全 文 ：第 23 卷　第 3 期　　　　　　　　　　　　　植　　　物　　　研　　　究 2003年　7 月
Vol.23　No.3　　　　　　　　　　　BULLETIN OF BOTANICAL RESEARCH July , 　2003
中国藓类植物孢子类型的初步研究
于　晶1 , 2　王全喜1　曹　同1
(1.上海师范大学生物系 ,上海　200234)
(2.复旦大学生物多样性科学研究所 ,上海　200433)
摘　要　在对中国藓类植物孢子形态结构的系统观察基础上 ,从孢子壁层结构及孢子形态角度将
中国藓类孢子类型初步划分为 9种类型 。每种类型以代表属名命名 ,对每种类型的大小 、颜色 、形
状 、极性 、萌发孔及孢壁结构进行了描述 。孢子的大小 、颜色和形状有时会随环境条件而改变 ,而
极性 、萌发孔及孢壁结构很稳定 ,在类型划分中更重要 。
关键词　藓类植物;孢子;中国
PRELIMINARY STUDYOF SPORE TYPESOF CHINESE MOSSES
YU Jing1 , 2　WANG Quan-Xi1　CAO Tong1
(1.College of Life and Environment Sciences , Shanghai Normal University , Shanghai　200234)
(2.I nstitute of Biodiversity Science, Fudan University , Shanghai　200433)
Abstract　The spores of mosses f rom China were preliminarily classified into 9 types based on sys-
tematic observation of the constitution of sporoderm and spore morphology.Each type is named af ter
the representative genus and described in spore size , colo r , shape , polarity , aperture and sporoderm
structure , the last three characters are more useful in classification.
Key words　mosses;spo res;China
1　Introduction
Morphology and ult rast ructure of moss spores
are stable and signif icant not only in theory but also
in practice , especially in taxonomy and phylogeny of
bryophy tes.Rich terms of spore mo rphology have
been accumulated by some experts , such as Erdt-
man
[ 1～ 3]
and Miyoshi
[ 4] , whose description mostly
came from the observat ion under light microscope.
Clarke[ 5] put fo rw ard that the ornamentation pat-
terns of many spores are i rregular and a verbal sys-
tem of terminology is unlikely to be adequate to de-
scribe them.Therefore , photog raphs are essential in
such cases.Punt et al.[ 6] summarized glossaries of
pollen and spore. Gupta[ 7] divided spo res of
bryophy te into different types f rom dif ferent charac-
ters including shape , cell numbers , polarity and
aperture respectively , with appended ideographs.
Till now , term s and types of bryophyte spores are
mostly named by superficial mo rphology w ithout
pho tographs.Sometimes w e feel puzzled w hen w e
used these terms.For example , most of moss spores
are g ranulate that can not be dif ferent iated by super-
ficial morphology (Plate I:7;Plate III:7), but un-
基金项目:上海市青年科技启明星项目(02QA14034),上海市高等学校教师青年基金(02DQ35)第一作者简介:于晶(1972—),女 ,博士 ,讲师 ,主要从事植物学 、生态学方面的研究 。收稿日期:2002-10-15
der t ransmission microscope , one is composed of
perispore (Plate II I:4)and another is const ituted by
exospore (Plate I:7).Similar condition can be
found in echinate o rnamentation (Plate II:1 , 3;
Plate II:7;Pl.II:9).So how to classify the type of
spores is still a question that needs clarification.
We have systematically observed spo res of
mosses(57 families , 178 genera , 388 species)from
China under both scanning electron microscope
(SEM ) and transmission elect ron microscope
(TEM)since 1990s .Based on the w orks of prede-
cessors and present study , we felt that it is insuf fi-
cient to classify the spo re type only by superficial
morphology .The sy stem of classification needs inte-
g rated characters , especially the sporoderm struc-
ture.Ornamentation constitution of spores is differ-
ent too.It is the perispore that makes up of orna-
mentation mostly , but exospore or exospo re together
w ith perispo re constitutes ornamentation in few
types and endospore seldom joins in the consti tut ion.
In our opinion , resumptively classify the spore
type based on the constitution of sporoderm in the
first place except spores w ith multi-cells , and then
classify in details based on mo rphology and superfi-
cial st ructure of spo res.Thus , the spores w ith simi-
lar morphology but wi th dif ferent sporoderm consti-
tution can be dif ferentiated firstly and it is easy to
classify further.In the present paper , 9 types were
primarily put forw ard and detailed classification w ill
be presented in another paper.
2　Material and methods
Spores used in this study were obtained from
mature sporophy tes of field collections and dried
herbarium specimens f rom IFSBH.The spo res to be
used for SEM were mounted on double sided adhe-
sive tape attached to metal stubs , sput ter coated
w ith gold and viewed w ith a H ITACHI-520 SEM at
an accelerating voltage of 15 to 25 kV.Fo r TEM ,
the material w as fixed in 4% buf fered glutaralde-
hyde fo r 2 hours , rinsed several times in 0.1M Na-
cacody late buf fer , and post fixed in 2%OsO4.After
dehydration to 100% ethanol , the material w as in-
f iltrated in spurr s mix ture and polymerized.Ultra-
thin sections w ere prepared using a diamond knife ,
post-stained wi th uranyl acetate and lead cit rate ,
and examined in a HITACHI-600 TEM operated at
80 kV.
3　Results and discussion
Based on spore mo rphology and st ructure , es-
pecially structure such as polarity , aperture and or-
namentation constitution , the spores of Chinese
mosses studied were classified into 9 types and each
w as named af ter the representative genus.The for-
mer 8 types have obvious characters to distinguish
f rom each o ther.Spores of the last one appear as
having dif ferent superficial mo rphology , but these
spores have similar const itution of sporoderm.The
ornamentation of different morphology is listed sim-
ply .The detailed info rmation w ill be provided in an-
o ther report.Each type is described from spore size ,
colo r , shape , polarity , aperture and ornamentation
st ructure.The representative pictures of SEM and
TEM were presented here wi th.The key to differ-
ent spore types w as also provided.We hope that our
preliminary division of spo re types of Chinese mosses
will provide useful info rmation for people to bet ter
understand spores of mosses.
3.1　Sphagnoid type(Plate I:1 ～ 3)
　　Size:20 ～ 50 μm in diameter.
　　Color:yellow to dark brow n.
　　Shape:tet rahedral.
　　Polarity:heteropolar and radiosymmetric.
　　Aperture:trilete , having a distinct t rilete mark
on the pro ximal face;trilete laesurae surrounded by
a wel-developed margo.
　　Ornamentation st ructure:sporoderm thick and
perispore thinner than exospore , smooth exospore
forming basic contour of spore such as having a dis-
tinctive w all st ructure of large , rounded , triradiate
ridges w ith or without bifurcate arms on distal face
and perispore varying f rom g ranulate to coarsely pa-
pillose o r rounded-verrucate.
　　Only spores of Sphagnaceae belong to it.
3.2　Hedw igioid type(Plate I:4 , 5)
　　Size:15 ～ 34 μm in diameter.
　　Color:yellow to light y ellow .
3133 期　　　　　　　　　　　　于　晶等:中国藓类植物孢子类型的初步研究
　　Shape:slight ly sub-tet rahedral.
　　Polari ty:heteropolar and radiosymmetric.
　　Aperture:t rilete , laesura arms narrow , fine ,
running out onto the edge of amb in proximal face.
　　Ornamentation structure:perispore fo rming the
superficial ornamentation(smooth exospo re exposed
besides arms), fine exospo re is covered w ith low ,
small , irregular verruca in the distal face and gradu-
ally diminishing both in size and number in the
proximal face.
　　Spores of Hedwigiaceae belong to i t.
3.3　Andreaeoid type(Plate I:6 , 9)
　　Size:21 ～ 34μm in diameter.
　　Colo r:dark brow n.
　　Shape:more or less tetrahedral.
　　Polari ty:cryptopolar and radiosymmetric.
　　Aperture:katalept.
　　Ornamentation st ructure:sporoderm thick and
perispore thinner than exospore;Perispore two lay-
ers and fractional , the outer layer forming the super-
f icial verruca , exospore thicker than perispo re and
insert into exospo re sometimes.
　　Spores of Andreaceae and Tarkakiaceae belong
to it.
3.4　Achidioid type (Plate I:7 ,8)
　　Size:130 ～ 220 μm in diameter , nearly the
big gest in moss spores.
　　Colo r:g ray-yellow .
　　Shape:irregularly spherical , amb often slightly
triangular.
　　Polari ty:isopolar.
　　Aperture:at reme.
　　Ornamentation st ructure:exospo re forming the
superficial g ranulate ornamentation w hich is encrust-
ed w ith thin perispo re.The unique character is that
multi-layered endospore not only is thick but also
stratifies and appears as light strands betw een dark
ones , especially at amb corner.
　　Spores of Archidiaceae belong to it.
3.5　Bruchioid type(Plate II:1 ～ 3)
　　Size:18 ～ 48 μm in diameter.
　　Colo r:yellow or green yellow .
　　Shape:spherical and amb slight ly t riangular.
　　Polari ty:heteropolar and radiosymmetric.
　　Aperture:katalept
　　Ornamentation st ructure:distal surface pilate ,
pilum on distal surface is longer than on proximal
face.In distal surface , exospore forming the super-
ficial ornamentation encrusted w ith perispore that
appears as g ranulate.In pro ximal surface , it is en-
dospo re that consists most parts of the girdle sur-
rounding the leptoma and over w hich covered ex-
ospo re and perispore.
　　Spores of Bruchiaceae belong to it.
3.6　Encalyptoid type(Plate II:4 ～ 6)
　　Size:12 ～ 40μm in diameter.
　　Color:yellow to brow n.
　　Shape:spherical or amb slightly t riangular.
　　Polarity:heteropolar or isopolar.
　　Aperture:katalept , sometimes seems trilete
under light microscope(LM)or SEM [ 8 ,9] , but obvi-
ously katalept under TEM.
　　Ornamentation structure:all of perispore , ex-
ospo re and endospore take part in the formation of
ornamentation , in heteropolar spore proximal face
wi th ridges that amalg amated into a large central
shield and in the middle of w hich is the posi tio n of
leptoma , surface appears as verrucate , clavate o r
carinate.
　　Spores of Encalyptaceae belong to i t.
3.7　Bart ramioid type (Plate II:7 ～ 9)
　　Size:17 ～ 40μm in diameter.
　　Color:yellow to brow n.
　　Shape:spherical or elliptical.
　　Polarity:heteropolar.
　　Aperture:kataplet , leptoma usually concave.
　　Ornamentation st ructure:coarsely echinate ,
ret iculate or clavate on both face , sporoderm thick ,
perispore forming the differently superficial process-
es.
　　Spores of Bartramiaceae , Physcom itrium and
Entosthodon belong to i t.
3.8　Drummondioid type(Plate III:1)
　　Size:45 ～ 80 μm in diameter , including multi
cells.
　　Color:brow n green.
　　Shape:elongated ellipt ical or spherical.
　　Polarity:isopolar.
314 　　　　　　植　　物　　研　　究　　　　　　　　　　　　　　　　　　23 卷
　　Aperture:katalept o r at reme.
　　Ornamentation st ructure:sparsely scabrate or
g ranulate.
　　Spores of Drummondia , Myurella and some
species of Glyphomitrium , all species w ith mult i-
cells spores belong to i t.
3.9　Bryoid type (Plate III:2 ～ 10)
　　Size:6 ～ 30 μm in diameter.
　　Colo r:yellow or brow n.
　　Shape:spherical or elliptical.
　　Polari ty:isopolar.
　　Aperture:katalept o r at reme.
　　Ornamentation st ructure:Ornamentation is
composed of perispore appearing as:foveolate(Plate
III:2 , 3), granulate (Plate III:4 , 7), clavate
(Plate III:5), rugulate (Plate III:6), baculate
(Plate III:8), tuberculate (Plate III:9), vermi-
formed(Plate III:10).
　　Spores except those involved in the former
types belong to i t.
Based on preliminary study and above descrip-
tion of spo res of mosses , the key to theses 9 types is
given here.Detailed classificat ion needs an in -
depth analy sis.
Key to the spore types of the mosses
1.Spo re multi-cells Drummondioid type……………………………………………………………………………………………
1.Spo re uni-cells 2…………………………………………………………………………………………………………………
　2.Spo re trilete 3…………………………………………………………………………………………………………………
　2.Spo re atreme , katalept or cryptoaper ture 4……………………………………………………………………………………
3.T rilete laesurae with margo Sphagnoid type………………………………………………………………………………………
3.T rilete laesurae without margo Hedwigioid type…………………………………………………………………………………
　4.Spo re cryptoaperture Andreaeoid type…………………………………………………………………………………………
　4.Spo re atreme or ka talept 5……………………………………………………………………………………………………
5.Spo re big , more than 100 μm , shape irregular Achidioid type…………………………………………………………………
5.Spo re small , less than 80 μm , shape regular 6…………………………………………………………………………………
　6.Spo re heteropolar or isopolar. 7…………………………………………………………………………………………………
　6.Spo re isopolar Bryoid type……………………………………………………………………………………………………
7.Spo re heteropolar , o rnamentation constituted by perispo re. Bartramioid type…………………………………………………
7.Spo re heteropolar or isopolar , ornamenta tion constituted by exospo re o r to gether with o ther two layers 8……………………
　8.Ornamentation structure constituted by exospore , perispo re only mantle the exospore Bruchioid type………………………
　8.Ornamentation constituted by perispore , exospore and endospo re Encalyptoid type…………………………………………
　　References
1.Erdtman G.On pollen and spore terminology.Paleob-
o tanist , 1952 , 1:169～ 176
2.Erdtman G.Pollen and spore morpholog y/P lant Taxono-
my.Gymnospermae , Pteridophy ta, Bryophy ta (Illustra-
tions). An introduction to palynology.Stockholm:
Almqvist and Wiksell , 1957
3.Erdtman G.Pollen and spore morpholog y/P lant Taxono-
my.Gymnosperma , Bryophyta(Tex t).An introduction to
Palynology.Stodkholm:Almqvist and Wiksell , 1965
4.Miyoshi N.Spore morpholog y of Hepaticae in Japan.Bull
Okayama Coll Sci , 1966 , 2:1 ～ 46
5.Clarke G C S.Spore morphology and bryophyte sy stematics ,
In:G C S Clarke , J G Duckett , eds.Bryophy te Sy stema t-
ics.London and New York:Academic Press , 1979.231 ～
250
6.Punt W , Blackmore , et al.Glossary of pollen and spo re ter-
minology.The Netherlands:University of U trecht , 1994
7.Gupta A.Spore types in bryophy tes.Chenia:[ s.n.] ,
1998.5:39～ 48
8.Boros A.An atlas o f recent European bryophy te spores(sec-
ond edition).Budapest:Academic Press , 1993.1～ 321
9.Hor ton D G.A revision of the Encalyptaceae(Musci), with
par ticular reference to the Nor th American taxa.J Hattori
Bo t Lab , 1982 , 53:36～ 418
Explanation of plates
Plate I.1 ～ 3.Sphagnum imbricatum , 1.SEM microg raph ,
distal face(×2 200);2.SEM micrograph , proximal face(×
3153 期　　　　　　　　　　　　于　晶等:中国藓类植物孢子类型的初步研究
2 200 ). 3. TEM microg raph , sporoderm structure
(×15 000).4 ～ 5.Braunia delavayi , 4.SEM microg raph ,
proximal face (×3 300);5.SEM microg raph , distal face
(×3 300).6 , 9.Andreaea rupestris , 6.SEM microg raph ,
proximal face (×1 700);9.TEM microg raph , spo roderm
structure(×10 000).7～ 8.Archidium ohioense , 7.SEM mi-
crograph(×400);8.TEM microg raph.sporoderm structure
(×10 000)
Plate II.1 ～ 3.Trematodon ambiguous , 1.SEM microg raph ,
lateral face (×3 000);2.SEM micrograph , prox imal face
(×3 000);3.TEM micrograph. spo roderm structure
(×3 500).4 ～ 6.Encalypta vulgaris , 4.SEM microg raph ,
distal face (×2 000);5.SEM micrograph , proximal face
(×2 000);6.TEM micrograph , sporoderm structure
(×4 000).7 ～ 9.Physcomitrium eurystom um , 7.SEM mi-
crog raph , distal face(×3 000);8.SEM microg raph , proximal
face (×3 000);9.TEM micro graph , sporoderm structure
(×15 000)
Plate III.1.Drummondia sinensis , SEM micrograph
(×1 000).2 , 3.Schistostega pennata;2.SEM micrograph
(×4 000);3. TEM micrograph , sporoderm structure
(× 40 000).4 , 7.Dicranum japonicum , 4.TEM micro-
g raph , spo roderm structure (×3 500);5. Dicranella
fukienensis , SEM microg raph (×3 300).6.Leptodontium
v iticulosoidesSEM micrograph(×3 300).7.SEM micrograph
(×4 000).8. Pohlia sphagnicola SEM micrograph
(×4 000).9.Regmatodon declinatus , SEM micrograph
(×2 200).10.Anoectangium thomsonii , SEM micrograph
(× 4 000)
316 　　　　　　植　　物　　研　　究　　　　　　　　　　　　　　　　　　23 卷