全 文 :第 23 卷 第 3 期 植 物 研 究 2003年 7 月
Vol.23 No.3 BULLETIN OF BOTANICAL RESEARCH July , 2003
黑龙江省东部山地湿地主要植被类型及其演替规律的研究
倪志英1 毛子军1 孙 龙1 戈忠达2 周明娜2
(1.东北林业大学 ,哈尔滨 150040)
(2.黑龙江省阿城市国有林场管理局 ,阿城 150300)
摘 要 首次系统论述了黑龙江省东部山地湿地植被类型 ,并在此基础上 ,对黑龙江省东部山地
湿地植被演替规律进行了全面探讨 。研究表明:黑龙江省东部山地湿地植被演替与发展过程 ,经
过三个阶段:即富营养沼泽※中营养沼泽※贫营养沼泽 。无论是森林沼泽 ,还是草甸 、沼生及水生
植被都是如此 。
关键词 湿地;植被;植被类型;演替
A STUDYON THE MAIN VEGETATION TYPES AND SUCCESSION
DISCIPLINEOF THE WETLAND IN THE EASTERN MOUNTANIOUS
REGION OF HEILONGJIANG PROVINCE
NI Zhi-Ying1 MAO Zi-Jun1 SUN Long1 GE Zhong-Da2 ZHOU M ing-Na2
(1.Northeast Forestry University ,Harbin 150040)
(2.A cheng Managment Adminstration of State Fo rest Farms ,Heilongjiang P rovince , A cheng 150300)
Abstract In this paper , the authors elao rated the w etland vegetation types in the first time , and
based on this , the authors completely explore the succession discipline of the w etland vegetations in
the eastern mountanious region of Heilongjiang Province.The investigation indicates that the process
of the w etland vegetation succession and development in the eastern mountanious region of Hei-
longjiang Province are three phrases w hich is eut rophic sw amp※meso trophic sw amp ※dyst rophic
sw amp.No matter w hich is helohy linm , sw amp , helo vegetation or aquatic vegetation.
Key words wetland;vegetation;vegetation types;succession
Wetland is the important ecosystem type on the
earth , constantly bounding the continent and the
w ater body , taking on both ecological function of ter-
restrial and aquatic ecosystem , having these charac-
ters:high biodiversity and productivity , providing
mankind wi th manifold food stuff ,medication , ener-
gy sources and industry raw material;wetland is also
the natural reservoir , which performs an important
function to regulate local climate , to keep river run
of f balance , to supply g roundw ater and to store
floodwater;otherw ise , wetland is the impo rtant
reservoir sink of carbon and ni trogen element and
natural factory of deg rading pollutants , which takes
important action on controlling the emission of
greenhouse gas and purifying air , so it is named “ the
kidney of the nature”.Especially , the wetland in the
forest regions of Heilongjiang forest industry is fa-
mous of habitat of rare water fow ls and the calving
ground of mig rant and it is eco tourism scenic spot.It
is obvious that the w etland in the forest regions of
第一作者简介:倪志英(1963—),女 ,副研究员 ,主要从事植物学研究 。收稿日期:2002-09-12
Heilong jiang Province is a special ecosystem which
possesses mult i-economy functions and ecological
service function , taking on important application val-
ue and scientif ic value.According to the national
forest ry bureau s related policy , we have made gen-
eral investig at ion on the wet land in the forest regions
of Heilongjiang Province f rom 2000 ~ 2001.This in-
vestigation is only a part of the investigations on
w etland animals and plant resources in the forest re-
gions of Heilong jiang Province.
1 The mostly wetland vegetation types of in the
eastern mountanious region of Heilongjiang
Province
The fo rest regions of Heilongjiang Province is
important fo rest regions in our country , with high
forest cover , large w ood reserves , more rare t ree
species and large w etlands.The fo rest regions of
Heilong jiang forest indust ry located betw een 43°30′
30″N and 49°08′45″N , 127°15′E and 134°05′E .
The total management area is 1006×104 hm2 which
is about one-forth of the area of Heilong jiang
Province , and wet land s area is about 310 593 hm2.
According to the classification principle of Chi-
nese wet land and combining the consti tutional char-
acteristic of vegetations and ecological difference of
the nature condi tions such as topography , soil , wa-
ter condition etc w hich are related with g rowing
closely , the w etland vegetations of this region can be
classified 2 vegetation type groups , 7 vegetation
types and 7 subtypes , 15 formation g roups , 31 for-
mations , 52 association g roups or associations;2
vegetation type groups are sw amp type g roup and
shallow water plants type g roup;7 vegetation types
are fo rest sw amp type , shrub sw amp type , brush-
wood sw amp type , moss sw amp type , phytoneuston
type , leaf-float plant type , submerged macrophyte
type;7 subtypes are sw amp conifer subtype , sw amp
broad-leaf forest subtype , swamp deciduous broad
leaf shrub fo rest subtype , ever-green broadleaved
shrub sw amp subtype , sedge sw amp subtype , grass
sw amp subtype , weed sw amp subtype;15 fo rma-
tion g roups are Larix formation group , Alnul for-
mation group , Betula formation g roup , Salix for-
mation group , Spiraea fo rmation g roup , Ledum
palustre fo rmation g roup , Carex swamp formation
g roup , Scirpus formation g roup , Carex lanceolata
formation g roup , Phragm ites communis formation
group , Glyceria fo rmation group , Typha orientalis
formation group , Equisetum hyemale formation
group , Sphagnum fo rmation g roup , Potamogeton
formation g roup;31 formations are Larix gmelin;
formation , Alnus sibirica formation , Betula oval i-
fol ia fo rmation , Betula frut icosa formation , Salix
rosmarini folia fo rmation , Spiraea salici folia for-
mation , Ledum palustre formation , Carex
schmidt ii format ion , C.meyeriana fo rmation , C.
appendiculata formation , C.lasiocarpa forma-
tion , C.pseudocuraica fo rmation , C.humida
formation , C.globularis formation , C.limosa
formation , Seripus tabernaemotani formation ,
Scirpus tabernaemontani formation , Eripherum
formation , Phragm ites australis formation , Glyce-
ria spiculosa formation , Typha angust ifolia forma-
tion , Equisteum f uv iat le formation , Sphagnum
magellanicum format ion , S .robustum formation ,
Salv inia natans formation , Nymphoides peltatum
formation , Nymphaea tatragona formation ,
Nelumbo nuci fera formation , Ranunculus gmelinii
formation , Potamogeton natans formation , P .
malaianus fo rmation , Utricularia vulgalis forma-
tion.
2 The succession of wetland vegetations in the
eastern mountanious region of Heilongjiang
Province
The succession of vegetation communities is not
accidental , but is especially a kind of anf ractuosity
relation among climate , hydrology , topog raphy and
soil etc natural facto rs w hich makes one community
replaced by another community.After the swamp
plants came into being , peat accumulates constantly
and sw amp s nutrition condition dilutes day by day
wi th the vegetation dying year af ter year , At last it
w as replaced by the dyst rophic sw amp.Some
swamps w ere inf luenced by physical geog raphy con-
dition , although one community is replaced by an-
o ther one , its t rophic ecolog y doesn t occur“virtuali-
ty” change , some sw amp s g row th w ill rest on the
eutrophic swamp stage fo r a long time.So the pro-
cess of sw amp vegetation s succession is more com-
plex .
The process of formation and development of
different sw amp will take three stages , that is eu-
3573 期 倪志英等:黑龙江省东部山地湿地主要植被类型及其演替规律的研究
trophic sw amp ※mesotrophic sw amp ※dyst rophic
sw amp.Tw o kinds of development courses of a
sw amp in our country :one is complete development
course , that is , f rom eutrophic sw amp to dyst rophic
sw amp coming f rom mesot rophic sw amp;the other
one is uncomplete development course , that is , rest-
ing on a certain development stage fo r a long time or
lacking a certain development stage.
2.1 The succession of fo rest sw amp
Some forests are evolving to sw amp , some for-
est swamps have came into being.The forming of
the forest sw amp most ly because of the natural suc-
cession w hich arises from the inf luence of the nature
condition or comes f rom the expanding of sw amping
in meadow and lake w hich locates near the forest.
Generally the horizontal distribution of all kinds
of communities often is a kind of reflection of vege-
tation development , so f rom the combination rela-
tionship among all kinds of sw amp vegetation com-
munities which distributed horizontally we can see
the process and tendency of development of the for-
est sw amp.For example , dyst rophic Larix gmelin
-Ledum palustre -Sphagnum sw amp and eu-
trophic Larix -Betula ovali folia -Carex sw amp
which locates in Hongqi fo restry center in Xiaoxing
an Mountain and high hirst in right bank of upper
reaches of Tang wang river , there w ere varied w idth
mesot rophic Larix -Betula ovali folia -Carex -
Moss sw amp(Fig .1)between these tw o kinds of
sw amps.At present , w ith the elapsing of time , the
sw amp developing , the peat thickening and the nu-
trient lacking , this spatial dist ribution w ill chang ,
dyst rophic sw amp plants w ill g radually int rude and
increase , eutrophic sw amp w ill develop to
mesot rophic sw amp , mesotrophic sw amp will devel-
op to dy st rophic swamp and dyst rophic sw amp w ill
continue grow and expand.Finally , the w hole
sw amp will be replaced by dy st rophic swamp.At the
same time the edge of the sw amp w ill enlarge to
mesot rophic sw amp or eutrophic sw amp.This re-
search results accords w ith Lang huiqing s[ 1] .
2.1.1 Fo restry natural swamping
Forest ry natural sw amping mostly takes place
in some fo rest regions w here topog raphy is plat ,
low , water table is high , drainage is bad , water is
often accumulated , such as flat ditch , floodplain ,
depression out of bank , terraces , lakeside , fountain
overflow zone etc.In these areas , water is easy to
accumulate , addition to gleyization of soil , soil qual-
i ty is stick , soil is hard to leak , fo r example there is
f rozen layer in some regions of Xiaoxing an Moun-
tain , it forms w ater-resisting f loor under the soils ,
and makes the earth s surface more wet or accumu-
late w ater because the surface w ater is hard to dis-
charge and permeate , which makes hyg rophy te
species invade constantly.
Firstly hygrophilous glomerate Carex sp.vege-
tation and shallow root system shrub invades , subse-
quently , Poly trichum of Bryum and Sphagnum of
Sphagnum family etc invade.All these g radually
form peat , because soil is bad of ventilation at a con-
dition of full of w ater and lack of ai r for a long peri-
od , the activities and reproductive ability of the mi-
crobes in soil is very faint , the dead plant residues
are hard to decompose.Peat keeps water , increases
soil s humidity , makes the ventilation not w ell , the
forests growing and development not well and de-
crease gradually .Fo rest succeeds to forest swamp
gradually.At the beginning of natural forest
swamping , it generally formed Larix - Carex
swamp o r Alnul -Carex sw amp in Xiaoxing an
Mountain , the process of sw amping can be con-
firmed from plant residues in peat at the bot tom of
forest sw amp at present.For example , dyst rophic
Larix -Ledum palustre -Sphagnum swamp in
Tangbei forest ry center Hongxing Forest Service Xi-
aoxing an Mountain , the peat layer thickness has
arrived 1.2 ~ 1.5 m , plant residues in peat at the
bot tom of fo rest sw amp are made of w ooden and
herbage peat , wooden plant residues are roots and
stems of Larix , herbage plant residues are roots and
leaves of Carex , which show that the sw amp is
formed from forest sw amping.
358 植 物 研 究 23 卷
Fig.1 The succession process of fo rest swamp in Xiaoxing an Mountain
1.Larix-Betula ovali fol ia-Carex sw amp.2.Carex sw am p.3.Larix -Ledum palust re-Sphagnum sw amp
4.Larix.5.Larix -Vaccin ium u liginosum -Moss sw amp.6.Rha comi tr ietum 7.Nym phaea tet ragona.8.muk
9.Carex -Sphagnum peat.10.Wooden plants-Herbage-Sphagnum peat.11.Herbage peat.12.Wooden plants-Herbage peat.13.
Slope materials:Ⅰ Lake sw amping.Ⅱ Forest sw am ping.Щ Forest sw amp.
2.1.2 Forest sw amping arised from expanding of
meadow swamping
There g row hyg rophyte or middle hyg rophyte
species because w ater table is high and the earth s
surface is more w et in moist meadow or hirst of for-
est regions.And there g radually grow s helophyte
and has seeper in sw ale betw een tussocks , especially
af ter glomerate Carex invading it , the leaves and
roots of dead plants are hard to decompose owing
lacking of air every year , finally formed peat.After
the peat formed , the humidity of the soil increased ,
accelerating the constant development of meadow
sw amping , expanding the area of seeper on the
earth s surface , at the same time , increasing the
w ater table around forest , and the forest soil become
more w et or accumulate.The vegetations under for-
est gradually w ere replaced by hygrophilous Carex
and shrubbery Betula , e.g Betula ovalifolia , B .
f ruticosa etc , the soil also constantly accumulated
the peat , the trees begun to g row bad.Finally , they
g radually succeed to fo rest sw amp.
This kind of phenomenon can be seen in the
Larix forest near the Carex sw amp on the shoal of
Tangwang river Xiaoxing an Mountain.Swamp s
surface accumulates water year by year , the peat
layer has 0.5 m around , Larix forest w hich nears
the Carex sw amp begins to sw amp , because there
w as hyg rophilous shrub Betula ovalifolia and
Lonicera edulis under the Larix forest , and the field
layer on the g round is made of Carex schmidtii and
Eriopho rum vaginatum , there is tempo rary seeper in
the rainy day s in small billabongs between tussocks ,
the peat layer of soil is 5 ~ 7 cm , it is thinner than
near C.schmidt ii sw amp , all these indicate that
forest swamping is related with adjacent Carex
swamp which comes from meadow sw amping.Be-
cause Carex sw amp accumulating w ater year by year
increases the water table under the forest , and leads
the soil mo re wet , and forest sw amping.
2.1.3 Forest sw amping arised f rom expanding of
lake sw amping
Forest sw amping of ten occur because of the
higher w ater table in the lake shoal during the lake
swamping in forest region.The process of all kinds
of lake sw amping for example , oxbow lake , Yansai
lake , caldera lake , rerong lake etc often cause adja-
cent forest sw amping .For example there is a Carex
lasiocarpa -sphagnum sw amp which is elliptic and
has a 150m diameter and no fo rest on the high hirst
of right bank of Tang w ang river Hongqi fo restry
center Hongx ing Forest Service Xiaoxing an Moun-
tain , which locates betw een Larix -Ledum palus-
tre-Sphagnum sw amp.The thick of peat layer is
1.9 m , under the peat layer is lacustrine sediments-
putrid slime , subsequently is lacustrine sediments
w hich is made of silver sand , proved it be a oxbow
3593 期 倪志英等:黑龙江省东部山地湿地主要植被类型及其演替规律的研究
lake in the pass time.The fo rest sw amp locates out-
sides of Carex lasiocarpa-Sphagnum sw amp , the
thick of the peat layer is 1.00 ~ 1.20 m , the peat
layer is thinner w ith the distance longer f rom cen-
ter , all these indicates that the Carex sw amp is
firstly caused in the oxbow lake , in the process of
lake swamping , it increases the w ater table around
i t , make the around forest sw amps form and devel-
op[ 2~ 4] .
2.2 Succession of the herbage vegetations(mead-
ow , helophy te and hydrophy te)
The chiefly condition of fo rming swamp is the
earth s surface is more wet or always has seeper.
The main ecological factors of sw amp combining
succession are the hydrological condition on the
earth s surface , the thickness of peat and the change
of its physics and chemist ry characters of peat after
sw amp vegetations fo rmed.
2.2.1 Meadow sw amping
M eadow swamping is sw amp which comes from
meadow , it mostly happens on the floodplain of riv-
er , terrace , forest grasslands between valleys.It has
the largest area sw amps in our country.Because the
topography is low-lying , the earth s surface is of ten
more wet , the w ater table is higher.Water and
g round w ater , soil porosity is full of w ater fo r a long
time , the condition of ventilation deteriorates , the
meadow swamping formed the envi ronment of lack-
ing air , and made the soil layer aggravate gleying ,
under the condition of lacking air , the plant residues
decomposed slow ly , so the plant residues can t be
mineralized.Under the condi tion of increasing w ater
and decreasing nutrient , it creates advantages for
the vegetation natural succession , makes some
meadow vegetations g radually decrease , demanding
for nutrient less , hyg rophyte increasing , especially
the glomerate Carex .After these vegetations dead ,
they can t be decomposed completely under the con-
di tion of lacking air , g radually fo rmed peat , f inally
succeed to swamp from meadow .For example ,
Carex schmidt ii swamp and C.appendiculata
which are on the mountain lands , cloughs and river
shoals of Xiaoaingan Mountain , C.meyeriana -
Deyeu xia angusti folia which is in the Sanjiang plain
etc.They mostly comes f rom meadow sw amping.
Black humus transit ion layer is product f rom decom-
posing of meadow vegetations under the peat layer
during these sw amps.
The process of development of meadow swamp
is also from eutrophic swamp to dyst rophic swamp
passing by meso trophic swamp.But then , the de-
velopment of sw amp in dif ferent areas is imbalance ,
some of them are f rom eutrophic sw amp to dys-
t rophic sw amp passing by meso trophic sw amp;some
of them were in mesot rophic swamp fo r a long t ime.
The majo rities are in the process of eut rophic
swamp , the mino rities are in the process of
mesot rophic sw amp o r dy strophic sw amp in the
swamps that forms from meadows at present in our
country.
2.2.2 Water body sw amping
Water body swamping is beginning w ith shal-
low w ater plants vegetation there grow ing hy-
drophy tes in the lake or river w hose velocity of flow
is very slow .The process of w ater sw amping can be
seen in most ly places in our country , most of them
are in the init ial stages at present because of differ-
ence of natural condit ion and the extent of sw amp-
ing .M inori ties of lake sw amping is complex , some
of them enters mesot rophic sw amp , even to the ini-
tial stages of dy strophic swamp.
The process of lake sw amping begins w ith f loat
grass.Process of low bank lake sw amping , firstly
there grow s hydrophy tes species on the bot tom of
the middle lake , e.g Potamogeton , Chara etc;
there grows phy toplankton belt on the lake surface ,
e.g Nymphoides pel tatum , Nymphaea tetragona ,
Menyanthestrifoliata or Leersia japonica etc;there
grows emerging plant belt on the shallow w ater of
the lakeside , e.g Phragm iees communities;there
grows Carex communities on the lake shoal.With
the development of the sw amp , the plant residues in
the lake w hich piled on the bot tom of the lake
formed peat , piled on the bo ttom and lakeside year
by year which made the lake more shallow ;the veg-
etation belt also invades to the center of the lake cor-
responding ly , during this course , Sphagnum in-
vades too , at the bot tom of the lake piled the peat
360 植 物 研 究 23 卷
layer w hich w as made of plant residues of dif ferent
vegetation belt co rrespondingly , finally the lake
stacks the peat , the w ater surface disappeared[ 5] .
This phenomenon of sw amping can be seen in San-
jiang plain and small Xingkai Lake also.The bot tom
of south bank of small Xingkai Lake has a slow
slope , inclines to the center of the Lake slow ly , the
vegetations have a regular belt dist ribution from
bank to center;the center is l ~ 2 m deep of water ,
there are submerged plants such as P.malaianus ,
Myriophyllum , Ceratophyl lum demersum , Hy-
dril la vertici llata etc in the bot tom of the water.
Floating leaf plant- Nymphaea tetragona , duck-
weed lotus grass and Trapa bicornis osbeck g row
outer of the lake w here the water is mo re shallow .
Submerged plant Potamogeton g row s on the bot tom
of the w ater.Emerg ing plant Zizania lati folia
g row s on the lake shoal w here the w ater is shallow .
Phragmiees communities g row in outmost of the lake
w here the water is 20 ~ 30 cm deep , accompanying
Typha orientalis , Equisteum fuviatle etc , and
Glyceris spiculosa enchased among the Phragmites
austral is in the parts area.It is Carex-Deyeu xia
angust ifolia communi ties in the seasonal hydrocele
region under 10 m in the lake.Deyeuxia angust ifo-
lia wet meadow is on the w et area of the lake bank.
At present , the lake shoal expands constantly , all
kinds of communit ies invades the center of the lake ,
the lake w ill shrink day by day.
The sw amping process of steep and deep lake is
different f rom the invasion type vegetation belt what
have said above , commonly there grows floating
plants e.g Carex lasiocarpa , Menyanthes tri foli-
ate , Comarum palustre etc.whose cervix inter-
weaves into web each other and forms “ floating
blanket” on the lake surface where the w ater move
feebly .The species w ill increase and the “ floating
blanket” will thicken gradually because sands and
plant seeds brought from blow ing and flow ing were
stopped and kept dow n by “ floating blanket” w ith
the elapsing of the time.The thin “ f loating blanket”
that can be f lew into many pieces w ill fo rm “ floating
island” which likes floating oasis and paints on the
lake s surface on the large lake.The “ f loating is-
land” is floating with the river , even connects with
the larger “ floating island” , the lake s surface w ill
reduce , and rudimental lake s surface becomes “ lake
window”.Because the thickening of “ f loating blan-
ket” constantly , under the action of g ravitation , the
plant residues of bot tom half of “ float ing blanket”
will drop on the bo ttom of the lake and form the
peat.There exists hollow pure water layer betw een
peat and “ f loating blanket” , there w ill be dangerous
if people step on the “ float ing blanket” .This stage
can be named eutrophic swamp.
With the further development of lake sw amp-
ing , the lake w indow reduces , pure w ater layer w ill
be thinner , till that up and down peat layer connect
wi th each other completely .The “ f loating blanket”
spread to the lake s center constantly , finally the
w hole lake was covered by the “ floating blanket” ,
the lake basin stacked peat , the whole lake evolved
to sw amp.The large amounts of sphagnum invade
the “ floating blanket” on the lake surface , the cover
deg ree arrived over 90%, this stage calls mesot roph-
ic sw amp.It is typical one in the lake groups the
middle reaches of Dulu river Xiaoxing an Mountain
Heilongjiang Province.The middle reaches of Dulu
river has developed meander , i t is named “ 72
curves” .Many oxbow lakes w ere disused af ter river
diversion , more than 20 in the range of 10 km ,
Quanpaozi lake is bigger among them , it has 400 m
long and 100 m w ide , there is “ floating blanket”
which is made of Carex pseudocuraica , Equisetum
heleocharis and Oenanthe javanica etc.on the steep
north-east bank.There grow s Nymphaea tetragona
on the w ater surface of inner of “ float ing blanket” ,
and Potamogeton , Ceratophyllum demersum etc on
the bo ttom of the lake.The outer of the “ floating
blanket” is connecting w ith the Carex sw amp on
the bankside.The “ floating blanket” is spreading to
the center of the lake at present.
The sw amp continues to develop , the nutrient
of sw amp is deficient day by day , sphagnum gradu-
ally has absolute advantages , flowering plant species
is decreasing , the sphagnum g radually forms moss
hillock in local regions , at the same time , the
swamp succeed to dyst rophic sw amp.e.g Carex
3613 期 倪志英等:黑龙江省东部山地湿地主要植被类型及其演替规律的研究
lasiocarpa -S .imbricatum communit ies in the
oxbow lake on the high f lood plain Hongqi forestry
center Hongxing Fo rest Service Xiaoxing an Moun-
tain
[ 6 , 7] .
All evolution principle is the same fo r the above
three different sw amps during developing course , al-
though they have different dist ributions , origins ,
the w ays of forming and successions.It is the peat
layer is thickening and ash decreasing , acidity in-
creasing w ith the developing of the sw amps.Re-
plenishment types of w ater source are g round w ater
and precipitation;Surface configuration of sw amp is
from low-lying negative physiognomy to f lat phys-
iognomy to raised positive physiognomy;Combina-
tion of swamp vegetation residues is f rom herbage
residues o r w ood combination to herbage residues
combination , to herbage and moss residues or wood
and moss combination , to S phagnum residues com-
bination.The swamp is f rom eutrophic sw amp to
mesot rophic sw amp , to dystrophic sw amp.It w ill
happen the retrog ressive succession that is the w et-
land will degenerate to farmland or wasteland if the
wetland suffered destroying.
The following figure is succession principle of
the main fo rmation of wetland vegetation on the
mountainous region of eastern Heilong jiang
Province.
Reference
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1980
4.Zhou Yiliang ed.Vegetation geog raph in the northeast of
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5.Changchun institute of Geog raphy of Chinese Academy of
Sciences.Mire in Sanjiang plain.Beijing:science press ,
1982
6.Zhao Kuiyi.Flora in swamp of China.Beijing:science
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7.Huang Xichou.Study on Chinese swamp.Beijing:science
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362 植 物 研 究 23 卷