全 文 ：Population Distribution and Dynamics of Quercus liaotungensis , Fraxinus
rhynchophylla and Acer mono in Dongling Mountain , Beijing
WU Xiao_Pu , ZHENG Yu , MA Ke_Ping＊
(Institute of Botany , The Chinese Academy of Sciences , Beijing 100093 , China)
Abstract:　The spatial distribution and population dynamics of Quercus liaotungensis Koidz., Fraxinus rhyn-
chophylla Hance and Acer mono Maxim.were studied in three warm temperate deciduous forests in Dongling
Mountain , Beijing.Clumped distributions appeared in most seedlings(≤0.4m), short saplings(0.4-1 m)
and tall saplings(1-2m).Random distributions appeared in adults of Q.liaotungensis in Plot 1 , F .rhyn-
chophylla in Plot 3.The LSD method was used to discern the differences of different aged population.The
class distribution of Q.liaotungensis was nearly a reverse_J shape for Plot 1 and Plot 3 , suggesting that the
population regenerates continuously.The age_structures of F .rhynchophylla and A.mono in Plot 1 and the
test results indicated that they would increase continuously.In Plot 2 , the class distribution and the test results
showed that all populations would not regenerate continuously.
Key words:　spatial distribution , population structure , Quercus liaotungensis
　　The spatial pattern of phytopopulation is the distribu-
tion way of a phytopopulation at the horizontal direction.
It is determined by the population characters , interspecies
relationship and the environment , which reflects the influ-
ence of some environmental factors to the individual s ac-
tion , survival and development[ 1] .The investigation to
the dynamic and population quantitative variations of dom-
inant species in forest community is of significance for us
to learn the structure and development of the ecosystem ,
community succession and the phytopopulation s ecologi-
cal characteristics.
Broad_leaved forest is the most common zonal vegeta-
tion in the area of Dongling Mountain.Quercus liaotun-
gensis , one of the most typical species of the vegetation ,
is important to the structure and composition of the whole
community[ 2] .Because it is distributed in high altitude
zone , where the disturbance of human being is relatively
insignificant , nevertheless there still have the well_devel-
oped secondary forests distributed in the middle mountain
area in the warm temperate zone.Q.liaotungensis ,
Fraxinus rhynchophylla and Acer mono are the main dom-
inant arbor species of Q. liaotungensis forests in
Dongling Mountain , Beijing.In this study , we chose
three Q.liaotungensis forests as samples , using contigu-
ous grid quadrats method and 4 coefficients to research
and compare the population distribution and dynamics of
the three species.The main issues to deal with in this pa-
per are:
1)The differences of population distributions at all
ages;
2)The dynamic structures of all populations;
3)The quantitative variations of different populations
at all ages at the same plot and their influence to the
whole community.
4) The quantitative variations of the individual
groups at all ages in the same population and their influ-
ence to the population dynamics.
1　The Study Site
The experiment was carried out at Beijing Forest
Ecosystem Research Station (BFERS)(40°00′N , 115°
26′E).This area belongs to Dongling Mountain Region ,
with a warm temperate continent monsoon climate.The
mean annual temperature averages 4.8-10.1 ℃ in Jan-
uary , and 18.3 ℃in July.The annual accumulated tem-
perature ranges from 2 300 to 3 600 ℃, and the annual
sunshine time is about 2 600 h.Precipitation amounts to
611.9 mm/a and 78%of annual rainfall occurs in June ,
July and August.Nonfrost period is shorter than 160 d.
The soil is dominated by mountain brown earth.
Three communities named Plot 1 , Plot 2 and Plot 3
were chosen , characteristics of which are listed in Table
1.The arbor dominants included Q.liaotungensis ,
Fraxinus rhynchophylla and Acer mono in Plot 1 , with a
canopy height of 8 to 10 m.The shrub layer was com-
posed of Abelia biflora , Spiraea pubescens and Lespedeza
bicolor etc.On the floor herbaceous species were abun-
dant , and mainly composed of Adenophora divaricata ,
Aster ageratoides , Calamagrostis arundinacea , Saussurea
nivea , Carex humilis var.nana and Dendranthema
chanetii etc., with a coverage of 90%.The arbor domi-
nants in Plot 2 included F .rhynchophylla , A.mono ,
Betula dahurica and Populus davidiana in addition to Q.
liaotungensis.The shrub layer was composed of Spiraea
pubescens , Lespedeza bicolor and Deutzia grandiflora
etc., with a coverage of about 70%.The majority of
herbaceous species were Calamagrostis arundinacea ,
Carex humilis var.nana , D .chanetii , S.nivea and
Spodiopogon sibiricus etc.The height of herbaceous layer
was 10 to 35 cm , and the coverage was about 60%.The
Received:2001-02-19　Accepted:2001-07-24
Supported by the National Natural Science Foundation of China (39893360).
＊Author for correspondence.
植　物　学　报　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　　

Acta Botanica Sinica　2002 , 44(2):212-223
Table 1　 Characteristics of communities
Plot no. Area(hm2) Altitude (m) Aspect Slope Slope site Canopy coverage(%) Soil depth(cm)
1 0.2 1 250 293 30 Middle 87 50
2 0.2 1 280 30 36 Upper 90 55
3 0.2 1 107 165 28 Middle 60 27
arbor species in Plot 3 included Q.liaotungensis , F .
rhynchophylla , Prunus davidiana and P.armeniaca
var.ansu , with a canopy height of 3 to 4 m.The shrub
layer was composed of Vitex negundo var.heterophylla ,
Spiraea trilobata , Ulmus macrocarpa and Syringa
pekinensis , with a height of 1 to 2 m and coverage of
65%.The herbaceous layer was mainly composed of
Artemisia gmelinii , Poa spp., Indigofera bungeana ,
Elsholtzia stauntoni and Scorzonera divaricata etc., with
a coverage of 35%.
2　Research Methods
2.1　Population characteristics and samplings in the
population distribution study
Investigation to the three communities beginning in
August , 1999 included the general condition of the com-
munities and setting the coordinate axis by prescribing two
adjacent edges of every plot and taking down the locations
of all individuals.Diameter at breast height (DBH),
height , and canopy area of those individuals taller than 2
m were recorded , while only the heigts were recorded for
the rest.
Samples were taken with the contiguous grid quadrats
method , and each grid was 5 m×5 m.
2.2　Measurement of spatial distribution
Population distributions were measured by counting
the number of individuals at all ages according to popula-
tions in each grid and calculating 4 coefficients as fol-
lows:C , Iδand IA , CA , m＊/m[ 6] .
2.3　Theoretical curve fitting to discrete distribution
We made age structure graphs of different popula-
tions , fit theoretical curves and tested them to see if they
were fit for the negative binomial distribution , Neyman A
distribution or Possion distribution[ 6] .
2.4 　Analysis and comparison of population struc-
tures and dynamics
2.4.1 　Analysis of population age structures　Age
classes were substituted by size classes to analyze the pop-
ulation structures and dynamics in the 3 communities.
According to two ways as follows , we made age structure
graphs:Divide the individuals shorter than 2 m into 3
groups:<0.4 m , 0.4-1 m and 1-2 m.Divide the
individuals taller than 2 m into several groups according to
DBH , adding a group every 2 cm.
2.4.2　Comparison of populations in the same com-
munity　After each population was divided into 4 classes
as seedling , short sapling , tall sapling and adult , we took
down the numbers of every class by grid.F_test and LSD
method were used respectively to examine if there existed
quantitative variations between two different populations at
the same age and the significances.The result suggested
the variations of the three populations at different horizon-
tal levels.
2.4.3 　Comparison of individuals in different age
class of the same population　F_test and LSD method
were also used to examine if there existed differences be-
tween two age classes of the same population and the sig-
nificances.The results suggested the variations of all age
classes of every populations , which were important for us
to learn the population dynamics.
3　Results
3.1　Population distribution
Figures 1-3 indicated the distributions of individu-
als belonging to different age group in three plots , using
Q.liaotungensis as a example.The coordinates were a-
gree with the former as we prescribed.These graphs were
used as evidences in theoretical curve fitting to the popu-
lation distributions.
Fig.1.　Spatial distributions of different individuals of Quercus liaotungensis in Polt 1.
WU Xiao_Pu et al:Population Distribution and Dynamics of Quercus liaotungensis , Fraxinus rhynchophylla and Acer mono 213　
Fig.2.　Spatial distributions of different individuals of Quercus liaotungensis in Plot 2.
Fig.3.　Spatial distributions of different individuals of Quercus liaotungensis in Plot 3.
　　The calculated results of 4 coefficients , C , Iδand
IA , CA , m＊/m , to the population distributions were
unanimous.F_test and χ2_test were used in calculated C
and Iδ.The results were illustrated in Table 2.Due to
their insignificant quantity , F.rhynchophylla and A.
mono in Plot 2 were not taken into account.
3.2　Curve fitting to the population distributions
The results of the theoretical curve fitting to all pop-
ulations are illustrated in Table 3.The fitness was tested
by χ2_test and the distribution was estimated from the
leastχ2 value.Thus , most distributions were fit for the
typical distributions except the seedling group of A.
mono in Plot 1(Table 1).
3.3　Age structures of the populations
Figure 4 illustrates the differences in age structures
of all populations.
3.3.1　Age structures and dynamics of Q.liaotun-
gensis populations　The age structure of Q.liaotungen-
sis population in Plot 1 , similar with that in Plot 3 , was
nearly reverse_J shape.Although the class distribution
was not quite regular , it was still a stable structure and
the population can maintain regeneration.The seedlings
and saplings were insufficient , which would be one of the
limited conditions of the regeneration of Q.liaotungensis
in Plot 2.The class distribution was more or less like a
declined structure.
3.3.2　Age structures and dynamics of F.rhyn-
chophylla and A.mono populations　The class struc-
ture of F .rhynchophylla in Plot 1 was similar with that
of A.mono.The seedlings and saplings were much more
than the adults , but few of them could reach maturity.
The class distribution was more like an increasing struc-
ture.Although population density of the two populations
in Plot 2 was minute , their age structures were similar
with Q.liaotungensis and like declining structures.The
class distribution of F .rhynchophylla in Plot 3 was a
typical increasing structure , for the number of individuals
reduced as the age was increased.
3.4　Comparison of quantitative population varia-
tions
3.4.1　Comparison of populations in the same com-
munity　The variations and the significance of variance
of populations in the same community , checked by F_test
and LSD methods , were illustrated in Table 4 and Table
5.The dominant conditions at every horizontal level could
be known positively by comparison of the quantity in dif-
ferent age class between populations in the same commu-
nity , which can inform us of the direction of community
succession.
3.4.2 　Comparison of individuals in different age
classes of the same population　The quantitative differ-
ences and their significance in different age groups of the
same population illustrated in Table 6 was significant to
obtain information of population development.
3.4.3 　Comparison of individuals in different age
classes of the same population　The quantitative differ-
ences and their significance in different age groups of the
same population illustrated in Table 6 and Table 7 were of
importance to obtain information of population develop-
ment.
4　Discussion
4.1　Population distribution
4.1.1　Sampling technique　The significance of popu-
lation distribution may be biased by deflection of the
species distribution to random distribution in vegeta-
tion[ 4 ,8] .The degree of deflection to random distribution
was used to being measured by taking samples with ran-
dom sized grids
[ 4] .The shortage of the method lies in that
the result may depend on the size of sample grids ,
214　 植物学报　Acta Botanica Sinica　Vol.44　No.2　2002
Table 2　Valus of indexes of dispersion for seedling , short sapling , tall sapling and adult of Quercus liaotungensis , Fraxinus rhynchophylla
and Acer mono
Plot Indexes of dispersion Seedling Short sapling Tall sapling Adult
Q.liaotungensis in Plot 1
C 2.06 0.97 1.75 0.78
Iδ 1.93 0.91 4.71 0.88
CA 0.94 -0.09 3.53 -0.12
m＊/m 1.94 0.91 4.53 0.88
IA 1.96 0.95 5.0 0.89
Dispersion Clump Random Clump Regular
χ2 165 77.68 140.04 62.73
F.rhynchophylla in Plot 1
C 4.21 1.91 1.89 2.16
Iδ 2.20 1.47 1.60 1.76
CA 1.21 0.47 0.61 0.77
m＊/m 2.21 1.47 1.61 1.77
IA 2.21 1.47 1.62 1.78
Dispersion Clump Clump Clump Clump
χ2 336.9 152.98 150.9 172.73
A.mono in Plot 1
C 3.52 2.64 5.72 1.3
Iδ 2.04 3.39 7.21 2.3
CA 1.05 2.38 6.19 1.2
m＊/m 2.04 3.38 7.19 2.2
IA 2.05 3.46 7.33 2.4
Dispersion Clump Clump Clump Clump
χ2 281.82 210.91 457.51 106
Q.liaotungensis in Plot 2
C 1.21 1.24 1.24 1.1
Iδ 1.52 1.69 1.66 1.0
CA 0.50 0.67 0.64 0
m＊/m 1.51 1.68 1.64 1.0
IA 1.56 1.76 1.71 1.0
Dispersion Clump Clump Clump Random
χ2 96.69 98.95 99.24 87
Q.liaotungensis in Plot 3
C 3.11 1.02 1.38 1.7
Iδ 2.98 1.07 1.69 1.6
CA 1.98 0.06 0.68 0.6
m＊/m 2.98 1.06 1.68 1.6
IA 3.02 1.11 1.73 1.6
Dispersion Clump Random Clump Clump
χ2 248 81.62 110.1 133
F.rhynchophylla in Plot 3
C 5.74 1.83 1.40 0.94
Iδ 2.78 1.77 1.48 0.93
CA 1.79 0.77 0.48 -0.07
m＊/m 2.79 1.77 1.48 0.93
IA 2.79 1.79 1.50 0.95
Dispersion Clump Clump Clump Random
χ2 459 146.53 112.31 75.55
χ20.05=101;F0.05=1.27.
especially to clumped distribution.How many grids
should be taken in different sizes may conform to statisti-
cal calculation prior to sample taking.To a size_fixed
sample , it is impossible to take prerequisite samples as
size without repetition
[ 6] .Improved by many ecologists ,
contiguous grid quadrats method has been an effective way
to take samples
[ 4 ,8-11] .To a sample with suitable grid
size , it not only overcomes the effect of grid size , but also
keeps advantages of taking samples by grids.In the ex-
periment , we chose the rectangular arrangement according
to the local terrain.
4.1.2　Coefficients of population distribution　In our
research , about 4 coefficients were calculated to express
the distributions of all populations:C , Iδand IA , CA ,
m＊ and m＊/m.C is the ratio of variance to mean.F_test
andχ2_test will be transacted to insure objective results on
condition that the sample quantity is enough.Although C
is a convenient index , it changes with the means even
though the distribution degree is the same when the means
change.So we used the Morisita indexes(Iδand IA)and
WU Xiao_Pu et al:Population Distribution and Dynamics of Quercus liaotungensis , Fraxinus rhynchophylla and Acer mono 215　
Table 3　Testing results of 3 population distribution patterns from the data of contiguous grid quadrats method
Plot Population Age Distribution type χ2 χ20.05
Plot 1
Quercus liaotungensis
Seedling Negative binomial distribution 1.92 7.81
Short sapling Possion distribution 1.98 9.49
Tall sapling Negative binomial distribution 2.76 5.99
Adult Possion distribution 7.43 9.49
Fraxinus rhynchophylla
Seedling Negative binomial distribution 14.04 18.31
Short sapling Neyman A distribution 8.11 11.07
Tall sapling Neyman A distribution 5.92 11.07
Adult Negative binomial distribution 11.35 12.59
Acer mono
Seedling - -
Short sapling Negative binomial distribution 4.35 9.49
Tall sapling Negative binomial distribution 8.62 9.49
Adult Neyman A distribution 5.89 5.98
Plot 2 Q.liaotungensis
Seedling Neyman A distribution 1.17 11.07
Short sapling Neyman A distribution 0.48 11.07
Tall sapling Neyman A distribution 1.81 11.07
Adult Neyman A distribution 2.61 11.07
Plot 3
F.rhynchophylla
Seedling Negative binomial distribution 16.2 22.36
Short sapling Neyman A distribution 6.78 11.07
Tall sapling Neyman A distribution 1.08 11.07
Adult Possion distribution 0.92 9.49
Q.liaotungensis
Seedling Negative binomial distribution 10.24 12.59
Short sapling Negative binomial distribution 1.41 5.99
Tall sapling Neyman A distribution 8.2 11.07
Adult Neyman A distribution 6.67 11.07
CA to overcome the defect.CA is the reciprocal of K , a
parameter of negative binomial distribution.CA is more
efficient to clumped distributions;nevertheless , it is
scarce of rigorous mathematical test.m , representing the
average of individuals , not grids , could reflect the influ-
ence of intraspecific competition and biotic factors.m＊ ,
the mean , impacted by the empty grids , can hardly re-
flect the influence of biotic factors when the individuals in
mosaics are crowded and competition of interspecies is
drastic while the value of m is minute.m＊ can supply
more accurate information in distribution analysis than
other coefficients when there are lots of empty grids
[ 13] .
Generally , every coefficient has its advantages and
disadvantages.Using various indexes in distribution re-
search may provide us an objective result to the fact.
4.1.3　Distribution patterns and types　Both theoret-
ical curve fitting and coefficients are aimed at expression
to the population distribution.The former can impress us
with vivid distribution condition , while the latter can em-
body the extent of the actual distribution deflected to ran-
dom distribution.So , the combination of both can give us
a comprehensive expression to actual distribution.
The distribution of a population is determined by its
biotic characteristics , the environment and their interac-
tion[ 8] .The results of the study indicated that clumped
distribution appeared frequently in groups of seedling and
sapling of all the tree populations , and the distribution
type was of the Neyman A distribution or negative binomi-
al distribution , while Q.liaotungensis , as the dominant
species , appeared to be regular distribution or random
distribution in adult group , and its distribution type was
Possion distribution.Figure 4 and significance test of
population quantity indicated that the adult groups of Q.
liaotungensis , were of great importance in Plot 2 and Plot
3.It is concluded that it was the results of spatial niche
redispersion and environmental heterogeneity , which
caused population inter_differentiation , that different aged
groups had different distributions[ 14] .The results of coef-
ficient calculation indicated that the distribution of adult
Q.liaotungensis in Plot 1 was a transition from regular
distribution to random distribution.As we know , regular
distribution appeared infrequently in natural condition ,
which suggested that it was probably the results of distur-
bance from human being and natural competition.The
population succession changes with the environmental
changing , which was one of the potential reasons that the
distribution of Q.liaotungensis adult group in Plot 2 was
different from Plot 1.The terrain in Plot 3 was so complex
that the microenvironment of individuals was obviously
different.To Q.liaotungensis , the environment of Plot 3
was much more rigorous than that of Plot 1 and Plot 2.
The investigation showed , the dominance of Q.liaotun-
gensis adults in canopy level of Plot 3 were not so signifi-
cant as Plot 1 and Plot 2.Therefore , clumped distribution
of a population seemed more likely the result of common
actions of environment and other populations.
Figure 4 also indicated that the population densities
216　 植物学报　Acta Botanica Sinica　Vol.44　No.2　2002
Fig.4.　Class distribution of all populations in Plots.
of F.rhynchophylla and A.mono in Plot 1 were less
than Q.liaotungensis , and most of the individual heights
were also shorter than those of Q.liaotungensis.Al-
though F . rhynchophylla is a common species in
Dongling Mountain region , the individuals often grow in
the shrub layer of various natural forests , not frequently
seen in canopy layers[ 5] .After a long period of competi-
tion with Q.liaotungensis , they were eliminated by Q.
liaotungensis eventually , and formed clumped distribu-
tions in the spare areas of adult Q.liaotungensis.In Plot
3 , its population density of adults was even higher than
Q.liaotungensis and became the dominant species.De-
velopment and dispersion took place towards random dis-
tribution , with which the population could make the best
use of resources.
Although both Neyman A distribution and negative
binomial distribution belong to clumped distribution , their
dispersal modes are different.Neyman A distribution con-
sists of several mosaics dispersing randomly in space , and
the individuals in each mosaic also disperse randomly.
Negative binomial distribution represents a striated disper-
sal way.Table 3 indicated that the seedling groups of
some populations belonged to negative binomial distribu_
tions , as the seedlings were impressionable to abiotic
WU Xiao_Pu et al:Population Distribution and Dynamics of Quercus liaotungensis , Fraxinus rhynchophylla and Acer mono 217　
Table 4　Comparison of quantity of different populations in the same plot
Plot Age　　 Difference origin f s 2 F F0.05 F0.01
Comparison in Plot 1
Seedlings
Interspecies 2 53.52 7.28＊＊ 3.03 4.7
Intraspecies 237 7.35
Total differece 239
Shortsaplings
Interspecies 2 53.52 29.05＊＊ 3.03 4.7
Intra species 237 7.35
Total differece 239
Tall saplings
Interspecies 2 31.40 12.58＊＊ 3.03 4.7
Intraspecies 237 2.50
Total differece 239
Adults
Interspecies 2 52.25 31.32＊＊ 3.03 4.7
Intraspecies 237 1.67
Total differece 239
Comparison in Plot 2
Seedlings
Interspecies 2 2.71 11.28＊＊ 3.03 4.7
Intraspecies 237 0.24
Total differece 239
Short saplings
Interspecies 2 2.15 10.54＊＊ 3.03 4.7
Intraspecies 237 0.20
Total differece 239
Tall saplings
Interspecies 2 2.55 10.77＊＊ 3.03 4.7
Intraspecies 237 0.24
Total differece 239
Adults
Interspecies 2 116.08 84.19＊＊ 3.03 4.7
Intraspecies 237 1.38
Total differece 239
Comparison in Plot 3
Seedlings
Interspecies 2 110.81 10.89＊＊ 3.90 6.80
Intraspecies 158 9.26
Total differece 159
Short saplings
Interspecies 2 23.26 20.33＊＊ 3.90 6.80
Intraspecies 158 1.14
Total differece 159
Tall saplings
Interspecies 2 3.30 3.42 3.90 6.80
Intraspecies 158 0.97
Total differece 159
Adults
Interspecies 2 4.56 3.35 3.90 6.80
Intraspecies 158 1.36
Total differece 159
The signif icance was tested by value of F_statistics.
environment.It seemed that it was the restriction of light ,
terrain or something else that led to the striated distribu-
tions of seedlings and saplings.Neyman A distribution
appeared in adult groups commonly.In addition to some
abiotic reasons , the interaction of individuals seemed to
be more efficient.A typical instance was that all groups
in Plot 2 belonged to the Neyman A distribution , which
had a close relation with the interaction of individuals , for
Q.liaotungensis had the dominant position in Plot 2 by
statistic test.
According to the result of coefficients calculated ,
most seedlings and saplings of all populations were of
clumped distribution , while the short saplings of Q.
liaotungensis in Plot 1 and Plot 3 were of random distri-
butions.To seedlings , living together was beneficial to
the individuals[ 15] .The distributions of seedlings and
saplings were associated with the parent trees , seed rain ,
microenvironment , population density and animal re-
moval[ 16] .To seedlings and saplings of Q.liaotungensis ,
the influence factors included moisture , light , nutrition
and animal removal , in addition to shield of the adjacent
adults and self_chemoreception[ 7] .From the record of
seed rain and distribution diagrams of Q.liaotungensis ,
the regeneration mostly depended on sprout in Plot 2 ,
while on seeds in Plot 3.The production of seed rain of
Q.liaotungensis in Plot 2 was obviously lower than that
in Plot 3 , despite of that the density of adults was as high
as 1 195/hm2.The seeds dropped on the ground were al-
most completely consumed and removed by animals , and
few dispersed near the parent tree.Therefore the actual
distribution probably resulted in sprout and self_chemore-
ception.In Plot 1 , the adult density of Q.liaotungensis
was only 710/hm2 , and the coverage of canopy layer was
also lower than that in Plot 2.Plenty of seedlings
218　 植物学报　Acta Botanica Sinica　Vol.44　No.2　2002
Table 5　Different comparisons by LSD method
Comparison
in Plot 1
Seedlings in Plot 1
Species Mean
Signif icance
LSD0.05=0.84 X i-1.14
LSD0.01=1.10 X i-2.40
Fraxinus rhynchophylla 2.66 1.52＊＊ 0.26
Acer mono 2.40 1.26＊＊
Quercus liaotungensis 1.14
Short saplings in Plot 1
Species Mean
Signif icance
LSD0.05=0.43
X i-1.14
LSD0.01=0.57
X i-2.40
F.rhynchophylla 1.94 1.60＊＊ 1.25＊＊
A.mono 0.69 0.35
Q.liaotungensis 0.34
Saplings in Plot 1
Species Mean
Signif icance
LSD0.05=0.49 X i-0.21
LSD0.01=0.64 X i-0.76
F.rhynchophylla 1.46 1.25＊＊ 0.7＊
A.mono 0.76 0.55＊＊
Q.liaotungensis 0.21
Adults in Plot 1
Species Mean
Signif icance
LSD0.05=0.40
X i-0.26
LSD0.01=0.53
X i-1.51
Q.liaotungensis 1.77 1.51＊＊ 0.26
F.rhynchophylla 0.76 1.25＊＊
A.mono 0.26
Comparison
in Plot 2
Seedlings in Plot 2
Species Mean
Signif icance
LSD0.05=0.15
X i-0.09
LSD0.01=0.20
X i-0.10
Q.liaotungensis 0.41 0.32＊＊ 0.31
A.mono 0.10 0.01
F.rhynchophylla 0.09
Short saplings in Plot 2
Species Mean
Signif icance
LSD0.05=0.14 X i-0.03
LSD0.01=0.18 X i-0.15
Q.liaotungensis 0.35 0.32＊＊ 0.20＊＊
F.rhynchophylla 0.15 0.13
A.mono 0.03
Saplings in Plot 2
Species Mean
Signif icance
LSD0.05=0.15
X i-0.03
LSD0.01=0.20
X i-0.14
Q.liaotungensis 0.38 0.35＊＊ 0.24＊＊
F.rhynchophylla 0.14 0.11
A.mono 0.03
Adults in Plot 2
Species Mean
Signif icance
LSD0.05=0.36 X i-0.39
LSD0.01=0.48 X i-0.7
Q.liaotungensis 2.61 2.23＊＊ 1.91＊＊
F.rhynchophylla 0.70 0.31
A.mono 0.39
Comparison
in Plot 3
Seedlings in Plot 3
Species Mean
Signif icance
LSD0.05=0.94 X i-1.06
LSD0.01=1.24 X i-1.06
Q.liaotungensis 2.65 1.58＊＊ 1.58＊＊
F.rhynchophylla 1.06
WU Xiao_Pu et al:Population Distribution and Dynamics of Quercus liaotungensis , Fraxinus rhynchophylla and Acer mono 219　
Table 5　(continued)
Comparison
in Plot 3
Short saplings in Plot 3
Species Mean
Signif icance
LSD0.05=0.33
X i-0.31
LSD0.01=0.44
X i-0.31
Quercus liaotungensis 1.08 0.77＊＊ 0.77＊＊
Fraxinus rhynchophylla 0.31
Saplings in Plot 3
Species Mean
Signif icance
LSD0.05=0.30 X i-0.55
LSD0.01=0.40 X i-0.55
Q.liaotungensis 0.84 0.29 0.29
F.rhynchophylla 0.55
Adults in Plot 2
Species Mean
Signif icance
LSD0.05=0.36 X i-0.83
LSD0.01=0.47 X i-0.83
Quercus liaotungensis 1.16 0.33 0.33
Fraxinus rhynchophylla 0.83
＊＊, the difference is very significant;＊, the difference is signif icant.
Table 6　Quantitative comparison of different aged populations of the same species in the same plot
Plot Species Difference origin f s 2 F F0.05 F0.01
Plot 1
Quercus liaotungensis
Interspecies 3 42.84 38.60＊＊ 2.63 3.85
Intraspecies 316 1.11
Total differece 319
Fraxinus rhynchophylla
Interspecies 3 24.65 4.71＊＊ 2.63 3.85
Intraspecies 316 5.24
Total differece 319
Acer mono
Interspecies 3 71.71 19.10＊＊ 2.63 3.85
Intraspecies 316 3.75
Total differece 319
Plot 2
Q.liaotungensis
Interspecies 3 99.80 94.06＊＊ 2.63 3.85
Intraspecies 316 1.06
Total differece 319
F.rhynchophylla
Interspecies 3 6.67 18.35＊＊ 2.63 3.85
Intraspecies 316 0.36
Total differece 319
A.mono
Interspecies 3 2.34 19.73＊＊ 2.63 3.85
Intraspecies 316 0.12
Total differece 319
Plot 3
F.rhynchophylla
Interspecies 3 61.43 12.84＊＊ 2.63 3.85
Intraspecies 316 4.79
Total differece 319
Q.liaotungensis
Interspecies 3 13.26 8.41＊＊ 2.63 3.85
Intraspecies 316 1.58
Total differece 319
Significance was shown by F_text.＊＊, the difference is very significant;＊, the diff erence is signifi cant.
germinated in the broad gaps in Plot 1 , where sunlight
was abundant.The coverage of canopy layer in Plot 3 was
the lowest.The density of adult Q.liaotungensis was on-
ly 465/hm2 , which meant that sunlight and interspecific
competition were not the limitations of seedling growth.
Most seedlings in Plot 3 dispersed under their parent
trees , and the more sparely , the farther they were from
parent trees , which implied that most of them came from
seeds.The densities of F.rhynchophylla and A.mono
in every plot were lower than 300/hm2 , which meant they
were major companying species.The clumped distribution
of seedlings and saplings were the result of integration of
self_regeneration and environmental impacts.
The assemblage of saplings was not the simple suc-
cession of seed bank or seedling cohort accumulation , but
an accumulation of different aging groups companioning
with the disruption of seedling cohort.Therefore it was
not an occasional phenomenon that random distributions of
short saplings of Q.liaotungensis appeared in Plot 1 and
Plot 3.Up to sapling stage , the impacts of adult trees on
saplings was not as strong as that on seedlings and the
adaptability of saplings to the environment increased in
the process , which resulted in such a distribution pattern.
220　 植物学报　Acta Botanica Sinica　Vol.44　No.2　2002
Table 7　Different comparisons by LSD method
Quercus liaotungensis in Plot 1
Age Mean
Signif icance
LSD0.05=0.33 LSD0.01=0.43
X i-0.21 X i-0.34 X i-1.14
Adults 1.76 1.56＊＊ 1.44＊＊ 0.64＊
Seedling 1.14 0.93＊＊ 0.80＊＊
Short saplings 0.34 0.13
Tall saplings 0.21
Fraxinus rhynchophylla in Plot 1
Age Mean
Signif icance
LSD0.05=0.71 LSD0.01=0.93
X i-1.46 X i-1.51 X i-1.94
Seedling 2.66 1.2＊＊ 1.15＊＊ 0.73＊
Short saplings 1.94 0.48 0.43
Adults 1.51 0.05
Tall saplings 1.46
Acer mono in Plot 1
Age Mean
Signif icance
LSD0.05=0.71 LSD0.01=0.93
X i-0.26 X i-0.69 X i-0.76
Seedling 2.41 2.15＊＊ 1.73＊＊ 1.65＊＊
Short saplings 0.76 0.50 0.01
Adults 0.69 0.43
Tall saplings 0.26
Q.liaotungensis in Plot 2
Age Mean
Signif icance
LSD0.05=0.32 LSD0.01=0.42
X i-0.35 X i-0.38 X i-0.41
Seedling 2.61 2.26＊＊ 2.24＊＊ 2.20＊＊
Short saplings 0.41 0.06 0.04
Adults 0.38 0.03
Tall saplings 0.35
F.rhynchophylla in Plot 2
Age Mean
Signif icance
LSD0.05=0.19 LSD0.01=0.25
X i-0.09 X i-0.14 X i-0.15
Seedling 0.70 0.61＊＊ 0.56＊＊ 0.55＊＊
Short saplings 0.15 0.06 0.01
Adults 0.14 0.05
Tall saplings 0.09
A.mono in Plot 2
Age Mean
Signif icance
LSD0.05=0.11 LSD0.01=0.14
X i-0.03 X i-0.03 X i-0.10
Adults 0.39 0.36＊＊ 0.36＊＊ 0.29＊＊
Seedling 0.10 0.08 0.08
Tall saplings 0.03 0.00
Short saplings 0.03
F.rhynchophylla in Plot 3
Age Mean
Signif icance
LSD0.05=0.68 LSD0.01=0.89
X i-0.83 X i-0.84 X i-1.08
Seedling 2.65 1.82＊＊ 1.81＊＊ 1.57＊＊
Short saplings 1.08 0.25 0.24
Adults 0.84 0.01
Tall saplings 0.83
Q.liaotungensis in Plot 3
Age Mean
Signif icance
LSD0.05=0.39 LSD0.01=0.51
X i-0.31 X i-0.55 X i-1.06
Seedling 1.16 0.85＊＊ 0.61＊＊ 0.1
Short saplings 1.06 0.75＊＊ 0.51＊＊
Adults 0.55 0.24
Tall saplings 0.31
＊＊, the difference is very significant;＊, the difference is signif icant.
WU Xiao_Pu et al:Population Distribution and Dynamics of Quercus liaotungensis , Fraxinus rhynchophylla and Acer mono 221　
4.2　Analysis on the development of populations
Comparison of the graphics of age structure of all
populations suggested that most age structures did not ap-
pear simplex ascending or descending tendencies , some
phase probably lower than the adjacent phases.In natural
condition , there were several possible causes , one of
which was the result of competition , including inter-
species and intraspecies.When a community has devel-
oped for a special period , the individuals of population
would reach some scale , which brings competitions to re-
source and space.Some strong individuals which have
grown for a period of time , get rid of the weaker ones by
shield or allelophathy , the results of which are self thin-
ning or interspecific competition.Another reason is the
special natural regeneration of species.Some species have
large seedling storage , but they should go through several
environment sieves if the seedlings grow to saplings.In
the process , plenty of seedlings would be eliminated by
one environmental sieve.The remains would go through
another environmental sieve after a stable period.Recur-
rently , they eventually grow up[ 13 ,17] .The fluctuations
happened during the population regeneration in our study
probably were the result of the interaction of the two fac-
tors described above.
Q.liaotungensis diperses widely in warm temperate
zone as a typical regional vegetation in its mountain re-
gions.Because it can bear both cold_moisture climate and
warm_dry climate and has wide ecological amplitude , Q.
liaotungensis has an excellent perspective.In the study ,
Q.liaotungensis was the main dominant plant in the
three plots and had superior dominance at the canopy lev-
els.The key factor that impacted on its distribution was
the competition to resource of the individuals , for the
adults had strong capability to environmental disturbance.
Q.liaotungensis population was declining in terms of its
population structure in Plot 2 , but the DBH of the trees at
the eldest age class in this plot was less than that in Plot
1.So , the declining phenomenon should be temporary ,
because sprouting was the major way for natural regenera-
tion at present.It was believed that the capability for seed
production would increase.
In Plot 1 and Plot 3 , both F .rhynchophylla and
A.mono appeared to be of increasing population;never-
theless , they were companion species , not superior to Q.
liaotungensis in competition to light and space at the
canopy level.The investigation indicated that , at the el-
der age class , their densities were much lower than Q.
liaotungensis.From this , it was concluded that the age
structure would probably maintain for a certain period of
time and not change into stable structure.In Plot 2 , the
densities of both were low , the reason of which probably
was their weak competition capability.Their declining
structure probably was affected by Q.liaotungensis.
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222　 植物学报　Acta Botanica Sinica　Vol.44　No.2　2002
北京东灵山地区辽东栎 、大叶白蜡和五角枫
种群分布格局与动态
吴晓莆　郑　豫　马克平
(中国科学院植物研究所 , 北京 100093)
摘要:　对北京东灵山地区 3 块样地中的辽东栎(Quercus liaotungensis Koidz.)、大叶白蜡(Fraxinus rhynchophylla
Hance)和五角枫(Acer mono Maxim.)种群的分布格局和种群动态进行了考察。 调查 3 块样地中每个树种的全部个
体 ,按高度将它们分为 4 个不同龄级 ,用相邻格子样方法进行取样。在研究分布格局时 , 计算了扩散系数 C 、扩散
型指数 Iδ与 IA 、CA指数和聚块指数 m＊/m 等各项指标 , 测定它们的分布格局 , 并通过计算拟合它们的分布类型。
结果表明 ,不同地点各种群的幼苗(<0.4 m)、小幼树(0.4 ～ 1 m)和大幼树(1～ 2 m)绝大多数呈聚集分布 , 少量呈随
机分布;成林辽东栎成熟个体(≥2 m)是一种介于均匀分布和随机分布之间的状态 , 幼林辽东栎成林和灌丛大叶白
蜡属于随机分布 ,成林大叶白蜡和五角枫 、灌丛辽东栎呈聚集分布。在研究种群动态时 ,分别作出每个种群的龄级
结构图 ,再对同一样地不同种群以及同一种群不同龄级个体的数量差异进行最小显著差法(LSD)检验和比较 ,结果
表明检验结果和龄级结构图显示出来的趋势是一致的 , 即成林辽东栎种群具有较稳定的结构 ,成林大叶白蜡和五
角枫表现为增长型结构 ,幼林辽东栎 、大叶白蜡和五角枫都表现为衰退型结构 , 灌丛辽东栎具有稳定型结构 ,灌丛
大叶白蜡具有增长型结构。研究表明 ,作为优势种的辽东栎种群个体之间的竞争很激烈 , 在乔木层中占有明显优
势;作为伴生种的大叶白蜡的幼苗对环境的适应能力很强 ,在较低位层中占有明显优势。
关键词:　分布格局;种群结构;辽东栎
中图分类号:Q948.12　　　文献标识码:A　　　文章编号:0577-7496(2002)02-0212-12
收稿日期:2001-02-19　接收日期:2001-07-24
基金项目:国家自然科学基金(39893360);中国科学院“十五”预研项目;中国科学院重点项目(KZ952_SI_127)。
(责任编辑:崔金钟)
WU Xiao_Pu et al:Population Distribution and Dynamics of Quercus liaotungensis , Fraxinus rhynchophylla and Acer mono 223