东亚草木菝葜的物种分化和亲缘地理学研究
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摘要
菝葜科草本菝葜组包括11个种,其中东亚有两种：牛尾菜(Smilax riparia)和白背牛尾菜(Smilax nipponica),北美有9个种,分别为：S. ecirrhata, S. herbace, S. biltmoreana, S. hugeri, S. illinoensis, S. lasioneuron, S. pseudo-china, S. pulverulenta和S.jamesii。东亚的草本菝葜广泛分布于中国中西部地区、东部和北部以及日本和韩国。本文选取了东亚和北美的草本菝葜的60个群体416个个体,利用2个叶绿体片段对东亚两个草本种及其之间的系统发育和亲缘地理学进行了研究,结果显示：
(1) rbcL/atpB和atpl/atpH基因间隔区联合分析产生了65种单倍型,具有64个变异位点；牛尾菜物种水平的单倍型多样性(h)为0.937,核苷酸多态性(Pi)为0.00739；高于白背牛尾菜物种水平的单倍型多样性(h=0.712)和核苷酸多态性(Pi=0.00279)；AMOVA分析结果显示牛尾菜和白背牛尾菜的遗传变异主要存在于群体间,群体间遗传分化(Fst)分别为0.68985和0.92952。
(2) cpDNA的单倍型多样性以及地理分布图表明东亚草本菝葜的多样性中心和冰期避难所有两个,一个是中国中西部地区,包括湖南天平山(TPS)、湖北大老岭(DLL)、神农架(SNJ)和咸丰(XF)群体；另一个避难所是中国东部地区,包括江苏宜兴(YX),安徽天堂寨(TTZ)和浙江天目山(TMS)群体。
(3) BEAST分析表明,东亚牛尾菜和白背牛尾菜的分化时间在16.2 mya,牛尾菜内群体间分化时间约在15.2 mya,白背牛尾菜种内群体分化时间约在8.5mya,处于晚第三纪草本植物大发展时期。
(4)推测东亚草本菝葜起源于中国西部,向东向北迁移形成现在的分布格局。本研究还揭示了中国台湾、日本以及朝鲜半岛的草本菝葜与大陆关系密切,起源于中国大陆。
(5) cpDNA数据构建的单倍型系统发育树表明牛尾菜和白背牛尾菜间不存在共有单倍型,物种分化明显；同时支持中国西南地区可能存在牛尾菜亚种。
There are 11 species in Smilax Sect, nemexia. They are disjuct in East Asia (EA) and North-America(NA). There are two species in Eastern Asia, Smilax riaparia and S. nipponica, while other nine in North-America. They are S. ecirrhata, S. herbacea, S. biltmoreana, S. hugeri, S. illinoensis, S. lasioneuron, S. pseudo-china, S. pulverulenta and S. jamesii. The two Asian herb Smilax are widely distributed in the Mid-Western China, Eastern China, Northern China, and also in Japan and Korea. To investigate the phylogeography and phylogeny of the two Asian species, we choose 60 populations (416 individuals) using two cpDNA fragements in this research, including 17 populations of North America as contrast. The results are as follows:
(1) The combined sequence of rbcL/atpB and atpl/atpH have detected 64 polymorphic sites and generated 65 haplotypes. The haplotype diversity (h) and the nucleotide diversity (Pi) in S. riparia are 0.937 and 0.00739, respectively, which is higher than that of S. nipponica'(Pi= 0.00279, h=0.712). AMOVA analysis revealed that most of the existing variation are mainly among populations, in S. riparia, Fst=0.68985 and in S. nipponia, Fst=0.92952.
(2) The haplotype diversity and the haplotype distributing map shows that there are two diversification centers and refuges in East Asia, one is the region of the Mid-Western China, including Mt.Tianping in Hunan province(TPS), and Dalaoling(DLL), Mt.Shennong(SNJ), Xianfeng (XF) in Hubei province; Another is in Eastern China,that is Yixing (YX) in Jiangsu province, Tiantangzhai (TTZ) in Anhui province and Mt. Tianmu (TMS) in Zhejiang province.
(3) The current pattern of herb Smilax in Asia came from the event of migrating eastward and northward from the west. It is also discovered that the herb Smilax in Taiwan, China, Japan and Korea has a close relationship with the mainland, which may be the original place.
(4) BEAST analysis shows that the divergence time of two species'is about 16.2 mya, the divergence time between populations in S. riparia is around 15.2 mya, while the S. nipponica populations differentiated 8.5 mya.
(5) The phylogenetic tree based on cpDNA indicates that there is no haplotype shared between S. riparia and S. nipponica, the differentiation of the two species is apparent.This research also surports the existence of var. acuminate in South-western China.
(1) rbcL/atpB和atpl/atpH基因间隔区联合分析产生了65种单倍型,具有64个变异位点；牛尾菜物种水平的单倍型多样性(h)为0.937,核苷酸多态性(Pi)为0.00739；高于白背牛尾菜物种水平的单倍型多样性(h=0.712)和核苷酸多态性(Pi=0.00279)；AMOVA分析结果显示牛尾菜和白背牛尾菜的遗传变异主要存在于群体间,群体间遗传分化(Fst)分别为0.68985和0.92952。
(2) cpDNA的单倍型多样性以及地理分布图表明东亚草本菝葜的多样性中心和冰期避难所有两个,一个是中国中西部地区,包括湖南天平山(TPS)、湖北大老岭(DLL)、神农架(SNJ)和咸丰(XF)群体；另一个避难所是中国东部地区,包括江苏宜兴(YX),安徽天堂寨(TTZ)和浙江天目山(TMS)群体。
(3) BEAST分析表明,东亚牛尾菜和白背牛尾菜的分化时间在16.2 mya,牛尾菜内群体间分化时间约在15.2 mya,白背牛尾菜种内群体分化时间约在8.5mya,处于晚第三纪草本植物大发展时期。
(4)推测东亚草本菝葜起源于中国西部,向东向北迁移形成现在的分布格局。本研究还揭示了中国台湾、日本以及朝鲜半岛的草本菝葜与大陆关系密切,起源于中国大陆。
(5) cpDNA数据构建的单倍型系统发育树表明牛尾菜和白背牛尾菜间不存在共有单倍型,物种分化明显；同时支持中国西南地区可能存在牛尾菜亚种。
There are 11 species in Smilax Sect, nemexia. They are disjuct in East Asia (EA) and North-America(NA). There are two species in Eastern Asia, Smilax riaparia and S. nipponica, while other nine in North-America. They are S. ecirrhata, S. herbacea, S. biltmoreana, S. hugeri, S. illinoensis, S. lasioneuron, S. pseudo-china, S. pulverulenta and S. jamesii. The two Asian herb Smilax are widely distributed in the Mid-Western China, Eastern China, Northern China, and also in Japan and Korea. To investigate the phylogeography and phylogeny of the two Asian species, we choose 60 populations (416 individuals) using two cpDNA fragements in this research, including 17 populations of North America as contrast. The results are as follows:
(1) The combined sequence of rbcL/atpB and atpl/atpH have detected 64 polymorphic sites and generated 65 haplotypes. The haplotype diversity (h) and the nucleotide diversity (Pi) in S. riparia are 0.937 and 0.00739, respectively, which is higher than that of S. nipponica'(Pi= 0.00279, h=0.712). AMOVA analysis revealed that most of the existing variation are mainly among populations, in S. riparia, Fst=0.68985 and in S. nipponia, Fst=0.92952.
(2) The haplotype diversity and the haplotype distributing map shows that there are two diversification centers and refuges in East Asia, one is the region of the Mid-Western China, including Mt.Tianping in Hunan province(TPS), and Dalaoling(DLL), Mt.Shennong(SNJ), Xianfeng (XF) in Hubei province; Another is in Eastern China,that is Yixing (YX) in Jiangsu province, Tiantangzhai (TTZ) in Anhui province and Mt. Tianmu (TMS) in Zhejiang province.
(3) The current pattern of herb Smilax in Asia came from the event of migrating eastward and northward from the west. It is also discovered that the herb Smilax in Taiwan, China, Japan and Korea has a close relationship with the mainland, which may be the original place.
(4) BEAST analysis shows that the divergence time of two species'is about 16.2 mya, the divergence time between populations in S. riparia is around 15.2 mya, while the S. nipponica populations differentiated 8.5 mya.
(5) The phylogenetic tree based on cpDNA indicates that there is no haplotype shared between S. riparia and S. nipponica, the differentiation of the two species is apparent.This research also surports the existence of var. acuminate in South-western China.
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