铁杉属花粉形态及其聚类分析研究

doi:10.12302/j.issn.1000-2006.202406025

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南京林业大学学报（自然科学版） ›› 2024, Vol. 48 ›› Issue (4): 37-45.doi: 10.12302/j.issn.1000-2006.202406025

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铁杉属花粉形态及其聚类分析研究

陈芷涵¹^,³(), 尚鑫¹^,², 谢淦², 张鑫¹^,^*()

1.西北农林科技大学林学院,陕西杨凌 712100
2.中国科学院植物研究所,北京 100093
3.中国科学院昆明植物研究所,云南昆明 650201

收稿日期:2024-06-17 修回日期:2024-07-10 出版日期:2024-07-30 发布日期:2024-08-05
通讯作者: *张鑫(xin.zhang@nwafu.edu.cn),讲师,博士。
作者简介:
陈芷涵(13657974778@163.com)。
基金资助:
国家植物标本资源库项目(E0117G1001)

Phylogenetic relationship and evolutionary patterns of Tsuga pollen morphology: a cluster analysis-based study

CHEN Zhihan¹^,³(), SHANG Xin¹^,², XIE Zin², ZHANG Xin¹^,^*()

1. College of Forestry, Northwest A&F University, Yangling 712100,China
2. Institute of Botany, Chinese Academy of Sciences, Beijing 100093, China
3. Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650201, China

Received:2024-06-17 Revised:2024-07-10 Online:2024-07-30 Published:2024-08-05

摘要/Abstract

摘要：

【目的】对铁杉属(Tsuga)现存所有10种植物花粉形态特征进行观察比较,结合花粉化石分析铁杉属系统发育关系及花粉形态的演化过程。【方法】通过扫描电子显微镜(SEM)观察铁杉属的花粉形态特征,并利用7个定量指标和1个定性指标进行系统聚类分析(HCA)。【结果】铁杉属花粉为N₁P₃C₁型花粉,远极面具薄壁区,赤道轴长度为21.20~58.00 μm,多无气囊,稀有气囊;花粉粒表面多疣状,稀微疣状,表面有棘突或无。系统聚类分析显示铁杉属花粉可以分类为3类:①有气囊,无棘突类:山地铁杉(T. mertensiana)与长苞铁杉(Nothotsuga longibracteata);②无气囊,无棘突类:卡罗莱纳铁杉(T. caroliniana)、加拿大铁杉(T. canadensis)与化石种2 (T. sp.2);③无气囊,有棘突类: 异叶铁杉(T. heterophylla)、郁林铁杉(T. ulleungensis)、米铁杉(T. diversifolia)、丽江铁杉(T. forrestii)、日本铁杉(T. sieboldii)、云南铁杉(T. dumosa)、铁杉(T. chinensis)、化石种1(T. sp.1 )和化石种3 (T. sp.3 )。【结论】铁杉属花粉是从有气囊向无气囊,无棘突向有棘突演化的,且花粉特征与分布地有密切关系,同一地区分布的花粉形态相似。基于花粉形态的聚类分析结果与分子系统发育树大体符合,可以通过花粉形态对铁杉属植物及化石进行物种鉴定,并为其系统发育研究提供参考。

关键词: 铁杉属, 花粉形态, 扫描电镜, 聚类分析

Abstract:

【Objective】To analyze the phylogenetic relationships and evolutional patterns of pollen morphology in Tsuga, we observed and compared the pollen characteristics of all 10 extant species of Tsuga along with pollen fossils.【Method】Pollen morphology was examined using scanning electron microscopy (SEM), and hierarchical cluster analysis (HCA) was conducted using seven quantitative indicators and one qualitative indicator.【Result】Tsuga pollens typically exhibits characteristics of the N₁P₃C₁ type, with a leptoma on the distal face. The equatorial length ranges from 21.20 to 58.00 μm, predominantly lacking sacci but occasionally with echinae saccate forms. The pollen surface is typically warty or sparsely micro-warty, with spines or without. According to the hierarchical cluster analysis, Tsuga could be divided into three categories: ①saccate but lacking echinae type: T. mertensiana and Nothotsuga longibracteata; ②lacking sacci and echinae type: T. caroliniana, T. canadensis and pollen fossil T. sp.2; ③lacking sacci but echinate type: T. heterophylla, T. ulleungensis, T. diversifolia, T. forrestii, T. sieboldii, T. dumosa, T. chinensis and pollen fossil T. sp.1, T. sp.3.【Conclusion】Pollen morphology in Tsuga has evolved from saccate to non-saccate forms, and from lacking spines to possessing spines. These pollen characteristics are closely related to geographic distribution, indicating similarity among pollen from the same regions. The clustering analysis based on pollen morphology largely aligns with molecular phylogenetic trees, offering a method to distinguish extant species and fossils of Tsuga and providing valuable insights for phylogenetic studies of Tsuga.

Key words: Tsuga, pollen morphology, scanning electron microscopy(SEM), cluster analysis

中图分类号:

Q944.5

陈芷涵,尚鑫,谢淦,等. 铁杉属花粉形态及其聚类分析研究[J]. 南京林业大学学报（自然科学版）, 2024, 48(4): 37-45.

CHEN Zhihan, SHANG Xin, XIE Zin, ZHANG Xin. Phylogenetic relationship and evolutionary patterns of Tsuga pollen morphology: a cluster analysis-based study[J].Journal of Nanjing Forestry University (Natural Science Edition), 2024, 48(4): 37-45.DOI: 10.12302/j.issn.1000-2006.202406025.

图/表 9

表1

表2

性状及编码"

性状 character	编码类型 code type	详细编码情况 code details
极轴长度(P) polar length	N	测量数据
赤道轴长度(E) equatorial length	N	测量数据
气囊长(L) length of saccus	N	测量数据
气囊宽(W) width of saccus	N	测量数据
气囊高(H) height of saccus	N	测量数据
气囊体积(V) volume of saccus	I	V= $43 × π × L × W × H$
棘突长(L_s) length of spine	N	测量数据
棘突宽(W_s) width of spine	N	测量数据
棘突长宽比(R_s) aspect ratio of spine	I	R_s=L_s/W_s
棘突密度(N_s) density of spine	N	N_s为单位面积(μm²)的棘突个数
外壁纹饰赋值(S) value of sexine sculpture	D	微疣状,疣状:0、1 无棘、具棘:0、1

表2

图1

表3

表4

图2

图3

图4

图5

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序号 No.	种 species	标本号 specimen barcode	采集号 collection number	采集地 collecting site
1	长苞铁杉N. longibracteata	NWFC 0047265	余恩萍00125	中国华南国家植物园
2	加拿大铁杉T. canadensis	MO 6400215	Gerrit Davidse 38706	美国密苏里植物园
3	卡罗莱纳铁杉T. caroliniana^[2]	—	B. K. Boom 39279	美国北卡罗莱纳州阿什维尔
4	铁杉T. chinensis	NWFC 0047266	尚鑫 00085	中国陕西宁陕火地塘
5	米铁杉T. diversifolia	PE 00145720	Debreczy,Biro,Racz 22618	美国哈佛大学阿诺德植物园
6	云南铁杉T. dumosa	MO 6061137	Gaoligong Shan Biodiversity Surevey 29176	中国云南腾冲明光镇
7	丽江铁杉T. forrestii	PYU 02000384	64250	中国云南昆明禄劝
8	异叶铁杉T. heterophylla	MO 6065231	A. D. E. Elmet 2482	美国华盛顿州克拉勒姆湾
9	山地铁杉T. mertensiana	MO 1255214	J. William Thompson 15036	美国斯诺夸尔米国家森林公园艾克斯山
10	日本铁杉T. sieboldii	MO 05089273	S. Tsugaru, Gen Murata &T. Takahashi 22360	日本京都府船井郡日吉町
11	郁陵铁杉T. ulleungensis	NWFC 0047268	Taewon Kim 231124	韩国郁陵岛
12	化石种1 T. sp.1^[33]	—	—	奥地利卡林西亚州拉万塔尔盆地
13	化石种2 T. sp.2^[33]	—	—	奥地利卡林西亚州拉万塔尔盆地
14	化石种3 T. sp.3	—	JH-0801	中国浙江宁海县

种 species	P/μm	E/μm	L/ μm	W/ μm	H/μm	V/μm³	L_s/μm	W_s/μm	R_s	N_s/ (个·μm^-2)	外壁纹饰类型 style of sexine sculpture	S
长苞铁杉 N. longibracteata	35.61±8.06	76.64±6.29	56.25±8.18	42.04±5.35	28.12±5.47	278541.02	—	—	—	—	微疣状、无棘	0
加拿大铁杉 T. canadensis	21.20±7.17	42.74±8.04	—	—	—	0	—	—	—	—	疣状、无棘	1
卡罗莱纳铁杉 T. caroliniana^[2]	58	95.3	—	—	—	0	—	—	—	—	疣状、无棘	1
铁杉 T. chinensis	33.64±10.92	60.31±6	—	—	—	0	1.17±0.41	0.49±0.15	2.39	0.36±0.09	疣状、具棘	2
米铁杉 T. diversifolia	28.6±8.01	61.71±10.69	—	—	—	0	1.32±0.40	0.48±0.13	2.75	0.20±0.09	疣状、具棘	2
云南铁杉 T. dumosa	23.17±6.28	57.43±9.79	—	—	—	0	1.16±0.34	0.54±0.12	2.15	0.39±0.13	疣状、具棘	2
丽江铁杉 T. forrestii	26.04±7.90	53.82±12.14	—	—	—	0	1.09±0.39	0.57±0.24	1.91	0.24±0.09	疣状、具棘	2
异叶铁杉 T. heterophylla	34.19±6.69	69.07±5.84	—	—	—	0	0.83±0.26	0.60±0.14	1.38	0.13±0.05	疣状、具棘	2
山地铁杉 T. mertensiana	25.27±5.62	53.23±8.52	30.37±7.60	22.19±2.34	26.98±10.80	76 161.00	—	—	—	—	微疣状、无棘	0
日本铁杉 T. sieboldii	49.50^[2]	65.30	—	—	—	0	1.25±0.55	0.58±0.31	2.16	0.40	疣状、具棘	2
郁陵铁杉 T. ulleungensis	23.48	69.74	—	—	—	0	1.12±0.34	0.61±0.17	1.84	0.18±0.05	疣状、具棘	2
化石种1 T. sp.1^[33]	47.2±6.97	70.98±1.88	—	—	—	0	0.86±0.31	0.48±0.2	1.79	0.25±0.12	疣状、具棘	2
化石种2 T. sp.2^[33]	42.26±0.91	63.43±13.76	—	—	—	0	—	—	—	—	疣状、具棘	2
化石种3 T. sp.3	—	66.81	—	—	—	0	1.67±0.64	0.66±0.19	2.53	0.31±0.06	疣状、具棘	2

性状 character	主成分 principal component
性状 character	PC1	PC2	PC3
赤道轴长度 equatorial length	-0.042	0.040	0.982
气囊长 length of saccus	-0.330	0.370	-0.018
气囊宽 width of saccus	-0.330	0.371	-0.013
气囊高 height of saccus	-0.329	0.317	-0.138
气囊体积 volume of saccus	-0.310	0.390	0.083
棘突长 length of spine	0.331	0.351	-0.010
棘突宽 wdth of spine	0.333	0.320	0.023
棘突长宽比 aspect ratio of spine	0.335	0.349	-0.022
棘突密度 density of spine	0.317	0.348	-0.049
纹饰赋值 value of sexine sculpture	0.377	0.056	0.080
特征值 latent root	6.726	1.941	1.018
贡献率/% contributor ratio	67.259	19.415	10.176
累计贡献率/% accumulative contributor ratio	67.259	86.674	96.850

铁杉属花粉形态及其聚类分析研究

Phylogenetic relationship and evolutionary patterns of Tsuga pollen morphology: a cluster analysis-based study

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