全 文 ：第 26卷第 2期
2006年 2月
生 态 学 报
ACTA ECOLOGICA SINICA
Vo1．26．No．2
Feb．．20()6
西双版纳尚勇亚洲象的食物组成与取食生态
陈 进 ，邓晓保 ，张 玲 ，白智林
(中国科学院西双版纳热带植物园，云南 勐腊 666303)
摘要：于 1998—2000在西双版纳国家级自然保护区尚勇子保护区的自然生境中，通过对亚洲象取食植物调查和粪便分析 ，了解
亚洲象的食物组成与取食习性。结果显示，野外跟踪调查共记录有 106种植物被亚洲象所取食 。其中有 83种出现在象的粪便
中，这些种类分别属于：禾本科 8种 (10．0％)、桑科 7种(9．9％)、蝶形花科 4种 (8．4％)、五加科 3种 (6．6％)、葡萄科 3种
(5．7％)、夹竹桃科3种(4．6％)、芭蕉科 1种(4．2％)、姜科3种(3．7％)、紫金牛科 3种(3．6％)、蔷薇科 3种(3．6％)、大戟科5种
(3．3％)、榆科2种(3．o％)、古羞草科4种(2．9％)13个植物科。根据食物中所占的比率，桑科的榕属(F／cus)、禾本科的竹类
(BambUSOideae)、小果野芭蕉(MusⅡacuminate)和莠竹(Microstegium ciliatum)是亚洲象的主要食物资源。在觅食过程中，亚洲象取
食包括乔木、藤本灌木和草本等各种生活型的植物，其中先锋种所占比率(59％)高于后续种；选择性啃食枝条的植物种类(77
种)高于牧草式取食的种类(6种)。亚洲象取食植物种类的月变化与月平均温度和月降雨量成负相关，旱季取食植物种类高于
雨季。本研究对制定保护区野生动物管理策略，以及解决保护区周边日趋恶化的人象矛盾，具有一定的参考价值。
关键词：亚洲象；食物结构；粪便分析；取食方式；人象矛盾
文章编号：1000．0933(2006)02．0309．08 中图分类号：Q958．1，Q959．845 文献标识码：A
Diet composition and foraging ecology of Asian elephants in Shangyong，
Xishuangbanna，China
CHEN Jin，DENG Xiao—Bao ，ZHANG Ling，BAI Zhi—Lin (Xishuangb M 叩 “ 0ln“ cnr， ， f， e Ac础 D，＆ ， ，
Mengla County，Yunnan Province 666300．China)Acta Ecologica Sinica，2006，26(2)：309—316．
Abstract：Diet composition and foraging ecology of Asian elephant were studied in its natural habitat in Shangyong National Natural
Reserve，Xishuangbanna，China，through field observation and dung analysis．from 1998 to 2000．A total of 106 species were
recorded as being eaten by Asian elephants，among them 83 species were identified in elephant’s dung．Plant families that
contributed a major proportion of elephant’s diet in the study area were：Gramineae(8 spp．，10．0％)，Moraceae(7 spp．，
9．9％)，Papilionaceae(4 spp．，8．4％)，Araliaceae(3 spp．，6．6％)，Vitaceae(3 spp．，5．7％)，Apocynaceae(3 spp．，
4·6％)，Musaceae(1 spp．，4．2％)，Zingiberaceae(3 spp．。3．7％)，Myrslnaceae(3 spp ，3．6％)，Rosaceae(3 spp．，
3．6％)，Euphorbiaceae(5 spp．，3．3％)，Ulmaceae(2 spp．，3．0％)and Mimosaceae(4 spp．，2．9％)．The most important
plants in elephants’diet are Ficus spp
． (Moraceae，9．0％)，Dendrocalamus spp．(Gramineae，4．5％)， n minntn
(Musaceae，4．2％)，Microstegium ciliatum (Gramineae，3．5％)and Amaloea ylH1J1．artens／s(Apocynaceae，3．1％)．Asian
基金项目：麦克阿瑟基金中老跨边界生物多样性保护资助项目(1998—2000年)；中国科学院资助项 VI(KSCX2．sw．t05)；the Joh D
． a d Cath i e T
MacArthur Foundation，USA资助项目(Grant No．97．49435)
收稿日期：2004．11 03；修订 日期：2005．04．02
作者简介：陈进(1965一)，男，江苏省太县人，博士，研究员，主要从事动植物相互关系研究 ．
*通讯作者 Author for corresp0nden‘!e．E-mail dxb@xtb异．ac．cn
Foundation item：The project was supported by the MicArthur Foundation"Sin0．L丑0 transboundary bi0diversity develoPn1ent and mna nlent．’(1998～200O)
Chinese Academy of Sciences(No．NSCX2-SW-lO5)，and the John D．and Catherine T．MacArthur Foundation．USA(Grant N0
．97．49435)
Received date：2004-1 1-03；Accepted da te：2005．04．02
Biography：CHEN Jin，Ph．D．，Professor，mainIv en aged
Acknowledgements：We thank Pmf．Z．-X
． Yang for his advice on meth0dol0gy，M8．Y．
Prof·Dr·K．。F．Cao，Mr．Andreas Wilkes
， Pwf．Q．．J．Li，Ms L． M．Li，Prof．J．-C
also thank Xishuanghanna National Natural Reserve Bureau for great suppOrt in fieldwork
Fu for her assistance in nutrient analysis．We thank Prof
． Dr．H．Zhu．
Xu for their helpful suggestions and help to improve the early draft；We
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3lO 生 态 学 报 26卷
elephants consumed a variety of plants in terms of life form ， including tree，vine， shrub and herb
． Early successional sDecies
comprise a higher pmportion of diet than late successional plants(42 spp．taking 59％ vs．32 spp．taking 37％)．Browse species
contributed a larger proportion of diet compared to grazing species(77 spp．taking 91％ s．6 spp．taking 9％)．The number 0f
plant taxa(species，genus，family)in elephant’s diet each month is negatively correlated with monthly rainfal and mean
temperature．The study may help to develop proper strategies for wildlife management especially referring to the human．elephant
conflict，which is now a serious issue in the conservation of Asian elephants in this area．
Key words：asian elephants；diet composition；dung analysis；foraging type；human·elephant conflict：Southwest China
Introduction
Asian elephant Elephas maximus L．is number about 45 000 in the wild and is distributed over South and Southeast
Asia⋯ ．As a mega—herbivore，Asian elephant consumes more than 1 10 plant species and the daily mass of an elephant’s
diet may be as much as 1．5％ 一1．9％ of its body weight in dry weight fodder ．Previous studies have suggested that
Asian elephants’feeding preferences might be related to plants’ palatability， phenophase【3]
， texture and protein
content ．Sukumar has also suggested that elephants’feeding selection were due to certain nutrients requirement，such as
crude protein。calcium and sodium ．Several studies indicate the Asian elephants’food selection behavior are highly
spatial-dependent and vary among diferent seasons 一 ．The investigation in different areas may enrich our knowledge on
feeding preference and forging ecology of Asian elephant．
Asian elephant ale often considered as the flagship species in their habitats，and efforts have been puting into its
conservation(
．
However．increasing conflicts between human and elephant occur in many areas in Asian countries[ ，
．
Elephants come out ot’the protective habitats，destroy farming lands and even injury people．Elephant human conflicts
become an important issue to the conservation of elephants in many areas ，as wel as in Xishuangbanna，Southwest of
China，where most wild elephants of China distributed ．Understanding of the diet composition and forging ecology of
Asian elephants in our study area may help to develop proper strategies concerning the human—elephant conflict．
Our study addressed the folowing questions．What kinds of plants do Asian elephant feed of in the study area?What
is the seasonal change in their diet patterns?Whether the diet composition correlates to certain nutritional items?
1 Study area
This study was conducted in Shangyong protected forest(21。15 ～21。25 N，101。20 ～101。37 E)of Xishuangbanna
National Natural Reserve，Yunnan Province of China 93
．
Average rainfall is about 1 500 mm per year．With the influence
of the southwest monsoon，the wet season starts in early May and ends in early October，so that the climate in the study
area can be characterized by two distinct seasons，the wet season(May October)and the dry season(November—April)．
Annual average temperature is 15．1～21．7qc．The total area is about 321．85 sq．km and elevation ranges from 630 m to
1 300 m above sea leve1．The floristic composition and structure of the forest have been described by Cao& Zhang⋯ ．The
forest is linked with Nam Ha National Biodiversity Conservation Area(20。33 一21。13N ，100。08 ～101。30 E，2224
km )in Luang Namtha province of Lao People’Democratic Republic．
The exact number of the wild Asian elephants in the forest is not yet known．Local villagers estimate there are about
60 individuaIs in the forest in Shangyong while the largest number implied by villagers’witness was 33 individuals·The
estimate of 1 30～1 50 elephants in that forest by Zhu＆ Santiapillai is more optimistic．Differences may be related to the
fact that wild elephants frequently across the China—Laos border ．
2 Methods
2．1 Field observation
We f0lo、ved fTesh elephant trace on 5～8 January，4～6 May and 27～28 August in 2000 respectively，and each
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期 陈 进等：西双版纳尚勇亚洲黎的食物组成与取食生态 311
time walked more than 3 km．During the walks，we recorded all plants eaten by elephants， dent fying±oraging tYPe
(grazing vs．browsing)，life form of the plants．The classifcation of life history was based 0n our± eld expenences
Voucher specimens were c0lected and brought to laboratory for identifcation and references of plant t ssues m elephant
dung that each species were consumed by elephants in the same are -
2．2 Dung analysis
We colected fresh e1ephant dung samples from the field every month from September 1998 to December 1999，and
everv two monfhs during 2000．All together we collected 22 samples．Dung was sampled from >1 0 diferent places each
time．afld the nearest distance between each sites was >30 m， so that a good representative sample is obtained- resh
dung samp1es were brought back to laboratory and dried at 40~C in an oven and were stored under dry condlflon tor
subsequent analysis．A smal part of each dung sample was randomly picked and then placed in a dish with distilled water,
and analvzed under a microscopy(25 times)work，with 30 views being taken per sample-We identified and c0unted the
appearance 0f eaeh plant species by using known plants vouchers as a reference· Microscopic charac ers， such 五Ls
epider pattem ，trichomes，cellular inclusions，and starch grains，as wel as shape，texture’and color 0士part cles were
used五Ls characters f0r identmcation．We excluded the views with unidentifiable plant tissues·Each dung samPle requ re
about 2 hours of microscope work．
The proportion of plant species i(P )in each month was calculated the folowing formula：
= s ／∑s ，
1
Where S．denotes the sum of occurrences of species i that appeared in the 30 views，and n means the total nun1ber
of species recorded in the sample．
The pmportion 0f species in different seasons was calculated by the average of尸 over the months in a glVen season·
The average proportion of species i was calculated using the average of Pi of 2 1 months·
2．3 Measurement of nutrients
Representative parts of plants foraged by Asian elephant were collected．The determination of sampling parts from th。
Dlants was based 0n auth0rs’field obsenration．Samples were sealed in plastic bag and brought back to laboratory within
tw0 davs for chemica1 analysis．The following nutritional parameters were determined： (1)Energy content per unit o±dry
mass was calcu1ated using the sum of energy from protein(4×4186 kJ／g)，carb。hydrate(4×4186 kj／g)and fat(9×
41 86 kJ／g) ’ ．(2)Warer content was measured by drying in an oven 40℃ until constant mass reached over a period of
about one week． (3) Content 0f protein was calculated by 6．25 × (nitrogen content)． (4) Crude faty acid was
determined bv using the Soxhlet—extI．acti0n method． (5)Carbohydrate was calculated using the folowing formula： Carbon
hvdr0gen=1一(warer content)一(ash)一(protein)一(crude faty acid)一(crude fiber)，(6)Crude fiber：the celulose
content、忸s qualifed by determining the ash—free dry weight of the residues after treating the samples with 1．25％ (w／v)
of t4，so and 1，25％ (WIV)of NaOH．(7)Ash was measured by firing under 525R2，(8)Mineral element contents：
For ana1ysis 0f nitrogen(N)，samples were digested with H2SO4一H2O2，and for phosphorus(P)，potasium(K)，sodium
(Na)，calcium(Ca)，magnesium (Mg)，zinc(Zn)，copper(Cu)and iron(Fe)，samples were digested with HNO3一
HC104．T0ta1．N was detem ined by diffusion，total—P by molybdenum—blue colorimetrie method，K，Na，Ca，Mg，Zn，Cu
and Fe were determ ined by Inductively Coupled Plasma
USA，1999．Type：IRIS Advantage—ER)，
Atomic Emission Spectrometry(Thermo Jarrell Ash Corporation，
3 Results
3．1 Plant species eaten by Asian elephants
Asian elephants consumed a wide range of plant species．In totaly 1 06 species were recorded by field observation and
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3l2 生 态 学 报 26卷
83 species by dung identification(see Appendix 1 A listof the plant species of>1％ in average diet proportion)．Plant
families that contributed a major proportion of elephant’s diet in the study area were：Gramineae(8 spp。，10。O％)，
Moraceae(7 spp．，9．9％)，Papilionaceae(4 spp。，8．4％)，Araliaceae(3 spp．，6．6％)，Vitaceae(3 spp．，5．7％)，
Apocynaceae(3 spp．，4．6％)，Musaceae(1 spp．，4。2％)，Zingiberaceae(3 spp。，3．7％)，Myrsinaceae(3 spp。，
3。6％)，Rosaeeae(3 spp．，3．6％)，Euphorbiaceae(5 spp。，3．3％)，Ulmaceae(2 spp。，3．O％)and Mimosaceae(4
spp．，2．9％)．The most important plants in elephants’diet are觑 spp。(Moraceae，9．O％)，Dendrocalamus spp．
(Gramineae，4．5％)， Musa acuminata (Musaceae，4．2％)， Microstegium ciliatum (Gramineae，3．5％)and
Amalocalyx yunnanensis(Apocynaceae，3．1％)(Appendix 1)．
Asian elephants consumed a variety of plants in terms of life form，including tree，vine，shrub and herb(Table 1)．
Eady successional species contain a higher proportion of diet than late successional plants．Forty—two early successional
species that accounted for 59％ in diet were recorded by dung analysis．while 32 species of late successional plants
accounted for 37％ of the diet(Table 1)。Asian elephants often visit secondary forest habitats or forest gaps where pioneer
plant species are more abundant。Browse species contributed a larger proportion of diet compared to grazing species(Table
1)。We also observed that elephants did not obviously avoid feeding on plants that have defense traits against herbivores，
such as thorns on leaves and twigs。
Table 1 The number of species fN)and pralmrtlon in diet(％)of diferent kinds of plants consumed by Asian elephants(The data is based on dung
analysis and numbers in parentheses me the number of species identified by field observation
* ES：Early successional plants，LS：Late successional plants
3．2 Seasonal change of plants consumed
The number of plants consumed by Asian elephants difered among seasons．During the cold dry season，elephants
consume more plant taxa at family，genus and species levels compared with those consumed in hot rainy season．The
number of plant species in diet was negatively con-elated with the monthly mean temperature and precipitation(Fig．1)．
Some species，such as Spondias pinnata (Anacardiaceae)，Microstegium ciliatum (Gramineae)，Curculigo
i
言
芎
缸
主
Monthly mead temp~ature Monthly rainfall
Fig．1 The number of plant speeies CONSnmed by Asian elephant in the diet of each month a8 identified through dung anaiysis was negatively related with th
monthly rabfal(ram)，monthly mean temperature(℃)；n 21
Nunlber。f plant taxa genu$and 脚i／y ar：e als。negatirely relate t。the m。nthly Ilean temperature(a)．(R =O．4942，P<0．00l；R =0．4136，P<0·O1)
and m0nthlv rainfall(b)．(R ：O．3597．P维普资讯 http://www.cqvip.com
2期 陈 进等：西双版纳尚勇亚洲象的食物组成与取食生态 3l3
capj m珊 (Hyp。xidaceae)，and ArnomMm glabrum(Zingiberaceae)were c。nsumed more in the wet seacon·While，other
plants such as Castanopsis mekongensis ro。t(Fagaceae)，Acacia sP．stem(Mimosaceae)，Leen f cn(Vitaceae)were
more consumed in the dry season(Appendix 1)．
3．3 Plant selection in relation to nutrients
The nutrient contents of 29 most important species consumed by elephants are given in Appendix 2．Iron content in
plant parts was highly corelated with the plant species proportion in the diet(R =O．4199，P=O．0006，N=25)·No
relationshiDs were found between the proportion and other nutrient items，such as energy，protein，protein to fiber ratio，
protein plus ash，the contents of phosphorus，potassium，sodium，calcium，magnesium，zinc and copper，etc·
4 Discussion
The diets of elephants in our study were very diferent from those previously reported in India．Among the most
important plant ramilies of elephant diets in our study， only Gramineae， Papilionaeeae and Mimosaceae have been
reeognized as important in India[
．
Comparing the two studies，among 39 plant genera of the elephants fodder recorded in
sjr Lanka，29譬enera occurred in our study，but only 6 ofthe genus(21％)were fed by elephants．These were Capparis，
Cordia，Croton，Malotus，Phylanthus and Sysigium_2J．Also，among 17 plant genera of the elephants’diets in
Indi aI ，14 genera exist in our study area，but only 4 of them(29％)recorded as elephants’food plants(Acacia，
Albizia．Capparis and Dendrocalam“s)．Janzen has pointed out that diet breadth cannot be defined usefully in terms of
plant taxa， as herbivores are unlikely to select food on the basis of L in binomials⋯
． Plant resources are spatialy’
dependent．AsJan elephants may eat contemporary plants to meet their nutritional requirements．Elephants need such large
口uantities of food it is to be expected that availability of plant species will be an important factor in their diet·
Asian elephants are mixed feeders eating both graze and browse．Early studies by direct field observation suggested
that food plants by grazing makes up a major part of elephants’diet Later Sukumar&Ramesh reported that intake of
plants by browsing constitutes a major part of diet as indicated by analysis of stable carbon isotope ratios in dung ·Our
study showed that 91％ of food plants were consumed by browsing·
A total of seven species were recorded as,consumed by grazing，of which six species appeared in dung．Comparing
nutrients of grazed species with browsed species did not reveal any significant diference．
The minimal amount of different nutritional components to maintain elephants’body is unknown．Sukumar discussed
elephants’requirements for Drotein，calcium and sodium，and suggested that the minimal daily amounts for an adult
elephant(supposed body weight 3 t)were 900g of digestible protein，and 60 g and 75—1 00 g of calcium and sodium
respectively．Supposed an elephant consumed plants amounting to 1．5％ in dry weight of its own body weight and that
40％ of crude Drotein are digestiblel3_
，
the minimal amount of average content of the consumed plants would be 5％ ，
0．13％ and 0．17％ ．O．22％ for protein，calcium and sodium respectively．The average amount of nutritive items in 29
plants identifed in our study suggested that the plant materials in the study habitat were easily sufficient to supply enough
Drotein and calcium，while there is some difficulty in meeting elephant’s requirements for sodium．This might be the
reason why elephants often seek sodium from salt pond．
Some plants were only consumed during certain seasons．This may be interpreted in term of their nutrient content-
Acacia spp．got intensively consumed in dry season compare that in wet season in our study(4．1％ vs．1．2％ in diet)·
This is also reported by several studies in other regions，and，the selection of Acacia is explained to be high protein
contentl
．
Our study indicated that the consumed part of Acacia species did not show a relatively high protein content
(3．2％～3．5％)but had a relatively high fiber content(40．4％ 一51．3％)．Elephants need to lose their body weight
during the dry season so as to avoid excessive fat deposits that inhibit eficient therm oregnlation ．During the dry sea,on，
excessive protein intake is also undesirable，because nitrogen excretion requires more water which may be in short
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314 生 态 学 报 26卷
supply ．Seeking for Acacia and Leea iadica as fiber rich plants may be critical for elephants to achieve a proper ratio of
protein to fiber and to avoid constipation during dry and hot season．
The present study had showed the iron content in consumed plants positively related to the composition percentage in
dung．Th e reasons for this pattern require further study．It might be that elephant prefer iron rich plants，or by chance
those intensive consumed plants have higher iron contents．
Understanding the diet composition and forging ecology may help conservationists and protective reserve managers to
develop proper strategies for a better wildlife management in this area，especially referring to the conflict between human
and elephants．The National Natural Reserve is strictly protected and al vilagers have been excluded from the reserve．
Secondary and early successional forests which were 0ften created by villagers’shifting cultivation in the past time．nOW
become less and less in Natural Reserve．This may cause the decrease of biomass of many important elephant food plants in
the forest， e．g．， Dendrocalamus spp． (Gramineae)， Musa acuminata (Musaceae)， Microstegium ciliatum
(Gramineae)，etc．With the help of the research team，forest managers are trying to reestablish some patches of early
successional forests and plant some elephant food plants in the protect area 80 as to solve the human elephant conflict．
Certainly the effects of such practice need eareful assessment．
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Araliaeeae
Heteropanaxfragrans
Macropaaax dispelmus
Trevesia pMmata
Bignoniaceae
Oroxyh~mindic~zn
Boraginaceae
Cordiafurcark~
C邮 etaeeae
Combretum prmctatum
Cypcraceae
Carex baccans
Eu曲orbiaeeae
Brldd tomentosa
Croton argyratus
Fagaceae
Castanopsis ∞
C．mckongensls
Gramineae
Bamboo
Microst Ibm ciliatum
T sfrr,o~aerra mn Hm
Hypoxidaceae
Curculigo capitulata
Marar,taceae
Phryrdum capitatum
Menispermaeeae
Parabaena sagitata
Mimosaceae
Acacia megaladena
Acacia pennata
Moraceae
Ficus atriculata
F． stulosa
F．佃 emDsn
F．semieordata
Musaceae
Mu Ⅱ acuminata
Myrsinaceae
Meo．~a ndica
Papilionaceae
Derfs caudatilimba
Milletia leptobotrya
Shuteria hirsu．ta
Ro$~ceae
Rubus elliptieus var．obcordatus
R ．p vzr．cordatus
Rubiaccae
l／ncaria laev~ a
U．scan&ns
Solanaeeae
Solanum tOl~3tm
Staphyleaeeae
Turpiniapordfera
Stemuliaeeae
2．1
2．1
1．1
2．5
1．8
0．7
1．9
1．5
1．8
1．3
O．8
0．9
2．2 2．2
1．5 0．4
1．8±0．9
2．3±0．7
2．5±1．0
2．0± 1．3
1．5±0．8
1．0±0．g
1．7±0 7
1．1-e 0．9
1．2± 1．1
2 3± 1．1
1．1± 1．2
4．5± l
3．5± l
1．8±、
1．6± l 1
l 5±0．7
2．3土1．1
1．4± 1 7
1．1± l 2
2 4±1．0
2．5±0。6
2．4±0．6
1 7±0．8
4．2±1．3
2．1士1．0
2．2±0．6
1．1±1．0
2．8±O．6
l 9±0．7
1．1±0．9
1．3±1．1
1．3±1．0
2．3±0．7
l、0±0．8
TRe
Tree
Ttee
Vine
Herb
Vinc
Tree
Vine
Shrub
Shmb
Vine
Vine
Shrub
yes
DO
yes
ye晕
yes
yes
n0
yes
yes
yes
yes
yes
yes
yes
yes
yes
Ls
LS
LS
Br
Br
Br
Br
Br
BT
Br
r
Br
曰r
Br
n0 ES
yes
yes
yes
yes
yes
yes
yes
ES
ES
ES
ES
ES
ES
ES
Br
Br
Br
Br
Br
Br
Br
yes ES Br
Tree no LS
耻 m m 瞰 研
髓 雎 隅 既 盼 皓 瞵 雎
}三 聃
№№ ‰ ‰ 胁胁
¨ ¨ ¨m ¨ 邮 ㈨
”
m m ㈨ ¨ ¨ ¨ ¨ ¨ 枷 l 2 l 2 1 l
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316 生 态 学 报 26卷
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●__ ●__-__ _-_●___ _。__ _____-_ __‘。●。 。。。。●_ 一
Bytneria grandifolia
Ulmaceae
Celtis timorensis
Urticaceae
Boehmeria dldemloides2．0
Vitaceac
Leea ndica
Tetr~tigma planlcaulum
Zingiberaceae
Amomum glabrurn
1．5 0．7 1．1±0 9 Vine yes LS
2．6 2．8 2．8±0．6 Tree
1．6 1．9±0 8 Shrub
2．3
2．6
2．5
1．9
2．4± 1 0
2．4±1．0
Shrub
Vine
ES Gr
yes
yes
Ls
LS
Br
Br
1．0 2．3 1．8±1．2 Herb yes LS Br
1．1 1．1 1．2．4-0．8 Herb yes LS Br
LF lifef0ITn；AR abm母0f re th，ifthe plant h e出d for a蛇xual propagation and re-gmw after bei“g eatenis‘yes’，。therwi8eitis’n。’；LH
l hist。ry，LS 【丑te 8ucce搴sional plant8，ES Early succe8sional plant8；FT ~raging type，Br browse，Gr graze；Dry sea8。n：N。vembe 一April，R i y
se n：Mav一 0ctober； Bambo。here includes Dinochl。。bannaensi ，Gigamochl。。nigrocillat。and DendrOC。lamus 。m “
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