Spore Morphology of Cyatheaceae in China

2007·2.2 MB·English

by  Wang, Zi-Juan

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ISSN 1346-7565 ActaPhytotax.Geobot.58(2136)9-:77 (2007)

SporeMorphologyofCyatheaceaeinChina

ZI-JUAN WANG XIAO CHENG
and
Kitnming institu tqef'Botap cC}Zleines Aecacienu 7ofSbienc 1e(s),inmin 6g50204,}itnnan ,enina
Scanning light in
electron microscopy and microscopy weTe used a palynological study of eight
species of Cyatheacea ein China :SPhaempte rbriusnoniana, Alsophila spinulos aA,. latebrosa A,. cos-
tularis ,A. denticulat aA,. gigantea ,A. austmpunnanensis and A. khasyana .The firs tthree species of
Alsophil aare in subgerrusAlsophila and the las ftou rare in subgerms Gymnosphae Trhae, spores ofnear-
ly al1 species ofAlsophita examined are characterized by a ridged or modified ridged perin eand a gran-
ular or verrucate exine, except the spores ofA. dentieulat aand A. austroyunnanensis, which have a
smooth exine, Specie swith a granula orr verrucate exine may be advanced in comparison with the species
with a smooth exine, implying that the species of Cyatheaceae in China are advanced. The speres of
Eiphaempteri sbrunoniana featur aen incipien gtranula routermost layer and a yerrucate exine. Species
ofAlsophila subgenus (lymnosphaera resemble species of subgenus Alsophil ain hayin ga granula ror
verrucate exine and a ridged perine s,upporting the placement of Gymnosphae writahin Alsophil arather
than recognizing it as a separate genus, Because spores in the species ofsubgenus Clymnosphaera are
more divers eand have a rnore cornplicated perine ,it is possibl etha tthis group is adranced when cem-
pared with other species ofAlsophita.
Key words: Alsophila, China, Cyatheaceae ,(l)smnosphaera, SPhaempte srpories ,morphology
Cyatheacea eKanlf are a pantropic aflamily of two genera ,E2Jhaemp tandeAlrsoiphsila, The genus
approximate 500-600 species with tree-lik etrunks, EPhaempte riis dsistinguish efdrom other Cyathe-
Since the family was established in 1827, consid- aceae by it sunifbrrn stipe scales. It has a pantropi-
erable attention has been paid to it smorphology, cal distributio bnu,t is absent from Africa and
anatomy, taxonomy and systematic position Madagascar (Larg &e Braggins 2004). Holttum &
(Holttu1m963,1965,HolttLmi& Edwards 1983, Edwards(1983) thaEtPhaempterisin
assumed arose
Holttum & Sen 1961, Tryon 1970, Tryon & Tryon Southeas tAsia, it scenter ofdistribution. There are
1982, Lellinger 1987, Lucansky 1974a, b, Lucansky two species of EiPhaeropteri sin China; S. Iepijlara
& White 1974, Conallt et al, 1994, 1995) ,Still, and S. brttnonian aA.lsophil ahas a wide distribu-
however, there is no consensus on it sdelimitation, tion, with its center in the Malesian region. Tryon
subdivisions and phylogery Holttum & Sen (1961),(1970 )C,hing (1978 )T,ryon and Tryon (1982),
Tryon (1970 )and Lellinge r(1987 d)ivided the Lellinge r(1987 a)nd Xia (1989 r)ecognized the
Cyatheaceae s. str, into one, six and fbur genera, genus, while Holttum & Edwards (1983 t)reated it
respectively. Ching (1978 c)lassified the 14 species as a section in byathe saubgenus Cyathe Taw.elve
and two varieties of Cyatheaceae in China into species and two varieties ofAlsophila are in China,
three genera; Sphaeropter iAsls,ophila and (:lymnosphaera is a controversial group. Copeland
(lymnosphaera ,while Xia (1989 p)laced them in (1947 an)d Ching (1978 re)cognized it at the gener-
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ic level w,hi]e Holttum (196 3t)reate dit as a section photographed with a KYKYLAMRAY 1000B
of subgenus dyathe aan,d Xia (1989 a)s a sub- EIectro nMicroscope at the Electro nMicroscopy
genus ofAlsophila. Although based on the same Centeg KIB, 1[Ih eSEM was operated at 25 KV In al1
series ofmorphological and anatomical characters, species in which the exine appears to be concealed
those classification systems are still controversial by a perine layer during any stage of spore devel-
due to varied taxonomic opinions. The classification opment, the spores were boiled sufficiently long
we use is according to Xia's two genus system, (examin eidntermitten tblyy ligh tmicroscope) in
In order to revalue the relationship ef 1 N hot sodium hydroxid eto remove the perine, as
{IPhaempter iAsl,sophila aiid Glymnosphaer ar,esort- describe bdy Gastony (1974 T)h.ese alkali-prepared
ing to the scanning electron microscopy (SEM) spores were then washed with distill weadter, sus-
and ligh tmicroscopy, we observed spore morphol- pended in a few drops of70% ethanel, and piped to
ogy of eight species of Cyatheace aien China, of doubl esided adhesive tape attached to stubs fbr
which one in the genus EPhaeropte Sr. ibsru;noni- SEM study, Micrographs of the proximal andror
ana, others in the genus Alsophila A;. spinulosa ,A. dista flace sand ef some omamentation detail swere
Iatebros aA,. costularis, A, denticulat Aa., gigantea, photographed to compare spore morphology at each
A, austroyunnannesis and A. khasyana (Thb l1)e. orlentatlon.
The las tfbur species ofAlsophila belon gto the
Gymnosphaera. Results
subgenus
Gastony (1974 1,979) and Gastony & Tryon
(197 6ha)ve done a let of researches on the spore The spores ofal1 species of Cyatheacea einves-
morphology of Cyatheaceae, but they seldom tigate dwere trilete r,adially symmetrical, heteropolar
focused on the Asian species of Cyatheaceae. and perinous (Fig s1.-24) .The polar outline is tri-
However, the percentage ofendemism ofthis fam- angular, usually with concave sides and rounded
ily in Asia is very high, fbr example, the species angles. In equatorial viewl the proximal face appears
endemism of the fatnil ayttains 90% in Southeast as a low pyramid, while the dista lfac eis hemi-
Asia (exc lMa.lay Peninsula )(Tryo &n Gastony spherical to convex,
1975) .So it is necessary to add new spore mor- Of the two species ofE??haeropteris in China,
phology data from species ofthis region. only S. brunoniana was accessible fbr SEM obser-
vation, The spores of S. brunoniana are perinous
Materials and Methods with a verrucate exine (Fi g1.) ,Figure 2 shows an
densely
alkali-untreated spore with a granular out-
The sources of the samples used in thi sstudy ermost layer,
are liste idn [[abl e1. All were obtained from wild The spores ofAlsqphila spinulosa featur ea
plants in August, 2005, exceptAlsqphiia spinulosa ridged perin eand a verrucate exine (Fig s3-,4) ,The
was from a livin gplant transplanted to the fern laesura eare wide, slightly elevated, are distinct
garden ofthe Kunming Institu otfBeotanM Chinese under the coarse perin eand each one is grooved
Academy ofSciences (KIB )andA. denticula twaas (Fi g3,). The ridge smore or les sconnect with each
from a herbarium specimen at KUN. Vbucher spec- other, Seme granules are scattered on the surface of
imens are deposit eidn KUN. the perine .Figur e5 shows a spore after･being treat-
The spore samples were mounted on aluminum ed with sodium hydroxide to demonstrat ethe dif
stubs with double sided adhesive tape, coated with ferenti aexlpansion and perinous nature ofthe out-
a thin laye rofgold-palladium, then examined and ermost layer,
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TABLE 1. Specimens ofCyatheaceae examined by scanning electron microscopy and ligh tmicroscopy and the results.
Vbucherin
Taxon Localit yand altitude KUN Perine surfaceExinesurface Figureno.
ophaenzpterisbrunonXiiaaonlaa highway ,Mengla, Xl Cheng iOI pointed-projections verrucate 1-2
(Hook.R).M,Tryon Xishuangbanna,1050m

Alsophila spinulosa (Wall.ftansplanted plant in KIB, 192gm Z eneng103 ridges verrucate 3-5
ex Hook,) R. M. TryDn
A, latebro sWaall. ex Daweishan, HeKou, 900m X Cheng i06 ridges granulaT 6-8
Hook.
A. costularis Baker Adeboxiang, Jinping ,1500m X eneng102 ridges verrucate 9-10
A, denticutat aBaker Caoguoshan, Xichou, 1520m IVb ,OO06272 ridgeswithrods smoeth 11-14
and echinates
A, gtgunt Welaa11 e,x Hook. Xiaela highway, Mengla, X eneng100 fused strands granuEar 15-18
Xishuangbanna, 1050m

A, attsmoyunnanensis S. G. Hekou 3 km point way to Pingbian, Z Cheng 108 ridges with smooth 19-21
LuA. 1460mHekeu echinates
nttasyan aT. Moore ex 3 km point way to Pingbian,X eneng109 ridges verrucate 22-24
Kuhn 1460rn
The spores ofAlsophila latebros aare charac- and on the rest of the surface of the perine ,The
terized by a typical ridged perin e(Fig s6-,7) and a laesura aere concealed by the perin eand could not
granula rexine (Fi g8). .lhe Iaesura aere covered by be observed. Figur e13 shows the exine ofaspore of
a perine and are obscure, The elongated and A. denticula tparepare din sodium hydroxide ,The
branched ridges are fewer on the proximal surface exine is nearly smooth, except for a few scattered
than on the dista slurface. Granule sare scattered on protuberances ,and the laesura eare distort esldight-
both the ridges and on the surface ofthe perine. ly inward.
The spores ofAlsophila costularis also share Alsophil agigantea represents a differe nktind
the typical ridged perine (Fi g9.), but the ridges of spore morphology (Fig s15,-18) D.ense woven or
are shorter and lower than those on the spores ofA, fused strands constitute the perin esurface and even
latebros aS,ome granule asre scattered on the perine the laesur raeegions and show no differe bnecteween
surface. The exine is verrucate, even on the laesurae the proxirn aalnd dista s1ur faces N,arrow laesura aere
regions (Fi g1,0), conspicuous. The exine also has granules (Fi g1.8)
Alsqphil adenticulat aA,, gigantea, A. aus- 1ik ethose ofA. Iatebros a(Fi g8,),
trtryunnanensis and A, khasyana are here placed in Alsophila austToyunnanensis, endemic to
subgerllis Clymnosphaera T.he spores ofA. ctentic- China, is characterized by it sdimorphic fronds.
ulata also have a ridged perine ,with the ridges Because of it sechinate stipes, this species was
somewhat connected to each other (Fig 1s 1.-12) b,ut include idn subgenus ALsophila (Lu 1998), Based on
ther eare some rod-shaped appendages (Fi g1.1) or melecular data ,A, austroyunnanensis is placed in
echinate or pointed projection (sFig .12) on the subgenus Gymnosphae r(aL iet al. 2004, Lu & Li
ridges, Some granules are scattered on the ridges 2005) .The perine of it sspores has short ridges,
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which are nearly absent in the laesura eregions perine depositio onccurring lat ein sporogenesis,
(Fi g1.9). The laesura eare wide and grooved .The Cao et al. (20e p7. ,I2, Figs. 21-24) reported tha tS.
ridges are somewhat connected with each other and branonian aspores develo ap short echinulate perine.
have (Fig2.0).Figure21 The be
echinate prejection s shows spores with shortly echinulate perine may
a spore with a nearly smooth exine, Perhaps the corrrparable to those with spaced pointe dprojections,
spore was treate dinsuMcient lwyith alkali; there are as Gastony & Tryon (197 6de)scribe idn the perine
some low verrucae and perin eremaining on the of all paleotropic asplecies of Eiphaeropte rainsd
surface ofthe exine. This patter ndiffe rfsrom that in for the neotropical S. horridn group. ophaempteris
the The brunoniana developed
species ofAlsophila with granular exine. possibly a perine with point-
spores ofA. aust7zo,unnanensis, howeve4 share the ed projection bsu,t additional infbrrnatio non spore
smooth exine type as describ beyd Gastony & Tryon wall development and a molecular phylogenetic
(1976 f)or the spores of all neotropical and most analysis is needed to funher clarify the systematic
paleotropical species ofALsophiia, The echinate positio onfthis species.
projections on perine ridges, however, recall some Most spores ofthe species ofAlsophila exam-
paleotropic saplecies ofEk)haempteris with similar ined feature a ridged or modified ridged perin eand
perine morphology (Gasto n&y Tryon 1976). a granular or vermcate exine, except that the spores
The spores ofA. khasyana are also very special ofA, clenticulata and A. austroyunnanensis develop
among the species of Cyatheaceae in China. The a type of smooth exine. The exine surface of the
spores represent a coarse perin ewith short, irregu- spores ofA. Iatebro asnad A. gtgant beeaar granules,
larly deposited ridges (Fig s22.-23). The exine is The spores ofA. spinulosa ,A. costularis and A,
verTucate CFi g2.4). The wide laesura aere also obvi- khasyana with verrucate exine recall the spore type
ous. of S. brunoniana. The spores of subgenus
(lymnosphaera resemble those of other species of
Discussion Alsqphii ain havin gridged perine, suggesting that
(lymnosphaera is closet to Alsophila than to
Gastorr &y Tryon (197 r6ep)orted the peime of EPhaeFtl pbtaseerd ions spore morphology. At this
the spores of the paleotropical species of point ,it supports the traditional classification that
EPhaeropt etor hiasve pointe dprojection asn,d usu- Glymnosphaer ais attached to Alsophil a(Holttum
ally The 1963,Tryon& Tryon1982,Xia 1989).Apartfrom
smooth exine without sculpturing. spores of
the neotropical species, excluding those in the S. this, the species of (lymnosphaera have diverse
horric ignroup ,haye hairLli okre semetimes incipientspore characters. Their perine morphologies are
(S.
granula r elongata) perine and pitte dor verrucate more complicated than those of other species of
exine, They also reported the spores ofAlsophila Alsophil aa,nd some perine appendages are similar
species to be characterized by a typical type of to those of many paleotropical species of
ridged perine and smooth exine. In the present fik)haero p(tGearsitso n&y Tryon 1976), implying
study, the species were not all the same, that (lymnosphaera is either an adyanced group
Ek)haerop tberruinosniana had a granular outer- derive dfrom the Alsophil agroup, with spores char-
most layer and a verrucate exine. Gastony & acterized by the simpler typic raidlged peime, or is
Tryon (197 6re)ported the presence of granules an interrnedi agtreoup between 52?haeizr pantderis
external to the exine in certain collection of some other species ofAlsophila in spore evolution and
species of E2)haerqpt seucrhi sas S. elongata (p. conservation ofthe rod-1ike and pointe dappendages
744, Fig ,38 and 39) to be an incipien sttage in on the ridges of the perine.
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FIGs.1-8 .Perine and exine morphology characteristics, - 1-2 ,Eiphaer(lp tberrtitsnonia n1a. D;ista f1ac eof epore safter treate din sedi-
um hydroxide showing verrucate exine; 2. Dista lface of untreated spores showing incipien gtranula rperine dispesiti oonver tlie
exine. rm 3-5. Alsophila spinulosa :3. Proximal face showing coarse perine surface; 4. Dista lface showing verrucate exine ef
a spore after chemical removal Qfperine; 5. Light micrograph of spere after preparatio nwith sodium hydroxide to demonstrate
-
differenti eaxlpansion and perinous nature ofthe outermost layer . 6-8. A. Iatebrosa :6. Proximal face; 7, Dista 1face show-
ing perine with ridges and grarrule s8;. Dista lfac eshowing granulate dexine befor eperine disposition,
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FIGs. 9-16. Perine and exine morphology characteristics. 9-1O. A. costularis: 9. Dista lface showing ridged perine ;1O, Proximal
r
face showing verrucate exine after chemical removal ofperine, 11-14 A.. clentic"lata: 11, Proximal face showing rod
appendages on perine ridges ;12. Distal face showing echmate prejection son perine ridges ;13, Preximal face showing
smooth exine after chemical rernoval ofperine; 14. Light micrograph of spore after alkali-treatment to demonstra tdeiffereritial
-
e)rpansien andperinous nature ofthe outermest layet 15-16 .A. gignntea :15, Proximal face showing perine ;16. Dista 1face
showingperine.
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FIGs.17-24. Peime and exine morphology characteristics. 17-18 A.. gr'gante a1:7 ,High rnagnification ofperine surface; 18, Lateral
side showing granulat eexidne. - 19-21 ,A. austroyunnanensis: 19. Proximal face showing fewer ridges ofperine on th elaesurae
regions; 20. Dista lface showing ridge pderine with echinate projection 2s1;. Dista flac eshowing nearly smooth exine after chem-
-
ica lremoval ofperine, 22-24. A. khasyana: 22. ?roximal face shffwing perine ;23. Dista 1face showing perine ;24. The round
angle showing vcrmcate exine,
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As to the inter-Antra-gen erreliactionship in We would like to thank Ms, Yir Jiao ,Sir YU Chen and
Cyatheaceae, there are several differe notpinions, Shae-Wen YU of KIB fbr their help in cellecting spores.
Holttum and his colleagues suggested that the The othcers of KUN have been generous in making
Alsophila dyathea based rnaterials available for our study, We also thanlc Xi-Kai
and clades are primitive ,
on feature sof the sori and indusia, and the Fan efKIB fbr his advice in the SEM study.
EPhaempteris indusiumisderived
elade without an References
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