全 文 :横断山区植物的花粉形态及生态意义∗
郑 鑫1ꎬ 闵运江1ꎬ 徐 波2ꎬ 孙 航3ꎬ 周忠泽1∗∗
(1 安徽大学资源与环境工程学院ꎬ 安徽省生物多样性信息中心ꎬ 合肥 230601ꎻ 2 西南林业大学
林学院ꎬ 昆明 650204ꎻ 3 中国科学院昆明植物研究所ꎬ 昆明 650201)
摘要: 应用光学显微镜和扫描电子显微镜对中国西南部横断山区夏季开花的 27 科 43 属 46 种 2 变种植物
的花粉形态进行了观察和研究ꎬ 并对花粉形态进行了细致的描述ꎮ 结果表明ꎬ 花粉类型以近长球形—长球
形和长球形为主ꎬ 分别为 27 1%和 25%ꎬ 还有少量的为球形ꎬ 近球形—近长球形ꎬ 近扁球形和四合花粉ꎮ
萌发孔以 3孔沟为主ꎬ 占 43 8%ꎬ 还有 3沟ꎬ 散孔ꎬ 多沟ꎬ 三拟孔沟ꎬ 6 ̄沟ꎬ 单沟等类型ꎮ 外壁纹饰以细
网状为主占 50%ꎬ 还有粗网状ꎬ 细颗粒状ꎬ 光滑ꎬ 刺状纹饰等ꎮ 这些物种的孢粉学特征为第四纪地层孢粉
研究提供了现代孢粉学依据ꎮ 此外在花粉形态性状基础上ꎬ 通过 SPSS软件聚类分析ꎬ 对部分同科物种间
的鉴定特征进行了讨论分析ꎬ 并根据植物的生态学特性讨论了它们的生态环境指示意义ꎮ
关键词: 花粉形态ꎻ 生态意义ꎻ 横断山区
中图分类号: Q 944 文献标志码: A 文章编号: 2095-0845(2015)06-657-26
Pollen Morphology of Plants from the Hengduan Mountains
and Their Ecological Significance∗
ZHENG Xin1ꎬ MIN Yun ̄jiang1ꎬ XU Bo2ꎬ SUN Hang3ꎬ ZHOU Zhong ̄ze1∗∗
(1 School of Resources and Environmental Engineeringꎬ Anhui Universityꎬ Anhui Biodiversity Information Centerꎬ
Hefei 230601ꎬ Chinaꎻ 2 College of Forestryꎬ Southwest Forestry Universityꎬ Kunming 650204ꎬ Chinaꎻ
3 Kunming Institute of Botanyꎬ Chinese Academy of Sciencesꎬ Kunming 650201ꎬ China)
Abstract: Pollen morphology of selected taxa from the Hengduan Mountainsꎬ Southwest China are investigated in the
present study. Forty ̄eight taxa (flowering in the summerꎬ 46 species and 2 varieties) of 43 genera belonging to 27
families were observed by light and scanning electron microscopy. Pollen morphology is described in detail. Two pol ̄
len shapes are mainly found in these species: subprolate to prolate (27 1%) and prolate (25%). Spheroidalꎬ sub ̄
spheroidal to subprolateꎬ suboblate and tetrad shapes can also be found in some species. Aperture type is mostly tri ̄
colporateꎬ with a percentage of 43 8ꎬ and also contains tricolpateꎬ pantoporateꎬ stephanocolpateꎬ 3 ̄colporoidꎬ 6 ̄
colpateꎬ monocolpate. The most common ornamentation is finely reticulateꎬ with a percentage of 50. Other exine or ̄
namentationsꎬ such as coarsely reticulateꎬ finely granulateꎬ smoothꎬ spinulose are also observed. The palynological
documentation of these species will provide the modern palynological basis for paleopalynological studies of the Qua ̄
ternary Strata. On the basis of pollen morphology charactersꎬ identification features of some species in the same fami ̄
lies were discussed according to the results of cluster analysis used by SPSS. The ecological significance was also dis ̄
cussed based on the ecological properties of these taxa.
Key words: Pollen morphologyꎻ Ecological significanceꎻ Hengduan Mountains
植 物 分 类 与 资 源 学 报 2015ꎬ 37 (6): 657~682
Plant Diversity and Resources DOI: 10.7677 / ynzwyj201515044
∗
∗∗
Funding: The Specialized Research Fund for the Doctoral Program of Higher Education of China (20123401110005)ꎬ the National Natural
Science Foundation of China (41072251) and the State Key Laboratory of Palaeobiology and Stratigraphy (Nanjing Institute of
Geology and Palaeontologyꎬ CASꎬ No. 113106)
Author for correspondenceꎻ E ̄mail: zhzz@ahu edu cn
Received date: 2015-03-16ꎬ Accepted date: 2015-09-10
作者简介: 郑鑫 (1990-)ꎬ 女ꎬ 硕士研究生ꎬ 主要从事生物多样性与保护生态学研究ꎮ E ̄mail: xin5a1102@163 com
The Hengduan Mountains have drawn much at ̄
tention in the climate and environmental change
studies ( Li et al.ꎬ 2012ꎻ Kramer et al.ꎬ 2010ꎻ Yao
et al.ꎬ 2015ꎻ Maꎬ 2013). Pollen analysis has been
one of the most widely useful and reliable tools to re ̄
construct paleovegetation and paleoclimate in this re ̄
gion. Studies of modern pollen are the indispensable
prerequisite for the fossil pollen interpretation in the
same area. The comparison between the modern pol ̄
len spectra and the fossil pollen records not only in ̄
dicates local climate changeꎬ but also provides a val ̄
uable basis for the restoration of the area’s ancient
environment. (Tangꎬ 2002ꎻ Tang et al.ꎬ 2009ꎻ Li et
al.ꎬ 2006ꎻ Zhao et al.ꎬ 2006).
Howeverꎬ few modern pollen studies have been
carried out on the relationship among pollenꎬ vegeta ̄
tion and climate in the Hengduan Mountains. Thus
we still lack the modern pollen spectra which could
be a background database for better identification of
fossil pollen in the Hengduan Mountainsꎬ to recon ̄
struct ancient vegetationꎬ palaeoclimate and palaeo ̄
environment there.
The Hengduan Mountains region which located
at the southeastern end of the Tibetan Plateauꎬ is the
general term for a chain of parallel mountains run ̄
ning south to north in Sichuanꎬ Yunnan and Eastern
Tibet. Covering an area of about 500 000 km2 (Shi
et al.ꎬ 1998)ꎬ their elevation gradient is from 2 000
to 6 000 m. With its highly complex geological struc ̄
tureꎬ the region constitutes the transition zone con ̄
necting China’s eastern Pacific and western ancient
Mediterranean areas (Panꎬ 1989).
The Hengduan Mountains region is the richest
area of China in endemic plants and one of the biodi ̄
versity hotspots in the world (Boufford et al.ꎬ 2004).
It is estimated that the Hengduan Mountains has more
than 9 000 plant species (Yang et al.ꎬ 2012). It is
also a well ̄known habitat for many modern temperate
and alpine plantsꎬ especially alpine plants in the gen ̄
era Gentiana L.ꎬ Saxifraga Tourn. ex L.ꎬ Saussurea
DC.ꎬ Primula L.ꎬ etc. (Liu et al.ꎬ 1986ꎻ Wuꎬ 1979ꎬ
1980ꎬ 1987ꎻ Wangꎬ 1992ꎻ Li and Liꎬ 1993).
In this studyꎬ both light microscopy (LM) and
scanning electron microscopy ( SEM) were used to
examine the pollen morphology of plants from the H ̄
D Mountains in order to:
— describe pollen and document them with LM
and SEM micrographsꎻ
— provide modern palynological information for
the reconstruction of the paleoclimate and paleoen ̄
vironment based on corresponding fossil pollen from
Quaternary strata.
1 Material and Methods
1 1 Observation of pollen morphology
In 2011ꎬ flowers ( flowering in summer) of 46
species and 2 variants of plants belonging to 43 gen ̄
era of 27 families from the Hengduan Mountains
were collected in the field or from ex ̄situ cultures
(Table 1) and then dried. All the species were listed
in Table 1 which was based on the concepts of Engler
System. The exsiccata were kept in the herbarium of
the Department of Ecologyꎬ School of Resources and
Environmental Engineeringꎬ Anhui Universityꎬ Chi ̄
na. The palynological terminology used is according
to Punt et al. (2007) and Hesse et al. (2009).
To prepare them for LMꎬ pollen grains were ace ̄
tolyzed according to the standard method (Erdtmanꎬ
1971): dry anthers were placed in glacial acetic acid
for 12 hꎬ incubated in a mixture of concentrated sul ̄
phuric acid and acetic anhydride (1∶9)ꎬ and placed
in a water bath at 95 ℃ for 5 min and then centri ̄
fuged. The supernatant was dischargedꎬ and the sedi ̄
ment was washed three times in distilled water and
placed in glycerine. The pollen was then embedded in
glycerin jelly on glass slides (Wodehouseꎬ 1959). A
Leica DME light microscope was used for observations
and measurements. Ten regularly shaped and fully ex ̄
panded pollen grains were measured in each sample.
All the measurements were taken under magnification
400 ×. Generallyꎬ for each speciesꎬ 20 pollen grains were
measuredꎬ and the meanꎬ maximum and minimum val ̄
ues of P (polar axis)ꎬ E (equatorial axis) and P / E
ratio were recorded to reveal the range of variation. LM
856 植 物 分 类 与 资 源 学 报 第 37卷
-images were created with a stereoscopic light micro ̄
scope OLYMPUS BX51 at 1 000 × magnification.
For observation under SEMꎬ pollen grains were
extracted from dry anthersꎬ transferred onto stubsꎬ
sealed with a drop of nail polishꎬ coated with gold or
alloy of platinum and palladium using JFC - 1100E
sputterꎬ and then observed under the scanning elec ̄
tronic microscopes JSM-6300 at 15-20 kV. The ter ̄
minology follows Hesse et al. (2009)ꎬ and Kupriy ̄
anova and Aleshina (1972).
Table 1 Collection locality information and vouchers for all samples
Order Family Species Vouchers Location Altitude / m
Farinosae Commelinaceae Cyanotis vaga 0908023 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2241
Liliflorae Liliaceae Tofieldia divergens 0908018 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2239
Polygonales Polygonaceae Polygonum chinense 09051 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2239
Centrospermae Caryophyllaceae Silene yunnanensis 0908106 Yulong Snow Mt.ꎬ Lijiangꎬ Yunnanꎬ China 2859
S baccifera 0908014 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2239
Ranales Ranunculaceae Delphinium delavayi 0908034 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2181
Thalictrum delavayi 0908107 Yulong Snow Mt.ꎬ Lijiangꎬ Yunnanꎬ China 2859
Anemone hupehensis 0908032 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2181
Rosales Saxifragaceae Astilbe rivularis 0908007 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2235
Rosales Rosaceae Sorbaria arborea 0908113 Baima Snow Mt.ꎬ Deqinꎬ Yunnanꎬ China 4292
Potentilia fulgens 0908008 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2235
Spenceria ramalana 0908052 Shangri ̄Laꎬ Yunnanꎬ China 3300
Rosales Fabaceae Cochlianthus montanus 0908103 Yulong Snow Mt.ꎬ Lijiangꎬ Yunnanꎬ China 2859
Pueraria peduncularis 0908003 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2235
Clitoria mariana 0908009 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2235
Vicia cracca 0908020 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2239
Sapindales Celastraceae Tripterygium hypoglaucum 0908013 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2239
Sapindales Balsaminaceae Impatiens uliginosa 0908031 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2181
I radiata 0908035 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2181
I procumbens 0908030 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2181
I delavayi 0908036 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2181
Guttiferae Hypericum addingtonii 0908012 Yulong Snow Mt.ꎬ Lijiangꎬ Yunnanꎬ China 2810
Begoniaceae Begonia grandis 0908100 Yulong Snow Mt.ꎬ Lijiangꎬ Yunnanꎬ China 2856
Melastomataceae Osbeckia crinita 0908012 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2239
Onagraceae Fuchsia hybrida 0908044 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2176
Oenothera rosea 0908041 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2176
Apiaceae Bupleurum longicaule 0908025 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2241
Heracleum franchetii 0908038 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2181
Ericaceae Lyonia ovalifolia 0908015 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2239
Primulales Primulaceae Lysimachia violascens 0908024 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2239
Contortae Loganiaceae Buddleja fallowiana 0908049 Yulong Snow Mt.ꎬ Lijiangꎬ Yunnanꎬ China 2875
B forrestii 0908037 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2181
Contortae Gentianaceae Halenia elliptica 0908047 Yulong Snow Mt.ꎬ Lijiangꎬ Yunnanꎬ China 2880
Gentianopsis paludosa 0908057 Dongda Mt.ꎬ Zuogongꎬ Xizangꎬ China 5008
Tubiflorae Labiatae Ajuga forrestii 0908048 Shangri ̄Laꎬ Yunnanꎬ China 3300
Nepeta wilsonii 0908101 Yulong Snow Mt.ꎬ Lijiangꎬ Yunnanꎬ China 2856
Clinopodium megalanthum 0908010 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2239
Elsholtzia rugulosa 0908111 Yulong Snow Mt.ꎬ Lijiangꎬ Yunnanꎬ China 2891
Tubiflorae Scrophulariaceae Verbascum thapsus 0908050 Yulong Snow Mt.ꎬ Lijiangꎬ Yunnanꎬ China 2868
Pedicularis tenuisecta 0908027 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2241
P gruina 0908113 Yulong Snow Mt.ꎬ Lijiangꎬ Yunnanꎬ China 2890
Tubiflorae Bignoniaceae Incarvillea arguta 0908040 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2176
Acanthaceae Pteracanthus forrestii 0908102 Yulong Snow Mt.ꎬ Lijiangꎬ Yunnanꎬ China 2856
Tubiflorae Valerianaceae Valeriana flaccidissima 0908004 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2235
Rubiales Cucurbitaceae Zehneria maysorensis 0908043 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2176
Campanulales Campanulaceae Campanula colorata 0908026 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2241
Lobelia doniana 0908033 Cangshan Mt.ꎬ Daliꎬ Yunnanꎬ China 2181
Campanulales Compositae Saussurea stella 0908055 Dongda Mt.ꎬ Zuogongꎬ Xizangꎬ China 5008
9566期 ZHENG Xin et al.: Pollen Morphology of Plants from the Hengduan Mountains and Their Ecological
1 2 Cluster analysis
A matrix with 48 species per 6 qualitative and
quantitative selected palynological characters (Table
2) was built basing on the compiled data (Table 3)
and laboratorial analyses. The selection of palynolog ̄
ical characters was used to describe pollen morpholo ̄
gy of species generally. In order to retain much more
informationꎬ the selected characters were encoded
according to their dimensional types. The quantita ̄
tive characters were used directly without transforma ̄
tion while the qualitative characters were endowed
with nominal values (Table 3). Then Hierachical clus ̄
ter analysis was conducted on the selected pollen
characters codes using the SPSS package (Version
19 0ꎬ IBM http: / / www.ibm.com).
2 Results
2 1 General trends of pollen morphology in the
Hengduan Mountains
After observation of the pollen morphology data
aboveꎬ statistical analysis was conducted. The results
from plants in the H ̄D Mountains show that pollen
grains are mainly subprolate to prolate and prolateꎬ
with the proportions of 27 1% and 25 0%ꎬ respec ̄
tively. Spheroidalꎬ subspheroidal to subprolateꎬ sub ̄
oblate and tetrad shapes are also found in a few spe ̄
cies. Aperatures are mostly tricolporateꎬ amounting
to 43 8%ꎬ rarely tricolpateꎬ pantoporateꎬ polyplicateꎬ
etc. The majority of ornamentations are finely reticu ̄
lateꎬ amounting to 50 0%ꎬ a few species having an
ornamentation that is coarsely reticulateꎬ finely gran ̄
ulateꎬ smoothꎬ spinuloseꎬ etc (Appendix).
2 2 Pollen morphology
Under LMꎬ pollen types are presented in plates
I- IVꎬ pollen types and their ornamentation under
SEM are presented in plates V-VII.
Commelinaceae
Cyanotis D. Don
Cyanotis vaga (Pl. I: 1ꎻ Pl. V: 1)
Pollen grains 46 6 (25 0-60 0)×23 1 (10 0
-32 5) μmꎬ prolate to perprolateꎬ P / E=2 01 (1 45
-2 86). Equatorial view ellipticalꎬ ends upwardsꎬ
nearly cymbiform. Polar view circular or elliptical.
Aperture monocolpateꎬ anacolpus narrow. Exine 2 0 μm
thickꎬ sexine 1 5 times as thick as nexine. Columel ̄
lae distinct. Ornamentation: finely reticulate under
LMꎻ regulate ̄perforate under SEM.
Liliaceae
Acorus L.
Tofieldia divergens Bur. and Franch. (Pl. I: 2ꎻ
Pl. V: 2)
Pollen grainsꎬ 21 5 (20 0-25 0)×16 8 (15 0
-17 5) μmꎬ prolateꎬ P / E = 1 28 (1 14-1 5). E ̄
quatorial view elliptical. Polar view circular. Aperture
monocolpateꎬ anacolpus narrow. Exine 1 0 μmꎬ sex ̄
ine equal to nexine. Columellae indistinct. Ornamen ̄
tation: finely reticulate under LM and under SEM.
Polygonaceae
Polygonum L.
P chinense L. var. paradoxum (Levl.) A J. Li
(Pl. I: 3ꎬ 4)
Pollen grains 52 5 (47 5-55 0) μm in diam ̄
eterꎬ spheroidal. Equatorial view circular. Polar view
3 ̄lobed circular. Apertures 3 ̄colpateꎬ colpus short.
Exine 7 0 μm thickꎬ sexine 4 times as thick as nex ̄
ine. Columellae developed. Ornamentation: coarsely
reticulate under LMꎬ reticulation big and distinctꎬ
lumina containing 4-10 smaller columellae.
Table 2 Pollen characters and the code of qualitative characters
Number Characters and code Measure
1 Length of polaraxis scale
2 With of equatorial axis scale
3 P / E scale
4 Type of aperture: monocolpate(0) / 3 ̄colporate(1) / 3 ̄colpate(2) / 3 ̄porate(3) / 4 ̄colpate(4) / 4 ̄colporate(5) / 5 ̄porate(6) / pantoporate(7) / 6 ̄colpate(8) / 3 ̄ colporoid(9) / 2 ̄ syncolpate(10) nominal
5 Ornamentation(LM): reticulate(0) / granulate(1) / smooth(2) / spinulose(3)microreticulato ̄ stri ̄ate(4) / granulate ̄perforate(5) nominal
066 植 物 分 类 与 资 源 学 报 第 37卷
Caryophyllaceae
Silene L.
Silene yunnanensis Franch. (Pl. I: 5)
Pollen grains 51 8 (47 5-57 5) μm in diam ̄
eterꎬ spheroidal. Aperture pantoporateꎬ porusꎬ usu ̄
ally 20-28ꎬ circularꎬ 4 0-5 0 μm in diameterꎬ in ̄
terporal distance 7 0-9 0 μm. Exine 4 0 μm thickꎬ
sexine 3 times as thick as nexine. Ornamentation:
Table 3 List of the selected pollen charactersꎬ followed by their respective codes
Species Length of polaraxis/ μm
With of equatorial axis
/ μm P / E
Type of
aperture
Ornamentation
(LM)
1 2 3 4 5
Cyanotis vaga 46 6 23 1 2 01 0 0
Tofieldia divergens 21 5 16 8 1 28 0 0
Polygonum chinense 52 5 52 5 1 2 0
Silene yunnanensis 51 8 51 8 1 7 1
S baccifera 41 8 41 8 1 7 1
Delphinium delavayi 40 30 1 33 2 1
Thalictrum delavayi 25 5 25 5 1 7 1
Anemone hupehensis 26 23 5 1 09 2 1
Astilbe rivularis 15 5 12 3 1 27 1 0
Sorbaria arborea 21 8 19 8 1 1 1 0
Potentilla fulgens 24 75 21 1 17 1 0
Spenceria ramalana 29 5 19 25 1 53 1 0
Cochlianthus montanus 34 6 26 7 1 3 1 0
Pueraria peduncularis 35 31 1 12 1 0
Clitoria mariana 42 7 42 7 1 6 0
Vicia cracca 34 6 26 7 1 44 1 0
Tripterygium hypoglaucum 30 27 1 1 1 1 0
Impatiens uliginosa 41 4 26 1 59 4 0
I radiata 35 5 22 5 1 56 4 0
I procumbens 41 4 27 1 53 4 0
I delavayi 41 3 29 5 1 4 4 0
Hypericum addingtonii 26 8 22 8 1 18 1 0
Begonia grandis 27 9 13 2 2 11 1 0
Osbeckia crinita 32 5 25 5 1 27 1 2
Fuchsia hybrida 55 55 1 3 0
Oenothera rosea 109 109 1 3 4
Bupleurum longicaule 24 14 1 71 1 0
Heracleum franchetii 46 5 18 8 2 48 1 0
Lyonia ovalifolia 35 35 1 1 0
Lysimachia violascens 30 8 20 5 1 5 1 0
Buddleja fallowiana 21 6 14 7 1 47 5 0
B forrestii 24 5 19 5 1 25 5 0
Halenia ellipticala 38 8 36 5 1 06 1 0
Gentianopsis paludosa 17 3 15 5 1 11 1 0
Ajuga forrestii 26 4 20 9 1 26 2 0
Nepeta wilsonii 56 8 44 3 1 28 8 0
Clinopodium megalanthum 47 8 36 3 1 31 8 0
Elsholtzia rugulosa 28 9 20 1 44 8 0
Verbascum thapsus 28 5 22 8 1 25 9 0
Pedicularis tenuisecta 28 3 16 3 1 74 10 2
P gruina 36 8 30 8 1 19 10 2
Incarvillea arguta 24 5 19 5 1 25 8 5
Pteracanthus forrestii 101 6 61 6 1 65 1 0
Valeriana flaccidissima 45 5 41 3 1 1 2 3
Zehneria maysorensis 58 49 1 18 1 0
Campanula colorata 25 5 25 5 1 3 3
Lobelia doniana 44 3 31 3 1 42 1 0
Saussurea stella 66 7 60 1 11 1 3
1666期 ZHENG Xin et al.: Pollen Morphology of Plants from the Hengduan Mountains and Their Ecological
finely granulate under LM.
Silene baccifera (L.) Roth (Pl. I: 6ꎬ Pl. V: 3)
Pollen grains 41 8 (40 0-45 0) μm in diam ̄
eterꎬ spheroidal. Apertures pantoporateꎬ porus usu ̄
ally 10-16ꎬ circularꎬ 3 0-4 0 μm in diameterꎬ in ̄
terporal distance 6 0-8 0 μm. Exine 6 0 μm thickꎬ
sexine 3 times as thick as nexine. Ornamentation:
finely granulate under LMꎬ microechinate under SEM.
Ranunculaceae
Delphinium L.
Delphinium delavayi Franch. (Pl. I: 7ꎬ 8ꎻ Pl.
V: 4)
Pollen grains 40 0 (37 5-42 5)×30 0 (27 5
-32 5) μmꎬ P / E= 1 33 (1 15-1 41)ꎬ subprolate
to prolate. Equatorial view oblong. Polar view 3 ̄lobed
circular. Apertures 3 ̄colpateꎬ colpus margins indis ̄
tinctꎬ membrane containing granule. Exine 2 0 m
thickꎬ sexine equal to nexine. Exine rather thick at
poles. Columellae indistinct. Ornamentation: finely
granulate under LMꎬ microechinate under SEM.
Thalictrum L.
Thalictrum delavayi Franch. (Pl. I: 9ꎬ 10ꎬ 11)
Pollen grains 25 5 (22 5-27 5) μm in diam ̄
eterꎬ spheroidal. Apertures pantoporateꎬ porus usu ̄
ally 6-7ꎬ circularꎬ 1 0-2 5 μm in diameterꎬ inter ̄
poral distance 4 0 - 4 5 μm. Exine 2 0 μm thickꎬ
sexine equal to nexine. Columellae distinct. Orna ̄
mentation: finely granulate under LM.
Anemone L.
Anemone hupehensis Lem. f. alba W T. Wang
(Pl. I: 12ꎬ 13ꎻ Pl. V: 5)
Pollen grains 26 0 (22 5-27 5)×23 5 (20 0
-25 0) μmꎬ P / E = 1 09 (1-1 25)ꎬ subspheroidal.
Equatorial view elliptical. Polar view 3 ̄lobed circu ̄
lar. Apertures 3 ̄ colpateꎬ colpus long and slenderꎬ
extending to the polesꎬ 0 5 - 1 0 μm wide. Exine
2 0 μm thickꎬ sexine slightly thicker than or equal
to nexine. Columellae distinct. Ornamentation: finely
granulate under LMꎬ microechinate under SEM.
Saxifragaceae
Astilbe Buch. ̄Ham. ex D. Don
Astilbe rivularis Buch. ̄Ham. ( Pl. I: 14ꎬ 15ꎬ
16ꎻ Pl. V: 6)
Pollen grains 15 5 (15 0-17 5)×12 3 (10 0
-15 0) μmꎬ P / E= 1 27 (1 0-1 75)ꎬ subspheroi ̄
dal to subprolate. Equatorial view elliptical. Polar
view 3 ̄lobed circular. Apertures 3 ̄colporate. Ectoap ̄
erture ̄colpusꎬ long and slenderꎬ extending to the
poles. Endoaperture ̄porusꎬ lalongateꎬ 1 0-1 5 μm
in diameter. Exine 1 5 μm thickꎬ sexine equal to
nexine. Columellae distinct. Ornamentation: faintly
reticulate under LMꎻ distinct reticulate under SEM.
Rosaceae
Sorbaria (Ser.) A. Br. ex Aschers.
Sorbaria arborea Schneid. var. subtomentosa Reh ̄
der (Pl. I: 17ꎬ 18)
Pollen grains 19 8 (15 0-25 0)×21 8 (17 5
-27 5) μmꎬ P / E = 1 10 (1 0-1 17)ꎬ suboblate.
Equatorial view elliptical. Polar view 3 ̄lobed circu ̄
lar. Apertures 3 ̄colporate. Ectoaperture ̄colpusꎬ long
and slenderꎬ extending to the poles. Endoaperture ̄
porusꎬ circularꎬ 2 0 μm in diameter. Exine 2 0 μm
thickꎬ sexine slightly thicker than nexine. Columel ̄
lae indistinct. Ornamentation: faintly reticulate un ̄
der LM.
Potentilla L.
Potentilla fulgens Wall. ex Hook. (Pl. I: 19ꎬ
20ꎬ 21ꎻ Pl. V: 7)
Pollen grains 24 75 (22 5-27 5)×21 (17 5-
22 5) μmꎬ P / E = 1 17 (1 11-1 29)ꎬ subprolate
to prolate. Equatorial view elliptical. Polar view 3 ̄
lobed circular. Apertures 3 ̄colporate. Ectoaperture ̄
colpusꎬ long and slenderꎬ extending to the poles. End ̄
oaperture ̄porusꎬ circularꎬ 2 0 μm in diameter. Ex ̄
ine 3 0 μm thickꎬ sexine 1 5 times as thick as nex ̄
ine. Columellae distinct. Ornamentation: finely re ̄
ticulate under LMꎬ striate ̄perforate under SEM.
Spenceria Trimen
Spenceria ramalana Trimen (Pl. I: 22ꎬ 23ꎻ Pl.
V: 8)
Pollen grains 29 5 (20-37 5)×19 25 (12 5-
22 5) μmꎬ P / E= 1 53 (1 25-1 75)ꎬ prolate. E ̄
quatorial view elliptical. Polar view 3 ̄lobed circular.
Apertures 3 ̄colporate. Ectoaperture ̄colpusꎬ long and
266 植 物 分 类 与 资 源 学 报 第 37卷
slenderꎬ extending to the poles. Endoaperture ̄porusꎬ
circularꎬ 2 5 μm in diameter. Exine 2 0 μm thickꎬ
sexine equal to nexine. Columellae indistinct. Orna ̄
mentation: finely reticulate under LMꎬ striate ̄perfo ̄
rate under SEM.
Fabaceae
Cochlianthus Benth.
Cochlianthus montanus (Diels) Harms (Pl. I:
24ꎬ 25)
Pollen grains 34 6 (25 0-67 5)×26 7 (17 5
-55 0) μmꎬ P / E= 1 30 (1 09-1 57)ꎬ subprolate
to prolate. Equatorial view elliptical. Polar view 3 ̄
lobed circular. Apertures 3 ̄colporate. Ectoaperture ̄
colpusꎬ long and slenderꎬ extending to the poles. En ̄
doaperture ̄porusꎬ lalongateꎬ 3 0 μm in diameter. Ex ̄
ine 2 5 μm thickꎬ sexine equal to nexine. Columellae
indistinct. Ornamentation: finely reticulate under LM.
Pueraria DC.
Pueraria peduncularis (Grah. ex Benth.) Benth.
(Pl. I: 26ꎬ 27ꎬ 28ꎻ Pl. V: 9)
Pollen grains 35 0 (22 5-42 5)×31 0 (17 5
-37 5) μmꎬ P / E = 1 12 (1 06-1 28)ꎬ subsphe ̄
roidal to subprolate. Equatorial view elliptical. Polar
view 3 ̄lobed circular. Apertures 3 ̄colporate. Ectoap ̄
erture ̄colpusꎬ rather long. Endoaperture ̄porusꎬ big
and lalongateꎬ 4 0 μm in diameter. Exine 2 5 μm
thickꎬ sexine slightly thicker than nexine. Columel ̄
lae distinct. Ornamentation: finely reticulate under
LM and SEM.
Clitoria L.
Clitoria mariana L. (Pl. II: 29ꎬ 30ꎻ Pl. V: 10)
Pollen grains 42 7 (44 0-63 0) μm in diam ̄
eterꎬ spheroidal. Aperture 5 ( ̄6) ̄porateꎬ porus cir ̄
cularꎬ 7 0-8 0 μm in diameter. Exine 2 5 μm thickꎬ
sexine thicker than nexine. Columellae distinct. Orna ̄
mentation: finely reticualte under LM and SEM.
Vicia L.
Vicia cracca L. (Pl. II: 31ꎬ 32ꎻ Pl. V: 11)
Pollen grains 34 6 (25 0-67 5)×26 7 (17 5
-55 0) μmꎬ P / E= 1 44 (1 29-2 0)ꎬ prolate. E ̄
quatorial view elliptical. Polar view nearly 3 ̄lobed
circular. Apertures 3 ̄colporate. Ectoaperture ̄colpusꎬ
rather longꎬ extending to the poles. Endoaperture ̄
porusꎬ circularꎬ 5 0 μm in diameter. Exine 2 5 μm
thickꎬ sexine slightly thicker than nexine. Columel ̄
lae indistinct. Ornamentation: finely reticualte under
LM and SEM.
Celastraceae
Tripterygium Hook. f.
Tripterygium hypoglaucum (Levl.) Hutch (Pl.
II: 33ꎬ 34ꎻ Pl. V: 12ꎬ 13)
Pollen grains 30 0 (25 0-32 5)×27 1 (25 0
-30 0) μmꎬ P / E= 1 1 (1 0-1 3)ꎬ subspheriodal
to subprolate. Equatorial view subcircular. Polar view
3 ̄lobed circular. Apertures 3 ̄colporate. Ectoaperture ̄
colpusꎬ long and slenderꎬ extending to the poles. En ̄
doaperture ̄porusꎬ lalongate. Exine 3 0 μm thickꎬ sex ̄
ine 4 times as thick as nexine. Columellae developed.
Ornamentation: finely reticulate under LM and SEM.
Balsaminaceae
Impatiens L.
Impatiens uliginosa Franch. ( Pl. II: 35ꎬ Pl.
V: 14ꎬ 15)
Pollen grains 41 4 (35 0-42 5)×26 0 (25 0
-28 8) μmꎬ P / E= 1 59 (1 33-1 7)ꎬ prolate. E ̄
quatorial view oblate. Polar view rectangular. Aper ̄
tures goniotremeꎬ 4 ̄colpateꎬ colpus slender and short.
Exine 1 0 μm thickꎬ hierarchy indistinct. Ornamen ̄
tation: finely reticulate under LMꎻ distinct reticulate
with granule under SEM.
Impatiens radiata Hook. f. (Pl. II: 36ꎻ Pl. V:
16ꎬ 17)
Pollen grains 35 5 (32 5-37 5)×22 5 (17 5
-32 5) μmꎬ prolateꎬ P / E=1 56 (1 08-1 86). Equ ̄
atorial view oblate. Polar view rectangular. Apertures
goniotremeꎬ 4 ̄colpateꎬ colpus slender and short. Ex ̄
ine 1 5 μm thickꎬ sexine slightly thicker than nex ̄
ine. Ornamentation: reticulate under LMꎻ distinct
reticulate with few granules under SEM.
Impatiens procumbens Franch. (Pl. II: 37ꎻ Pl.
V: 18)
Pollen grains 41 4 (37 5-45 0)×27 0 (25 0
-30 0) μmꎬ prolateꎬ P / E = 1 53 (1 42-1 7). E ̄
quatorial view oblate. Polar view rectangular. Aper ̄
3666期 ZHENG Xin et al.: Pollen Morphology of Plants from the Hengduan Mountains and Their Ecological
tures goniotremeꎬ 4 ̄colpateꎬ colpus slender and short.
Exine 2 0 μm thickꎬ sexine slightly thicker than nex ̄
ine. Ornamentation: distinct coarsely reticulate under
LM and SEM.
Impatiens delavayi Franch. (Pl. II: 38ꎻ Pl. V: 19)
Pollen grains 41 3 (40 0-45 0)×29 5 (25 0
-32 5) μmꎬ prolateꎬ P / E = 1 40 (1 31-1 6). E ̄
quatorial view oblate. Polar view rectangular. Aper ̄
tures goniotremeꎬ 4 ̄colpateꎬ colpus slender and short.
Exine 2 0 μm thickꎬ sexine slightly thicker than nex ̄
ine. Ornamentation: reticulate under LMꎻ distinct ret ̄
iculate with granule under SEM.
Guttiferae
Hypericum L.
Hypericum addingtonii N. Robson (Pl. I: 39ꎬ
40ꎬ 41)
Pollen grains 26 8 (20 0-35 0)×22 8 (17 5
-30 0) μmꎬ P / E = 1 18 (1 0-1 3)ꎬ subprolate.
Equatorial view elliptical. Polar view 3 ̄lobed circu ̄
lar. Apertures 3 ̄colporate. Ectoaperture ̄colpusꎬ ex ̄
tending to the polesꎬ 3 0 μm in width. Endoaper ̄
ture ̄ porusꎬ lalongateꎬ 2 0 - 2 5 μm in diameter.
Exine 2 0 μm thickꎬ sexine equal to nexine. Colu ̄
mellae distinct. Ornamentation: faintly reticulate un ̄
der LM.
Begoniaceae
Begonia L.
Begonia grandis Dryand. subsp. sinensis (A. DC.)
Irmsch. (Pl. II: 42ꎻ Pl. VI: 20)
Pollen grains 69 7 (62 5-75 0)×33 1 (25 0
-37 5) μmꎬ P / E= 2 11 (1 86-2 5)ꎬ prolate. E ̄
quatorial view elliptical. Polar view triangle. Aper ̄
tures 3 ̄colporate. Ectoaperture ̄colpusꎬ long and slen ̄
derꎬ extending to the poles. Endoaperture ̄porusꎬ la ̄
longateꎬ 4 0 μm in diameter. Exine 1 5 μm thickꎬ
sexine equal to nexine. Columellae indistinct. Orna ̄
mentation: faintly reticulate under LMꎻ striate ̄perfo ̄
rate under SEM.
Melastomataceae
Osbeckia L.
Osbeckia crinita Benth. ex C B. Clarke (Pl. II:
43ꎬ 44ꎻ Pl. VI: 21)
Pollen grains 32 5 (30 0-35 0)×25 5 (22 5
-27 5) μmꎬ P / E = 1 27 (1 18-1 44)ꎬ prolate.
Equatorial view elliptical. Polar view 3 ̄lobed circu ̄
lar. Apertures 3 ̄colporate. Ectoaperture ̄colpusꎬ rather
long. Endoaperture ̄porusꎬ circularꎬ 5 0 μm in di ̄
ameter. Exine 1 5 μm thickꎬ sexine slightly thicker
than nexine. Ornamentation: smooth under LMꎻ gran ̄
ulate under SEM.
Onagraceae
Fuchsia L.
Fuchsia hybrida Hort. ex Sieb. and Voss. (Pl.
II: 45ꎬ 46ꎬ Pl. VI: 22)
Pollen grains 55 0 (32 5-82 5) μm in polar
diameterꎬ spheroidal. Equatorial view oblate. Polar
view triangularꎬ angles obtuse. Apertures 3 ̄porateꎬ
porus big and roundꎬ 15 0-18 0 μm in diameter.
Exine 2 5 μm thickꎬ sexine and nexine rather thic ̄
ker at the apertures. Columellae distinct. Ornamenta ̄
tion: finely reticulate under LMꎻ finely reticulate
with silk under SEM.
Oenothera L.
Oenothera rosea L. Herpt. ex Ait. (Pl. III: 47ꎻ
Pl. VI: 23)
Pollen grains 109 0 (100 0-120 0) μm in polar
diameterꎬ spheroidal. Equatorial view oblate. Polar
view triangularꎬ angles obtuse. Apertures 3 ̄porateꎬ
porus big and roundꎬ 45 0-50 0 μm in diameter.
Exine 7 0 μm thickꎬ sexine and nexine rather thic ̄
ker at the apertures. Columellae distinct. Ornamenta ̄
tion: faintly microreticulato ̄striate under LMꎻ gran ̄
ulate ̄perforate with silk under SEM.
Apiaceae
Bupleurum L.
Bupleurum longicaule Wall. & DC. ( Pl. III:
48ꎬ 49)
Pollen grains 24 0 (20 0-25 0)×14 0 (10 0
-15 0) μmꎬ P / E= 1 71 (1 67-2 0)ꎬ prolate. E ̄
quatorial view elliptical. Polar view triangleꎬ angles
round. Apertures 3 ̄colporate. Ectoaperture ̄colpusꎬ
long and slenderꎬ extending to the poles. Endoaper ̄
ture ̄porusꎬ lalongateꎬ circularꎬ 2 0-2 5 μm in di ̄
ameter. Exine 2 0 μm thickꎬ sexine slightly thicker
466 植 物 分 类 与 资 源 学 报 第 37卷
than nexine. Columellae distinct. Ornamentation: finely
reticulate under LM.
Heracleum L.
Heracleum franchetii M. Hiroe (Pl. III: 50ꎬ 51ꎻ
Pl. VI: 24)
Pollen grains 46 5 (42 5-52 5)×18 8 (17 5
-20 0) μmꎬ P / E = 2 48 (2 25-2 86)ꎬ perprolate.
Equatorial view elliptical. Polar view triangleꎬ angles
round. Apertures 3 ̄colporate. Ectoaperture ̄colpusꎬ
long and slenderꎬ extending to the poles. Endoaper ̄
ture ̄porusꎬ lalongateꎬ circularꎬ 2 5 μm in diameter.
Exine 2 5 - 3 0 μm thickꎬ sexine slightly thicker
than nexine. Columellae distinct. Ornamentation: finely
reticulate under LMꎻ striate ̄perforate under SEM.
Ericaceae
Lyonia Nutt.
Lyonia ovalifolia (Wall.) Drude (Pl. III: 52ꎬ
53ꎻ Pl. VI: 25)
Pollen grains 35 0 (22 5-45 0) μm in diam ̄
eterꎬ tetrahedral tetrad or decussate tetrad. Every
single pollen grain are subspheroidal with 3 ̄colpor ̄
ateꎬ which shaped as half ̄colpi due to two neighbor ̄
ing grains of tetrahedral tetrad link together. Exine
1 5 μm thickꎬ sexine slightly thicker than nexine.
Columellae indistinct. Ornamentation: finely reticu ̄
late under LMꎻ distinct reticulate under SEM.
Primulaceae
Lysimachia L.
Lysimachia violascens Franch. (Pl. III: 54ꎬ 55ꎻ
Pl. VI: 26)
Pollen grains 30 8 (25 0-37 5)×20 5 (17 5
-22 5) μmꎬ P / E= 1 5 (1 22-1 75)ꎬ prolate. E ̄
quatorial view elliptical. Polar view 3 ̄lobed circular.
Apertures 3 ̄colporate. Ectoaperture ̄colpusꎬ long and
slenderꎬ extending to the polesꎬ 1 0 μm in width.
Endoaperture ̄porusꎬ lalongateꎬ cross with colpusꎬ
3 0 μm in diameter. Exine 1 5 μm thickꎬ sexine
slightly thicker than nexine. Columellae distinct. Or ̄
namentation: finely reticulate under LM and SEM.
Loganiaceae
Buddleja (Buddleia auct.) L.
Buddleja fallowiana Balf. F. and W. W. Sm. (Pl.
III: 56ꎬ 57ꎻ Pl. VI: 27)
Pollen grains 21 6 (17 5-22 5)×14 7 (12 5
-17 5) μmꎬ P / E = 1 47 (1 14-1 8)ꎬ subprolate
to prolate. Equatorial view elliptical. Polar view 4 ̄
lobed circular. Apertures 4 ̄colporate. Ectoaperture ̄
colpusꎬ long and slenderꎬ extending to the polesꎬ
1 0 μm in width. Endoaperture ̄porusꎬ lalongateꎬ
margins at ends diffuse. Exine 1 5 μm thickꎬ indis ̄
tinct in layer. Exine at poles thicker than that at e ̄
quatorꎬ nexine slightly thicker than exine. Columel ̄
lae distinct. Ornamentation: finely reticulate under
LM and SEM.
Buddleja forrestii Diels ( Pl. III: 58ꎬ 59ꎻ Pl.
VI: 28)
Pollen grains 24 5 (22 5-27 5)×19 5 (17 5
-22 5) μmꎬ P / E= 1 25 (1 11-1 42)ꎬ subprolate
to prolate. Equatorial view rectangular. Polar view 4 ̄
lobed circular. Apertures 4 ̄colporate. Ectoaperture ̄
colpusꎬ short. Endoaperture ̄porusꎬ lalongateꎬ 5 0-
6 0 μm in diameter. Exine 1 5 μm thickꎬ indistinct
in layerꎬ exine at poles thicker than that at equatorꎬ
nexine slightly thicker than exine. Columellae indis ̄
tinct. Ornamentation: finely reticulate under LM and
SEM.
Gentianaceae
Halenia Borkh.
Halenia ellipticala D. Don ( Pl. III: 60ꎬ 61ꎻ
Pl. VI: 29ꎬ 30)
Pollen grains 38 8 (35 0-45 0)×36 5 (35 0
-37 5) μmꎬ P / E= 1 06 (1 0-1 2)ꎬ spheroidal to
subprolate. Equatorial view elliptical. Polar view 3 ̄
lobed circular. Apertures 3 ̄colporate. Ectoaperture ̄
colpusꎬ rather longꎬ extending to the poles. Endoap ̄
erture ̄porusꎬ circularꎬ 3 0 μm in diameterꎬ with one
crack respectively on the two sides. Exine 2 5 μm
thickꎬ sexine slightly thicker than nexine. Columel ̄
lae indistinct. Ornamentation: finely reticulate under
LMꎬ distinct reticulate under SEM.
Gentianopsis Ma
Gentianopsis paludosa (Hook f.) Ma (Pl. III:
62ꎬ 63)
Pollen grains 17 3 (10 0-22 5)×15 5 (10 0
5666期 ZHENG Xin et al.: Pollen Morphology of Plants from the Hengduan Mountains and Their Ecological
-22 5) μmꎬ P / E= 1 11 (1 0-1 5)ꎬ spheroidal to
subprolate. Equatorial view elliptical. Polar view 3
(4) ̄lobed circular. Apertures 3 (4) ̄colporate. Ec ̄
toaperture ̄colpusꎬ rather longꎬ colpus membrane with
granules or tumors. Endoaperture ̄porusꎬ indistinct.
Exine 3 0 μm thickꎬ sexine 2 times as thick as nex ̄
ine. Columellae developed. Ornamentation: distinct
reticulate under LM.
Labiatae
Ajuga L.
Ajuga forrestii Diels (Pl. III: 64ꎻ Pl. VII: 31)
Pollen grains 26 4 (15 0-45 0)×20 9 (12 5
-35 0) μmꎬ P / E=1 26 (1 12-1 6)ꎬ subprolate to
prolate. Equatorial view elliptical. Polar view 3 ̄lobed
circular. Apertures 3 ̄colpateꎬ colpus longꎬ extending
to the polesꎬ ends narrow. Exine 2 0 μmꎬ sexine 3
times as thick as nexine. Columellae developed. Ex ̄
ine much thicker at the poles by equatorial view. Or ̄
namentation: finely reticulate under LM and SEM.
Nepeta L.
Nepeta wilsonii Duthie (Pl. III: 65ꎬ 66)
Pollen grains 56 8 (50 0-62 5)×44 3 (32 5
-55 0) μmꎬ P / E = 1 28 (1 1-1 77)ꎬ subprolate
to prolate. Equatorial view elliptical. Polar view 6 ̄
lobed circular. Apertures 6 ̄colpateꎬ colpus longꎬ ex ̄
tending to the poles. Exine 2 0 μm thickꎬ sexine e ̄
qual to nexine. Columellae distinct. Ornamentation:
distinct reticulate under LM.
Clinopodium L.
Clinopodium megalanthum (Diels) C Y. Wu and
Hsuan ex H W. Li (Pl. III: 67ꎬ 68ꎻ Pl. VII: 32)
Pollen grains 47 8 (20 0-57 5)×36 3 (15 0
-47 5) μmꎬ P / E= 1 31 (1 17-1 89)ꎬ subprolate
to prolate. Equatorial view elliptical. Polar view 6 ̄
lobed circular. Apertures 6 ̄colpateꎬ colpus longꎬ ex ̄
tending to the poles. Exine 2 5 μm thickꎬ sexine
slightly thicker than or equal to nexine. Columellae
developed. Ornamentation: finely reticulate under LM
and SEM.
Elsholtzia Willd.
Elsholtzia rugulosa Hemsl. ( Pl. III: 69ꎬ 70ꎻ
Pl. VII: 33)
Pollen grains 28 9 (25 0-32 5)×20 0 (17 5
-22 5) μmꎬ P / E = 1 44 (1 33-1 71)ꎬ prolate.
Equatorial view elliptical. Polar view 6 ̄lobed circu ̄
lar. Apertures 6 ̄colpateꎬ colpus long and slenderꎬ
extending to the polesꎬ 1 μm wideꎬ ends narrow.
Exine 2 0 μm thickꎬ sexine equal to nexine. Colu ̄
mellae distinct. Ornamentation: finely reticulate un ̄
der LM and SEM.
Scrophulariaceae
Verbascum L.
Verbascum thapsus L. (Pl. IV: 71ꎬ 72ꎬ 73ꎻ Pl.
VII: 34ꎬ 35)
Pollen grains 28 5 (25 0-32 5)×22 8 (15 0
-25 0) μmꎬ P / E= 1 25 (1 1-1 83)ꎬ subspheroi ̄
dal to subprolate. Equatorial view elliptical. Polar
view 3 ̄lobed circular. Apertures 3 ̄colporoid. Exine
2 0 μm thickꎬ sexine equal to nexine. Columellae
distinct. Ornamentation: finely reticulate under LMꎻ
distinctly reticulate under SEM.
Pedicularis L.
Pedicularis tenuisecta Franch. ex Maxim. ( Pl.
IV: 74ꎬ 75ꎻ Pl. VII: 36ꎬ 37)
Pollen grains 28 3 (25 0-30 0)×16 3 (10 0
-20 0) μmꎬ P / E= 1 74 (1 5-3 0)ꎬ flatꎬ bilater ̄
ally symmetrical. Apertures 2 ̄syncolpate with the
ends of two colpi anastomose at the polesꎬ colpus
cutting pollen grain into equal halves. Exine 2 0 μm
thick. Sexine slightly thicker than nexine. Ornamenta ̄
tion: almost smooth under LMꎻ perforate under SEM.
Pedicularis gruina Franch. ex Maxim. (Pl. IV:
76ꎬ 77)
Pollen grains 36 8 (33 0-42 0)×30 8 (27 5
-34 0) μmꎬ P / E = 1 19 (1 06-1 31)ꎬ flatꎬ bi ̄
laterally symmetrical. Apertures 2 ̄syncolpate with
the ends of two colpi anastomose at the polesꎬ colpus
cutting pollen grain into equal halves. Exine 2 0 μm
thickꎬ sexine slightly thicker than nexine. Columel ̄
lae indistinct. Ornamentation: almost smooth under
LMꎻ perforate under SEM.
Bignoniaceae
Incarvillea Juss.
Incarvillea arguta (Royle) Royle (Pl. IV: 78ꎬ
666 植 物 分 类 与 资 源 学 报 第 37卷
Pl. VII: 38ꎬ 39)
Pollen grains 24 5 (22 5-27 5)×19 5 (17 5
- 22 5) μmꎬ subprolate to prolateꎬ P / E = 1 25
(1 11-1 42). Equatorial view elliptical. Polar view
6 ( ̄7) ̄lobed circular. Apertures 6 ( ̄7) ̄colpateꎬ
colpus longꎬ extending to the polesꎬ colpus mem ̄
brane beset with fine and distinct granules. Exine
2 0 μm thickꎬ sexine equal to nexine. Columellae
distinct. Ornamentation: finely granulate ̄perforate un ̄
der LMꎻ microechinate ̄perforate under SEM.
Acanthaceae
Pteracanthus (Nees) Bremek.
Pteracanthus forrestii (Diels) C Y. Wu (Pl. IV:
79)
Pollen grains 101 6 (90 0-130 0)×61 6 (42 5
-77 5) μmꎬ P / E = 1 65 (1 35-2 12)ꎬ prolate.
Equatorial view elliptical. Polar view 18 ( ̄21) ̄lobed
circular. Apertures 3 ̄colporate and 15 - 18 pseud ̄
ocolporate. Ectoaperture ̄colpusꎬ long and slender.
Endoaperture ̄porusꎬ circularꎬ 3 μm in diameter. Ex ̄
ine 2 5 μm thickꎬ sexine 2 times as thick as nexine.
Columellae distinct. Ornamentation: coarsely reticu ̄
late under LMꎬ sexine semitectateꎬ reticulate with
developed lumina and muriꎬ muri densely compri ̄
sing equirotal granules.
Valerianaceae
Valeriana L.
Valeriana flaccidissima Maxim. (Pl. IV: 80ꎬ 81ꎬ
82ꎻ Pl. VII: 40)
Pollen grains 45 5 (30 0-62 5)×41 3 (27 5
- 57 5) μmꎬ subspheroidal to subprolateꎬ P / E =
1 1 (1 0-1 18). Equatorial view elliptical. Polar
view 3 ̄lobed circular. Apertures 3 ̄colpateꎬ colpus
long and wide. Exine 3 5 μm thickꎬ sexine equal to
nexine. Columellae distinct. Ornamentation: micro ̄
echinate under LMꎻ sparsely spinuloseꎬ 1 3-1 6 μm
long under SEM.
Cucurbitaceae
Zehneria Endl.
Zehneria maysorensis (Wight and Arn.) Arn.
(Pl. IV: 83ꎬ 84ꎻ Pl. VII: 41)
Pollen grains 58 0 (50 0-75 0)×49 0 (40 0
-55 0) μmꎬ P / E = 1 18 (1 0-1 67)ꎬ subprolate
to prolate. Equatorial view elliptical. Polar view 3 ̄
lobed circular. Apertures 3 ̄colporate. Ectoaperture ̄
colpusꎬ colpus short and narrow. Endoaperture ̄porusꎬ
circularꎬ 6 0 μm in diameter. Exine 2 0 μm thickꎬ
sexine slightly thicker than nexine. Columellae dis ̄
tinct. Ornamentation: finely reticulate under LMꎻ
distinct finely reticulate under SEM.
Campanulaceae
Campanula L.
Campanula colorata Wall. ( Pl. IV: 85ꎬ 86ꎻ
Pl. VII: 42ꎬ 43)
Pollen grains 25 5 (22 5-27 5) μmꎬ spheroi ̄
dal. Apertures 3 ̄porateꎬ porus membrane beset with
granules. Exine 2 0 μm thickꎬ sexine equal to nex ̄
ine. Ornamentation: spinulose under LMꎻ prominent ̄
ly spinuloseꎬ surface between spinules striate ̄perfo ̄
rate under SEM.
Lobelia L.
Lobelia doniana Skottsb. (Pl. IV: 87ꎬ 88ꎬ 89ꎻ
Pl. VII: 44ꎬ 45)
Pollen grains 44 3 (32 5-60 0)×31 3 (25 0
-35 0) μmꎬ P / E = 1 42 (1 29-1 71)ꎬ prolate.
Equatorial view elliptical. Polar view 3 ̄lobed circu ̄
lar. Apertures 3 ̄colporate. Ectoaperture ̄colpusꎬ longꎬ
extending to the poles. Endoaperture ̄porusꎬ lalon ̄
gate. Exine 2 5 μmꎬ sexine slightly thicker than or
equal to nexine. Columellae distinct. Ornamentation:
finely reticulate under LM and SEM.
Compositae
Saussurea DC.
Saussurea stella Maxim. (Pl. IV: 90)
Pollen grains 66 7 (65 0-70 0)×60 0 (55 0
-65 0) μmꎬ P / E = 1 11 (1 07-1 18)ꎬ subsphe ̄
roidal to subprolate. Equatorial view elliptical. Polar
view 3 ̄lobed circular. Apertures 3 ̄colporate. Ectoap ̄
erture ̄colpusꎬ short. Endoaperture ̄porusꎬ narrow.
Exine 9 0 μm thick ( not including spine length).
Columellae distinct. Ornamentation: sexine tectate with
4 μm long spinules under LM.
2 3 Q ̄type Hierarchical cluster and classification
Q ̄type Hierarchical cluster analysis were con ̄
7666期 ZHENG Xin et al.: Pollen Morphology of Plants from the Hengduan Mountains and Their Ecological
ducted using Between ̄groups Linkage method. Clus ̄
ter dendrogram of 48 species classified on the basis
of the code matrix for 3 quantitative traits and 2
qualitative traits was shown in Fig 1ꎬ which indica ̄
ted that four major clades (Aꎬ Bꎬ Cꎬ D) were iden ̄
tified at the dissimilarity coefficient of L2 (0 292).
Clade Aꎬ Clade B and Clade C consisted of the spe ̄
cies which belonged to Archichlamydeae mostly.
Clade D consisted of the species of Metachlamydeae
and Archichlamydeae. Besidesꎬ we could found that
at the dissimilarity coefficient of L1 ( 0 242)ꎬ 6
small clades are all comprised of species in the same
familyꎬ such as Ranunculaceaeꎬ Balsaminaceaeꎬ Log ̄
aniaceaeꎬ Caryophyllaceaeꎬ Scrophulariaceaeꎬ Labi ̄
atae. Howeverꎬ some species in the same family were
divided into different cladesꎬ such as the Ranuncu ̄
laceaeꎬ Fabaceaeꎬ Labiatae etc.
3 Discussion
3 1 Cluster analysis
Pollen morphology has long been used as the
evidence for species classification by phytogeogra ̄
phers (Zhang et al.ꎬ 1990ꎻ Carrijo et al.ꎬ 2013).
Pollen grains were not affected by environment easilyꎬ
and inherited from generation to generationꎬ which
could objectively reflect the interspecific relationship
(Zhangꎬ 2004ꎻ Wu et al.ꎬ 2011). It was suggested
that Ranalesꎬ Rosalesꎬ Sapindalesꎬ Parietalesꎬ Myr ̄
tifloraeꎬ Umbelliflorae and Cucurbitales could have
the closed relationship according to the clade A in the
dendrogram aboveꎬ the pollen grains of which had 3 ̄
colporate or 3 ̄colpate apertures mostly. It was sup ̄
ported by relationships in the Engler System. Howeverꎬ
the species of other clades didn’t show much consis ̄
tency with phylogentic relationships. It was likely that
we didn’t get enough characters data and couldn’t
reveal the relationships between species in different
families with pollen morphology method only. We
needed to analyze syntheticly combining with pollen
morphologyꎬ characters of botany and molecular bi ̄
ology methods to discuss relationships between dif ̄
ferent families further. Fig 1 Dendrogram of 48 species
866 植 物 分 类 与 资 源 学 报 第 37卷
3 2 Pollen identification features in the same family
Based on the cluster dendrogram aboveꎬ some
species in the same family are divided into different
clades. Thus the similarities and dissimilarities of
pollen morphology among these species in the same
family are provided.
Ranunculaceae pollen is divided into four types
basing on apertures. These four types are inaperturate
typeꎬ 3 ̄colpate typeꎬ pantoporate type and stephano ̄
colpate ̄pantocolpate typeꎬ respectively (Wang et al.ꎬ
1995). Pollen grains of Delphinium delavayi and A ̄
nemone hupehensis both belong to 3 ̄colpate typeꎬ
and they assemble in a small clade. While pollen
grains of Thalictrum delavayi have pantoporate aper ̄
tures and consist in another clade. Description of the
three species apertures accord with the results of past
studies (Xi and Zhangꎬ 1964ꎻ Wang et al.ꎬ 1995ꎻ
Yang et al.ꎬ 1995). Besidesꎬ there are some simi ̄
larities among pollen morphology of the three spe ̄
cies: pollen grains subspheroidal to spheroidalꎬ or ̄
namentation finely granulate under LM.
Fabaceae contains three subfamilies. All the four
species in this research belong to Papilionoideae (Chen
et al.ꎬ 1994). Their pollen morphology are similar:
pollen grains prolateꎬ ornamentation distinct reticu ̄
late in LM. Aperture types are the main differences
to divide the samples into two clades. Pollen grains
of Cochlianthus montanusꎬ Pueraria peduncularis.
and Vicia cracca are 3 ̄colporate. While pollen grains
of Clitoria mariana are 5 ( ̄6) ̄porate. Wang et al.
(1995) found that there are also other aperture
types such as 3 ̄colpateꎬ 3 ̄porateꎬ and 6 ̄porate ex ̄
cept for 3 ̄colporate which is the aperture type of
most Fabaceae species. In additionꎬ Ferguson and
Skvarla (1979) showed that the Cranocarpus genus
species have 5 ̄porate aperture.
Labiatae pollen in this research are divided into
two main types: 3 ̄colpate type and 6 ̄colpate type.
Pollen grains of Ajuga forrestii Diels belong to 3 ̄col ̄
pate typeꎬ while Nepeta wilsoniiꎬ Clinopodium mega ̄
lanthum and Elsholtzia rugulosa belong to 6 ̄colpate
type. In additionꎬ Elsholtzia rugulosa is separated
from Nepeta wilsonii and Clinopodium megalanthum
because of their different pollen grains size. Orna ̄
mentation of the four species pollen grains are simi ̄
lar under LMꎬ distinctly reticulate. Hu et al. (2012)
compared and studied pollen morphology of three
genera of Labiatae. They suggested that based on the
differences of pollen grains ornamentations and shapesꎬ
species of different genera of Labiatae can be classi ̄
fied. Thereforeꎬ according to their taxonomic meth ̄
odsꎬ exine ornamentation of pollen grains under SEM
of Clinopodium megalanthumꎬ Ajuga forrestii and
Elsholtzia rugulosa have shown heteromorphic char ̄
acteristicsꎬ they are reticulateꎬ irregularly reticulate
and dually reticulate ( small reticulum nested within
big lumia) respectively.
Besidesꎬ some species in the same family were
not classified basing on our selected characters in the
cluster dendrogramꎬ since their pollen grains were
similar generallyꎬ i e. Balsaminaceae. All the four
Balsaminaceae species in this research belong to Im ̄
patiens L. They possess the same pollen features:
pollen grains goniotreme and zygomorphicꎬ 4 ̄col ̄
pate. But they are different in the shape and orna ̄
mentation. Thereforeꎬ according to two taxonomic
methods of Cai et al. (2007)ꎬ different types in the
species we tested can be identified. Based on the
shape of pollenꎬ two types are identified. Pollen
grains of Impatiens uliginosa are elliptic typesꎬ and
pollen grains of the other three are rectangular types.
Based on the width of muri and luminaꎬ two types
are aslo identified. Ornamentation of Impatiens uligi ̄
nosa and Impatiens radiata belong to finely reticulate
typeꎬ and that of Impatiens delavayi and Impatiens
procumbens belong to coarsely reticulate type.
3 3 Ecological significance of main pollen types
Ma et al. (2009) made taxonomic identification
of Coniferopsida and Compositae pollens which exist
broadly in sedimentary strataꎬ and inferred the eco ̄
logical significance of pollen types according to the
ecological properties of these taxa. Yang et al. (2013)
observed the pollen morphology of nine hygrophyte
species from Seven Star Lake area and discussed the
9666期 ZHENG Xin et al.: Pollen Morphology of Plants from the Hengduan Mountains and Their Ecological
environmental indication significance based on the
habitatsꎬ origin and distribution of these species.
Thus using the similar methodsꎬ the ecological sig ̄
nificance of the main pollen types are discussed.
Impatiens L. in Balsaminaceae are mainly dis ̄
tributed across Asian tropical and subtropical zones
and in Africa. There are about 220 species distribu ̄
ted in China. Impatiens radiata grows mostly on shady
hillsides or in humid places under forest cover with
the altitude of 2 100-3 500 m in the southwestern
provinces of China. Because the southwestern China
climate is mainly subtropical monsoonal and I radiate
grows in a humid and shady habitat of mountains.
Thereforeꎬ pollen of I radiate can be an indicator for
warm and dank climates of the north subtropical zone
and also for a medium ̄ and high ̄ mountain environ ̄
ment in southwestern China.
Oenothera L. in Onagraceae are native to America
temperate and subtropical zones. There are 19-20 spe ̄
cies introduced into China. Oenothera rosea grows most ̄
ly on grassland and in semi ̄shady places along ditches
with the altitude of 1 000-2 000 mꎬ and this species
naturalizes well in Zhejiangꎬ Jiangxi (Lushan Moun ̄
tain)ꎬ Yunnan (Kunming)ꎬ Guizhou Provinces. Be ̄
cause these regions climate are mainly subtropical
monsoonal and O rosea grows along ditches. Thus pol ̄
len of O rosea may serve as an indicator for warm and
humid climates. It can also indicate a medium ̄ and
low ̄mountain environment due to the habitat of O rosea
is on the mountains with the altitude of 1 000-2 000 m.
Lyonia Nutt. in Ericaceae are mainly distributed
in eastern Asia. There are about 6 speciesꎬ 5 vari ̄
ants in China. Lyonia ovalifolia grows mainly in the
forests with the altitude of 700-2 800 m and is dis ̄
tributed in Fujianꎬ Guangdongꎬ Guangxiꎬ Sichuanꎬ
Guizhou Provincesꎬ etc. Because these regions cli ̄
mate are mainly subtropical monsoonal and humid.
Pollen of L ovalifolia may serve as an indicator for
warm and humid climates. Description of L ovalifolia
pollen can provide identification characteristic of Eri ̄
caceae in Quaternary strata.
Clinopodium L. in Labiatae are mainly distribu ̄
ted in Europeꎬ Central Asia and eastern Asia. There
are about 11 speciesꎬ 6 varieties in China. Clinopodium
megalanthum grows mainly on hillsidesꎬ grasslandsꎬ
roadsidesꎬ and among shrubs and underbrush with the
altitude of 1 300-3 200 mꎬ and is distributes in Yun ̄
nan Provinceꎬ southwest of Sichuan Provinceꎬ south ̄
west of Hubei Province and north of Guizhou Prov ̄
ince. Because these regions climate are mainly sub ̄
tropical monsoonal and humid. Thus pollen of C meg ̄
alanthum may serve as an indicator for warm and
humid climates of the southern subtopical zones.
Pedicularis L. in Scrophulariaceae contains more
than 500 speciesꎬ mainly distributed in the Frigid
Zone and high mountains of the Northern Hemi ̄
sphere. There are about 340 species in China. Pedic ̄
ularis tenuisecta grows mainly in grasslands and on
the edges of cypress forests with the altitude of 1 500
-3 660 m and is distributes in southwest of Sichuan
Provinceꎬ northwest of Yunnan Province and west of
Guizhou Province. Because these regions are mainly
high mountains and climate of these region are humid.
Thus pollen of P tenuisecta may serve as an indicator
for cool and humid climates of the subtropical zoneꎬ
and also for a cool and humid alpine environment.
Saussurea DC. in Asteraceae are mainly distrib ̄
uted in Asia and Europe. There are about 264 spe ̄
cies in China. Saussurea stella grows mainly in al ̄
pine meadows and on rocky beaches in a cold or dry
environment with the altitude of 2 000-5 400 mꎬ and
is distributed in areas at high elevation of Gansuꎬ Si ̄
chuanꎬ Yunnan and Xizang Province. Thereforeꎬ pol ̄
len of S stella may act as an indicator for the cool
climatesꎬ and also for a high mountain environment.
Cyanotis D. Don in Commelinaceae are mainly
distributed in the tropical and subtropical zones of
Asia and Africa. Cyanotis vaga grows mainly on hill ̄
side grasslands or under open forest with the altitude
below 3 300 m and is distributed in Guangdongꎬ
Guizhouꎬ Sichuanꎬ Yunnan Provincesꎬ etc. These
regions are mainly subtropical monsoonal and humid.
Thereforeꎬ pollen of C vaga may serve as an indica ̄
tor for warm and humid climates in the tropical zone.
076 植 物 分 类 与 资 源 学 报 第 37卷
Acknowledgements: We should like to thank Mr. John Ols ̄
en for his help with the English language.
References:
施雅风 (Shi YF)ꎬ 李吉均 (Li JJ)ꎬ 李炳元 (Li BY)ꎬ 1998. 青藏
高原晚新生代隆起与环境变化 [M]. 广州: 广东科学技术
出版社
吴学尉 (Wu XW)ꎬ王丽花 (Wang LH)ꎬ张艺萍 (Zhang YP) 等ꎬ
2011. 8种野生百合的花粉形态及聚类分析 [ J] . 江苏农业
科学ꎬ 39 (3): 197—199
席以珍 (Xi YZ)ꎬ 张金谈 (Zhang JT)ꎬ 1964. 银莲花属 (Anemone
L.) 花粉形态的研究 [J] . 植物学报
张金谈 (Zhang JT)ꎬ 王萍莉 (Wang PL)ꎬ 郝海平 (Hao HP)ꎬ
1990. 现代花粉应用研究 [M] . 北京: 科学出版社
Boufford DEꎬ Dijk PPVꎬ Zhi Lꎬ 2004. Mountain of Southwest China
[A] / / Mittermeier RAꎬ Robles ̄Gil Pꎬ Hoffmann M et al eds.ꎬ
Hotspots Revisited: Earth’s Biologically Richest and Most Endan ̄
gered Ecoregionsꎬ 2nd [M]. Mexico: Cemexꎬ 159—164
Cai XZ (蔡秀珍)ꎬ Liu KM (刘克明)ꎬ Cong YY (丛义艳) et al.ꎬ 2007.
SEM Observation on the pollen grains of ten species in Impatiens L.
(Balsaminaceae) [J]. Bulletin of Botanical Researchꎬ 279—283
Carrijo TTꎬ Garbin MLꎬ Leite WP et al.ꎬ 2013. Pollen morphology of
some related genera of Vernonieae (Asteraceae) and its taxonomic
significance [ J ] . Plant Systematics & Evolutionꎬ 299 ( 7 ):
1275—1283
Chen DZ (陈德昭)ꎬ Chen BY (陈邦余)ꎬ Fang YY (方云忆) et
al.ꎬ 1994. Fabaceae [A] / / Flora Reipubliceae Popularis Sini ̄
caeꎬ Volume 40 [M]. Beijing: Sciences Press
Erdtman Gꎬ 1971. Pollen Morphology and Plant Taxonomy: Angio ̄
sperms. An Introduction to Palynology. I [M]. New York: Hafner
Publishing Coꎬ 1—539
Ferguson IKꎬ Skvarla JJꎬ 1979. The pollen morphology of Cranocarpus
martii Bentham ( Leguminosae: Papilionoideae) [ J] . Granaꎬ
18: 15—20
Hesse Mꎬ Halbritter Hꎬ Zetter R et al.ꎬ 2009. Pollen Terminology: An
Illustrated Handbook [M]. New York: Springer Wienꎬ 1—264
Hu Y (胡彦)ꎬ Ding YF (丁友芳)ꎬ Wen CX (温春秀) et al.ꎬ
2012. Study on the pollen morphology of three Genera of Lami ̄
aceae [J] . Northern Horticultureꎬ 11: 60—64
Kramer Aꎬ Herzschuh Uꎬ Mischke S et al.ꎬ 2010. Holocene treeline
shifts and monsoon variability in the Hengduan Mountains
(southeastern Tibetan Plateau)ꎬ implications from palynological
investigations [J]. Palaeogeographyꎬ Palaeoclimatologyꎬ Palae ̄
oecologyꎬ 286 (1): 23—41
Kupriyanova LAꎬ Aleshina LAꎬ 1972. Pollen and Spores of Plants
From the Flora of European Part of the USSR. I [M]. Lenin ̄
grad: Naukaꎬ 1—171
Li CH (李春海)ꎬ Tang LY (唐领余)ꎬ Feng ZD (冯兆东)ꎬ 2006.
A high ̄resolution late Pleistocene record of pollen vegetation and
climate change from Jingningꎬ NW China [J] . Science in China
Series D (中国科学 D辑)ꎬ 36 (5): 453—460
Li XW (李锡文)ꎬ Li J (李捷)ꎬ 1993. A preliminary floristic study
on the seed plants from the region of Hengduan Mountain [ J] .
Acta Botanica Yunnanica (云南植物研究)ꎬ 15 (3): 217—231
Li Zꎬ Zhang Qꎬ Ma Kꎬ 2012. Tree ̄ring reconstruction of summer tem ̄
perature for A. D. 1475-2003 in the central Hengduan Mountainsꎬ
Northwestern Yunnanꎬ China [J] . General Informationꎬ 110 (1 ̄
2): 455—467
Liu SZ (刘淑珍)ꎬ Chai ZX (柴宗新)ꎬ Chen JL (陈继良)ꎬ 1986.
Function of Quaternary Glaciation in the northern Hengduan
Mountains [A] / / Special Issue of Hengduan Mounatins Scientific
Expedition ( II) (横断山考察专集 II) [M]. Beijing: Science
and Technology Press
Ma HJ (马宏杰)ꎬ 2013. Research on Cenozoic stratigraphy and
Palaeoenvironmental change in the Hengduan Mountainsꎬ South ̄
west China [D]. Kunming: Kunming University of Science and
Technology (昆明理工大学)
Ma YZ (马玉贞)ꎬ Meng HW (蒙红卫)ꎬ Sang YL (桑艳礼) et
al.ꎬ 2009. Pollen keys for identification of coniferopsida and
compositae classes under light microscopy and their ecological
significance [J] . Acta Palaeontologica Sinica (古生物学报)ꎬ
48 (2): 240—253
Pan YS (潘裕生)ꎬ 1989. Division of geologic structure in the Hengd ̄
uan mountainous region [J] . Journal of Mountain Research (山
地研究)ꎬ 7 (1): 3—12
Punt Wꎬ Hoen PPꎬ Blackmore S et al.ꎬ 2007. Glossary of pollen and
spore terminology [J] . Review of Palaeobotany and Palynologyꎬ
143: 1—81
Tang LYꎬ 2002. Temporal ̄spatial distribution of vegetation in the
Qinghai ̄Xizang plateau during the past 12 ka BP [ J] . Journal
of Integrative Plant Biologyꎬ 44 (7): 872—877
Tang LY (唐领余)ꎬ Shen CM (沈才明)ꎬ Li CH (李春海) et al.ꎬ
2009. Pollen record of vegetation and environmental changes in
the central Tibetan Plateau since the Holocene [ J] . Science in
China Series D: Journal of Earth Science (中国科学 D 辑: 地
球科学)ꎬ 39 (5): 615—625
Wang WC (王文采)ꎬ 1992. On some distribution patterns and some
migration routes found in the Eastern Asiatic region [ J] . Acta
Phytotaxonomica Sinica (植物分类学报)ꎬ 30: 1—24
Wang FX (王伏雄)ꎬ Qian NF (钱南芬)ꎬ Zhang YL (张玉龙) et
al.ꎬ 1995. Pollen Flora of China (中国植物花粉形态) [M].
Beijing: Science Press
Wodehouse RPꎬ 1959. Pollen Grains: Their Structureꎬ Identificationꎬ
and Significance in Science and Medicine [ M]. New York:
Hafnerꎬ 1—1935
Wu ZY (吴征镒)ꎬ 1979. The regionalization of Chinese flora [ J] .
Acta Botanica Yunnanica (云南植物研究)ꎬ 1 (1): 1—22
Wu ZY (吴征镒)ꎬ 1980. The Vegetation of China (中国植被)
[M]. Beijing: Science Press
1766期 ZHENG Xin et al.: Pollen Morphology of Plants from the Hengduan Mountains and Their Ecological
Wu ZY (吴征镒)ꎬ 1987. Origin and Evolution of Flora of Tibet [A]
/ / Wu CY ed.ꎬ Flora of Tibet (Vol 5) ( 西藏植物志第 5卷)
[M]. Beijing: Science Press
Yang YH (杨永红)ꎬ Yang CG (杨灿光)ꎬ Cui RZ (崔汝正) et
al.ꎬ 1995. SEM observation of the pollen of genus Delphinium in
China [ J] . Journal of Yunnan Agricultural University (云南农
业大学学报)ꎬ 10 (4): 263—267
Yang Zꎬ Yi Tꎬ Pan Y et al.ꎬ 2012. Phylogeography of an alpine plant
Ligularia vellerea ( Asteraceae ) in the Hengduan Mountains
[J] . Journal of Systematics and Evolutionꎬ 50 (4): 316—324
Yang Xꎬ Zhang Rꎬ Jiang H et al.ꎬ 2013. Pollen morphology and eco ̄
logical significance of nine hygrophyte species at sevenstar lake
area in the bashang area of Hebei Province [ J] . Agricultural
Science & Technologyꎬ 14 (7): 962—965
Yao YFꎬ Song XYꎬ Wortley AH et al.ꎬ 2015. A 22 570 ̄year record of
vegetational and climatic change from Wenhai Lake in the Hengd ̄
uan Mountains biodiversity hotspotꎬ Yunnanꎬ Southwest China
[J] . Biogeosciencesꎬ 12 (5): 1525—1535
Zhang YM (张元明)ꎬ 2004. Cluster analysis on pollen morphology of
the Tamaricaceae from China [ J] . Acta Botanica Boreali ̄Occi ̄
dentalia Sinica (西北植物学报)ꎬ 24 (9): 1702—1707
Zhao XX (赵秀霞)ꎬ Zhou ZZ (周忠泽)ꎬ Wang WG (汪文革)ꎬ
2006. Pollen morphology and ecological factors of Alnus trabecu ̄
losa [J] . Acta Micropalaeontologica Sinica (微体古生物学
报)ꎬ 23 (4): 419—424
Explanation of Plates
Plate I Pollen morphology and exine ornamentation under LM
1. Cyanotis vagaꎻ 2. Tofieldia divergensꎻ 3 - 4: P chinense L. var.
paradoxumꎻ 5. Silene yunnanensisꎻ 6. Silene bacciferaꎻ 7-8. Delphini ̄
um delavayiꎻ 9-11. Thalictrum delavayiꎻ 12-13. Anemone hupehensis
Lem f. albaꎻ 14-16. Astilbe rivularisꎻ 17-18: Sorbaria arborea var.
subtomentosaꎻ 19-21. Potentilla fulgensꎻ 22- 23. Spenceria ramala ̄
naꎻ 24- 25. Cochlianthus montanusꎻ 26- 28. Pueraria peduncularis.
Scales bar 10 μm
Plate II Pollen morphology and exine ornamentation under LM
29-30. Clitoria marianaꎻ 31-32. Vicia craccaꎻ 33-34. Tripterygium
hypoglaucumꎻ 35. Impatiens uliginosaꎻ 36. Impatiens radiataꎻ 37. Im ̄
patiens procumbensꎻ 38. Impatiens delavayiꎻ 39-41. Hypericum addin ̄
gtoniiꎻ 42. Begonia grandis subsp. sinensisꎻ 43-44. Osbeckia crinitaꎻ
45-46. Fuchsia hybrida. Scales bar 10 μm
Plate III Pollen morphology and exine ornamentation under LM
47. Oenothera roseaꎻ 48-49: Bupleurum longicauleꎻ 50-51. Heracle ̄
um franchetiiꎻ 52-53. Lyonia ovalifoliaꎻ 54- 55. Lysimachia violas ̄
censꎻ 56-57. Buddleja fallowianaꎻ 58-59. Buddleja forrestiiꎻ 60-61.
Halenia ellipticalaꎻ 62 - 63: Gentianopsis paludosaꎻ 64: Ajuga for ̄
restiiꎻ 65-66. Nepeta wilsoniiꎻ 67-68. Clinopodium megalanthumꎻ 69
-70. Elsholtzia rugulosa. Scales bar 10 μm
Plate IV Pollen morphology and exine ornamentation under LM
71-73. Verbascum thapsusꎻ 74 - 75. Pedicularis tenuisectaꎻ 76 - 77.
Pedicularis gruinaꎻ 78. Incarvillea argutaꎻ 79. Pteracanthus forrestiiꎻ
80-82. Valeriana flaccidissimaꎻ 83-84. Zehneria maysorensisꎻ 85-
86. Campanula colorataꎻ 87-89. Lobelia donianaꎻ 90. Saussurea stel ̄
la. Scales bar 10 μm
Plate V Pollen morphology and exine ornamentation under SEM
1. Cyanotis vagaꎻ 2. Tofieldia divergensꎻ 3. Silene bacciferaꎻ 4. Del ̄
phinium delavayiꎻ 5. Anemone hupehensis f. albaꎻ 6. Astilbe rivularisꎻ
7. Potentilla fulgensꎻ 8. Spenceria ramalanaꎻ 9. Pueraria peduncu ̄
larisꎻ 10. Clitoria marianaꎻ 11. Vicia craccaꎻ 12 - 13. Tripterygium
hypoglaucumꎻ 14-15. Impatiens uliginosaꎻ 16-17. Impatiens radia ̄
taꎻ 18. Impatiens procumbensꎻ 19. Impatiens delavayi
Plate VI Pollen morphology and exine ornamentation under SEM
20. Begonia grandis Dryand. subsp. Sinensisꎻ 21. Osbeckia crinitaꎻ
22. Fuchsia hybridaꎻ 23. Oenothera roseaꎻ 24. Heracleum franchetiiꎻ
25. Lyonia ovalifoliaꎻ 26. Lysimachia violascensꎻ 27. Buddleja fallowi ̄
anaꎻ 28. Buddleja forrestiiꎻ 29-30. Halenia ellipticala
Plate VII Pollen morphology and exine ornamentation under SEM
31. Ajuga forrestiiꎻ 32. Clinopodium megalanthumꎻ 33. Elsholtzia rug ̄
ulosaꎻ 34-35. Verbascum thapsusꎻ 36-37: Pedicularis tenuisectaꎻ 38
-39. Incarvillea argutaꎻ 40. Valeriana flaccidissimaꎻ 41. Zehneria
maysorensisꎻ 42-43. Campanula colorataꎻ 44-45. Lobelia doniana
276 植 物 分 类 与 资 源 学 报 第 37卷
郑鑫等: 图版Ⅰ ZHENG Xin et al.: Plate Ⅰ
3766期 ZHENG Xin et al.: Pollen Morphology of Plants from the Hengduan Mountains and Their Ecological
郑鑫等: 图版Ⅱ ZHENG Xin et al.: Plate Ⅱ
476 植 物 分 类 与 资 源 学 报 第 37卷
郑鑫等: 图版Ⅲ ZHENG Xin et al.: Plate Ⅲ
5766期 ZHENG Xin et al.: Pollen Morphology of Plants from the Hengduan Mountains and Their Ecological
郑鑫等: 图版Ⅳ ZHENG Xin et al.: Plate Ⅳ
676 植 物 分 类 与 资 源 学 报 第 37卷
郑鑫等: 图版Ⅴ ZHENG Xin et al.: Plate Ⅴ
7766期 ZHENG Xin et al.: Pollen Morphology of Plants from the Hengduan Mountains and Their Ecological
郑鑫等: 图版Ⅵ ZHENG Xin et al.: Plate Ⅵ
876 植 物 分 类 与 资 源 学 报 第 37卷
郑鑫等: 图版Ⅶ ZHENG Xin et al.: Plate Ⅶ
9766期 ZHENG Xin et al.: Pollen Morphology of Plants from the Hengduan Mountains and Their Ecological
书书书
!
#
$ %
&
(
) *
+ + #
$
, -
. /
. ,
0 #
/ & 1
0 & ,
1
* 2
& $
3 #
1 0 &
4 .
0 #
%
1
# ,
& #
1
2 / *
5
0 -
#
6
# $
4 %
7 .
$
8
* 7
$ 0
. &
$ 1
9 .
5
& + :
; #
$ 7
1
<
# ,
& #
1
< -
.
#
< &
= #
> !
5
? :
#
* 2
.
# /
0 7
/ #
@ -
. /
. ,
0 #
/ & 1
0 & ,
1
* 2
#
& $
#
< 0
/ .
0 & 2
& ,
. 0
& *
$
? -
& ,
A $
# 1
1
> !
5
(
(
B /
$ .
5
# $
0 .
0 & *
$
(
(
C $
% #
/
D 8
C $
% #
/
< E
8
(
) +
. 0
#
@ *
5
5
# +
& $
. ,
# .
#
!
# $
% &
(
F G
F *
$
!
# $
% &
(
) #
* #
/
* +
. 0
#
0 *
#
/
/ *
+ .
0 #
H I
G I
! J
K G
L M
I L
G L
N
J O
G P
! P
L G
L M
O J
G K
5
* $
* ,
* +
.
0 #
J Q
+ .
: #
/ #
% #
1 #
&
$ #
P G
K
0 & 5
# 1
. 1
0 -
& ,
A
. 1
$ #
&
$ #
J G
L
2 & $
# +
:
/ #
0 & ,
7 +
. 0
#
/ #
4 7
+ .
0 #
Q
#
/ 2 *
/ .
0 #
) +
G R $
P %
) +
G S
$
P
D &
+ & .
, #
. #
+ ,
% -
. (
D G
/ %
0 1
2 3
#
3
) 1
- *
1 $
(
T 7
/ G
. $
%
9 /
. $
, -
G
/
* +
. 0
#
J P
G K
! J
L G
L M
J K
G L
N
P I
G U
! P
K G
L M
P V
G K
5
* $
* ,
* +
.
0 #
J Q
+ .
: #
/ #
% #
1 #
&
$ #
# W
7 .
+
0 *
$ #
&
$ #
P G
L
2 & $
# +
:
/ #
0 & ,
7 +
. 0
#
2 & $
# +
:
/ #
0 & ,
7 +
. 0
#
) +
G R $
J %
) +
G S
$
J
) *
+ :
4 *
$ .
, #
. #
4 %
2
* %
$ .
5
D G
4 %
2
* %
$ .
5
, 6
$
1 $
( 1
D G
3 .
/ G
7 #
- #
3 %
8 .
5
! D
# 3
+ G
! X
Y G
D &
1
- #
/ *
& %
. +
K J
G K
! H
V G
K M
K K
G L
O Q
, *
+
. 0
#
J Q
+ .
: #
/ #
% #
1 #
&
$ #
H
0 & 5
# 1
. 1
0 -
& ,
A
. 1
$ #
&
$ #
V G
L
, *
. /
1 #
+ :
/ #
0 & ,
7 +
. 0
#
) +
G R $
O #
H
@ .
/ :
*
- :
+ + .
, #
. #
9
2 1 $
1
D G
9
2 1 $
1
.
$ $
# $
1 $
( (
9 /
. $
, -
G
1
- #
/ *
& %
. +
K P
G U
! H
V G
K M
K V
G K
.
$ 0
*
* /
. 0
#
J Q
+ .
: #
/ #
% #
1 #
&
$ #
O
0 & 5
# 1
. 1
0 -
& ,
A
. 1
$ #
&
$ #
H G
L
2 & $
# +
:
4 /
. $
7 +
. 0
#
) +
G R $
K
9 X
: #
, ,
0 1
- #
! D
G
Z *
0 -
1
- #
/ *
& %
. +
H P
G U
! H
L G
L M
H K
G L
.
$ 0
*
* /
. 0
#
% *
I G
L
2 & $
# +
:
4 /
. $
7 +
. 0
#
5
& ,
/ *
# ,
- &
$ .
0 #
) +
G R $
I #
) +
G S
$
O
Z .
$ 7
$ ,
7 +
. ,
# .
#
; 1
2 7
6
$
. 5
D G
; 1
2 7
6
$
. 5
3 1
2 #
) #
9 /
. $
, -
G
1 7
[
/ *
+ .
0 #
0 *
/
* +
. 0
#
H L
G L
! O
V G
K M
H J
G K
N
O L
G L
! J
V G
K M
O J
G K
O Q
, *
+
. 0
#
J Q
+ .
: #
/ #
% #
1 #
&
$ #
# W
7 .
+
0 *
$ #
&
$ #
J G
L
2 & $
# +
:
4 /
. $
7 +
. 0
#
5
& ,
/ *
# ,
- &
$ .
0 #
) +
G R $
V #
U %
) +
G S
$
H
/ 6
# 2
, &
- .
5
D G
/ 6
# 2
, &
- .
5
3 1
2 #
) #
9 /
. $
, -
G
1
- #
/ *
& %
. +
J K
G K
! J
J G
K M
J V
G K
.
$ 0
*
* /
. 0
#
J Q
+ .
: #
/ #
% #
1 #
&
$ #
# W
7 .
+
0 *
$ #
&
$ #
J G
L
2 & $
# +
:
4 /
. $
7 +
. 0
#
) +
G R $
\ #
P L
# P
P
+ $
1 5
% $
1
D G
+ $
1 5
% $
1
6 .
7 1
6 1
$ (
(
D #
5
G
2 G
# 2
: #
]
X ?
G ]
. $
4
1 7
[ 1
-
# /
* &
% .
+
J I
G L
! J
J G
K M
J V
G K
N
J O
G K
! J
L G
L M
J K
G L
O Q
, *
+
. 0
#
J Q
+ .
: #
/ #
% #
1 #
&
$ #
1 + &
4 -
0 + :
0 -
& ,
A #
/
0 -
. $
* /
# W
7 .
+ 0
*
$ #
&
$ #
J G
L
2 & $
# +
:
4 /
. $
7 +
. 0
#
5
& ,
/ *
# ,
- &
$ .
0 #
) +
G R $
P J
# P
O %
) +
G S
$
K
< .
&
2 / .
4 .
, #
. #
+ (
& 2
: 1
T 7
, -
G Q
6
. 5
G #
F G
F *
$
+ (
& 2
: 1
- )
. 2
# -
(
T 7
, -
G Q 6
. 5
G
1 7
[ 1
-
# /
* &
% .
+
0 *
1 7
[
/ *
+ .
0 #
P K
G K
! P
K G
L M
P V
G K
N
P J
G O
! P
L G
L M
P K
G L
O Q
, *
+
* /
. 0
#
J Q
+ .
: #
/ #
% #
1 #
&
$ #
# W
7 .
+
0 *
$ #
&
$ #
P G
K
2 .
& $
0 + :
/ #
0 & ,
7 +
. 0
#
% &
1 0 &
$ ,
0
/ #
0 & ,
7 +
. 0
#
) +
G R $
P H
# P
K #
P I
%
) +
G S
$
I
Z *
1 .
, #
. #
9 %
- :
# -
#
! <
# /
G
! G
T /
G #
! 1
, -
# / 1
G
9 %
- :
# -
#
# -
: %
- 1 #
< ,
- $
# & %
G
3 .
/ G
( .
: & %
5 1
$ &
% (
#
Z #
- %
# /
1 7
[ *
[ +
. 0
#
P G U
! P
K G
L M
J K
G L
N
J P
G U
! P
V G
K M
J V
G K
O Q
, *
+
* /
. 0
#
J Q
+ .
: #
/ #
% #
1 #
&
$ #
1 + &
4 -
0 + :
0 -
& ,
A #
/
0 -
. $
$ #
&
$ #
J G
L
2 .
& $
0 + :
/ #
0 & ,
7 +
. 0
#
) +
G R $
P V
# P
U
4 %
& 1 $
& 2
2 #
D G
4 %
& 1 $
& 2
2 #
0 .
2 *
1 $
(
]
. +
+ G
#
6
* *
A G
1 7
[
/ *
+ .
0 #
0 *
/
* +
. 0
#
J H
G V
K
! J
J G
K M
J V
G K
N
J P
! P
V G
K M
J J
G K
O Q
, *
+
* /
. 0
#
J Q
+ .
: #
/ #
% #
1 #
&
$ #
P G
K
0 & 5
# 1
. 1
0 -
& ,
A
. 1
$ #
&
$ #
O G
L
2 & $
# +
:
/ #
0 & ,
7 +
. 0
#
1 0 /
& .
0 #
Q
#
/ 2 *
/ .
0 #
) +
G R $
P # J
L #
J P
%
) +
G S
$
V
9 7
1 $
, 1
- #
? /
& 5
# $
9 7
1 $
, 1
- #
- #
5
# 2
# $
#
? /
& 5
# $
/
* +
. 0
#
J G K
! J
L M
O V
G K
N
P G J
K
! P
J G
K M
J J
G K
O Q
, *
+
* /
. 0
#
J Q
+ .
: #
/ #
% #
1 #
&
$ #
# W
7 .
+
0 *
$ #
&
$ #
J G
L
2 & $
# +
:
/ #
0 & ,
7 +
. 0
#
% *
) +
G R $
J J
# J
O %
) +
G S
$
U
9 .
[ .
, #
. #
! %
, 6
2 #
$ &
6 .
(
T #
$ 0
- G
! %
, 6
2 #
$ &
6 .
(
5
% $
& #
$ .
(
! F
& #
+ 1
6
. /
5
1
1 7
[
/ *
+ .
0 #
0 *
/
* +
. 0
#
O H
G I
! J
K G
L M
I V
G K
N
J I
G V
! P
V G
K M
K K
G L
O Q
, *
+
* /
. 0
#
J Q
+ .
: #
/ #
% #
1 #
&
$ #
# W
7 .
+
0 *
$ #
&
$ #
J G
K
2 & $
# +
:
/ #
0 & ,
7 +
. 0
#
) +
G R $
J H
# J
K
4 .
1 -
# -
#
F @
G
4 .
1 -
# -
#
7 1
3 .
$ ,
. 2
# -
(
! ;
/ .
- G
#
T #
$ 0
- G
T #
$ 0
- G
1 7
[ 1
-
# /
* &
% .
+
0 *
1 7
[ *
[ +
. 0
#
O K
G L
! J
J G
K M
H J
G K
N
O P
G L
! P
V G
K M
O V
G K
O Q
, *
+
* /
. 0
#
J Q
+ .
: #
/ #
% #
1 #
&
$ #
1 + &
4 -
0 + :
0 -
& ,
A #
/
0 -
. $
$ #
&
$ #
J G
K
% *
2 & $
# +
:
/ #
0 & ,
7 +
. 0
#
) +
G R $
J I
# J
V #
J U
%
) +
G S
$
! 2
& %
- #
D G
! 2
& %
- #
5
# -
#
$ #
D G
1
- #
/ *
& %
. +
H J
G V
! H
H G
L M
I O
G L
K !
Q I
Q
*
/ .
0 #
% *
J G
K
% *
% *
) +
G R R
$
J # O
L %
) +
G S
$
P L
<
, #
D G
<
, #
, -
# ,
, #
D G
/
* +
. 0
#
O H
G I
! J
K G
L M
I V
G K
N
J I
G V
! P
V G
K M
K K
G L
O Q
, *
+
* /
. 0
#
% *
J G
K
% *
% *
) +
G R R
$
O P
# O
J %
) +
G S
$
P P
086 植 物 分 类 与 资 源 学 报 第 37卷
书书书
!
# $
% &
( )
$ *
& $
+ #
%
!
#
$ % &
(
) *
+
, -
( .
$ (
+
, /
-
(
, $
0 (
1 !
#
2 &
- (
3 4
-
( 5
6 *
5 (
7 /
5
.
6 (
5 $ +
6 $ .
+
3 4
( 8
$ )
(
, 6
5
6 $ 4
$ .
6
$ 3
)
2 /
$ .
9 )
( +
+
1 !
#
:
:
; 5
)
#
( )
6
6 $ 3
)
:
:
< )
= (
5
> ?
< )
= (
5
, @
?
:
A %
6
(
7 (
%
+ 6 5
.
(
(
!
# $
% &
(
# )
*
B
3 3
9 C
4 C
!
# $ %
&
( #
) *
+
$ ,
( -
. )
/ )
*
! >
( D
% C
B
* 6
. /
+ *
E +
- /
( 5
$ 3
=
%
6 3
+ *
E -
5 3
%
6 (
F G
C G
! H
I C
G J
F H
C I
K
H L
C M
! H
I C
G J
F G
C G
F N
. 3
% -
3 5
6
(
H N
%
& (
5 (
= #
+ (
8 $
) (
O
6 $ #
( +
+
6 /
$ .
9
+
) (
8 $
) (
F C
G
= 3
= 3
A %
C P P
$
F F
# F
O %
A %
C Q
$
M H
# M
F
R
% +
#
$ )
.
(
(
0 *
$ .
% # &
1 2
> C
0 *
$ .
% # &
1 2
) -
# (
# 1
, 2
.
! 5
)
. /
C
- 5
3 %
6
(
O M
C O
! F
I C
G J
O H
C I
K
H S
C G
! H
I C
G J
H T
C T
O N
. 3
% -
6
(
/ $
( 5
5
. /
&
$ )
= $
+ 6 $
) .
6
M C
G
= 3
= $
+ 6 $
) .
6
5 (
6 $ .
* %
6
(
A %
C P P
$
F I
#
A %
C Q
$
M O
# M
I
0 U
.
3 #
. %
.
B
3 3
9 C
4 C
- 5
3 %
6
(
F I
C I
! F
H C
I J
F L
C I
K
H H
C I
! M
L C
I J
F H
C I
O N
. 3
% -
6
(
H N
%
& (
5 (
= #
+ (
8 $
) (
+ % $
V /
6 % &
6 /
$ .
9 (
5
6 /
)
) (
8 $
) (
M C
I
5 (
6 $ .
* %
6
(
= $
+ 6 $
) .
6
5 (
6 $ .
* %
6
(
W $
6 /
4 (
W
V 5
)
* %
( +
A %
C P P
$
F S
%
A %
C Q
$
M S
# M
L
0 U
$
, /
) *
4 &
1 2
! 5
)
. /
C
- 5
3 %
6
(
O M
C O
! F
L C
I J
O I
C G
K
H L
C G
! H
I C
G J
F G
C G
= 3
= 3
H C
G
. 3
5
+ (
% &
5 (
6 $ .
* %
6
(
A %
C P P
$
F L
%
A %
C Q
$
M T
0 U
3 &
- .
5 .
#
! 5
)
. /
C
- 5
3 %
6
(
O M
C F
! O
G C
G J
O I
C G
K
H X
C I
! H
I C
G J
F H
C I
= 3
H N
%
& (
5 (
= #
+ (
8 $
) (
+ % $
V /
6 % &
6 /
$ .
9 (
5
6 /
)
) (
8 $
) (
H C
G
5 (
6 $ .
* %
6
(
= $
+ 6 $
) .
6
5 (
6 $ .
* %
6
(
A %
C P P
$
F T
%
A %
C Q
$
M X
*
6 6 $
4 (
5
(
6
$ &
# /
) *
> C
6
$ &
# /
) *
. 3
3 #
1 (
% ,
1 #
#
Y
C Z
3 E
+ 3
)
+ *
E -
5 3
%
6 (
H S
C T
! H
G C
G J
F I
C G
K
H H
C T
! M
L C
I J
F G
C G
F N
. 3
% -
3 5
6
(
H N
%
& (
5 (
= #
+ (
8 $
) (
( [
*
%
6 3
) (
8 $
) (
H C
G
4
$ )
6 % &
5 (
6 $ .
* %
6
(
A %
C P P
$
F X
# O
G #
O M
R (
V 3
) $
.
(
(
7 &
( ,
1 #
.
> C
7 &
( ,
1 #
.
(
. 1
3 #
2
\ 5
&
) =
C
+ *
E +
- C
2 # 1
& 1
2 # 2
! ]
C 7 C
P 5 #
+ .
/ C
- 5
3 %
6
(
S X
C L
! S
H C
I J
L I
C G
K
F F
C M
! H
I C
G J
F L
C I
= 3
= 3
M C
I
= 3
+ 6 5
$
6 (
N
- (
5 4 3
5
6 (
A %
C P P
$
O H
%
A %
C Q
P $
H G
?
( %
+
6 3
#
6
.
(
(
8 2
4 &
/ 9
# .
> C
8 2
4 &
/ 9
# .
/
# 1
# % .
R (
) 6
/ C
( 8
7 C
R C
7 %
5
9 (
- 5
3 %
6
(
F H
C I
! F
G C
G J
F I
C G
K
H I
C I
! H
H C
I J
H L
C I
= 3
H N
%
& (
5 (
= #
+ (
8 $
) (
+ % $
V /
6 % &
6 /
$ .
9 (
5
6 /
)
) (
8 $
) (
M C
I
+ #
3 3
6 /
V 5
)
* %
6
(
A %
C P P
$
O F
# O
O %
A %
C Q
P $
H M
; )
V
5
. (
(
: )
/ +
2 # .
> C
: )
/ +
2 # .
+
4
# 3
.
B
3 5
6 C
( 8
, $
( E
C
) =
Q 3
+ +
C
+ -
/ (
5 3
$ =
%
I I
C G
! F
H C
I J
T H
C I
F N
- 3
5
6 (
H N
%
& (
5 (
= #
+ (
8 $
) (
)
=
) (
8 $
) (
5
6 /
( 5
6 /
$ .
9 (
5
6
6 /
(
-
( 5
6 *
5 (
+
H C
I
4 $ )
( %
&
5 (
6 $ .
* %
6
(
4 $ )
( %
&
5 (
6 $ .
* %
6
(
W $
6 /
+ $ %
9
A %
C P P
$
O I
# O
S #
A %
C Q
P $
H H
; )
V
5
. (
(
8 &
1 ,
% +
&
.
> C
8 &
1 ,
% +
&
.
,
2 &
.
> C
B
( 5
- 6
C (
8
] $
6 C
+ -
/ (
5 3
$ =
%
M G
X C
G
! M
G G
C G
J M
H G
C G
F N
- 3
5
6 (
H N
%
& (
5 (
= #
+ (
8 $
) (
)
=
) (
8 $
) (
5
6 /
( 5
6 /
$ .
9 (
5
6
6 /
(
-
( 5
6 *
5 (
+
L C
G
4
$ )
6 % &
# $
. 5 3
5 ( 6
$ . *
% 6
3 N
+ 6 5
$
6 (
V 5
)
* %
6
( N
- (
5 4 3
5
6 (
W $
6 /
+ $ %
9
A %
C P P
P $
O L
%
A %
C Q
P $
H F
] -
$
. (
(
7 )
$ -
& )
)
*
> C
7 )
$ -
& )
)
*
- ,
1 (
# / .
) -
&
^
%
% C
_
\ 7
C
- 5
3 %
6
(
H O
C G
! H
G C
G J
H I
C G
K
M O
C G
! M
G C
G J
M I
C G
F N
. 3
% -
3 5
6
(
H N
%
& (
5 (
= #
+ (
8 $
) (
+ % $
V /
6 % &
6 /
$ .
9 (
5
6 /
)
) (
8 $
) (
H C
G
4 $ )
( %
&
5 (
6 $ .
*
% 6 (
A %
C P P
P $
O T
# O
X
6 &
.
/ - &
) *
> C
6 &
.
/ - &
) *
; .
1 /
+ &
% # #
?
C B
$ 5 3
(
- (
5 -
5 3
%
6 (
O S
C I
! O
H C
I J
I H
C I
K
T C
T
! M
L C
I J
H G
C G
F N
. 3
% -
3 5
6
(
= 3
H C
I J
F C
G
= 3
+ 6 5
$
6 (
N
- (
5 4 3
5
6 (
A %
C P P
P $
I G
# I
M %
A %
C Q
P $
H O
@ 5
$ .
.
(
(
<
, 1
# .
Y *
6 6 C
<
, 1
# .
, 5
. -
# ; ,
- # .
! ^
%
% C
\ 5
* =
(
6 (
6 5
/ (
= 5
%
6 (
6 5
=
3 5
= (
. *
+ +
6 (
6 ( 6
5
=
F I
C G
! H
H C
I J
O I
C G
F N
. 3
% -
3 5
6
(
= 3
M C
I
= 3
= $
+ 6 $
) .
6
5 (
6 $ .
* %
6
(
A %
C P P
P $
I H
# I
F %
A %
C Q
P $
H I
A 5
$ #
* %
.
(
(
<
2 # *
. /
+ #
.
> C
<
2 # *
. /
+ #
.
5 # ,
- .
2 /
& 1
2
! 5
)
. /
C
- 5
3 %
6
(
F G
C T
! H
I C
G J
F L
C I
K
H G
C I
! M
L C
I J
H H
C I
F N
. 3
% -
3 5
6
(
= 3
M C
I
= 3
4 $ )
( %
&
5 (
6 $ .
* %
6
(
A %
C P P
P $
I O
# I
I %
A %
C Q
P $
H S
> 3
V
) $
.
(
(
7 )
3 3
- & =
.
! R
* =
N
= %
( $
*
. 6
C
> C
7 )
3 3
- & =
.
; .
- - ,
> #
. 1
.
R
% 4 C
! C
)
=
^
C^
C ,
#
C
+ *
E -
5 3
%
6 (
6 3
- 5
3 %
6
(
H M
C S
! M
L C
I J
H H
C I
K
M O
C L
! M
H C
I J
M L
C I
O N
. 3
% -
3 5
6
(
H N
%
& (
5 (
= #
+ (
8 $
) (
+ % $
V /
6 % &
6 /
$ .
9 (
5
6 /
)
) (
8 $
) (
M C
I
= 3
= 3
A %
C P P
P $
I S
# I
L %
A %
C Q
P $
H L
7 U
; ,
&
2 % #
# $ (
% +
+ *
E -
5 3
%
6 (
6 3
- 5
3 %
6
(
H O
C I
! H
H C
I J
H L
C I
K
M X
C I
! M
L C
I J
H H
C I
O N
. 3
% -
3 5
6
(
H N
%
& (
5 (
= #
+ (
8 $
) (
+ % $
V /
6 % &
6 /
$ .
9 (
5
6 /
)
) (
8 $
) (
M C
I
4 $ )
( %
&
5 (
6 $ .
*
% 6 (
4 $ )
( %
&
5 (
6 $ .
*
% 6 (
A %
C P P
P $
I T
# I
X %
A %
C Q
P $
H T
1866期 ZHENG Xin et al.: Pollen Morphology of Plants from the Hengduan Mountains and Their Ecological
书书书
!
# $
% &
( )
$ *
& $
+ #
%
!
#
$ % &
(
) *
+
, -
( .
$ (
+
, /
-
(
, $
0 (
1 !
#
2 &
- (
3 4
-
( 5
6 *
5 (
7 /
5
.
6 (
5 $ +
6 $ .
+
3 4
( 8
$ )
(
, 6
5
6 $ 4
$ .
6
$ 3
)
2 /
$ .
9 )
( +
+
1 !
#
:
:
; 5
)
#
( )
6
6 $ 3
)
:
:
< )
= (
5
> ?
< )
= (
5
, @
?
:
A %
6
(
(
) 6
$
)
. (
(
!
# $ %
&
B 3
5 9
/ C
!
# $ %
&
$ # #
&
( & )
#
D
C D
3 )
+ *
E +
- /
( 5
$ 3
=
%
6 3
+ *
E -
5 3
%
6 (
F G
C G
! F
H C
I J
K H
C I
L
F M
C H
! F
H C
I J
F N
C H
F O
. 3
% -
3 5
6
(
P O
%
& (
5 (
= #
+ (
8 $
) (
K
6 $ #
( +
+
6 /
$ .
9
+
) (
8 $
) (
P C
H
= 3
= $
+ 6 $
) .
6
5 (
6 $ .
* %
6
(
A %
C Q Q
Q $
M I
# M
R %
A %
C S
Q $
P T
# F
I
* $
% (
&
% +
,
& ,
?
* $
% (
&
% +
,
& ,
# -
. +
,
! U
3 3
9
4 C
?
+ -
/ (
5 3
$ =
%
6 3
+ *
E -
5 3
%
6 (
M T
C N
! M
P C
H J
N H
C I
L
F F
C R
! P
H C
I J
F N
C H
F !
K
O
. 3
% -
3 5
3 $
=
P O
%
& (
5 (
= #
+ (
8 $
) (
P
6 $ #
( +
+
6 /
$ .
9
+
) (
8 $
) (
F C
I
= $
+ 6 $
) .
6
5 (
6 $ .
* %
6
(
A %
C Q Q
Q $
M P
# M
F
>
E $
6
(
/ 0
- 1
> C
/ 0
- 1
2 +
3 3 $
, ( &
& D
$ (
% +
+ *
E -
5 3
%
6 (
6 3
- 5
3 %
6
(
P M
C K
! R
H C
I J
K H
C I
L
P I
C T
! R
P C
H J
F H
C I
F O
. 3
% -
6
(
P O
%
& (
5 (
= #
+ (
8 $
) (
F
6 $ #
( +
+
6 /
$ .
9
+
) (
8 $
) (
P C
I
4 $ )
( %
&
5 (
6 $ .
* %
6
(
4 $ )
( %
&
5 (
6 $ .
* %
6
(
A %
C Q Q
Q $
M K
%
A %
C S
Q Q
$
F R
4 $
$
(
> C
4 $
$
(
5 &
# , +
% &
& D
* 6
/ $
(
= 3
H M
C G
! H
I C
I J
M P
C H
L
K K
C F
! F
P C
H J
H H
C I
M O
. 3
% -
6
(
P O
%
& (
5 (
= #
+ (
8 $
) (
( V
*
%
6 3
) (
8 $
) (
P C
I
5 (
6 $ .
* %
6
(
A %
C Q Q
Q $
M H
# M
M
6 #
& %
+
+ .
& -
7
> C
6 #
& % +
+
. & -
7
7 $
1
# %
( 8 -
7
! D
$ (
% +
7 C
W
C X
*
)
=
U
+ *
)
( 8
U
C X
C >
$
+ *
E -
5 3
%
6 (
6 3
- 5
3 %
6
(
K N
C G
! P
I C
I J
H N
C H
L
F M
C F
! R
H C
I J
K N
C H
= 3
P O
%
& (
5 (
= #
+ (
8 $
) (
+ % $
Y /
6 % &
6 /
$ .
9 (
5
6 /
)
3 5
( V
*
% 6
3
) (
8 $
) (
P C
H
4 $ )
( %
&
5 (
6 $ .
* %
6
(
4 $ )
( %
&
5 (
6 $ .
* %
6
(
A %
C Q Q
Q $
M N
# M
G %
A %
C S
Q Q
$
F P
9 #
, 8
+ #
( : &
X
$ % %
= C
9 #
, 8
+ #
( : &
3 -
1 -
# +
,
U
( #
+ %
C
- 5
3 %
6
(
P G
C T
! P
H C
I J
F P
C H
L
P I
C I
! R
N C
H J
P P
C H
M O
. 3
% -
6
(
P O
%
& (
5 (
= #
+ (
8 $
) (
( V
*
%
6 3
) (
8 $
) (
P C
I
4 $ )
( %
&
5 (
6 $ .
*
% 6 (
4 $ )
( %
&
5 (
6 $ .
*
% 6 (
A %
C Q Q
Q $
M T
# N
I %
A %
C S
Q Q
$
F F
, .
5 3
- /
* %
5
$
. (
(
; $
3 <
,
) -
7
> C
; $
3 <
,
) -
7
( 8
, -
,
> C
+ *
E +
- /
( 5
3 $
=
%
6 3
+ *
E -
5 3
%
6 (
P G
C H
! P
H C
I J
F P
C H
L
P P
C G
! R
H C
I J
P H
C I
F O
. 3
% -
3 5
3 $
=
P O
%
& (
5 (
= #
+ (
8 $
) (
F
6 $ #
( +
+
6 /
$ .
9
+
) (
8 $
) (
P C
I
= 3
= 3
A %
C Q S
$ N
R #
N P
# N
F %
A %
C S
Q Q
$
F K
# F
H
= $
. &
) -
#
3 & ,
> C
= $
. &
) -
#
3 & ,
( $ %
- &
, $
) (
! 5
)
. /
C (
8
?
8
$ #
C
4 %
6
P G
C F
! P
H C
I J
F I
C I
L
R M
C F
! R
I C
I J
P I
C I
P O
+ &
) .
3 %
-
6 (
P O
%
& (
5 (
= #
+ (
8 $
) (
+ % $
Y /
6 % &
6 /
$ .
9 (
5
6 /
)
) (
8 $
) (
P C
I
+ #
3 3
6 /
- (
5 4 3
5
6 (
A %
C Q S
$
N K
# N
H %
A %
C S
Q Q
$
F M
# F
N
= Z
1 3
- &
%
! 5
)
. /
C
( 8
?
8
$ #
C
= 3
F M
C G
! F
F C
I J
K P
C I
L
F I
C G
! P
N C
H J
F K
C I
= 3
= 3
P C
I
= 3
= 3
A %
C Q S
$
N M
# N
N
B $
Y )
3 )
$
. (
(
> %
)
3 ? &
# # $
[ *
+ +
C
> %
)
3 ?
& # #
$
3
1 -
(
! 3 &
% (
\ 3
& %
(
+ *
E -
5 3
%
6 (
6 3
- 5
3 %
6
(
P K
C H
! P
P C
H J
P N
C H
L
R T
C H
! R
N C
H J
P P
C H
M !
O N
O
. 3
% -
6
(
P O
%
& (
5 (
= #
+ (
8 $
) (
( V
*
%
6 3
) (
8 $
) (
P C
I
4 $ )
( %
&
Y 5
)
* %
6
( O
- (
5 4 3
5
6 (
# $
. 5
3 (
. /
$ )
6 (
O
- (
5 4 3
5
6 (
A %
C Q S
$
N G
%
A %
C S
Q Q
$
F G
# F
T
] .
)
6 /
.
(
(
= (
$ 3
)
%
( 8
- ,
!
(^ (
+
B 5
( #
( 9
C
= (
$ 3
)
%
( 8
- ,
2 +
3 3 $
, ( &
&
! D
$ (
% +
7 C
W
C X
*
- 5
3 %
6
(
R I
R C
M
! T
I C
I J
R F
I C
I
L
M R
C M
! K
P C
H J
N N
C H
F O
. 3
% -
3 5
6
(
)
=
R H
O R
G
- +
( *
= 3
. 3
% - 3
5 6
(
P O
%
& (
5 (
= #
+ (
8 $
) (
P
6 $ #
( +
+
6 /
$ .
9
+
) (
8 $
) (
P C
H
. 3
5
+ (
% &
5 (
6 $ .
* %
6
(
A %
C Q S
$
N T
S
% (
5 $
)
. (
(
;
# $ 3
&
%
> C
;
# $ 3
&
%
2 #
) )
& .
& , ,
& 7
?
8
$ #
C
+ *
E +
- /
( 5
3 $
=
%
6 3
+ *
E -
5 3
%
6 (
K H
C H
! F
I C
I J
M P
C H
L
K R
C F
! P
N C
H J
H N
C H
F O
. 3
% -
6
(
P O
%
& (
5 (
= #
+ (
8 $
) (
( V
*
%
6 3
) (
8 $
) (
F C
H
# $
. 5
3 (
. /
$ )
6 (
+ -
5
+ (
% &
+ -
$ )
* %
3 +
(
A %
C Q S
$ G
I #
G R
# G
P %
A %
C S
Q Q
$
K I
7 *
. *
5 E
$ 6
. (
(
@ $
8 %
$ 3
&
@ )
= %
C
@ $
8 %
$ 3
&
7
A
, +
3 $
% ,
& ,
! X
$ Y
/ 6
)
=
] 5
) C
] 5
) C
+ *
E -
5 3
%
6 (
6 3
- 5
3 %
6
(
K P
C N
! K
K C
I J
M F
C I
F O
. 3
% -
3 5
6
(
P O
%
& (
5 (
= #
+ (
8 $
) (
+ % $
Y /
6 % &
6 /
$ .
9 (
5
6 /
)
) (
8 $
) (
P C
I
4 $ )
( %
&
5 (
6 $ .
*
% 6 (
= $
+ 6 $
) .
6 4
$ )
( %
&
5 (
6 $ .
*
% 6 (
A %
C Q S
$
G F
# G
K %
A %
C S
Q Q
$
K R
7
#
-
) *
%
. (
(
6
7
% -
#
> C
6
7
% -
#
> C
+ -
/ (
5 3
$ =
%
P H
C H
! P
P C
H J
P N
C H
F O
- 3
5
6 (
= 3
P C
I
+ -
$ )
* %
3 +
(
+ 6 5
$
6 (
O
- (
5 4 3
5
6 (
A %
C Q S
$
G H
# G
M %
A %
C S
Q Q
$
K P
# K
F
B +
< $
# &
> C
B +
< $
# &
. +
% &
%
, 9
3 6
6 + E
C
+ *
E -
5 3
%
6 (
6 3
- 5
3 %
6
(
K K
C F
! F
P C
H J
M I
C I
L
F R
C F
! P
H C
I J
F H
C I
F O
. 3
% -
3 5
6
(
P O
%
& (
5 (
= #
+ (
8 $
) (
+ % $
Y /
6 % &
6 /
$ .
9 (
5
6 /
)
3 5
( V
*
% 6
3
) (
8 $
) (
P C
H
4 $ )
( %
&
5 (
6 $ .
* %
6
(
4 $ )
( %
&
5 (
6 $ .
* %
6
(
A %
C Q S
$ G
N #
G G
# G
T %
A %
C S
Q Q
$
K K
# K
H
7 3
#
- 3
+ $ 6
(
C
- ,
, -
3 $
D 7
C
C
- ,
, -
3 $
, ( $
# #
?
8
$ #
C
+ *
E +
- /
( 5
3 $
=
%
6 3
+ *
E -
5 3
%
6 (
M M
C N
! M
H C
I J
N I
C I
L
M I
C I
! H
H C
I J
M H
C I
F O
. 3
% -
3 5
6
(
P O
%
& (
5 (
=
T C
I
+ -
$ )
* %
3 +
(
A %
C Q S
$
T I
286 植 物 分 类 与 资 源 学 报 第 37卷