性抗血清技术，采用环状界面凝集法可早期检测杨树冰核
细菌溃疡病。采用链霉素等诱导剂浸泡杨树插穗进行育苗，
有明显的诱导抗病效果。在早春杨树萌芽前喷洒防冻保护剂
对该病有一定防治效果。
关键词:杨树；冰核细菌溃疡病；主导因素；防治技术
杨树冰核细菌溃疡病流行规律的研究/董爱荣，项存悌，刘学
峰（东北林业大学，哈尔滨 150040，中国），李淳（黑龙江
省病虫害防治站），王传伟，林海波（黑龙江省肇东市林业局），
张景华（黑龙江省阿城市林业局），潘淑英，邓立文（黑龙江
省大庆市林业局）//Journal of Forestry Research, -2001,
12(3):165-168.
利用相关性分析和主成分分析的方法，分析了冰核细菌
溃疡病发生特点与环境条件的之间的关系，选出了影响杨树
病害发生的主要影响因子。同时综合分析病害流行的主因和
诱因，提出东北地区杨树冰核细菌溃疡病危险等级划分标准，
并将东北地区分为重发区、常发区、偶发区和安全区 4个不
同等级冰核细菌溃疡病发生区。6种模型曲线的非线性回归
分析表明，冰核细菌溃疡病流行的时间动态以理查德生长模
型为最优生长模型。运用逐步回归分析方法，建立了多元线
性的中期预报方程，可以预测第二年的病情指数。根据病情
指数随时间增长的趋势，选用灰色方法建立 GM(1,1)数
学模型，可做出中长期预报。
关键词:杨树；冰核细菌溃疡病；流行规律；预测预报
中国神农架国家自然保护区鄂西箬竹(Indocalamus wilsoni)
的 群 落 特 征 /李 兆 华 ,Manfred Denich(C ter for
Development Researches (ZEF), Bonn University, D-53113
Bonn, Germany)//Journal of Forestry Research,
-2001,12(3): 169-175.
通过对含有鄂西箬竹的 4种植被，即针阔混交林、次生
阔叶林、疏灌丛及草甸的样地调查，比较分析了箬竹群落的
区系构成，生活型、群落结构及竹类生存状况。20年前的针
叶树种的采伐提升了样地的物种多样性，但影响了其下层植
物箬竹的生存。疏灌丛比其背景植被草甸含有更高的物种多
样性和更多的地下芽植物，然而疏灌丛下的箬竹长势更好。
箬竹偏喜于缓坡上具有中等盖度的植被环境。发生于次生阔
叶林中的箬竹群体开花预示着在整个神农架亚高山生境的箬
竹将在未来的几年内同时开花死亡。这不仅会改变现存竹类
Chinese abstracts ii
立地的区系构成和群落结构，同时也会波及到这一稀有物种
的生存问题。
关键词：箬竹；Indocalamus wilsoni；群落结构；生存
刺槐种子萌发生热曲线的测定及其萌发生长的热动力学研究
/胡云楚（中南林学院， 株洲 412006），周培疆，王存信，
屈松生（武汉大学化学与环境科学学院，武汉 430072）
//Journal of Forestry Research, -2001, 12(3):176-178.
种子的萌发与物质转化和能量交换紧密相关，对种子萌
发过程中的热效应进行研究，有助于认识种子的萌发机理及
其影响因素。采用武汉大学研制的新型热导式微量量热计测
定了刺槐种子的萌发生热曲线。分析了其萌发生热规律与种
子萌发生理之间的关系。并根据动力学原理对其生长阶段的
热谱进行分析，建立了刺槐萌发生长的动力学模型，25℃
P=208.77/[0.1937+0.8063exp(-0.06563t)]
关键词：生热曲线；热动力学；种子萌发；刺槐
长白落叶松纸浆林培育措施对经济效益的影响/李凤山（东北
林业大学，哈尔滨 150040）//Journal of Forestry Research,
-2001, 12(3):179-182.
根据长白落叶松纸浆林的培育实践，以经济指标内部收
益率和净现值为基础，分析了培育措施对经济效益的影响。
结果表明：轮伐期越短，经济效益越高，立地级越高，经济
效益亦越高。造林密度对经济效益的影响也较明显。在保证
造林成活率和保存率的基础上，用于整地和幼林抚育方面的
经费越少，经济效益则越高。选定的 6项措施对财务指标的
影响顺序为：轮伐期→立地条件→造林密度→管理费水平→
幼林抚育强度→整地方式
关键词：长白落叶松纸浆林；培育措施；经济效益
基于遥感信息的中国耕地动态变化和典型区的土地覆盖变化
/张佳华，董文杰（中国科学院大气物理所全球变化东亚研究
中心， 北京 10029），王长耀，刘纪远（中国科学院遥感应
用研究所，北京 100101），姚凤梅（北京林业大学生态工程
学院，北京 100083）//Journal of Forestry Research. -2001,
12(3):183-186.
利用多时相的Landsat-TM数据和国家资源调查数据，对
我国耕地动态变化和典型区的土地覆盖变化进行了研究，结
果显示：我国耕地的分布向东北和西北推移，移动的结果导
致耕地的生态背景质量下降。黄河河口的耕地和海岸带扩大，
扩大的速度是0.73 kma-1，沉积速率是2.1 kma-1。 对科尔沁
沙地的研究结果表明，该区沙化面积从70年代的60.02%上升
到80年代的64.82%,但是90年代后沙化面积缩小为54.90%；对
藏北的湖泊动态变化研究结果显示，湖泊的面积下降，下降
的速度为2.14 km2a-1。
关键词：遥感信息；耕地；土地覆盖变化；典型生态区；中
国
沙地云杉(Picea mongolica)在中国的形成、分布和特性/邹
春静，韩士杰，徐文铎(中科院应用生态研究所，沈阳)，苏
宝玲（沈阳大学生物与环境工程学院，沈阳 110044）
//Journal of Forestry Research, -2001, 12(3): 187-191.
沙地云杉是中国的珍稀濒危树种。沙地云杉林是陆地上
非常特殊的森林生态系统类型。它存在于我国森林草原过渡
带，也是我国的农牧交错区。在本文中，我们讨论了沙地云
杉的形成和分布，研究了沙地云杉的性质和特性。沙地云杉
属于白扦云杉系列，是白扦云杉长期适应当地气候而演化的
地方宗，然后形成异域的半种，最后形成分类种。沙地云杉
林是森林草原过渡带上的超地带性顶极群落。
关键词：沙地云杉；形成；分布；性质；沙地森林；过渡带
红松高生长模型的研究/邓红兵，郝占庆，王庆礼（中
国科学院沈阳应用生态研究所，沈阳 110016）
//Journal of Forestry Research, -2001, 12(3):92-194.
高生长模型是定量研究树木生长过程的有效途径，本文
应用双曲线方程、Logistic方程、三个参数的 Richards方
程及四个参数的Richards方程对红松的高生长进行了研究。
结果表明四个参数的 Richards方程在模拟红松高生长时最
为合适，且当其参数取不同的数值时，该方程可转化为其它
理论方程。在使用四个参数的Richards方程时，可根据立地
条件事先给定树高最大值，这样可使模型更符合实际情况。
此外，文中还对一种以固定年龄时实际树高为参数值的模型
进行了讨论，该模型对于计算给定树木的高生长将更为有效。
关键词：红松；高生长；模型
中国自然保护区经费与管理问题探讨/李景文，崔国发，李俊
清(北京林业大学自然保护区研究中心，北京 100083，中
国)//Journal of Forestry Research, -2001, 12(3):95-200.
在2000年，作者对50个自然保护区进行了访问和调查，
获取大量相关数据。在分析中国自然保护区发展与管理的基
础上，提出了保护区收支与管理中存在的缺乏统一的管理、
特殊的指导和有效的监督不健全、资金收入与投资困乏、各
省在资金投入的不平衡等主要问题。这些问题导致了保护区
基本建设与运行资金的缺乏，以及保护区资源与环境的破坏，
进而导致许多保护区保护转向以赢利为目的的商业运营。结
合上述问题提出了初步的解决方案。
关键词：保护区；经费；管理；中国
华北落叶松初级种子园物候观测的研究/张新波（山西省林业
科学研究院，中国林业科学研究院华北林业研究所，太原
030012，中国），任建茹（山西省林业技术推广站），张旦儿
（山西省林木种苗站）//Journal of Forestry Research,
-2001, 12(3):201-204.
通过对华北落叶松初级种子园 5年的物候观测，编制了
年物候周期表。从种子园群体和无性系个体两方面，揭示出
华北落叶松年物候周期规律。并分析阐明了物候期与气温和
活动积温之间的相关性。还对无性系个体间物候的异同性及
雌雄球花的相遇问题作了剖析。本项研究对种子园制订良种
生产计划和经营管理具有重要价值。
关键词：华北落叶松；初级种子园；物候观测
半散放条件下梅花鹿春季个体行为生态学研究/刘振生（东北
林业大学野生动物资源学院，哈尔滨 150040），周晓禹（东
北虎林园），滕丽微（中科院动物研究所），王晓明（东北林
业 大 学 ） //Journal of Forestry Research, -2001,
12(3):205-208.
采用扫描取样法、随机取样法和所有事件取样法于1998
年春季研究了梅花鹿(Cervus nippon)的行为。结果表明：春
季梅花鹿的各种行为具有较强的规律性，其行为由取食、反
刍、卧息、移动、站立、警戒、争斗、饮水、护茸和其它行
为构成。其中取食构成春季梅花鹿的主要行为。
关键词：半散放；梅花鹿；个体行为