中国扁桃亚属四种野生扁桃的系统发育与物种分化

doi:10.17521/cjpe.2020.0366

植物生态学报 ›› 2021, Vol. 45 ›› Issue (9): 987-995.DOI: 10.17521/cjpe.2020.0366

中国扁桃亚属四种野生扁桃的系统发育与物种分化

王春成¹, 张云玲², 马松梅³^,^*(), 黄刚¹, 张丹¹, 闫涵³

¹石河子大学生命科学学院, 绿洲城镇与山盆系统生态兵团重点实验室, 新疆石河子 832000
²新疆维吾尔自治区草原总站, 乌鲁木齐 830049
³石河子大学理学院, 绿洲城镇与山盆系统生态兵团重点实验室, 新疆石河子 832000

收稿日期:2020-11-09 接受日期:2021-06-02 出版日期:2021-09-20 发布日期:2021-11-18
通讯作者: 马松梅
作者简介:ORCID: *马松梅: 0000-0002-3107-2256(shzmsm@126.com)
基金资助:
国家自然科学基金(41261011);国家自然科学基金(41561007);新疆维吾尔自治区草原生态修复治理补助项目(XJCYZZ202007)

Phylogeny and species differentiation of four wild almond species of subgen. Amygdalus in China

WANG Chun-Cheng¹, ZHANG Yun-Ling², MA Song-Mei³^,^*(), HUANG Gang¹, ZHANG Dan¹, YAN Han³

¹Key Laboratory of Ecological Corps for Oasis City and Mountain Basin System, College of Life Sciences, Shihezi University, Shihezi, Xinjiang 832000, China
²General Grassland Station of Xinjiang, Ürümqi 830049, China
³Key Laboratory of Ecological Corps for Oasis City and Mountain Basin System, College of Science, Shihezi University, Shihezi, Xinjiang 832000, China

Received:2020-11-09 Accepted:2021-06-02 Online:2021-09-20 Published:2021-11-18
Contact: MA Song-Mei
Supported by:
National Natural Science Foundation of China(41261011);National Natural Science Foundation of China(41561007);Grassland Ecological Restoration and Management Grant Project of Xinjiang Uygur Autonomous Region(XJCYZZ202007)

摘要/Abstract

摘要：

基于核基因ITS序列研究中国4种野生扁桃: 新疆野扁桃(Amygdalus ledebouriana)、蒙古扁桃(A. mongolica)、长柄扁桃(A. pedunculata)和西康扁桃(A. tangutica)的系统发育关系和物种分化, 为4种植物的遗传与演化研究提供数据支撑。利用单倍型网络和主坐标分析揭示单倍型的聚类关系; 利用最大似然树和贝叶斯系统树分析单倍型的系统发育关系; 利用R语言“ecospat”包分析4种扁桃的生态位分化及其环境驱动因子。4种扁桃ITS1-ITS4片段总长为634 bp, 鉴别出27个核苷酸变异位点, 共定义了28个单倍型。4种扁桃种间最小遗传距离均大于种内最大遗传距离, 种间存在显著的遗传分化。4种扁桃的单倍型聚为两支: 新疆野扁桃、蒙古扁桃和西康扁桃聚为一支, 长柄扁桃为另一支; 单倍型网络和主坐标分析揭示的单倍型聚类关系与系统树一致。西康扁桃与蒙古扁桃、与长柄扁桃之间均表现出了显著的生态位分化, 最暖月最高气温、年平均气温、最冷月最低气温和最暖季降水量是驱动物种生态位分化的关键因子。

关键词: 野生扁桃, ITS序列, 系统发育, 物种分化, 生态位分化

Abstract:

Aims Based on the ITS sequences, we aimed to analyze the spatial genetic structure, genealogy relationship, and species differentiation of the Amygdalus ledebouriana, A. mongolica, A. pedunculata, A. tangutica in China, and provide data for the future studies on the four species' genetics and evolution.

Methods The median-joining network and principal coordinate analysis (PCoA) were used to reveal haplotype clustering. The maximum likelihood method and Bayesian method were used to analyze the phylogenetic relationships of haplotypes. The “ecospat” package in R 4.0.2 was used to analyze the ecological niche divergence of four almond species and their environmental drivers.

Important findings The total length of the ITS1-ITS4 fragment after corrected alignment was 634 bp, 27 nucleotide variants detected, and a total of 28 haplotypes were identified. The minimum genetic distance among the four almond species is greater than the maximum genetic distance within species, and there are significant genetic differentiations among species. The haplotypes of the four almond species clustered into two branches: A. ledebouriana, A. mongolicaand A. tanguticafor one clade, and A. pedunculatafor the other. The revealed dendrogram relationship of haplotype network and PCoA analysis is consistent with the phylogenetic tree. The significant niche divergence was observed between A. tangutica and A. mongolica, as well as between A. tangutica and A. pedunculata, with annual mean temperature, max temperature of warmest month, min temperature of coldest month and precipitation of warmest quarter as key drivers of niche divergence.

Key words: wild almond species, ITS sequences, phylogeny, species differentiation, niche divergence

王春成, 张云玲, 马松梅, 黄刚, 张丹, 闫涵. 中国扁桃亚属四种野生扁桃的系统发育与物种分化. 植物生态学报, 2021, 45(9): 987-995. DOI: 10.17521/cjpe.2020.0366

WANG Chun-Cheng, ZHANG Yun-Ling, MA Song-Mei, HUANG Gang, ZHANG Dan, YAN Han. Phylogeny and species differentiation of four wild almond species of subgen. Amygdalus in China. Chinese Journal of Plant Ecology, 2021, 45(9): 987-995. DOI: 10.17521/cjpe.2020.0366

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2020.0366

https://www.plant-ecology.com/CN/Y2021/V45/I9/987

图/表 6

图1 4种扁桃亚属野生植物的野外采样点。BEC, 布尔津县; BTE, 大青山2; BTQ, 大青山1; DBL, 迭部县1; DBZ, 迭部县2; DMB, 达茂旗3; DMD, 达茂旗1; DMN, 达茂旗2; GQD, 固阳1; GQX, 固阳3; GX, 固阳2; HBK, 哈巴河县; MZS, 马鬃山; SPQ, 松潘县; TBB, 塔城2; TBT, 塔城1; TLB1, 托里县1; TLB2, 托里县2; WD, 乌拉山; WG, 乌盖苏木乡; WLB, 大坝口; WLJ, 乌加禾镇; WSH, 苏海图; WSM, 大麦力沟; YCQ, 龙首山; YLD1, 榆林1; YLD2, 榆林2; YMB1, 裕民县1; YMB2, 裕民县2; YMF, 阴山2; YMM, 阴山1; ZQK, 图克木嘎查; ZQL, 贺兰山; ZQX, 舟曲县; ZTL, 特莫乌拉嘎查; ZTM, 苏力图嘎查; ZYS, 祁连山。

Fig. 1 Field sampling points of four subgen. Amygdalus wild plants. BEC, Buerjin County; BTE, Group 2 in Daqing Mountain; BTQ, Group 1 in Daqing Mountain; DBL, Group 1 in Têwo County; DBZ, Group 2 in Têwo County; DMB, Group 3 in Damao Banner; DMD, Group 1 in Damao Banner; DMN, Group 2 in Damao Banner; GQD, Group 1 in Guyang County; GQX, Group 3 in Guyang County; GX, Group 2 in Guyang County; HBK, Habahe County; MZS, Mazong Mountain; SPQ, Songpan County; TBB, Group 2 Tacheng City; TBT, Group 1 Tacheng City; TLB1, Group 1 in Toli County; TLB2, Group 2 in Toli County; WD, Wula Mountain; WG, Ugai Sumu Township; WLB, Dam Mouth; WLJ, Wujiahe Town; WSH, Suhaitu; WSM, Damaili Furrow; YCQ, Longshou Mountain; YLD1, Group 1 in Yulin City; YLD2, Group 2 in Yulin City; YMB1, Group 1 in Yumin County; YMB2, Group 2 in Yumin County; YMF, Group 2 in Yinshan Mountains; YMM, Group 1 in Yinshan Mountains; ZQK, Tukemu Gazha; ZQL, Helan Mountain; ZQX, Zhugqu County; ZTL, Temowula Gazha; ZTM, Su Litu Gazha; ZYS, Qilianshan Mountain.

图2 扁桃亚属4种野生植物28个单倍型(R1-R28)的地理分布及其单倍型网络。图中种群采样点编码与图1一致, 饼图表示各种群的单倍型频率。A, 单倍型网络, 图中圆圈大小与单倍型频率成正比, 节点间的分支长度大致与单倍型的突变数成正比, 相应分支附近附有步长; 稠李和山桃作为外类群。

Fig. 2 Geographical distribution and the haplotype network of 28 haplotypes (R1-R28) of four subgen. Amygdalus wild plants. The population field sampling point codes in the figure are consistent with the ones in Fig. 1. Pie graphs indicate the frequency of each haplotype of these populations. A, In the median-joining haplotypes network, the sizes of the circles in the network are proportional to the haplotype frequencies. Branch lengths are roughly proportional to the number of mutation steps between haplotypes and nodes; the true number of steps is shown near the corresponding branch sections. Padus racemosa and Amygdalus davidiana was used as outgroup.

图3 基于ITS序列计算的扁桃亚属4种野生植物种间和种内的遗传距离。

Fig. 3 Genetic distance among four subgen. Amygdalus wild plants based on ITS sequences.

图4 扁桃亚属4种野生植物基于种群水平的前3个坐标的主坐标分析(PCoA)。

Fig. 4 Plots of the first three coordinates of the principal coordinates analysis (PCoA) at the population level for four subgen. Amygdalus wild plants.

图5 扁桃亚属4种野生植物基于ITS序列的单倍型系统发育树。A, 最大似然(ML)树。分支点上方的数字为大于等于70的自展支持率。B, 贝叶斯树。分支节点右侧的数字表示大于0.70的后验概率值, 两系统发育树右侧的中括号表示为4个物种相应的亚支。

Fig. 5 Haplotype phylogenetic trees of four subgen. Amygdalus wild plants based on ITS sequences. A, Maximum likelihood (ML) tree. Bootstrap values equal to or greater than 70 are shown above the corresponding branching points. B, Bayesian tree. The values on the right of the branching points represent the posterior probability greater than 0.70; the brackets on the right of the two phylogenetic trees indicate the corresponding subclades of the four species.

图6 扁桃亚属4种野生植物生态位相似性与驱动因子贡献率。A-F, 生态位可视化。绿色表示第一个种的生态位空间, 红色表示第二个种的生态位空间, 蓝色表示重叠空间。D为生态位重叠分数。G, 驱动因子主成分分析。箭头表示相关的方向(同一方向表示高度相关)。Bio1, 年平均气温; Bio4, 温度季节性变化标准差; Bio5, 最暖月最高气温; Bio6, 最冷月最低气温; Bio15, 降水量的季节性变异系数; Bio17, 最干季降水量; Bio18, 最暖季降水量。环境因子后括号里的数字为贡献率排名。

Fig. 6 Ecological niche differentiation and the contribution of driving factors of four subgen. Amygdalus plants. A-F, Visualization of ecological niches. The green colour depicts the niche space of the first species, red of the second species and the overlapping range is shown in blue. D is the ecological niche overlap score. G, Driver factors principal component analysis. The arrow depicts the direction of correlation (same direction indicates a high correlation). Bio1, annual mean temperature; Bio4, temperature seasonality (standard deviation × 100); Bio5, max temperature of warmest month; Bio6, min temperature of coldest month; Bio15, precipitation seasonality (coefficient of variation); Bio17, precipitation of driest quarter; Bio18, precipitation of warmest quarter. The numbers in brackets after the environmental factors are the contribution rankings.

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中国扁桃亚属四种野生扁桃的系统发育与物种分化

Phylogeny and species differentiation of four wild almond species of subgen. Amygdalus in China

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