全 文 ：假瘤蕨属 (水龙骨科)植物叶表皮特征的比较研究＊
邵　文1 , 陆树刚 1＊＊ , 商清春 2
(1云南大学地植物学与生态学研究所 , 云南 昆明　650091;2牡丹江医学院影像系 , 黑龙江 牡丹江　157011)
摘要:在光镜和扫描电镜下观察了假瘤蕨属 2系 、 5亚系的 24种植物的叶表皮。结果表明 , 叶表皮细胞
形状不规则 , 垂周壁波曲状。下生气孔 , 气孔类型有极细胞型 、 共环极细胞型 、 腋下细胞型 、 聚腋下细胞
型和不规则型 , 常伴生出现。在扫描电镜下 , 叶表皮气孔器外拱盖内缘为浅波状 、 少平滑;一些种的保卫
细胞两端有 “T” 型加厚;角质膜波状有条纹, 有时附有颗粒。叶表皮的一些特征对该属部分种的区分有一
定的参考价值。叶表皮的微形态特征, 如气孔器类型和垂周壁类型特征不能用来界定假瘤蕨属的系和亚系。
关键词:水龙骨科;假瘤蕨属;叶表皮;系统学意义
中图分类号:Q944　　　　　　文献标识码:A　　　　　　　　文章编号:2095 -0845(2011)02-174
-09
ComparativeMorphologyofLeafEpidermisintheFern
GenusPhymatopteris(Polypodiaceae)
SHAOWen1 , LUShu-Gang1＊＊ , SHANGQing-Chun2
(1InstituteofEcologyandGeobotany, YunnanUniversity, Kunming650091, China;
2DepartmentofPhysics, MudanjiangMedicalCollege, Mudanjiang157011, China)
Abstract:Leafepidermisesof24 taxa, representing2seriesand5subseriesofthegenusPhymatopteris(Polypodi-
aceae), wereexaminedbylightmicroscopy(LM)andscanningelectronmicroscopy(SEM)todeterminethesys-
tematicallyinformativecharacters.Theleafepidermalcellswereusuallyirregularinshape.Thepaternsofanticlinal
walsweresinuolate, sinuous, andsinuate.Thestomatalapparatuswererestrictedinabaxialepidermisesforalspe-
cies, andgenerallycouldbeassignedtopolocytic, copolocytic, coaxilocytic, axilocytic, andanomocytictypes.
UnderSEMobservation, theinnermarginoftheouterstomatalrimwasoftensinuousandseldomsmooth.Strikingly
tal, uprightouterstomatalrimsoccurredinmostspecies.Thecuticularmembraneoftheleafepidermiswassinuous
andoccasionalygranular.LeafepidermalfeaturesinPhymatopterisappearedtobeconstantwithinspecies, andthus
couldbeusedfordistinguishingsomespecies.Themicrocharactersofleafepidermises(thepatternsofanticlinal
walsandthestomatalapparatustypes)couldnotbeusedasevidencefortheclassificationoftheseriesandsub-
seriesinPhymatopteris.
Keywords:Polypodiaceae;Phymatopteris;Leafepidermis;Systematicsignificance
　　PhymatopterisPichi-Serm.(Polypodiaceae)was
anaturalgroup, characterizedbysimple, trifid,
palmatifid, orpinnatifidleaveswithusualycharta-
ceoustexture, glaucousunderneath, single-rowed
butusualysuperficialorslightlyimmersedlargeso-
ri, andthedrynarioidtypeofvenation(Ching,
1964).ThisgenushasbeenconfusedwithMicroso-
rum, CrypsinusandSeligueabyCopeland(1947),
Holtum(1954)andHovenkamp(1998);however
thereareseveralcharacteristicsthatcanbeusedto
植 物 分 类 与 资 源 学 报　 2011, 33 (2):174 ～ 182
PlantDiversityandResources　　　　　　　　　　　　　　　　DOI:10.3724/SP.J.1143.2011.09238
＊
＊＊
Foundationitems:TheNationalNaturalScienceFoundationofChina(30770164, 30970186)
Authorforcorespondence;E-mail:shuganglu@ 163.com
Receiveddate: 2009-11-27, Accepteddate: 2010-06-11
作者简介:邵文 (1979-)女 , 博士研究生 , 主要从事蕨类植物系统分类学研究。
distinguishthesefourgenera(ShaoandLu, 2009).
Phymatopterishasmorethan60 speciesandma-
jorityofthemarefromthemainlandofAsia.Ching
(1964)recognized2seriesand5subserieswithinPhy-
matopterisbasedoncharacteristicsofthefrondshape
andthefrondmargin, i.e.seriesHastatae, including
subseriesGrifithianaeandsubseriesHastatae;and
seriesOxylobae, includingsubseriesOxylobae, sub-
seriesEbenipedes, andsubseriesMalacodontes.
Leafepidermalcharacteristicssuchasdeposi-
tionofwaxcrystaloids, hairtypesandstomatatypes
areofpotentialtaxonomicalimportance.However,
fewstudiesontheleafepidermisofthisgenushave
beendone(Wang, 2007).Noobservationsunder
SEMhavebeenreported.Theaimsofthispaperare
topresentamoredetailedstudyontheleafepider-
malfeaturesofthe24 speciesofPhymatopterisfor
thefirsttime.
1　Materialsandmethods
Adultleavesof24 taxaweregatheredfromthe
specimensofPYU(Table1).Samplesweretaken
fromfulyexpandedsun-exposedleaves.Themateri-
alsforLMobservationwereboiledinwaterbefore
beingmaceratedinJefery′ssolution.Piecesofleaf
epidermiseswerestainedinasolutionofsafraninin
50% alcoholbeforebeingmountedinglyceringel.
Tochecktheconstancyofepidermalstructure, at
leastfiveslidesweremadefromdiferentpartsofa
singleleaforfromdiferentleavesforeachspecies.
ThematerialforSEM observationwasdirectly
mountedonstubswithoutanytreatment, andsputer
coatedwithgold-paladium.Thespecimenswereex-
aminedandphotographedusinganAMARY-1000B
SEM.Descriptiveterminologyofleafepidermisfol-
lowsZhang(1999), Dilcher(1974), andFryns-
ClaesensandvanCothem(1973).
2　Results
ThecharacteristicsofleafepidermisofthePhy-
matopterisarelistedinTables2 and3.Itappears
thatstomatalandotherepidermalfeaturesarecon-
stantwithinspeciesandthusmaybeusedinanalysis
anddiscusion.
Table1　VouchersforleafepidermisofPhymatopteris
Taxon Locality Voucher
Phymatopterisalbopes(C.Chr.etChing)Pichi-Serm. Pingbian, Yunnan, China(云南屏边) W.Shao(邵文)017
P.chrysotricha(C.Chr.)Pichi-Serm. Lushui, Yunnan, China(云南泸水) W.Shao(邵文)003
P.crenatopinnata(C.B.Clarke)Pichi-Serm. Xinping, Yunnan, China(云南新平) W.Shao(邵文)004
P.dactylina(Christ)Pichi-Serm. Deqin, Yunnan, China(云南德钦) W.Shao(邵文)009
P.daweishanensisS.G.Lu Pingbian, Yunnan, China(云南屏边) S.G.Lu(陆树刚)28885
P.ebenipes(Hook.)Pichi-Serm. Lushui, Yunnan, China(云南泸水) W.Shao(邵文)021
P.engleri(Luerss.)Pichi-Serm. YakushimaIsland, Japan(日本屋久岛) S.Mitsuta267
P.glaucopsis(Franch.)Pichi-Serm. Eryuan, Yunnan, China(云南洱源) W.M.Chu(朱维明)23013
P.griffithiana(Hook.)Pichi-Serm. Lushui, Yunnan, China(云南泸水) W.Shao(邵文)001
P.hastata(Thunb.)Pichi-Serm. Pingbian, Yunnan, China(云南屏边) W.Shao(邵文)002
P.majoensis(C.Chr.)Pichi-Serm. Yongshan, Yunnan, China(云南永善) W.M.Chu(朱维明)4898
P.nigrovenia(Christ)Pichi-Serm. Deqin, Yunnan, China(云南德钦) W.M.Chu(朱维明)26458
P.omeiensis(Ching)Pichi-Serm. Daxiangling, Sichuan, China(四川大相岭) X.X.Kong(孔宪需)3971
P.oxyloba(Wal.exKunze)Pichi-Serm. Xinping, Yunnan, China(云南新平) W.Shao(邵文)013
P.pianmaensisW.M.Chu Lushui, Yunnan, China(云南泸水) W.M.Chu(朱维明)11350
P.rhynchophylla(Hook.)Pichi-Serm. Pingbian, Yunnan, China(云南屏边) W.Shao(邵文)029
P.shensiensis(Christ)Pichi-Serm. Kangding, Sichuan, China(四川康定) X.X.Kong(孔宪需)6235
P.stewarti(Bedd.)Pichi-Serm. Lushui, Yunnan, China(云南泸水) W.Shao(邵文)019
P.stracheyi(Ching)Pichi-Serm. Lushui, Yunnan, China(云南泸水) W.Shao(邵文)018
P.subebenipes(Ching)Pichi-Serm. Dali, Yunnan, China(云南大理) W.Shao(邵文)005
P.tenuipes(Ching)Pichi-Serm. Mt.Jinfoshan, Chongqing, China(重庆金佛山) Z.Y.Liu(刘正宇)3547
P.tibetana(Ching)W.M.Chu Deqin, Yunnan, China(云南德钦) W.M.Chu(朱维明)23725
P.trisecta(Baker)Pichi-Serm. Kunming, Yunnan, China(云南昆明) W.Shao(邵文)024
P.yakushimensis(Makino)Pichi-Serm. YakushimaIsland, Japan(日本屋久岛) Muratas.n.
1752期　　　　SHAOWenetal.:ComparativeMorphologyofLeafEpidermisintheFernGenusPhymatopteris　　　　　　
Table2　LeafepidermalcharactersofPhymatopterisunderLM(surfaceview)
Taxa
Adaxialepidermis Abaxialepidermis
Shape
of
cels
Paternof
anticlinal
wals
Shape
of
cels
Paternof
anticlinal
wals
Sizerange
ofstomata
(μm2)
Stom-
atal
index
Stomatotype(%)
Pol Axi Coa Ano Cop
　PlateⅠ
SeriesHastatae
SubseriesGrifithianae
P.grifithiana ir sinuous ir sinuous 48.2×42.3 25.8 46.7 39.4 — 13.9 — 1, 2
P.chrysotricha ir sinuous ir sinuolate 46.1×38.9 27.8 61.6 25.7 — 5.7 7 3, 4
SubseriesHastatae
P.rhynchophyla ir sinuate ir sinuous 43.6×36.7 19.0 45.5 45 — 9.5 — 5, 6
P.dactylina ir sinuous ir sinuolate 46.6×38.9 14.8 25 50.6 — 24.4 — 11, 12
SeriesOxylobae
SubseriesOxylobae
P.oxyloba ir sinuate ir sinuate 47.9×41.7 27.1 48.6 26.9 — 16.5 8 7, 8
P.trisecta ir sinuolate ir sinuous 49.5×45.6 22.4 43.8 41.4 — 9.2 5.4 9, 10
SubseriesEbenipedes
P.albopes ir sinuous ir sinuous 48.7×39.8 24.3 44.6 32.7 — 12.7 10 13, 14
P.ebenipes ir sinuous ir sinuous 48.9×44.6 29.7 32.7 37 5.8 14.8 9.7 15, 16
SubseriesMalacodontes
P.crenatopinnata ir sinuous ir sinuate 52.1×39.9 24.3 50.6 29.4 4 10 6 17, 18
P.stewarti ir sinuolate ir sinuous 48.8×43.4 30.1 39 37 — 34 — 19, 20
irr=iregular;Pol=polocytic;Axi=axilocytic;Coa=coaxillocytic;Ano=anomocytic;Cop=copolocytic
Table3　LeafepidermalcharactersofPhymatopterisunderSEM(surfaceview)
Taxa Cuticularlayer Shapeofguardcels Outerstomatalrims Plates
SeriesHastatae
SubseriesGrifithianae
P.majoensis apophysisandgranules, manyinAb;fewinAd suborbiculate sinuolate Ⅱ:25, 26
P.chrysotricha manygranules(Ad＆Ab), fewapophysis(Ab) suborbiculate sinuolate Ⅱ:27, 28
SubseriesHastatae
P.engleri fewsinuousstripes(Ad), fewapophysisandgranules(Ab) suborbiculate sinuolate Ⅱ:21, 22
P.yakushimensis manysinuolatestripes(Ad), manysinuousstripes(Ab) suborbiculate nearlysmooth Ⅱ:23, 24
P.tenuipes manyapophysisandgranules(Ad＆Ab) suborbiculate sinuolate Ⅱ:29, 30
P.hastata manygranules(Ad＆Ab), fewapophysis(Ab) wideeliptic unknown Ⅱ:31, 32
P.omeiensis fewroundedapophysis(Ada＆Aba) suborbiculate nearlysmooth Ⅱ:33, 34
P.dactylina manygranules(Ad＆Ab), fewapophysis(Ad) suborbiculate unknown Ⅱ:35, 36
SeriesOxylobae
SubseriesOxylobae
P.trisecta manysinuousstripesandmanyhairs(Ad＆Ab) suborbiculate unknown Ⅱ:37, 38
P.oxyloba manysinuousstripes(Ad＆Ab) suborbiculate unknown Ⅱ:39, 40
SubseriesEbenipedes
P.tibetana manysinuousstripes(Ad), manygranules(Ab) suborbiculate unknown Ⅲ:41, 42
P.subebenipes fewsinuousstripes(Ad), manygranules(Ab) suborbiculate unknown Ⅲ:43, 44
P.pianmaensis fewsinuolatestripesandmanyhairs(Ad＆Ab) suborbiculate unknown Ⅲ:45, 46
P.nigrovenia fewsinuousstripesandmanyhairs(Ad＆Ab) suborbiculate sinuolate Ⅲ:47, 48
P.daweishanensis fewsinuolatestripes(Ad＆Ab) wideeliptic sinuolate Ⅲ:49, 50
SubseriesMalacodontes
P.shensiensis fewsinuousstripes(Ad), fewsinuolatestripes(Ab) wideeliptic sinuolate Ⅲ:51, 52
P.stracheyi fewsinuousstripes(Ad＆Ab) wideeliptic unknown Ⅲ:53, 54
P.glaucopsis fewsinuousstripes(Ad＆Ab) suborbiculate sinuolate Ⅲ:55, 56
P.crenatopinnata fewsinuousstripesandhairs(Ad＆Ab) wideeliptic unknown Ⅲ:57, 58
P.stewartii fewsinuousstripes(Ad), manyapophysis(Ab) wideeliptic nearlysmooth Ⅲ:59, 60
Ad=adaxialepidermis;Ab=abaxialepidermis
176　　　　　　　　　　　　 　　　　　植 物 分 类 与 资 源 学 报　　　　　　　 　　　　　　　第 33卷
2.1　CharacteristicsofleafepidermisunderLM
Epidermalcels.Theepidermalcelsofthe
PhymatopterisasseenunderLMareusualyirregular
inshape, withanticlinalwalssinuolate, sinuousor
sinuate.Thepaternsofanticlinicalcelsmayvary
indiferentspeciesorbetweenadaxial(Ad)andaba-
xial(Ab)epidermisesofthesamespecies:sinuo-
latewalsarefoundinP.chrysotrichaandP.trisecta
(Ad＆Ab, PlateⅠ :4, 9), sinuouswalsinP.
grifithianaandP.albopes(Ad＆Ab, PlateⅠ:1,
2, 13, 14), andsinuatewalsinP.oxylobaandP.
crenatopinnata(PlateⅠ :7, 8, 18).
Stomatalapparatus.Alspeciesstudiedhere
havethestomatalapparatusrestrictedinabaxialepi-
dermises.Theshapeofthestomataisusualysubor-
biculate.Thestomatacanbepolocytic, copolocytic,
axilocytic, anomocyticorlessoftencoaxilocytic.
However, thesetypesofstomataoccurofteninasin-
gleleaf.
Thesizerangeofstomatais(43.6-52.1)×
(36.7-45.6)μm2.Thestomataldensityisdif-
fe-
rentamongspecies.Thestomatalindexrangedfrom
30.1%inP.stewarti, withthestomatabeingdense-
lydistributed, to14.8% inP.dactylina, withthe
stomatabeingsparselydistributed.
2.2　Characteristicsofleafepidermisunder
SEM
Guardcels.Theoutlinesofthepairofguard
celsofthePhymatopterisarevariable, fromusualy
suborbiculatetowideelipticalasseeninthesurface
view, withlength/width(L/W)ratio1.1 -1.3∶
1.Guardcelsareoftenthickenedtoacertainex-
tent, aremadeupofouterstomatalledgesorrims,
andarenearlysmoothorsinuolate.Nearlysmooth
outerstomatalrimsoccurinP.stewartiandP.
omeiensis(PlateⅡ:34, PlateⅢ:60), andsinuo-
lateouterstomatalrimsarepresentinP.glaucopsis
andP.nigrovenia(PlateⅢ:48, 56).
Strikinglytalanduprightouterstomatalrims
occurinalspecies, exceptforP.omeiensis, P.tri-
secta, andP.glaucopsis(PlateⅡ:34, 38, Plate
Ⅲ:56), inwhichthestomatalrimsarealmostat
thesamelevelwiththeepidermis.Furthermore, the
cuticularthickeningofthecommonwalbetweenthe
guardcelsisfoundinsomespecies, suchasP.
chrysotrichaandP.dactylina(PlateⅡ:28, 36).
Cuticularandwaxornamentation.Stripes,
hairs, andwaxgranulesarepresentinbothsidesof
epidermisesofthespecimensinvestigated.Two
typesofthecuticularstripesarefound:sinuolate
andsinuous.Theoutersurfaceofthecuticleis
smoothtosinuolateinP.pianmaensisandP.daweis-
hanensis(PlateⅢ:45, 46, 49, 50);whilethe
sinuousstripesarefoundinP.stracheyi(PlateⅢ:
53, 54).Waxgranules(d≈20μm)arepresentin
P.dactylinaandP.tibetana(PlateⅡ:35, 36,
PlateⅢ:41, 42).Afewhairsaredistributedon
theabaxialepidermisofP.crenatopinnataandthe
adaxialepidermisofP.nigrovenia(PlateⅢ:47),
alsoalsobothsidesofepidermisesofP.trisecta.
2.3　KeytothePhymatopterisspeciesstudiedwithspecialreferencetofoliarmorphologicalcharacters
1a.Frondssimple, trilobedorpalmatelydivided 2⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
1b.Frondspinnateorpinnatifid 11⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
2a.Frondssimple 3⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
2b.Frondstrilobedorpalmatelydivided 10⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
3a.Frondsvenationissimple, costalareoleshavenoforkedveinlets, andtheexcurrentincludedveinletsdominateinmost
areoles 4⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
3b.Frondsvenationisvariable, veinletsareoftenforked, rarelysimpleoncostalareoles, andtherecurrentincludedveinlets
dominateinmostareoles 6⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
4a.Frondsdimorphic, thesterilemuchshorter, ovate, obtuseatapex;thefertilemuchlonger, lanceolate
P.rhynchophyla⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
4b.Frondssubdimorphic, sterilefrondsrarelypresent, fertilepartssimilartosterileparts 5⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
5a.Rhizomefewpruinose, frondmembranaceous, obtuseatapex, roundedatbase P.tenuipes⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
1772期　　　　SHAOWenetal.:ComparativeMorphologyofLeafEpidermisintheFernGenusPhymatopteris　　　　　　
5b.Rhizomepruinose, frondchartaceous, acuteatapex, cordateatbase P.chrysotricha⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
6a.Frondsdimorphic, thesterilemuchshorter, ovate, obtuseatapex;thefertilemuchlonger, lanceolate P.griffithiana⋯
6b.Frondsmonomorphic, withnodiferentiationoffertileandsterilefronds 7⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
7a.Manywaxgranulesonabaxialandadaxialepidermis P.majoensis⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
7b.Fewornogranulesonabaxialoradaxialepidermis 8⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
8a.Obtuseatapex, fewroundedapophysisonbothsidesofepidermis P.omeiensis⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
8b.Acuminateatapex, noroundedapophysisonabaxialoradaxialepidermis 9⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
9a.Laminaoblanceolate, outerstomatalrimssinuolate P.engleri⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
9b.Laminalanceolate, outerstomatalrimsnearlysmooth P.yakushimensis⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
10a.Frondspalmatelydivided P.dactylina⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
10b.Frondstrilobed, manywaxgranulesonbothsidesofepidermis P.hastata⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
11a.Frondstrilobedtopinnatifid 12⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
11b.Frondspinnatifidtopinnate 13⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
12a.Anticlinalwallsofepidermissinuolateorsinuous, morehairsonbothsidesofepidermis P.trisecta⋯⋯⋯⋯⋯⋯⋯⋯
12b.Anticlinalwalsofepidermisoffrondssinuate, fewhairsonbothsidesofepidermis P.oxyloba⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
13a.Notchesusuallypresent, withashalowlyserrateateachpairofvein 14⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
13b.Notchesusualyabsent, withbiserrateateachpairofvein 20⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
14a.Manywaxgranulesonabaxialandadaxialepidermis 15⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
14b.Fewornowaxgranulesonabaxialoradaxialepidermis 17⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
15a.Laterallobesallascending P.tibetana⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
15b.Lowestlobesdeflexedorslightlydeflexed 16⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
16a.Laminaobtuseatapex, pinnae3-5(7)pairs P.subebenipes⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
16b.Laminaacuminateatapex, pinnae4-10(13)pairs P.ebenipes⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
17a.Manyhairsonadaxialepidermis 18⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
17b.Fewornohairsonabaxialoradaxialepidermis 19⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
18a.Fewsinuolatestripesonbothsidesofepidermis, morehairsonbothsidesofepidermis P.pianmaensis⋯⋯⋯⋯⋯⋯⋯
18b.Fewsinuousstripesonbothsidesofepidermis, morehairsonadaxialepidermis P.nigrovenia⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
19a.Lowestlobesascending P.daweishanensis⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
19b.Lowestlobesdeflexedorslightlydeflexed P.albopes⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
20a.Laterallobesacuminateorcaudatelyacuminateatapex 21⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
20b.Laterallobesobtuseoracuteatapex 22⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
21a.Lowestlobesdeflexed, fewapophysisonabaxialepidermis P.stracheyi⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
21b.Lowestlobesascending, manyapophysisonabaxialepidermis P.stewartii⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
22a.Fewsinuolatestripesonabaxialepidermis, pinnae2-3(4)pairs P.shensiensis⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
22b.Fewsinuousstripesonabaxialepidermis, pinnae2-5 (7)pairs 23⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
23a.Lobescrenateorundulateatmargin, shapeofguardcelswideelliptic P.crenatopinnata⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
23b.Lobesdiserrateatmargin, shapeofguardcelssuborbiculate P.glaucopsis⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯⋯
3　Discussion
TheleafepidermalcelsintheferngenusPhy-
matopterisareusualyiregularinshape, andthesto-
matalapparatusarerestrictedinabaxialepidermises.
Themainstomatalapparatustypesinalspeciesexami-
nedinthispaperarepolocyticandaxilocytic.The
cuticularmembraneoftheleafepidermisisgeneraly
sinuolatetosinuous.Theleafepidermalfeaturesa-
mongspeciesarefairlysimilartoeachother, which
indicatetheafinityofPhymatopteris(Sen, 1981).
Polocyticandaxilocyticstomatalapparatus
typesareindicatedasmoreprimitivetypes(Pant,
1965), butasmoreevolutionaltypesinPolypodi-
aceae(Thurston, 1969).Furthermore, Wang(2007)
acceptedThurston′spoint, andfoundtheevidence
inPhymatopteriscrenatopinnata, whichalsohas
beenfoundinthispaper(PlateⅠ :18).
Thepaternsofanticlinalwalsandthestomata
178　　　　　　　　　　　　 　　　　　植 物 分 类 与 资 源 学 报　　　　　　　 　　　　　　　第 33卷
typesareconstantwithinspecies, andthuscanbe
usedtodistinguishcloselyrelatedspecies.Wealso
findthesetwocharacteristicsdistributedinalsub-
seriesrandomly, andthuscannotbeusedasevi-
dencefortheclasificationoftheseriesandsub-
seriesinthisgenus.
Thecuticularandwaxornamentation(stripes,
hairs, andwaxgranules)arepresentinsomeepi-
dermisesofthespecimensinvestigated.Waxgran-
ulesareinP.dactylina, P.subebenipesandP.tibet-
ana, hairsinP.crenatopinnata, P.nigroveniaand
P.trisecta, andsmalgranulesinP.tenuipes, P.
hastate, P.majoensis.Thoughitmaybeafectedby
thelivingenvironmentinsomedegree, thewaxand
hairappendagesareconstantwithinspeciesandthus
providesomeanatomicalevidencefortheclassifica-
tionofsomespecies.
Theoriginalfunctionofstomataisthelimitation
ofwaterlossinlandplantsandthemaintenanceof
homoiohydry, whilealowinggasexchange, which
placesconstraintsonstomatalpaterningwithinthe
epidermises.Theguardcelsofstomatalapparatus
inthispaperarealslightlysunken, andmoststoma
poresareclosed, whichmaybesuitabletotheirepi-
phytichabitus.
Ourresultssuggestthatthecharacteristicsofthe
leaves, i.e.theshapeoffrond, themarginandap-
pendagecharacteroffrondareconstantwithinspecies
andthuscanprovidesomeanatomicalevidenceforthe
clasificationofPhymatopterisinsomespecies.How-
ever, themicrocharactersofleafepidermises, suchas
thepaternsofanticlinalwalsandthestomatalap-
paratustypescannotserveasacriterionofdistin-
guishingtheseriesandsubseriesinPhymatopteris.
Acknowledgements:WewouldliketothankProf.W.M.
Chuforprovidingaccesstothespecimenscitedinthispaper,
andProf.R.X.WangandengineerX.K.Fanforassistance
ontheobservationofLMandSEM.Thefirstauthorwouldal-
sothankProf.JerryLiandProf.DamianSheaofNorthCaro-
linaStateUniversityforhelpfulsuggestionsandcriticalread-
ingofthemanuscript.
Reference:
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CrypsinusPresl.[ J] .ActaPhytotaxonomicaSinica(植物分类
学报), 9(2):194
CopelandEB, 1947.GeneraFilicum [ M] .Waltham:ChronicaBo-
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DilcherDL, 1974.Approachestotheidentificationofangiospermleaf
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Fryns-ClaessensE, vanCothemWRJ, 1973.Anewclassificationof
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structurespit[ J] .ScienceSeriesAlahabad, 1:1— 24
SenU, HennipmanE, 1981.Structureandontogenyofstomatain
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ExplanationofPlates
PlateⅠ 　1-20.ThecharactersofleafepidermisofPhymatopteris
underLM.　 1 -2.Phymatopterisgrifithiana, 3 -4.P.chry-
sotricha, 5-6.P.rhynchophyla, 7-8.P.oxyloba, 9-10.P.tri-
secta, 11-12.P.dactylina, 13-14.P.albopes, 15-16.P.ebeni-
pes, 17-18.P.crenatopinnata, 19-20.P.stewarti.
PlateⅡ　21-40.ThecharactersofleafepidermisofPhymatopteris
underSEM.　 21-22.Phymatopterisengleri, 23-24.P.yakushi-
mensis, 25-26.P.majoensis, 27-28.P.chrysotricha, 29-30.P.
tenuipes, 31-32.P.hastata, 33-34.P.omeiensis, 35 -36.P.
dactylina, 37-38.P.trisecta, 39-40.P.oxyloba.
PlateⅢ 　 41-60.ThecharactersofleafepidermisofPhymatopteris
underSEM.　 41-42.Phymatopteristibetana, 43-44.P.subebe-
nipes, 45-46.P.pianmaensis, 47-48.P.nigrovenia, 49-50.P.
daweishanensis, 51-52.P.shensiensis, 53-54.P.stracheyi, 55-
56.P.glaucopsis, 57-58.P.crenatopinnata, 59-60.P.stewarti.
1792期　　　　SHAOWenetal.:ComparativeMorphologyofLeafEpidermisintheFernGenusPhymatopteris　　　　　　
　邵　文等:图版Ⅰ SHAOWenetal.:PlateⅠ 　
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　邵　文等:图版Ⅱ SHAOWenetal.:PlateⅡ　
1812期　　　　SHAOWenetal.:ComparativeMorphologyofLeafEpidermisintheFernGenusPhymatopteris　　　　　　
　邵　文等:图版Ⅲ SHAOWenetal.:PlateⅢ　
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