山茶科及紫茎属的系统发育和生物地理研究
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摘要
山茶科(Theaceae)隶属于杜鹃花目(Ericales),共9属,约有460多种。以前的形态和分子研究已经将其分为3个族,分别是山茶族(Theeae),大头茶族(Gordonieae)和紫茎族(Stewartieae)。紫茎属(Stewartia)和折柄茶属(Hartia)是紫茎族的两个属,其中紫茎属呈东亚一北美(EA-NA)间断分布,北美有两种,其余分布在东亚；折柄茶属分布在中国及老挝和越南地区。两属的分合关系一直是争论和关注的焦点。本研究通过叶绿体基因组小单拷贝(SSC)和反向重复(IR)区域长度约为40,000bp的序列和ITS,trnL-F以及psbA-trnH总长度约为2400bp的序列,结合相应的形态特征,分别研究了山茶科和紫茎族的系统发育关系。利用地理学,古气候学,古植物学,分子等方面证据,探讨了山茶科和紫茎族现代地理分布格局形成机制。主要研究结果如下：
1)山茶科的形态演化及分类系统
本研究通过SSC和IR序列对山茶科及其邻近的杜鹃花目主要支系进行了系统发育研究,结果强烈支持山茶科为不包括厚皮香科(Ternstroemiaceae, APG III系统中该类群归入五列木科Pentaphyllaceae)的单系类群,并且支持Prince&Park提出的山茶科3个族的划分,并明确了紫茎族是山茶科中最早分化(约48mya)的类群,山茶族和大头茶族具有较近的亲缘关系。通过3个族主要形态差异的祖先特征重建表明,山茶科的形态特征演化路线：种子中胚乳的丰富度从极度丰富直至完全消失；子房室的数目由5个减少到3个；中轴由几乎无中轴进化为具有中轴；导管穿孔方式由多个孔穴的复穿孔演变为少数穿孔。在习性的演化过程中,大头茶族中由常绿演变为落叶；紫茎族中由落叶衍生出常绿特征。
2)大头茶族的属间系统发育关系
大头茶族共包括3个属：木荷属(Schima), Franklinia和Gordonia。其中木荷属分布在亚洲的热带和亚热带,Franklinia和Gordonia为北美分布的单型属。SSC和IR数据表明Franklinia和木荷属具有较近的亲缘关系,两者在形态上具有较相似的果实和种子,Gordonia在该族早期(约14mya)已分化形成。
3)紫茎族的系统发育问题
本研究利用ITS, trnL-F和psbA-trnH片段对紫茎族的折柄茶属和紫茎属进行了系统发育分析,探讨了两者的分合问题。结果显示所有折柄茶属是一个具有高支持率的单系类群,而所有紫茎属的种则没能成功的聚为一个分支。SSC和IR数据结果显示,折柄茶属的单系分支镶嵌在紫茎属的分支内部。通过形态特征比较,发现两者除了习性和有无芽苞的区别外,不存在显著的差异。因此,本研究支持将紫茎属和折柄茶属合并处理,并将合并后的紫茎属分为3个组。通过对紫茎属落叶和常绿类群的形态观察和测量统计,提出了紫茎属新的分类处理,新拟了4个新异名,3个新组合。
4)生物地理和分布格局
山茶科的3个族都呈洲际间断分布,其中大头茶族和紫茎族呈东亚-北美间断分布。在大头茶族内,北美的Franklinia和东亚的木荷属的祖先在中新世中期(约11.32mya)通过白令陆桥向东亚传播；紫茎族由北美起源,在中新世中期(约14.99mya)通过白令陆桥向亚洲迁移,并形成北半球广泛分布的植物类群。第二次分歧为东亚紫茎在中新世晚期(约7.8mya)通过白令陆桥向北美迁移,并各自演化,由于北美西部的造山运动和第四纪冰期影响,形成紫茎族的现代地理分布格局。
通过现代紫茎属的地理分布统计,推断广东和广西地区是常绿类群的现代多样性分布中心,而日本列岛和中国大陆未分离前的连接区域曾是落叶类群的多样性分布中心,现代落叶类群主要分布在日本,中国的华东地区。
Theaceae is included in order Ericales by Angiosperm Phylogeny Group (APG). It comprises of nine genera, over460species. Several phylogenetic studies of Theaceae have been published in recent years based on morphological and molecular data. Three major lineages or tribes have been recognized including Stewartieae (Stewartia and Hartia), Gordonieae (Gordonia, Franklinia, and Schima), and Theeae (Camellia, Laplacea, Apterosperma, Polyspora, and Pyrenaria). In this study we used ca.40,000base pairs of chloroplast genes to gain a better understanding of the phylogeneics and biogeography of Theaceae. Stewartia is disjuncted distribution in eastern Asia and eastern North America (EA-ENA). It has two species in southeastern United States and the remaining species in eastern Asia. we used ITS, trnL-F and psbA-trnH sequences to reveal the relationship between Hartia and Stewartia. Combination with the geographic, paleoclimatic, fossil evidences, we studied the implication of biogeography pattern of EA-ENA. There are four main conclusions of our research:
1) Evolution of morphological characteristics of Theaceae and its classification system
Our broad sampling of Theaceae as well as other major lineages of the Ericales provides strong support for the monophyly of Theaceae excluding Ternstroemiaceae (Pentaphyllaceae in APG2009). Our sequence data from the chloroplast genome further support the three clades of Theaceae. The first-diverging position is Stewartieae. Theeae and Gordonieae are closer related to each other.
The trace of evolution of morphological characteristics in Theaceae is endosperm may have evolved from copious layers in Stewartieae, thin layers in Gordonieae, to total absence in Theeae. The number of capsule valves decreased from5in Stewartieae and Gordonieae to3in Theeae. Carpels may have evolved from free in Stewartieae, basally connate in Gordonieae to well developed central columnella in Theeae. Perforation bars in vessel have probably evolved in the direction of reduction from Stewartieae to Gordonieae and Theeae.
In habit character, the deciduousness has evolved at least twice in Theaceae. The deciduousness in Franklinia is a specialized feature in the tribe due to global temperature has decreased from the Mid-Miocene. Therefore, adaptation to increasingly cold climate might be associated with the transformation of leaf habit from evergreen to deciduous in the ancestral population of Franklinia. In contrast, Hartia diverged from deciduous lineages of Stewartia becoming evergreen at ca.14.99mya, when evergreen monsoon forests may have taken shape in Southeast Asia.
2) Phylogenetic relationship of Gordonieae.
There are three genera of Gordonieae:Schima, Franklinia and Gordonia. Generic relationships of Gordonieae have not been fully resolved by previous molecular studies. Our chloroplast data show strong support for the early divergence of Gordonia (ca.14mya) in Gordonieae and for the sister relationship of Franklinia and Schima. Comparing to Gordonia, Franklinia and Schima share more similar morphological characters in fruit and seed.
3) Phylogenetic relationship of Stewartieae.
The North American species Stewartia ovata is more closely related to Asian species of Stewartia than to the other North American species S. malacodendron. Species of Hartia form a strongly supported monophyletic clade, which is sister to the branch containing S. ovata and Asian species of Stewartia. Therefore, the Hartia clade is embedded within Stewartia in the phylogenetic trees. There're no significant morphological differences between two genera except the leave habit and the numbers of buds scales. Our data support the combination of Hartia and Stewartia. Combining molecular data (ITS, trnL-F and psbA-trnH) and morphological characters, a new classification of Stewartia is proposed. Three new combinations and four new synonmy are included. Furthermore, the genus Stewartia is divided into three sections because of the color of anthers and habits.
4) Biogeography and distribution pattern
The three tribes of Theaceae are all disjuncted distributed between eastern Asia and America. Gordonieae and Stewartieae show disjuncted distribution between eastern Asia (EA) and North America (NA). Franklinia and Gordonia are monotypic, North American taxa. Schima has a wide distribution in subtropical and tropical areas of southern Asia. In Stewartieae, Species of evergreen species (Hartia) occur in central and southern China, whereas deciduous species (Stewartia) have two species in southeastern United States and the remaining species in eastern Asia. Divergence times of eastern Asian-eastern North American disjunct taxa:Franklinia and Schima in Gordonieae is ca.11.32million years ago (mya), from North America to Asia through the Bering land bridge; the two divergence time between S. malacodendron to all other Stewartia species and Stewartia ovata to Asian deciduous species are ca.14.99mya and ca.7.8mya, respectively. The estimated time is from the Mid-Miocene to late Miocene, which agrees with recent estimates of the disjunction in other angiosperm taxa.
There are about half of evergreen species of Stewartia distributed in Guangdong and Guangxi province. And two thirds of deciduous species distributed in the region of eastern China and Japanese archipelago before it's formatted. Those regions are/or used be the diversity centers of evergreen and deciduous species of Stewartia, respectively.
1)山茶科的形态演化及分类系统
本研究通过SSC和IR序列对山茶科及其邻近的杜鹃花目主要支系进行了系统发育研究,结果强烈支持山茶科为不包括厚皮香科(Ternstroemiaceae, APG III系统中该类群归入五列木科Pentaphyllaceae)的单系类群,并且支持Prince&Park提出的山茶科3个族的划分,并明确了紫茎族是山茶科中最早分化(约48mya)的类群,山茶族和大头茶族具有较近的亲缘关系。通过3个族主要形态差异的祖先特征重建表明,山茶科的形态特征演化路线：种子中胚乳的丰富度从极度丰富直至完全消失；子房室的数目由5个减少到3个；中轴由几乎无中轴进化为具有中轴；导管穿孔方式由多个孔穴的复穿孔演变为少数穿孔。在习性的演化过程中,大头茶族中由常绿演变为落叶；紫茎族中由落叶衍生出常绿特征。
2)大头茶族的属间系统发育关系
大头茶族共包括3个属：木荷属(Schima), Franklinia和Gordonia。其中木荷属分布在亚洲的热带和亚热带,Franklinia和Gordonia为北美分布的单型属。SSC和IR数据表明Franklinia和木荷属具有较近的亲缘关系,两者在形态上具有较相似的果实和种子,Gordonia在该族早期(约14mya)已分化形成。
3)紫茎族的系统发育问题
本研究利用ITS, trnL-F和psbA-trnH片段对紫茎族的折柄茶属和紫茎属进行了系统发育分析,探讨了两者的分合问题。结果显示所有折柄茶属是一个具有高支持率的单系类群,而所有紫茎属的种则没能成功的聚为一个分支。SSC和IR数据结果显示,折柄茶属的单系分支镶嵌在紫茎属的分支内部。通过形态特征比较,发现两者除了习性和有无芽苞的区别外,不存在显著的差异。因此,本研究支持将紫茎属和折柄茶属合并处理,并将合并后的紫茎属分为3个组。通过对紫茎属落叶和常绿类群的形态观察和测量统计,提出了紫茎属新的分类处理,新拟了4个新异名,3个新组合。
4)生物地理和分布格局
山茶科的3个族都呈洲际间断分布,其中大头茶族和紫茎族呈东亚-北美间断分布。在大头茶族内,北美的Franklinia和东亚的木荷属的祖先在中新世中期(约11.32mya)通过白令陆桥向东亚传播；紫茎族由北美起源,在中新世中期(约14.99mya)通过白令陆桥向亚洲迁移,并形成北半球广泛分布的植物类群。第二次分歧为东亚紫茎在中新世晚期(约7.8mya)通过白令陆桥向北美迁移,并各自演化,由于北美西部的造山运动和第四纪冰期影响,形成紫茎族的现代地理分布格局。
通过现代紫茎属的地理分布统计,推断广东和广西地区是常绿类群的现代多样性分布中心,而日本列岛和中国大陆未分离前的连接区域曾是落叶类群的多样性分布中心,现代落叶类群主要分布在日本,中国的华东地区。
Theaceae is included in order Ericales by Angiosperm Phylogeny Group (APG). It comprises of nine genera, over460species. Several phylogenetic studies of Theaceae have been published in recent years based on morphological and molecular data. Three major lineages or tribes have been recognized including Stewartieae (Stewartia and Hartia), Gordonieae (Gordonia, Franklinia, and Schima), and Theeae (Camellia, Laplacea, Apterosperma, Polyspora, and Pyrenaria). In this study we used ca.40,000base pairs of chloroplast genes to gain a better understanding of the phylogeneics and biogeography of Theaceae. Stewartia is disjuncted distribution in eastern Asia and eastern North America (EA-ENA). It has two species in southeastern United States and the remaining species in eastern Asia. we used ITS, trnL-F and psbA-trnH sequences to reveal the relationship between Hartia and Stewartia. Combination with the geographic, paleoclimatic, fossil evidences, we studied the implication of biogeography pattern of EA-ENA. There are four main conclusions of our research:
1) Evolution of morphological characteristics of Theaceae and its classification system
Our broad sampling of Theaceae as well as other major lineages of the Ericales provides strong support for the monophyly of Theaceae excluding Ternstroemiaceae (Pentaphyllaceae in APG2009). Our sequence data from the chloroplast genome further support the three clades of Theaceae. The first-diverging position is Stewartieae. Theeae and Gordonieae are closer related to each other.
The trace of evolution of morphological characteristics in Theaceae is endosperm may have evolved from copious layers in Stewartieae, thin layers in Gordonieae, to total absence in Theeae. The number of capsule valves decreased from5in Stewartieae and Gordonieae to3in Theeae. Carpels may have evolved from free in Stewartieae, basally connate in Gordonieae to well developed central columnella in Theeae. Perforation bars in vessel have probably evolved in the direction of reduction from Stewartieae to Gordonieae and Theeae.
In habit character, the deciduousness has evolved at least twice in Theaceae. The deciduousness in Franklinia is a specialized feature in the tribe due to global temperature has decreased from the Mid-Miocene. Therefore, adaptation to increasingly cold climate might be associated with the transformation of leaf habit from evergreen to deciduous in the ancestral population of Franklinia. In contrast, Hartia diverged from deciduous lineages of Stewartia becoming evergreen at ca.14.99mya, when evergreen monsoon forests may have taken shape in Southeast Asia.
2) Phylogenetic relationship of Gordonieae.
There are three genera of Gordonieae:Schima, Franklinia and Gordonia. Generic relationships of Gordonieae have not been fully resolved by previous molecular studies. Our chloroplast data show strong support for the early divergence of Gordonia (ca.14mya) in Gordonieae and for the sister relationship of Franklinia and Schima. Comparing to Gordonia, Franklinia and Schima share more similar morphological characters in fruit and seed.
3) Phylogenetic relationship of Stewartieae.
The North American species Stewartia ovata is more closely related to Asian species of Stewartia than to the other North American species S. malacodendron. Species of Hartia form a strongly supported monophyletic clade, which is sister to the branch containing S. ovata and Asian species of Stewartia. Therefore, the Hartia clade is embedded within Stewartia in the phylogenetic trees. There're no significant morphological differences between two genera except the leave habit and the numbers of buds scales. Our data support the combination of Hartia and Stewartia. Combining molecular data (ITS, trnL-F and psbA-trnH) and morphological characters, a new classification of Stewartia is proposed. Three new combinations and four new synonmy are included. Furthermore, the genus Stewartia is divided into three sections because of the color of anthers and habits.
4) Biogeography and distribution pattern
The three tribes of Theaceae are all disjuncted distributed between eastern Asia and America. Gordonieae and Stewartieae show disjuncted distribution between eastern Asia (EA) and North America (NA). Franklinia and Gordonia are monotypic, North American taxa. Schima has a wide distribution in subtropical and tropical areas of southern Asia. In Stewartieae, Species of evergreen species (Hartia) occur in central and southern China, whereas deciduous species (Stewartia) have two species in southeastern United States and the remaining species in eastern Asia. Divergence times of eastern Asian-eastern North American disjunct taxa:Franklinia and Schima in Gordonieae is ca.11.32million years ago (mya), from North America to Asia through the Bering land bridge; the two divergence time between S. malacodendron to all other Stewartia species and Stewartia ovata to Asian deciduous species are ca.14.99mya and ca.7.8mya, respectively. The estimated time is from the Mid-Miocene to late Miocene, which agrees with recent estimates of the disjunction in other angiosperm taxa.
There are about half of evergreen species of Stewartia distributed in Guangdong and Guangxi province. And two thirds of deciduous species distributed in the region of eastern China and Japanese archipelago before it's formatted. Those regions are/or used be the diversity centers of evergreen and deciduous species of Stewartia, respectively.
引文
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