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Diversity and Vertical Distribution Characteristics of Vascular Epiphytes in Bulong Nature Reserve Mengsong Section, Xishuangbanna

西双版纳布龙自然保护区勐宋片区附生维管植物多样性与分布特征



全 文 :西双版纳布龙自然保护区勐宋片区附生维管
植物多样性与分布特征∗
赵明旭1ꎬ2ꎬ3ꎬ Geekiyanage Nalaka4ꎬ5ꎬ Harrison Rhett Daniel1ꎬ3∗∗ꎬ 许建初1ꎬ3ꎬ
Khin Myo Myo6ꎬ Ridwan Nurdiana Dian7ꎬ Paudel Ekananda1ꎬ2
(1 中国科学院东亚植物多样性与生物地理学重点实验室ꎬ 昆明  650201ꎻ 2 中国科学院大学ꎬ 北京  100049ꎻ
3 世界农用林业中心东亚和中亚区域办公室ꎬ 昆明  650201ꎻ 4 Biomaterial Scienceꎬ Graduate School of Agricultureꎬ
Kyoto Universityꎬ Kitashirakawa Oiwakeꎬ Kyoto Prefectureꎬ Japanꎻ 5 Department of Plant Sciencesꎬ Faculty of Agricultureꎬ
Rajarata University of Sri Lankaꎬ Anuradhapuraꎬ Sri Lankaꎻ 6 Wildlife Conservation Societyꎬ Yangonꎬ Myanmarꎻ
7 Cibodas Botanic Gardenꎬ Indonesian Institute of Sciencesꎬ Cianjurꎬ West Javaꎬ Indonesia)
摘要: 利用地面观测和单绳上树法初次对布龙自然保护区勐宋片区开展的附生维管植物调查表明: 1) 在
6个样地 77株宿主上 (共调查 96株乔木ꎬ 占地约 0􀆰 2 ha)ꎬ 共有 1 756 株、 丛个体ꎬ 隶属 14 科 47 属 103
种ꎻ 相比世界其他区域ꎬ 物种丰富度处于旧世界热带水平区间ꎬ 高于温带ꎬ 但明显低于新世界热带水平ꎻ
2) 兰科植物为最丰富的类群 (60%)ꎬ 其次为蕨类植物 (24%)ꎬ 其他类群占 16%ꎻ 3) 垂直分布特征研究
表明ꎬ 距地面 10~15 m的中等高度带为物种最丰富的区间ꎬ 约有 51%的物种ꎻ 0 ~ 5 m 高度带为个体数量
最多的区间ꎬ 共有约 24%个体ꎬ 揭示了除中等高度带以外的另一个重要附生生境ꎻ 4) 常见的绞杀型榕属
植物未见ꎬ 而半附生植物密脉鹅掌柴 (Schefflera elliptica) 和多蕊木 (Tupidanthus calyptratus) 数量较多ꎮ
关键词: 附生维管植物ꎻ 垂直分布ꎻ 生物多样性ꎻ 布龙自然保护区ꎻ 西双版纳ꎻ 中国
中图分类号: Q 948              文献标志码: A              文章编号: 2095-0845(2015)03-327-12
Diversity and Vertical Distribution Characteristics of
Vascular Epiphytes in Bulong Nature Reserve
Mengsong Sectionꎬ Xishuangbanna
ZHAO Ming ̄xu1ꎬ2ꎬ3ꎬ Geekiyanage Nalaka4ꎬ5ꎬ Harrison Rhett Daniel1ꎬ3∗∗ꎬ XU Jian ̄chu1ꎬ3ꎬ
Khin Myo Myo6ꎬ Ridwan Nurdiana Dian7ꎬ Paudel Ekananda1ꎬ2ꎬ3
(1 Key Laboratory for Plant Diversity and Biogeography of East Asiaꎬ Kunming Institute of Botanyꎬ Chinese Academy of Sciencesꎬ
Kunmingꎬ Yunnan 650201ꎬ Chinaꎻ 2 University of Chinese Academy of Sciencesꎬ Beijing 100049ꎬ Chinaꎻ 3 World Agroforestry Centreꎬ
East and Central Asiaꎬ Kunmingꎬ Yunnan 650201ꎬ Chinaꎻ 4 Biomaterial Scienceꎬ Graduate School of Agricultureꎬ Kyoto Universityꎬ
Kitashirakawa Oiwakeꎬ Kyoto Prefectureꎬ Japanꎻ 5 Department of Plant Sciencesꎬ Faculty of Agricultureꎬ Rajarata
University of Sri Lankaꎬ Anuradhapuraꎬ Sri Lankaꎻ 6 Wildlife Conservation Societyꎬ Yangonꎬ Myanmarꎻ
7 Cibodas Botanic Gardenꎬ Indonesian Institute of Sciencesꎬ Cianjurꎬ West Javaꎬ Indonesia)
Abstract: The first survey of vascular epiphytes was conducted using ground based inventory assisted by single rope
technique in the recently ̄established Bulong Nature Reserveꎬ Xishuangbannaꎬ China. Results indicated that vascular
epiphytes were abundant and diverse there. On a total of 77 phorophytes in six plots (96 trees were examined in to ̄
植 物 分 类 与 资 源 学 报  2015ꎬ 37 (3): 327~338
Plant Diversity and Resources                                    DOI: 10.7677 / ynzwyj201514110

∗∗
Funding: National Key Basic Research Program of China (2014CB954100)ꎻ The Applied Fundamental Research Foundation of Yunnan
Province (2014GA003)
Author for correspondenceꎻ E ̄mail: r􀆰 harrison@cgiar􀆰 org
Received date: 2014-08-13ꎬ Accepted date: 2014-10-28
作者简介: 赵明旭 (1984-) 男ꎬ 博士ꎬ 从事植物分类学和附生植物群落生态学研究ꎮ E ̄mail: zhaomingxu􀆰 sfa􀆰 kmy@gmail􀆰 com
talꎬ covered ca. 0􀆰 2 ha area)ꎬ 1 756 individuals were recorded and were identified to 103 species (47 generaꎬ 14
families) . Compared with other regionsꎬ the epiphytes were as diverse as Paleotropicsꎬ and more diverse than tem ̄
perate zoneꎬ but significantly less than the Neotropics. Orchids and ferns comprised 60%ꎬ 24% of the total floraꎬ re ̄
spectivelyꎬ while others only took up 16%. The highest species richness and richest life ̄form diversity was found in
the middle canopy zone from 10 to 15 m (51% of total species)ꎬ where also supported high individual abundance
(19% of total individuals) . Besides the middle canopyꎬ the most abundant zone of epiphyte individuals was detected
at the base of the trunk (zone 0-5 mꎬ 24% of total individuals and 37% of total species)ꎬ indicating another impor ̄
tant niche for epiphytes in this forest environment. Primary hemiepiphytic figs were rare in the area and were not
found on the surveyed host treesꎬ while hemiepiphytic Araliaceae species (Schefflera elliptica and Tupidanthus ca ̄
lyptratus) were popular.
Key words: Vascular epiphyteꎻ Vertical distributionꎻ Biodiversityꎻ Bulong Nature Reserveꎻ Xishuangbannaꎻ China
  A high diversity of epiphytes is one of the gen ̄
erally ̄recognized and distinct characteristics of tropi ̄
cal rain forests. Epiphytes make up 25% or more of
the vascular plant diversity in certain Central Amazo ̄
nian forests (Küper et al.ꎬ 2004)ꎬ and a study in
Venezuela found that over 50% of all vascular plant
species were epiphytes (Kelly et al.ꎬ 1994). Howev ̄
erꎬ when extra ̄tropical regions are includedꎬ a large
number of studies confirmed thatꎬ globallyꎬ about
8%-10% of vascular plants are epiphytes (Gentry
and Dodsonꎬ 1987ꎻ Benzingꎬ 1990ꎻ Lowman and
Rinkerꎬ 2004ꎻ Zotzꎬ 2013).
Most studies on epiphyte have been conducted
in the Neotropics and tropical Africa ( Hsu and
Wolfꎬ 2009ꎻ Table 3). Relatively few surveys of ep ̄
iphyte diversity have been made in tropical Asiaꎬ es ̄
pecially in Chinaꎬ although some exist for subtropi ̄
cal China (Wang et al.ꎬ 1996ꎻ Xu and Liuꎬ 2005ꎻ
Yangꎬ 2008). The diversity of epiphyte community
in Hainan Island was recently investigatedꎬ and
found to be much lower than in the Neotropicsꎬ but
other aspects of epiphyte ecologyꎬ such as their ver ̄
tical stratificationꎬ appeared similar ( Liuꎬ 2010ꎻ
Liu et al.ꎬ 2010).
Xishuangbanna comprises the largest area of
tropical forest in China (Zhuꎬ 2006) and forms part
of the Indo ̄Burma biodiversity hotspot (Myers et al.ꎬ
2000). More than 4 000 species of seed plant occur in
the area (19 690 km2ꎬ Zhu and Yanꎬ 2012). Al ̄
though Wang and Zhu suggested that vascular epi ̄
phytes were a prominent characteristic of tropical
montane rainforest in Xishuangbanna (Wang et al.ꎬ
2001ꎻ Zhuꎬ 2006)ꎬ quantitative surveys of epiphyte
diversity and vertical distribution characteristics have
not been conducted.
The vertical gradient is a defining feature of for ̄
estsꎬ because habitat structure and microclimate fac ̄
tors (lightꎬ waterꎬ and mineral nutrition) (Benzingꎬ
1990ꎬ 2012) are vertically organized (Dhanmanon ̄
daꎬ 1996ꎻ Steege and Cornelissenꎬ 1989ꎻ Benzingꎬ
1990ꎻ McCune et al.ꎬ 1997ꎻ Lowman and Rinkerꎬ
2004). With increasing heightꎬ the humidityꎬ light
availabilityꎬ and substrate conditions (including pH)
all vary ̄which defines different microhabitats for epi ̄
phyte communities (Pittendrighꎬ 1948ꎻ Johanssonꎬ
1974ꎻ Kellyꎬ 1985ꎻ Cornelissen and Steegeꎬ 1989ꎻ
Parkerꎬ 1995ꎻ Dhanmanondaꎬ 1996ꎻ Freibergꎬ 1997ꎻ
Krömer et al.ꎬ 2007). For exampleꎬ Dhanmanonda
(1996) found that light availability increased expo ̄
nentially with increasing height above the forest
floorꎬ and epiphytes vertical distribution characteris ̄
tics had been shown to be influenced by the photon
flux density (Pittendrighꎬ 1948ꎻ Steege and Cornel ̄
issenꎬ 1989). Humidity is also believed to be one of
the most important factors in determining epiphyte
community assembly (Steege and Cornelissenꎬ 1989ꎻ
Pittendrighꎬ 1948ꎻ Freibergꎬ 1997ꎻ Nieder et al.ꎬ
1999). The adaptation of epiphytesꎬ in terms of
their anatomy and physiologyꎬ to vertical microhabi ̄
tat heterogeneityꎬ has been comprehensively reviewed
by Benzing (1990).
Our aims in this research are to describe the di ̄
823                                  植 物 分 类 与 资 源 学 报                            第 37卷
versity and vertical distribution characteristics of epi ̄
phyte community in the tropical montane forest of Xi ̄
shuangbannaꎬ addressing this knowledge gap in tropi ̄
cal China. Thusꎬ our study is an observational study
aimed at documenting diversityꎬ andꎬ henceꎬ can on ̄
ly be suggestive of the community assembly processes
(Johanssonꎬ 1974ꎻ Kellyꎬ 1985ꎻ Zotzꎬ 2007).
1  Methods
1􀆰 1  Study site
Our study was conducted in the recently ̄estab ̄
lished Bulong Nature Reserveꎬ Mengsong Section
(MS ̄BNRꎬ 2009)ꎬ in Xishuangbannaꎬ China (Fig􀆰 1).
The vegetation type has been categorized as tropical
montane rainforest and monsoon evergreen broadleaf
forestꎬ the former located in relatively humid regions
(montane valleyꎬ riparian)ꎬ while the latter usually
is found on the dry slopes. The elevation ranges from
1 110 m to 2 039 m (the peak of Sanduogeque)ꎬ and
mean monthly temperatures fluctuates between 15 -
21 ℃ (at 1 600 m asl) . The study site has a typical
monsoon climate ( greatly influenced by the India
Ocean monsoons) and the annual precipitation ran ̄
ges between 1 800-2 379 mmꎬ 80% of which occurs
between the months of May and October. The atmos ̄
phere has an annual relative humidity of 83% (Zhu
et al.ꎬ 2004).
1􀆰 2  Field observations
Field work was conducted during the dry season
November to Decemberꎬ 2012ꎬ when the crown lay ̄
ers were clear and epiphytes were easy to observe.
Six previously ̄establishedꎬ one hectare ( 100 m ×
100 m)ꎬ permanent sampling plots in the old growth
area of the forest ( ca. 35 m of canopy height) in
MS ̄BNR were selected for this present studyꎬ three
of which were located in a tropical montane rainforest
and three in an evergreen broadleaf forest ( Fig􀆰 1ꎬ
Table 1). Plots were separated by a minimum straight
Fig􀆰 1  Location of sampling plots and the MS ̄BNRꎬ Xishuangbanna. The map showing the old growth forest area of MS ̄BNRꎬ
the remaining blank areas are mainly covered by secondary forest or open land
9233期        ZHAO Ming ̄xu et al.: Diversity and Vertical Distribution Characteristics of Vascular Epiphytes in 􀆺       
Table 1  Basic information of sampling plots in MS ̄BNRꎬ Xishuangbanna
Plot Elevation/ m Forest type Dominate tree
Host occupancy
/ %
Basal area
/ Plot area(m2 / ha)
Epiphyte No. of Ind.
/ No. of species
107 1450 MEBF CASMEKꎬ STYTON 56􀆰 3 24􀆰 4 217 / 24
189 1700 MEBF CASMEKꎬ LITMAR 87􀆰 5 22􀆰 2 264 / 31
192 1785 MEBF CASMEKꎬ ANNFRA 100􀆰 0 25􀆰 6 389 / 30
214 1670 TMRF SYZBRAꎬ LITBAC 81􀆰 3 45􀆰 6 399 / 48
217 1700 TMRF ALAKURꎬ CRYBRA 93􀆰 8 22􀆰 2 310 / 36
240 1750 TMRF CALPOLꎬ CASCAL 62􀆰 5 34􀆰 1 177 / 21
Note: MEBF=Monsoon Evergreen Broadleaf Forestꎬ TMRF=Tropical Montane Rain Forest. Dominate trees were selected by the first two importance
value in a plot (nine sub plots tree data was appliedꎬ respectively) . CASMEK=Castanopsis mekongensisꎬ STYTON=Styrax tonkinensisꎬ LITMAR=
Litsea martabanicaꎬ ANNFRA=Anneslea fragransꎬ SYZBRA=Syzygium brachythyrsumꎬ LITBAC=Lithocarpus bacgiangensisꎬ ALAKUR=Alangium
kurziiꎬ CRYBRA=Cryptocarya brachythyrsaꎬ CALPOL=Calophyllum polyanthumꎬ CASCAL=Castanopsis calathiformis. Host occupancy was calcu ̄
lated by 100 × epiphyte occupied trees / total trees (16) in a plot
line distance of 700 m and a maximum of 3 600 m.
Nine 10 m ̄radius circle subplots were set inside a
big plotꎬ and trees (dbh≥10 cmꎬ dbh = diameter at
breast height ) were measured and identified. At
each big plotꎬ a random point was marked near the
central subplot of nineꎬ and the 16 nearest trees
(dbh≥10 cmꎬ covering an area of approximately 10 m
radius subplot) were examined for epiphytesꎬ and
96 trees were checked in total.
Binoculars and a spotting ̄scope were first used
to check host tree roughlyꎬ only if epiphyte was de ̄
tected on the groundꎬ we would access the crown u ̄
sing single rope technique at least to the main fork
site to assist specimen collecting and the latter epi ̄
phytes countingꎬ attaching height measurement work
(Perryꎬ 1978). A telescopic pruning shears with
maximum 5 m was also implemented to assist the col ̄
lecting process. Digital photographs were also taken
to document observations and assist in the identifica ̄
tion process. Due to identification barriersꎬ all small
seedlings were omitted and orchids that were hard to
identified without flower were all kept alive in green ̄
houses until identifiedꎬ then herbarium specimen
were made.
Epiphyte attaching height above ground was
measured using a 5 m poleꎬ orꎬ for those species
distributed in the outer of crownꎬ where could not be
accessedꎬ by estimating the height using the 5 m
pole as a reference. When estimating heightsꎬ the
observer stood at least 20 m away from the tree. Epi ̄
phytes that covered a substantial portion of the host
crown (creeping or clustered speciesꎬ like Pyrrosia
lingua and Cylindrolobus marginatus)ꎬ the height
was taken from the lowest to the highest points.
Two schemes of forest canopy zonation were ap ̄
plied in the former studiesꎬ six vertical tree zones
(Steege and Cornelissenꎬ 1989) and equal height in ̄
terval vertical tree zones (Zotz and Schultzꎬ 2008)ꎬ
both were proved to be efficient approachesꎬ we fol ̄
lowed the latter and forest canopy was divided into
seven height zones using a 5 m interval (the highest
crown layer was about 35 m) as several other studies
did ( Zotz and Schultzꎬ 2008ꎻ Liu et al.ꎬ 2010)ꎬ
hence our results could be compared both in the tropi ̄
cal forest domestically and abroadꎬ and the micro en ̄
vironmental factors were more consistent in the paral ̄
lel upper zones than the former arc ̄shaped scheme.
To define epiphyte individualsꎬ we referred to
the individual definition of ‘stand’ (Sanfordꎬ 1968).
A stand was defined as a cluster of pseudobulbs (or
the same epiphyte species leaves) was spatially sep ̄
arated from another—either by an area devoid of ep ̄
iphytes or occupied by other species. When the same
area was occupied by an intermingling of more than
one speciesꎬ one stand was counted for each species
present. If a stand of epiphyte covered two or more
vertical zonesꎬ especially those with long rhizomes or
large ̄area clustered pseudobulbsꎬ both individual
033                                  植 物 分 类 与 资 源 学 报                            第 37卷
number and its name were repetitively recorded in
each zone. Epiphyte species were registered either by
scientific or morphological names in the field tableꎬ
and the individual number of each species was also
recorded. Specimens were identified by comparing to
species at the herbarium at the Xishuangbanna Tropi ̄
cal Botanical Garden. Vouchers were lodged at the
Kunming Institute of Botany. The plant names and
families followed Flora of China (eFlorasꎬ 2008).
The epiphyte life forms were defined following
Benzing ( 1990 ) scheme Iꎬ and categories were
based on relationships to the host (A. Autotrophs):
true epiphyteꎬ hemiepiphyteꎬ facultative and acci ̄
dental epiphyte. Howeverꎬ when we applied these
rules in fieldꎬ distinguishing of epiphytic life forms
was difficult in some cases for the lack of former
quantitative information. Hereꎬ besides the definition
in the schemeꎬ some additional rules were set: we
grouped the species can live on treesꎬ rocks or any
surfaces where the substrate layers was thin or absent
(for exampleꎬ the moss mat or the thin layer of ca ̄
nopy soil) into true epiphyteꎬ like most of the epi ̄
phytic orchids and ferns. Hemiepiphyte referred to
those species both had epiphytic and terrestrial pha ̄
ses in their whole life processesꎬ and according to
first living on tree or groundꎬ they could be divided
into primary and secondary hemiepiphyte sub ̄catego ̄
ries. To avoid any confusion of lianas and secondary
epiphyte (Zotzꎬ 2013)ꎬ we excluded the secondary
epiphyte from hemiepiphyte categoryꎬ such as Ar ̄
oidsꎬ Piperoids and climber Ficus spp.ꎬ only species
like epiphytic Schefflera spp. would root in ground fi ̄
nallyꎬ were retained as primary hemiepiphyte. Facul ̄
tative epiphyte refers to species could inhabit forest
canopies and the ground interchangeablyꎬ such as
Peperomia spp.ꎬ Medinilla spp. and Hedychium spp.
Those species only few individuals anchored in the
canopy occasionallyꎬ but most of their individuals of
rooting in the groundꎬ were assigned to accidental
epiphyte category. The species records from Flora of
China and Flora of Yunnan (Wuꎬ 2006) were con ̄
sulted during the life from determination.
2  Results
2􀆰 1  Floristics
Across the six plotsꎬ 96 trees were surveyed.
Nineteen trees were without any epiphytesꎬ and the
remaining 77 trees (80%) were colonized by 1 756
epiphyte individualsꎬ which were identified to 103
species in 46 generaꎬ 14 families. The epiphyte com ̄
munity in MS ̄BNR was dominated by Orchidaceae
(60%) and ferns ( 24%)ꎬ while other epiphytic
species only took up 16% of the total flora recorded
(Fig􀆰 2ꎬ Appendix I) .
Fig􀆰 2  Epiphyte community floristic composition of
MS ̄BNRꎬ Xishuangbanna
2􀆰 2  Life form composition
Ninety ̄Six species of true epiphytes were recor ̄
ded. All 62 orchid species and all 25 ferns belonged
to this group. The most common species were Mycar ̄
anthes pannea (215 individuals) and Davallia trich ̄
omanoides (171 individuals). Other true epiphytes
included five Asclepiadaceaeꎬ three Gesneriaceaeꎬ
and one Ericaceae species. Surprisinglyꎬ we did not
observe any strangler figs in this tropical areaꎬ which
are normally a prominent characteristic of tropical
rain forests ( Harrison et al.ꎬ 2003). Tupidanthus
calyptratus and Schefflera elliptica were the hemiepi ̄
phytes we recorded. Five epiphytes were facultative
in characterꎬ occurring as both terrestrial and epi ̄
phytic plants and no accidental epiphyte were found.
1333期        ZHAO Ming ̄xu et al.: Diversity and Vertical Distribution Characteristics of Vascular Epiphytes in 􀆺       
2􀆰 3  Species and life forms vertical distribution
A summary of the vertical profile of epiphytes
observed in MS ̄BNR is given in Table 2 and Fig􀆰 3.
The highest species richness was found in 10-15 m
zoneꎬ which was in the mid ̄canopy of the forestꎬ
and the lowest zone was >30 m tree height zoneꎬ on
the topmost canopy layer. Howeverꎬ in terms of the
abundance of individualsꎬ the base of host trees was
the most important nicheꎬ where about 24% of epi ̄
phyte individuals were found. The highest attached
speciesꎬ Hoya chinghungensisꎬ was observed at a
height of 32 m.
Fig􀆰 3  Vertical distribution profile of epiphyte community
of MS ̄BNRꎬ Xishuangbanna
As to the vertical profile of life ̄formsꎬ true epi ̄
phytes dominated in terms of abundance in each
height zoneꎬ and above 20 m of tree heightꎬ the pro ̄
portion approached 100%. In the 15-20 m zone and
the 25-30 m zoneꎬ Hedychium villosum and Micholit ̄
zia obcordata were the only facultative epiphytes sha ̄
ring the upper canopy with true epiphytes. True epi ̄
phytes showed a humped shape of distribution of
species richness along the vertical zonesꎬ while fa ̄
cultative epiphytes declined in species richness with
increasing height. The hemiepiphytes were abundant
in the mid ̄zonesꎬ but only limited around the main
fork area.
2􀆰 4  Vertical zone details of the epiphyte community
0-5 m zone: Usuallyꎬ this is a moist area with
thick mossesꎬ very similar to the habitat of moss cov ̄
ered rocks or hard soil surfacesꎬ and is a transitional
zone for terrestrial to epiphytic lives. It is typified by
sufficient water and mineral nutrients supplyꎬ but
light availability is usually low. Overallꎬ we observed
421 epiphyte individuals from 38 species in 11 fami ̄
lies in this zone. About 60􀆰 5% of these epiphytes
were ferns (23 spp.)ꎬ mainly Polypodiaceae species
(15 spp.)ꎬ and the most abundant species were Pyr ̄
rosia lingua and Lepisorus scolopendrium. Other prom ̄
inent epiphytes in this zone were Orchidaceae species
(eight spp.). Most of the orchids growing here were
species that were generalist ( widely distributed
throughout the vertical zones)ꎬ like Pholidota articu ̄
lataꎬ Dendrobium falconeri and Dendrobium chryso ̄
toxum. Howeverꎬ Liparis cespitosa and Dendrobium
compactum were specialists (limited distribution spe ̄
cies) in this zone. The remaining epiphytes found in
this zone included species of Aspleniaceaeꎬ Davalli ̄
aceaeꎬ Hymenophyllaceae and so on. Excluding true
epiphytes ( 33 spp.)ꎬ facultative epiphyte was the
most abundant life ̄form. Most of these facultative ep ̄
iphytes had succulent functional organs to survive in
dry seasonꎬ such as leavesꎬ stemsꎬ and roots ( for
instanceꎬ the stem and leaves of Peperomia blandaꎬ
Pellionia heteroloba and Medinilla himalayanaꎬ and
Table 2  Occurrence of vascular epiphytes in the vertical profile of the MS ̄BNR forest
Height / m Indi. No. Spp. No. Indi.(% total)
Spp.
(% total) Typical species (>10% of total height zone)
  0-5 421 38 24 37 Haplopteris flexuosaꎬ Davallia trichomanoides
5􀆰 1-10 330 40 19 39 Davallia trichomanoides
10􀆰 1-15 325 53 19 51 Mycaranthes pannea
15􀆰 1-20 349 39 20 38 Mycaranthes panneaꎬ Coelogyne viscosaꎬ Coelogyne fuscescens
20􀆰 1-25 117 28 7 27 Bulbophyllum levineiꎬ Mycaranthes pannea
25􀆰 1-30 200 25 11 24 Mycaranthes panneaꎬ Cylindrolobus marginatus
>30􀆰 1    14 7 1 7 Mycaranthes panneaꎬ Bulbophyllum pectinatum
233                                  植 物 分 类 与 资 源 学 报                            第 37卷
Pseudostems of Hedychium villosum).
5-10 m zone: This height zone forms the tran ̄
sition from the trunk to the main fork. Compared to
the lower part of the tree trunkꎬ because most of the
stem flowꎬ this region is drier than ̄but still not as
dry as ̄the upper layers of crown. 40 species in 10
families of 330 individuals were recorded in this
zone. The flora composition of the epiphyte commu ̄
nity in zone 5-10 m was similar to that in zone 0-5
mꎬ but orchids (17 spp.) became more abundantꎬ
and the most common species were Coelogyne viscosa
and Dendrobium falconeri. Ferns were also a promi ̄
nent part of the epiphyte community in this zone (18
spp.)ꎬ still mainly Polypodiaceae species (11 spp.).
The most abundant ferns were Davallia trichoman ̄
oides and Polypodiastrum argutum. Hemiepiphyte
(Schefflera elliptica) made an appearance hereꎬ and
the remaining nine species belonged to Aspleniaceaeꎬ
Davalliaceae Hymenophyllaceaeꎬ Vittariaceaeꎬ Gesn ̄
eriaceaeꎬ Araliaceaeꎬ Piperaceae and Urticaceae.
10-15 m zone: The micro habitat environmental
factors here are all at a moderate levelꎬ together with
sufficient surfaces for adheringꎬ the epiphyte commu ̄
nity diversity was expected high in this zone. To that
endꎬ just as we anticipatedꎬ epiphyte species diversi ̄
ty was highest in this zoneꎬ with 53 species in nine
families and 325 individuals represented. The domi ̄
nant species were Mycaranthes pannea and Davallia
trichomanoides. Orchids (36 spp.) took up the most
proportion of the community. Ferns were also a sub ̄
stantial component (11 spp.)ꎬ and again were mainly
Polypodiaceae species (7 spp.)ꎬ like Pyrrosia lin ̄
guaꎬ Lepisorus henryiꎬ and Lepisorus sinensis. Other
ferns included Asplenium antrophyoidesꎬ Humata grif ̄
fithianaꎬ and Haplopteris flexuosa. Another character ̄
istic of this zone was the prevalence of hemiepiphytes
Schefflera elliptica and Tupidanthus calyptratusꎬ the
two hemiepiphytes were both found in this zone. The
remaining epiphytes of this zone included Asclepia ̄
daceaeꎬ Piperaceaeꎬ Gesneriaceae and Davalliaceae
speciesꎬ such as Dischidia tonkinensisꎬ Peperomia
blandaꎬ and Aeschynanthus austroyunnanensis.
15 - 20 m zone: The microhabitat in this zone
becomes drierꎬ and branches have smoother bark
and relatively ̄smaller adherence surfacesꎬ but light
conditions improve. A total of 349 individuals among
39 species in seven families were recorded. Domi ̄
nant species here were Mycaranthes pannea and
Coelogyne viscosa. Orchids still comprised the majori ̄
ty of species (23 spp.)ꎬ followed by ferns (10 spp.).
Species like Coelogyne fuscescensꎬ Polypodiastrum ar ̄
gutumꎬ and Pholidota yunnanensis were all prevalent
in this zone. The six remaining species were from As ̄
clepiadaceaeꎬ Gesneriaceae and Zingiberaceae. Epi ̄
phyte life forms diversity decreased in this zoneꎬ be ̄
sides the true epiphytesꎬ only one facultative species
(Hedychium villosum) were recorded.
20-25 m zone: We recorded 117 individuals a ̄
mong 28 species in five families in this zone. Com ̄
pared to the typical densely clustered appearances of
epiphyte community observed in lower zonesꎬ epi ̄
phytes here were scattered throughout on smaller
branches and forks. The dominant epiphytes were or ̄
chidsꎬ Bulbophyllum levinei and Mycaranthes panneaꎬ
and the proportion of orchids in the community was
high ( 20 spp. or 71%)ꎬ while there were fewer
ferns (6 spp. or 21%)ꎬ including Pyrrosia linguaꎬ
Araiostegia perdurans and Lepisorus scolopendrium.
Agapetes mannii here was the only Ericaceae epi ̄
phytic species in the study area.
25-30 m zone: Most epiphytes in this zone were
distributed on the relatively ̄larger branchesꎬ only
very few species occurred on the smaller forks or
twigs inside the canopy. 25 speciesꎬ 200 individuals
in four families were observed hereꎬ and the commu ̄
nity was dominated by Mycaranthes pannea and Cy ̄
lindrolobus marginatus. Most species were orchids
(19 spp.)ꎬ and four Polypodiaceaeꎬ one Asclepia ̄
daceaꎬ and one Ericaceae species constituting the
remaining community members.
30-35 m zone: This zone covers the uppermost
layer of canopyꎬ and was the most instable of habitat
among all zonesꎬ and could be described as the
harshest environment for most epiphytes. Only 14 in ̄
3333期        ZHAO Ming ̄xu et al.: Diversity and Vertical Distribution Characteristics of Vascular Epiphytes in 􀆺       
dividuals of seven species in four families occurred
here. Most of these species were generalists occur ̄
ring throughout the vertical profileꎬ including or ̄
chids (four spp.) like Mycaranthes pannea and Bul ̄
bophyllum pectinatumꎬ and ferns (two spp.)ꎬ Dava ̄
llia trichomanoides and Pyrrosia lingua. Hoya ching ̄
hungensis (Asclepiadaceae) six individuals were on ̄
ly found in upper zones (16-32 m of canopy) and
seemed to be a specialist to high canopy layers. Den ̄
drolirium tomentosum was also only found in this
zoneꎻ howeverꎬ as only one individual was observedꎬ
we cannot deduce anything about the vertical niche
of this species.
3  Discussion
3􀆰 1  Floristics
This is the first study to document the vascular
epiphyte community in MS ̄BNRꎬ and the first such
study performed in Xishuangbannaꎬ which is other ̄
wise well ̄known for its high plant diversity (Zhu and
Yanꎬ 2012). Although it was still an incomplete in ̄
ventory of the areaꎬ we recorded 103 species in 47
genera and 14 families on 77 host trees ( total plot
area ca. 0􀆰 2 ha)ꎬ confirming the assertion that epi ̄
phytes are abundant and diverse in the tropical area
(Küper et al.ꎬ 2004).
The survey of epiphytes in Huanglian mountain
cloud forest found 151 species (including lianas like
Piperoidsꎬ Aroids and Vitis) on 233 trees (Yangꎬ
2008). Liu (2010) found 120 species in Bawang
Ridge in Hainan Island among six forest types over a
3􀆰 6 ha area. Other studies from China reported from
subtropical or warm temperate areas all recorded
lower levels of species richness than we found at MS ̄
BNR. For exampleꎬ Xu and Liu (2005) only found
32 species on 80 host trees in a montane moist ever ̄
green broad ̄leaved forest on Ailao Mountain (about
200 km north of MS ̄BNR)ꎬ and in a semi ̄humid
evergreen broad ̄leaved forest only nine species of
epiphyte were found in 0􀆰 1 ha area ( Xu et al.ꎬ
2006). Similarlyꎬ Liu et al. (2010) found 27 spe ̄
cies in a 0􀆰 6 ha natural tropical coniferous forestꎬ
while Wang et al. (1996) found 41 species of ob ̄
ligate vascular epiphytes in a subtropical evergreen
broadleaf forest. Hsu (2009) recorded a high level
of epiphyte species richness for the whole of Taiwan
(336 speciesꎬ including lianas like Figsꎬ Piperoids
and Aroids)ꎬ not only that was based on the total
flora for a large island in many ecological zonesꎬ but
also included the confusing secondary hemiepiphytes
(Hsu and Wolfꎬ 2009).
Epiphyte research in tropical Africa suggests
epiphyte species richness is similar to the levels we
found in MS ̄BNR and for tropical China in generalꎬ
much higher than the Temperate zone ( Hsu and
Wolfꎬ 2009ꎬ Table 2)ꎬ but much lower than the
Neotropics. This may be explained by the lack some
important epiphyte families like Bromeliaceae ( ca.
1 770 epiphytic spp.)ꎬ Cactaceae (ca. 125 epiphytic
spp.) and some Orchid genera like Pleurothallis (1500
spp.) in the Paleotropics (Zotzꎬ 2013).
3􀆰 2  Vertical structure
Different micro ̄environmental factors are thought
to structure the vertical distribution of epiphytesꎬ as
described in several studies (Johanssonꎬ 1974ꎻ Stee ̄
ge and Cornelissenꎬ 1989ꎻ Benzingꎬ 1990ꎻ Lowman
and Rinkerꎬ 2004ꎻ Zotzꎬ 2007ꎻ Krömer et al.ꎬ 2007).
Moreoverꎬ investigations into microclimatic variation
have confirmed the expected patterns of vertical het ̄
erogeneity (Dhanmanondaꎬ 1996ꎻ Freibergꎬ 1997).
We found that the middle canopy had the highest
species richnessꎬ which is consistent with earlier
studies ( Steege and Cornelissenꎬ 1989ꎻ Freibergꎬ
1996ꎻ Zotz and Schultzꎬ 2008ꎻ Pos and Sleegersꎬ
2010). The humped shape distribution profile throug ̄
hout the forest canopy may be explained by the mod ̄
erate water and light conditions in the middle cano ̄
pyꎬ combined with the relative larger effective sur ̄
face area and more suitable substrate ( Nadkarniꎬ
1984ꎻ Freiberg and Freibergꎬ 2000). Krömer et al.
(2007) found thatꎬ besides the diverse and abun ̄
dant epiphyte flora of the middle canopyꎬ there was
a conspicuous epiphyte flora in the understory (40%
aroidsꎬ 35%-40% piperoids and 25%-30% ferns).
433                                  植 物 分 类 与 资 源 学 报                            第 37卷
We did not check the epiphyte flora on smaller trees
(dbh<10 cm)ꎬ but obtained a similar resultꎬ that
the highest abundance of epiphyte individuals was
observed in 0-5 m zone.
True epiphytes are the overwhelming majority in
every zoneꎬ and they mix with little proportion with
other epiphytic life ̄forms in and under middle zones.
Facultative and hemiepiphytes could be hardly found
above 15 m of canopy. The tree base areaꎬ corre ̄
sponding to 0 - 5 m zoneꎬ is a transition area from
terrestrial to epiphytic lives. Much of the host tree
base area is covered by a thick moss mat in humid
environments (Freibergꎬ 1997)ꎬ which could be a
good explanation of abundant facultative epiphytes
here. In 5-10 m and 10-15 m zonesꎬ the appear ̄
ance and prevailing of hemiepiphytes was the most
apparent characteristic. The upper zones of canopy
(15-35 m)ꎬ where almost only true epiphytes were
foundꎬ had more than half of the true epiphytes com ̄
prised of orchids. Epiphytic orchids are generally re ̄
garded as drought ̄enduring plantsꎬ and have succu ̄
lent structures ( pseudobulbꎬ terete leafꎬ or fleshy
root) (Benzingꎬ 1990)ꎬ thus can be used as a good
explanation for this phenomenon.
4  Conclusion
Our survey recorded a high diversity of epiphytesꎬ
as is generally reported in studies of tropical forestsꎬ
and confirmed a similar humped vertical structuring
of the epiphyte community around mid canopy. The
MS ̄BNR could potentially provide a protected habi ̄
tat for a large diversity of epiphytes. We suggest fu ̄
ture studies focus on species of conservation con ̄
cernꎬ as many other forests in China ̄and even in
Xishuangbanna ̄have been over ̄harvested for epi ̄
phytes ( in particularꎬ orchidsꎬ such as medicinal
and ornamental Dendrobium spp.ꎬ Vanda spp. and
Cymbidium spp.) .
Acknowledgement: We thank Xishuangbanna Tropical Bo ̄
tanical Garden and the Program for Field Studies in Tropical
Asia for supporting the “Advanced Field Course in Ecology
and Conservation” in Xishuangbanna (AFEC ̄X) during which
the field research for this study took place. We wish to thank
the resource staff and colleagues on the AFEC ̄X 2012 for
critical comments and suggestions. Funding for ZM and KMM
was provided by the Centre for Mountain Ecosystem Studies.
Other authors were funded by AFEC ̄X.
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833                                  植 物 分 类 与 资 源 学 报                            第 37卷