全 文 ：第 38 卷 增刊
2014 年 9 月
南京林业大学学报(自然科学版)
Journal of Nanjing Forestry University (Natural Sciences Edition)
Vol． 38，Suppl．
Sept．，2014
doi:10． 3969 / j． issn． 1000 － 2006． 2014． S1． 003
收稿日期:2014 － 09 － 10
基金项目:国家自然科学基金项目(30972406)
第一作者:臧德奎，教授，博士。E-mail:zhangdk@ sdau． edu． cn。
引文格式:臧德奎，马燕． 木瓜属及其近缘属的花粉形态及系统学意义［J］． 南京林业大学学报:自然科学版，2014，38(S1):13 － 16．
木瓜属及其近缘属的花粉形态及系统学意义
臧德奎，马 燕
(山东农业大学林学院，山东 泰安 271018)
摘要:对木瓜属(Pseudocydonia)、贴梗海棠属(Chaenomeles)和榅桲属(Cydonia)植物的花粉亚显微形态进行了研
究。结果表明，其花粉均属于 N3 P4C3型，为辐射对称的长球形，极少为超长球形。贴梗海棠属花粉大小(极轴长
(P) ×赤道轴长(E))为 33. 24 ～ 48． 27 μm × 18． 93 ～ 25． 58 μm;木瓜属为 37． 15 ～ 42． 50 μm × 19． 63 ～ 22． 83
μm;榅桲属为 36． 85 ～ 43． 95 μ m × 19． 63 ～ 22． 83 μm。贴梗海棠属的和木瓜属花粉的表面纹饰为条纹状，前者
的条纹极不规则而后者条纹近于平行，榅桲属花粉外壁近光滑。花粉亚显微形态对于 3 个近缘属的分类具有较
大意义，支持将木瓜单独成属。另外，对所有种类的花粉形态进行了描述，并编制了分类检索表。
关键词:木瓜属;贴梗海棠属;榅桲属;花粉形态;系统学意义
中图分类号:Q948 文献标志码:A 文章编号:1000 － 2006(2014)S1 － 0013 － 04
Pollen morphology and phylogenetic significance of Pseudocydonia and related genera
ZANG Dekui，MA Yan
(Forestry College，Shandong Agricultural University，Taian 271018，China)
Abstract:The pollen morphology of Pseudocydonia，Chaenomeles and Cydonia were studied with the aid of scanning e-
lectron microscope (SEM)． Pollen grains were N3 P4C3 type according to G． Erdtmans N． P． C system，radio symmetric
prolate or rarely perprolate，with the sizes of 33． 24 － 48． 27 μm (P)× 18. 93 － 25． 58 μm (E)in Chaenomeles，37． 15
－ 42． 50 μm (P)× 21． 74 － 24． 86 μm (E)in Pseudocydonia，and 36． 85 － 43． 95 μm (P)× 19． 63 － 22． 83 μm (E)
in Cydonia． The exine sculpture was striate in Chaenomeles and Pseudocydonia (stripes irregular in the former but nearly
parallel in the latter)，while nearly smooth in Cydonia． The results demonstrated that palynological characters were valu-
able in recognition of taxa in specific and genera level and agreed with the separation of Chaenomeles and Pseduocydonia．
The pollen submicroscopic characters of six species in three genera were described，and a key was compiled for them．
Key words:Pseudocydonia;Chaenomeles;Cydonia;pollen morphology;phylogenetic significance
Chaenomeles Lindley is a small genus of the fami-
ly Ｒosaceae distributed in East Asia，comprising four
species，i． e． C． cathayensis (Hemsley) C． K．
Schneider，C． speciosa (Sweet)Nakai，C． tibetica
T． T． Yu，and C． japonica (Thunberg)Lindley ex
Spach． The first three species distributed in China
and only C． japonica found in Japan． It is known as
flowering quince and used as ornamental shrubs and
traditional medicine in Japan and China for a long
time． Cydonia Miller is a monotypic genus found in
Asia． Pseudocydonia Schneider，which was estab-
lished as a section in the genus Chaenomeles at first，
but Schneider gave it a generic status in 1906，com-
prising only one species， Pseudocydonia sinensis
(Thouin)C． K． Schneider ［Chaenomeles sinensis
(Thouin)Koehne)］，and was endemic to China．
These three genera，Chaenomeles，Cydonia and
Pseudocydonia have close relationships each other．
Pseudocydonia，however，is a doubtful taxon． Some
authors recognized two genera， Chaenomeles and
Cydonia，with Pseudocydoniai as a synonymy of the
former ［1 － 3］，or as a synonymy of the latter ［4 － 6］．
However，Ｒobertson Ｒ K， et al［7 － 12］ recognized
Pseudocydonia as a separate genus．
It has been shown that pollen morphology is use-
ful for phylogenetic analysis． Pollen morphology of
南 京 林 业 大 学 学 报 (自 然 科 学 版 ) 第 38 卷
these three genera has never been investigated system-
atically． The aim of the present study was to clarify
the differences of the pollen grains among them，and
to evaluate significance of the micromorphology of pol-
len grains in taxonomy．
1 Materials and Methods
Our samples represent all the six species in three
genera:Chaenomeles cathayensis (Hemsley)C． K．
Schneider， C． japonica (Thunberg) Lindley ex
Spach，C． speciosa (Sweet)Nakai，C． tibetica T． T．
Yu， Cydonia oblonga Miller， and Pseudocydonia
sinensis (Thouin)C． K． Schneider．
Pollen grains from mature but not dehiscent an-
thers were collected from flowering quince germplasm
repository of Shandong Agricultural University in Ta-
ian City，China． Voucher specimens are deposited in
Shandong Agricultural University Herbarium
(SDAUH)． For scanning electron microscope (SEM)
observations，pollen grains were dusted onto stubs and
coated with gold． Pollen grains were examined with a
Hitachi S － 570 scanning electron microscope operated
at 15 kV voltages．
For measurements“Image Tools”software with
high accuracy and confidence degree were used． The
mean of 20 pollen grain characters were considered
and measured for each sample．
2 Ｒesults and Discussion
The pollen of the three genera were，usually，
isopolar，radio symmetric prolate，or rarely perprolate
(in Chaenomeles speciosa and C． japonica)，tri-col-
porate，medium size． They are circular or 3 － lobed
circular in polar view and elliptical in equatorial view．
Their apertures are N3 P4 C3 type according to G．
Erdtmans N． P． C system ［13］．
No significant differences in pollen shapes and
aperture types were observed among the three genera，
while the pollen sizes，exin sculpture，densities and
sizes of tectum perforations，and striate width varied
depending on species and genera (Table 1，Fig． 1)．
Table 1 Pollen morphological characters for each species
表 1 不同植物种花粉的特征
taxa
P /μm E /μm P /E value
mean min max mean min max mean min max
pollen
shape exine sculpture Fig． 1 No．
Chaenomeles
cathayensis 39． 59 38． 30 40． 72 22． 65 21． 87 24． 06 1． 75 1． 68 1． 82 prolate
roughly striate，clear，very
irregular， ca． 0． 39 μm
wide; tectum perforations
few，ca． 0． 25 μm in di-
ameter．
1 － 3
Chaenomeles
speciosa 41． 15 37． 86 43． 23 20． 56 19． 62 23． 04 2． 00 1． 87 2． 16
prolate，
perprolate
striate，irregular，ca． 0． 18
m wide;tectum perforations
few，ca． 0． 20 μm in diame-
ter．
4 － 7
Chaenomeles
japonica 44． 17 40． 51 48． 27 22． 61 20． 68 25． 58 2． 00 1． 84 2． 11
prolate，
perprolate
roughly striate， irregular，
ca． 0． 36 μm wide;tectum
perforations many， big，
ca． 0． 45 μm in diameter．
8 － 10
Chaenomeles
tibetica 35． 57 33． 24 37． 56 21． 48 18． 93 22． 81 1． 66 1． 55 1． 76 prolate
striate，irregular，ca． 0．
18 μm wide;tectum per-
forations rare or none．
11 － 13
Pseudocydonia
sinensis 40． 73 37． 15 42． 50 23． 45 21． 74 24． 86 1． 75 1． 61 1． 88 prolate
striate， mainly parallel
each other，ca． 0． 13 μm
wide; tectum perforations
many，ca． 0． 17 μm in di-
ameter．
14 － 17
Cydonia
oblonga 40． 90 36． 85 43． 95 21． 57 19． 63 22． 83 1． 90 1． 86 1． 92 prolate
very unconspicuous striate
or nearly smooth; tectum
perforations sparse， ca．
0. 15 μm in diameter．
18 － 20
The polar axis is the longest in Chaenomeles ja-
ponica (44． 17 μm)and the shortest in C． tibetica
(35． 57 μm)． P /E values range from 1． 55 in C． ti-
betica to 2． 16 in C． speciosa．
41
2014 年 9 月 臧德奎，等:木瓜属及其近缘属的花粉形态及系统学意义
Fig． 1 Pollen morphology of the genus Chaenomeles，Pseudocydonia，and Cydonia under SEM
图 1 贴梗海棠属、木瓜属及榅桲属植物花粉的显微形态
Chaenomeles cathayensis:1． group view (× 600) ;2． equatorial view (× 2 000) ;3． ornamentation (× 10 000)．
Chaenomeles speciosa:4． group view (× 600) ;5． equatorial view (× 2 000) ;6． ornamentation (× 10 000) ;7． polar view (× 2 500)．
Chaenomeles japonica:8． group view (× 600) ;9． equatorial view (× 2 000) ;10． ornamentation (× 10 000)．
Chaenomeles tibetica:11． group view (× 600) ;12． equatorial view (× 2 000) ;13． ornamentation (× 10 000)．
Pseudocydonia sinensis:14． group view (×600) ;15． equatorial view (×2 000) ;16． ornamentation (×10 000) ;17． polar view (×2 500)．
Cydonia oblonga:18． group view (× 600) ;19． equatorial view (× 2 000) ;20． ornamentation (× 10 000)．
Key to the genera and species
1． Exine sculpture striate，conspicuous．
2． Stripes wide and irregular． ［Chaenomeles］
3． Tectum perforations few，small，(0． 25 μm in di-
ameter．
4． Pollen grain big，polar axis 38． 30 －48． 27 μm long，
tectum perforations few，ca． 0. 25 μm in diameter．
5． Stripes very clear，ca． 0． 39 μm wide …………
Chaenomeles cathayensis…………………………
5． Striate，irregular，ca． 0． 18 μm wide;tectum per-
forations few，ca． 0． 20 μm in diameter ………
Chaenomeles speciosa……………………………
4． Pollen grain small，polar axis 35． 57(33． 24 －
37. 56)μm long;tectum perforations rare or none
Chaenomeles tibetica
3． Tectum perforations numerous，big，ca． 0． 45 μm
in diameter． Chaenomeles japonica………………
2． Stripes fine，nearly parallel each other ［Pseudo-
cydonia］ Pseudocydonia sinensis…………………
1． Exine sculpture very unconspicuous striate or near-
ly smooth［Cydonia］ Cydonia oblonga…………
The pollens from the same sample do not show
difference in their surface morphology． This indicates
that pollen morphology is stable within species． How-
ever，among species and genera pollen morphology
demonstrates a variation，which may indicate taxo-
nomic similarity or dissimilarity．
Cydonia is different from Chaenomeles and
Pseudocydonia in pollen characteristics． The exine
sculpture of pollen grain of Cydonia is almost smooth，
quite different from that of Chaenomeles and Pseudo-
cydonia． Difference also exists in morphological char-
51
南 京 林 业 大 学 学 报 (自 然 科 学 版 ) 第 38 卷
acteristics． In Cydonia，styles free，calyx persistent
in fruit，leaf blade entire，while styles connate at
base， calyx caducous， leaf blade serrate in
Chaenomeles and Pseudocydonia． This suggests the al-
ienation between Cydonia and the other two genera．
The pollen of Pseudocydonia and Chaenomeles
are both striate in exine sculpture，but stripes in
Pseudocydonia are fine and almost parallel each oth-
er， perforations small and numerous， while in
Chaenomeles，stripes are wide and irregular，with few
or rare perforations (except in C． japonica，which has
big perforations)． Also，these two genera differ in
their morphological characteristics． Pseudocydonia is
a tree，branchlets unarmed，flowers solitary，calyx
lobes reflexed，but the other is a shrub，branchlets
armed，flowers in clusters，and calyx lobes erect．
Considering these characters，separation of Pseudo-
cydonia from Chaenomeles is reasonable．
In the genus Chaenomeles， C． cathayensis
(Hemsley)C． K． Schneider is similar to C． speciosa
(Sweet)Nakai in exine sculpture． Similarity is also
shown in their morphology． C． japonica (Thunberg)
Lindley ex Spach，with dense and big perforations on
exine sculpture，is special and different from other
species．
In conclusion，the morphological dissimilarity a-
mong Chaenomeles and its relatives is also reflected in
their pollen morphology． The results demonstrate that
palynological characters are useful in recognition of
species and genera and agreed with the separation of
Chaenomeles and Pseduocydonia．
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