全 文 ：广 西 植 物 Guihaia 30(3)：339— 342 2Ol0年 5月
假瘤蕨属 (水龙骨科)植物叶片
附属结构的生态意义
邵 文1，商清春2，陆树刚1
(1．云南大学 地植物学与生态学研究所，昆明 650091；2．牡丹江医学院 影像系 ，黑龙江 牡丹江 157011)
摘 要：电镜下观察并研究 了假瘤蕨属植物的叶片附属结构 。报道了蜡质颗粒、片层、柔毛和鳞片。叶片附
属结构在数量及性质上的差别显示出种类与环境的适应性不同。结果表明，假瘤蕨属植物中土生的种类具有
较多的蜡质颗粒、柔毛及鳞片，而附生的种类具有较多的颗粒及片层 ，这些叶片特征均与亚热带常绿阔叶林的
生态环境相适应。
关键词 ：叶片附属结构 ；生态适应性 ；假瘤蕨属
GLC Number：Q942 Document Code：A Article ID：1000—3142(2O10)03—0339—04
Ecological significance of leaf appendage in the fern
SHAO Wen ，SHANG Qing-Chun ，LU Shu-Gang
(1．Institute of Ecology and Geobotany，Yunnan University，Kunming 650091，China：2．Department
of Physics，Mudangjiang Medical College，Mudanjiang 15,7011，China)
Abstract：Detailed morphological study of the leaf appenages Of the fern genus Phymatopteris in China under SEM is
presented．The study reveals the wax granules，stripes，apophysis and hairs．Leaf appendages are constant in some
species and variable in others in the aspects of quanlities and characteristics and thus of great significance in under—
standing the relationships between the species and the ecological environments adaptable．The results support that
there are more wax granules，hairs and scales on the terrestrial species，and more granules and stripes on the epiphytic
ones of this genus．Al l the frond appendage characters reveal an adaptation to ecological conditions of the fern genus
P， natopteris to the subtropical evergreen broad-leaf forest．
Key words：leaf appenages；ecological adaptable；Phymatopteris
Ph ymatopteris Pichi—Serm．(Polypodiaeeae)is
characterized by simple，trifid，palmatifid，pinnatifid
leaves of usually chartaceous texture，superficial or
slightly immersed large sori．This genus is of consider—
able size，more than 60 species，the majority of which
are from the mainland of Asia，terrestrial，lithophytic or
epiphytic in evergreen broad—leaved forest(Ching，
1964)．|『)^ ， 7“￡D r (Pichi，1973)is a natural group，
which has been confused with Microsorum，Crypsinus
and Selliguea by Copeland(1947)，Hohtum(1954)and
Hovenkamp(1998)，but there are several eharacteris—
tics(Table 1)that can be used tO distinguish these four
genera(Shao＆ Lu，2009)．
From Table 1，we can distinguish these four gene—
ra by several characteristics．However，some taxonomic
revisions in the infrageneric range and basic experi—
Received dale：2009—08-0 7 Accepted date：2010-01—18
Foundation item：Suplmrted by the National Natural Science Foundation of China(30770164)
Biography：SHAO Wen《l979一)．Ph．I)，female．mainly engaged in Systematiel~xonomy of Ferns，(E-mail)shaowenl9792005@163．corn．
Author for eorreslxmdence．E-mail：shuganglu@163．corn
340 广 西 植 物 3O卷
Table 1 Comparison among Crypsinus，Phymatopteris，Selliguea and Microsorium
ments need to be done．
Leaf appendage such as depositions of wax gra—
nums，hairs，stripes and scales are of potenial ecological
significance，which provides an important source of envi—
ronmental adaptability． No research about this genus，
describing the characteristics of leaf appendages，has
been reported．The purpose of this paper is to provide a
detailed study under SEM ，and that might assist in un—
derstanding of the ecological adaptation of the fern genus
Pkymatopteris to the evergreen broad_leaved forest．
1 Materials and methods
Total of 20 taxa of Phymatopteris were gathered
from the specimens of PYU (in Addendix)，materials
for SEM observation were directly affixed on stubs by
double-sided adhesive tape without any treatment．Af—
ter gold sputtering by IB-5 particle spraying apparatus，
the specimens were examined and photographed under
AM ARY一】000B SEM．
2 Results
The characteristics of leaf epidermal appendages
are listed in ~tbles 2．It appears that epidermal ap—
pendage feat ures are constant within species and thus
may be used in analysis and discussion．
Granules，st ripes and apophyses，hairs and
scales are presented in the leaf epidermis under
SEM observation．
Granules．Lots small granules(d≈ 2 m)are scat—
tered on both sides of epidermis of Phymatopteris
chrysotricha(Fig．1：1)，P．tenuipes(Fig．1：2)and P．
hastata(Fig．1：3)．Granules(d~ 20 m)are evident on
both sides of the leaf epidermis of P．dactylina(Fig．1：
4)；on the abaxial epiderm is of P．ebenipes(Fig．1：5)
and P．s“6 6 n pe (Fig．1：6)．
Table 2 Leaf epidermal characters of
Phymatopteris under SEM
Ad= adaxial epidermis；Ab=abaxial epidermis
Stripes and apophyses．Stripes and apophysis are
presented in most species of this genus．Many stripes
are found in Phymatopteris engleri(Fig．1：7)and P．
oxyloba(Fig．1：12)．Few stripes are presented in P．
yakushimensis(Fig．1：8)，P．daweishanensis(Fig．1：
14)，P．shensiensis(Fig．1：16)，P．stracheyi(Fig．1：17)
and P．glaucopsis(Fig．1：19)． Few apophyses are
found in P． oensis(Fig．1：9)and P．stewartii(Fig．
1：20)．Rounded apophyses occur in Phymatopteris
3期 邵文等：假瘤蕨属(水龙骨科)植物叶片附属结构的生态意义 34l
Fig．1 2O．CI／aracters of leaf epidermis of PhymatopLeris under SEM． I．1iz3rⅢatopteris chr3，sotricha；2．P．tenuipes；3．P．hastata
4．P．clactylina；5．P．ebenipes；6．P．subebenipes；7．P．engleri；8．P．yakushimensis；9．P．majoensis；10．P．omeiensis；l1．P．trisecta；12．P．oxylo
：13．P．piamnaensis；14．P．da~ ishanensis；15．P．nigrovenia；16．P．shensiensis；17．P．stracheyi；18．P．comnixta；19．P．glaucopsis；20．P
stewartii．
omeiensis(Fig．1：10)．
Hairs．Lots of long hairs are found on both
sides of the epidermis of Phyrnatopteris trisecta
(Fig．1：11)and P．nigrovenia(Fig．1：15)．A few
342 广 西 植 物 3O卷
l1airs are distributed on the abaxial epidermis of P．
pianmaensis(Fig．1：l3)and P．conmixta(Fig．1：
l8)．
Scales．Considering the occurance of scales not
on】v on the frond but also on the rhizome，we
would not present the figures of scales here．How—
ever．the value of scales would be discussed later．
3 Discussions
Some species have appendage on the one or two
sides of leaf epidermal cells，such as granules(Phy—
matopteris dactylina，P．subebenipes and P．ebenipes)，
small granules(P．tenuipes，P．hastate)，hairs(P．C07一
~niJcta，P．nig roTJe~lia and P．trisecta)，and scales(P．
subebenipes and P．ebenipes)． Considering the living
envimnment of the above mentioned species，we notice
that they all spread mainly in the dry regions of south·-
west China，and are terrestrial or epiphytic on rocks or
trunks of evergreen broad-leaved forest under alt．2 500
m．To prevenl lhe cold lemperature(≤O℃)in winter，
the species should have some dress to adapt such envi—
ronment． We also notice that the terrestrial species
have more appendages than the epiphytic ones；per—
haps the terrestrial ones need more protections to adapt
the colder environment．
The original function of stomata is the limitation
of water Ioss in Iand plants and the maintenance of ho—
moiohydry，while allowing gas exchange． The guard
cells of stomatal apparatus in this paper are all slightly
sunken，and most stoma pores are closed under the lay—
er of wax granules and stripes，which may be limited to
the waler loss and keep it warmer，and thus could en—
dure the cold winter safely．
A special terrestrial species is Phymatopteris tri—
sectu，distributing in the shade area of Yun-Gui Plateau
In short，the leaf appendages of ferns are of great
existed significance and can provide an important
source of environmental adaptability，and thus have
some ecological adapations of the fern genus Phy—
matopteris to the evergreen broad-leaved forest．
Addendix：Voucher informations of materials used for study
Phymatopteris chrysotricha(C．Chr．)Piehi-Serm．
CHINA：Yunnan，Pianma，Shao W ．025
P．conmixta(Ching)Pichi—Serm．
CHlNA：Yunnan，Xinping，M t．Ailaoshan，Shao W．004
P．dactv 西ta(Christ)Pichi—Serm．
CHINA：Yunnan，Deqin，Shao W．009
P．dazt,eishanensis S．G．Lu
CHINA：Yunnan，Pingbian，Mt．Daweishan，Lu S—G．28885
P．ebenipes(Hook．)Pichi—Serm．
CH1NA：Yunnan，Pianma，Shao W ．021
P．engleri(I uerss．)Pichi-Serm．
J APAN：Yakushima Island，S．Mitsuta 267
P．glaucopsis(Franch．)Pichi—Serm．
CHINA：Yunnan，Dali Eryuan，Chu W ．一~L 23013
P．hastata(Thunb．)Pichi—Serm．
CH1NA：Yutman，Pingbian，M t．Daweishan，Shao W．002
尸． ，ajoensis(C．Chr．)Pichi—Serm．
CHINA：Yunnan，Dali Yongshan，Chu W．一M．4898
P．，tigrovenia(Christ)Pichi—Serm．
CH1NA：Yutlnan，Deqin，Chu W．一M ．26458
P．Ollleiensis(Ching)Pichi—Serrn．
CH1NA：Sichuan，Daxiangling，Kong X．一X．3971
P．oxyloba(Wal1．ex Kunze)Pichi—Serm．
CHINA：Yunnan，Xinping，M t．Ailaoshan，Shao W．013
P．pianmaensis W ．M．Chu
CH1NA：Yunnan，Pianma，Chu W ．一M ．11350
P．shensiensis(Christ)Pichi—Serm．
CHINA：Sichuan，Kangding，Kong X．一X．6235
P．steztwrtii(Bedd．)Pichi—Sem1．
CHINA：YulIlarl，Pianma，Shao W．019
P．stracheyi(Ching)Pichi—Serm．
CHINA：Yunnan，Pianma，Shao W．018
P．subebenipes(Ching)Pichi—Serrn．
CHINA：Yunnan，Dali，M t．Cangshan，Shao、 005
P．tenuipes(Ching)Pichi-Serm．
CH1NA：Sichuan，Chongqing．Mt．Jinfoshan，I，iu Z．一Y．3547
P．tl—isecta(Baker)Pict1i Serm．
CHINA：Yunnan，Kunming，Shao W．024
P．yakushimensis(Makino)Pichi—Scrm．
J APAN：Yakushima Island，Murata s．n．
Ackamwledgement We would Iike to thank Prof．W．
M．Chu for providing access to the specimens cited in
this paper，and Prof．R．X．Wang and engineer X．K．
Fan for assistance on the observation of LM and SEM．
The first author would also thank Prof．Damian Shea of
North Carolina Slate University for helpful suggeslions
and critical reading of the manuscript．
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