Micromorphological Characteristics of Pollen and Its Taxonomic Significance of Saussurea in Sedgera Mountains of Tibet

doi:10.7525/j.issn.1673-5102.2023.05.011

Abstract

Abstract:

The pollen of 22 species of Saussurea and 1 species of Himalaiella were observed and compared by scanning electron microscopy to understand the degree of differentiation at the level of pollen morphology， and the pollen cluster analysis was carried out to provide a sporological basis for the classification of this genus， and further provided germplasm resources utilization of Saussurea in the region. The results showed that：（1）There were 20 species of 23 plants with subspherical pollen shape and three were oblate spherical； most of the pollen volume belonged to medium-sized pollen， and the overall type was relatively evolved. （2）Except for Saussurea Nephrolepis， which was a four-pore pollen， the pollen of the other 22 species had three pore grooves. （3）The outer wall ornamentation of pollen of 23 species could be divided into seven types： seven types of reticulate， six types of reticular and granular， six types of cavernous， and there were one smooth， one spine-based cavernous and one interspinous with wrinkled waves， one granular and one ruffled. （4）Cluster analysis based on pollen micromorphology of 23 plant species showed some consistency with the plant classification system and was even relatively clear among some subgenera. The results of this paper suggested that pollen micromorphological features might provide a basis for species identification in this genus.

Key words: Saussurea, the mountainous region of sedgera, pollen micromorphology, cluster analysis, Tibet

CLC Number:

Q949

Shujun ZHNAG, Xinxin YANG, Jian LUO. Micromorphological Characteristics of Pollen and Its Taxonomic Significance of Saussurea in Sedgera Mountains of Tibet[J]. Bulletin of Botanical Research, 2023, 43(5): 741-755.

Figures/Tables 9

Table 1

Basic information of experimental materials

编号

No.

种名

Species name

分类^［5-6］

Classification

生境

Habitat

海拔

Elevation/m

凭证标本

Voucher specimen

拉萨雪兔子

Saussurea kingii

雪兔子亚属

Subgen. Eriocoryne

河滩沙地、沙丘、山坡沙地

Beach sand，sand dunes，

and hillside sand

3 611.00

20141003315

绵头雪兔子

Saussurea laniceps

雪兔子亚属

Subgen. Eriocoryne

高山流石滩

Rugged rocks and pebbly shores

4 800.00

627

鼠曲雪兔子

Saussurea gnaphalodes

雪兔子亚属

Subgen. Eriocoryne

山坡流石滩

Rocky slope and pebbly shore

4 830.00

LJ-006-028

硕首雪兔子

Saussurea grandiceps

雪兔子亚属

Subgen. Eriocoryne

高山流石滩

Pebble beach on a high mountain

3 400.00

487

星状雪兔子

Saussurea stella

雪兔子亚属

Subgen. Eriocoryne

沼泽草地、河滩地

Marsh grassland，riverbank

3 385.00

SJ381

苞叶雪莲

Saussurea obvallata

雪莲亚属

Subgen. Amphilaena

山坡多石处

Rocky area on a hillside

4 660.12

QTP-LUO-1402

宝璐雪莲

Saussurea luae

雪莲亚属

Subgen. Amphilaena

山坡多石处

Rocky area on a hillside

5 200.00

QTP-LUO-2241

多鞘雪莲

Saussurea polycolea

雪莲亚属

Subgen. Amphilaena

山坡草地

Grassy slope

4 600.00

520

毛背雪莲

Saussurea pubifolia

雪莲亚属

Subgen. Amphilaena

高山流石滩

Rocky shore on a high mountain

5 200.00

2222

白背须弥菊

Himalaiella auriculata

须弥菊属

Himalaiella

河谷针阔混交林

Mixed coniferous and broad-

leaved forest in a river valley

2 100.00

980

尖叶风毛菊

Saussurea ciliaris

风毛菊亚属裸托组

Subgen. Saussurea（Sect. Depressae）

草坡、山地开阔处

Grassy slope，open area

in the mountains

4 629.48

QTP-LUO-1382

黑苞风毛菊

Saussurea melanotricha

风毛菊亚属无茎组

Subgen. Saussurea（Sect. Acaules）

开阔石质山坡

Open rocky slope

4 623.75

QTP-LUO-1398

锥叶风毛菊

Saussurea wernerioides

风毛菊亚属无茎组

Subgen. Saussurea（Sect. Acaules）

山坡草地或砾石山坡

Grassy slope or rocky slope

4 500.00

QTP-2193

薄苞风毛菊

Saussurea leptolepis

风毛菊亚属羽裂组

Subgen. Saussurea（Sect. Cyathidium）

高山草甸

Alpine meadow

4 830.00

LJ-006-059

川滇风毛菊

Saussurea wardii

风毛菊亚属羽裂组

Subgen. Saussurea（Sect. Cyathidium）

山坡草地

Grassy slope

4 180.00

倒披针叶风毛菊

Saussurea nimborum

风毛菊亚属羽裂组

Subgen. Saussurea（Sect. Cyathidium）

山坡草丛中、河滩草地

In a grassy slope，on a riverbank

4 624.28

QTP-LUO-1399

革苞风毛菊

Saussurea coriacea

风毛菊亚属羽裂组

Subgen. Saussurea（Sect. Cyathidium）

阶地

Terrace

4 629.58

QTP-LUO-1400

蒲公英风毛菊

Saussurea taraxacifolia

风毛菊亚属羽裂组

Subgen. Saussurea（Sect. Cyathidium）

山坡灌丛

Shrubbery on a hillside

4 630.17

QTP-LUO-1401

狮牙草状风毛菊

Saussurea leontodontoides

风毛菊亚属羽裂组

Subgen. Saussurea（Sect. Cyathidium）

山坡砾石地、草地

Rocky slope and grassy slope.

4 000.00

587

松潘风毛菊

Saussurea sungpanensis

风毛菊亚属羽裂组

Subgen. Saussurea（Sect. Cyathidium）

山坡草地或砾石山坡

Grassy slope or rocky slope.

4 500.00

512

弯齿风毛菊

Saussureaprzewalskii

风毛菊亚属羽裂组

Subgen. Saussurea（Sect. Cyathidium）

山坡灌丛草地、流石滩

Shrubbery and grassy slope，

pebble beach.

4 300.00

2235

紫苞风毛菊

Saussurea purpurascens

风毛菊亚属羽裂组

Subgen. Saussurea（Sect. Cyathidium）

山坡灌丛中

In shrubbery on a hillside.

4 666.96

QTP-LUO-652

粗裂风毛菊

Saussurea grosseserrata

风毛菊亚属风毛菊组

Subgen. Saussurea（Sect. Saussurea）

山坡草地或杜鹃灌丛中

Grassy slope or Rhododendron

shrubbery on a hillside.

4 665.19

QTP-LUO-651

Table 1

Table 2

Morphological characteristics of pollen from 23 plant species（1）

物种名称及编号 Species name and number	极轴长P/μm		赤道轴长E/μm		P/E	萌发孔宽G/μm		萌发沟长S/μm		体积 V/μm³
物种名称及编号 Species name and number	均值±标准误 Mean±SE	偏差系数 Cv/%	均值±标准误 Mean±SE	偏差系数 Cv/%	P/E	均值±标准误 Mean±SE	偏差系数 Cv/%	均值±标准误 Mean±SE	偏差系数 Cv/%	体积 V/μm³
1.拉萨雪兔子 Saussurea kingii	39.560±0.578i	1.500	37.998±0.842i	2.200	1.042	11.509±0.992b	8.600	29.520±0.795f	2.700	29 906.456
2.绵头雪兔子 Saussurea laniceps	43.236±0.893e	2.100	38.499±0.784i	2.000	1.123	7.922±0.818i	10.300	26.323±0.917j	3.500	33 553.901
3.鼠曲雪兔子 Saussurea gnaphalodes	43.291±0.949e	2.200	41.867±0.827e	2.000	1.034	12.588±0.880a	7.000	27.580±0.695i	2.500	39 731.423
4.硕首雪兔子 Saussurea grandiceps	44.490±0.028c	0.100	42.610±0.269e	0.600	1.044	9.613±0.379c	3.900	22.158±0.735l	3.300	42 294.519
5.星状雪兔子 Saussurea stella	42.602±0.774f	1.800	41.415±0.916f	2.200	1.029	10.081±0.893c	8.900	28.044±0.677h	2.400	40 547.279
6.苞叶雪莲 Saussurea obvallata	41.223±0.662g	1.600	41.843±0.232e	0.600	0.985	7.479±0.853h	11.400	30.124±0.508e	1.700	37 789.243
7.宝璐雪莲 Saussurea luae	11.927±0.810m	6.800	10.970±0.303n	2.800	1.075	2.709±0.268p	9.900	7.680±0.548p	7.100	751.503
8.多鞘雪莲 Saussurea polycolea	40.495±0.007h	0.100	40.075±0.035h	0.100	1.010	6.883±0.817k	11.900	24.853±0.868k	3.500	34 052.349
9.毛背雪莲 Saussurea pubifolia	47.233±1.322a	2.800	45.307±1.66c	3.700	1.044	11.331±0.750b	6.600	30.407±0.900e	3.000	50 765.823
10.白背须弥菊 Himalaiella auriculata	41.682±0.663f	1.600	40.452±0.911h	2.300	1.031	9.950±0.962f	9.700	27.550±0.912i	3.300	35 713.068
11.尖叶风毛菊 Saussurea ciliaris	28.903±0.894l	3.100	34.397±0.885l	2.600	0.840	2.603±0.059p	2.300	13.305±0.436o	3.300	17 905.204
12.黑苞风毛菊 Saussurea melanotricha	46.581±0.882b	1.900	43.239±0.800d	1.900	1.078	8.344±0.769g	9.200	31.825±0.373c	1.200	45 599.356
13.锥叶风毛菊 Saussurea wernerioides	42.020±0.990f	2.400	40.523±0.826g	2.000	1.037	9.102±0.867d	9.500	18.510±0.991n	5.400	36 129.158
14.薄苞风毛菊 Saussurea leptolepis	45.123±0.614c	1.400	43.207±0.618d	1.400	1.044	9.628±0.674c	7.000	30.947±0.590d	1.900	44 106.370
15.川滇风毛菊 Saussurea wardii	41.528±0.845f	2.000	60.068±0.820b	1.400	0.692	3.880±0.704o	18.100	46.893±0.948a	2.000	78 456.015
16.倒披针叶风毛菊 Saussurea nimborum	41.270±0.354g	0.900	41.655±0.035f	0.100	0.991	8.479±0.932g	11.000	25.540±0.389j	1.500	37 494.482
17.革苞风毛菊 Saussurea coriacea	41.687±0.909f	2.200	40.163±0.977h	2.400	1.038	6.177±0.297m	4.800	32.695±0.120c	0.400	35 209.129
18.蒲公英风毛菊 Saussurea taraxacifolia	34.080±0.711k	2.100	31.703±0.922m	2.900	1.076	3.708±0.432o	11.700	21.603±0.713m	3.300	17 935.276
19.狮牙草状风毛菊 Saussurea leontodontoides	43.793±0.654d	1.500	43.623±0.304c	0.700	1.004	8.800±0.706e	8.000	28.854±0.907g	3.100	43 635.938
20.松潘风毛菊 Saussurea sungpanensis	36.987±0.542j	1.500	38.397±0.316i	0.800	0.963	8.310±0.610g	7.300	28.528±0.800h	2.800	28 551.627
21.弯齿风毛菊 Saussurea przewalskii	37.200±0.752j	2.000	36.017±0.929k	2.600	1.033	6.575±0.411l	6.200	22.033±0.960l	4.400	25 266.704
22.紫苞风毛菊 Saussurea purpurascens	43.200±0.658e	1.500	62.303±0.644a	1.000	0.693	5.487±0.842n	15.300	38.035±0.967b	2.500	87 802.105
23.粗裂风毛菊 Saussurea grosseserrata	39.613±0.615i	1.600	37.207±0.484j	1.300	1.065	7.204±0.960j	13.300	22.404±0.727l	3.200	28 713.196

Table 2

Table 3

Morphological characteristics of pollen from 23 plant species（2）

物种名称及编号 Species name and number	网眼宽M/μm		网脊宽W/μm		刺宽W_S/μm		刺长L_S/μm		刺数 N_S/400 μm²	纹饰 Ornamentation
物种名称及编号 Species name and number	均值±标准误 Mean±SE	偏差系数 Cv（%）	均值±标准误 Mean±SE	偏差系数 Cv（%）	均值±标准误 Mean±SE	偏差系数 Cv（%）	均值±标准误 Mean±SE	偏差系数 Cv（%）	刺数 N_S/400 μm²	纹饰 Ornamentation
1.拉萨雪兔子 Saussurea kingii	0.443±0.070c	15.800	0.347±0.051c	14.800	4.211±0.991d	23.500	3.759±0.673b	17.900	17	网状-颗粒状 Reticular-granular
2.绵头雪兔子 Saussurea laniceps	0.540±0.078b	14.400	0.278±0.048c	17.100	4.597±0.839d	18.200	3.742±0.853h	22.800	21	网状-颗粒状 Reticulated-granular
3.鼠曲雪兔子 Saussurea gnaphalodes	0.486±0.074c	15.200	0.381±0.092c	24.000	4.008±0.723d	18.100	2.748±0.499i	16.100	17	网状 Reticulated
4.硕首雪兔子 Saussurea grandiceps	0.388±0.067c	17.200	0.366±0.080c	21.900	4.668±0.763e	16.300	2.512±0.595j	23.700	13	网状 Reticulated
5.星状雪兔子 Saussurea stella	一	一	一	一	2.129±0.384j	18.100	1.871±0.416j	22.300	20	粗糙具褶皱 Rough with folds
6.苞叶雪莲 Saussurea obvallata	0.365±0.038f	10.400	0.266±0.051c	19.100	5.154±0.763c	14.800	3.656±0.543b	14.800	9	穴状 Cavernous
7.宝璐雪莲 Saussurea luae	一	一	一	一	15.150±0.990a	6.500	10.105±0.880a	8.700	20	粗糙具颗粒 Coarsely granular with surface particles
8.多鞘雪莲 Saussurea polycolea	1.200±0.211a	17.600	0.849±0.193b	22.800	4.845±0.559c	11.500	3.522±0.474d	13.500	17	刺基穴状刺间具皱波状 Spiny basal apertures with wrinkled and undulating ridges between the spines
9.毛背雪莲 Saussurea pubifolia	0.364±0.037d	10.300	0.294±0.042c	14.400	4.992±0.627b	12.600	3.573±0.679h	19.000	11	网状 Reticulated
10.白背须弥菊 Himalaiella auriculata	0.297±0.044i	15.000	0.202±0.020h	9.800	5.846±0.837c	14.300	3.707±0.562e	15.200	10	穴状 Cavernous
11.尖叶风毛菊 Saussurea ciliaris	0.313±0.047h	15.000	0.224±0.031g	13.800	4.407±0.787g	17.900	3.210±0.378f	11.800	20	网状 Reticulated
12.黑苞风毛菊 Saussurea melanotricha	一	一	一	一	5.766±0.589b	10.200	3.870±0.523b	13.500	13	光滑状 Smooth in texture
13.锥叶风毛菊 Saussurea wernerioides	0.314±0.047g	15.000	0.222±0.032f	14.600	5.556±0.759b	13.700	4.205±0.491b	11.700	10	穴状 Cavernous
14.薄苞风毛菊 Saussurea leptolepis	0.401±0.054c	13.600	0.321±0.051c	16.000	4.819±0.681c	14.100	3.525±0.246c	7.000	19	网状 Reticulated
15.川滇风毛菊 Saussurea wardii	0.450±0.065c	14.400	0.331±0.055c	16.800	3.330±0.755i	22.700	1.356±0.208k	15.300	12	网状 Reticulated
16.倒披针叶风毛菊 Saussurea nimborum	0.358±0.034d	9.600	0.317±0.082c	25.700	4.979±0.699b	14.000	3.443±0.357e	10.400	16	网状-颗粒状 Reticulated-granular
17.革苞风毛菊 Saussurea coriacea	0.428±0.054c	12.600	0.351±0.057c	16.300	4.875±0.914b	18.700	3.759±0.353c	9.400	17	网状 Reticulated
18.蒲公英风毛菊 Saussurea taraxacifolia	0.312±0.027h	8.800	0.257±0.028d	11.000	4.253±0.535h	12.600	3.178±0.286g	9.000	20	穴状 Cavernous
19.狮牙草状风毛菊 Saussurea leontodontoides	0.341±0.050c	14.600	0.268±0.034c	12.700	5.293±0.850c	16.100	3.667±0.334c	9.100	18	网状-颗粒状 Reticulated-granular
20.松潘风毛菊 Saussurea sungpanensis	0.346±0.037e	10.600	0.271±0.028d	10.200	5.482±0.362b	6.600	3.768±0.360c	9.600	11	网状-颗粒状 Reticulated-granular
21.弯齿风毛菊 Saussurea przewalskii	0.378±0.073c	19.300	1.104±0.139a	12.600	4.320±0.729f	16.900	3.298±0.389e	11.800	25	网状-颗粒状 Reticulated-granular
22.紫苞风毛菊 Saussurea purpurascens	0.454±0.061c	13.400	0.279±0.040d	14.500	2.882±0.596j	20.700	1.892±0.345j	18.300	9	穴状 Cavernous
23.粗裂风毛菊 Saussurea grosseserrata	0.416±0.058c	13.900	0.226±0.042e	18.800	4.594±0.875c	19.100	3.625±0.424e	11.700	14	穴状 Cavernous

Table 3

Fig.1

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