东亚特有间断分布植物蛛网萼属的生物地理学研究
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摘要
蛛网萼属(Platycrater arguta Sieb. et Zucc)隶属于绣球花科(Hydrangeaceae),是中国—日本间断分布的特有单型属植物。在中国仅零星分布于浙江、福建、江西等地,在日本分布于本州中、南部以及九州和四国。属下仅包括两个变种：中国变种(Platycrater arguta var. sinensis H. Hara)与日本原变种(var. arguta)。本研究在标本查阅、野外调查的基础上,运用群体遗传学的原理与方法对中国和日本的14个蛛网萼群体(314个个体)进行了生物地理学研究。联合分析了cpDNA (psbA-trnH, trnD-trnE)、nrDNA ITS.单拷贝核基因(Tpi)以及微卫星标记(nSSRs)的群体遗传变异和遗传结构,结合生态位模拟(ENM),分析了物种/谱系进化历史与间断地理分布式样的形成原因,根据遗传变异的地理分布阐明了其优先保护点与迁地保护的取样策略。主要研究结果如下：
1基于叶绿体DNA片段的分析
蛛网萼群体具有相对较高的遗传多样性(RT=4.02,hT=0.88,πT=0.00475)和明显的谱系地理结构(NST=0.857>GST=0.787)。两个变种分化明显(FST=0.915),这可能是地理隔离造成的。TCS单倍型谱系关系分析表明,中国和日本群体相对独立,无共享单倍型。系统发育分析表明中国和日本单倍型构成一个并系类群。中国和日本群体的失陪分布分析不拒绝扩张模型(P>0.05)。中国和日本群体的cpDNA单倍型分化时间(0.89Ma)。分化以后,日本群体的扩张比中国群体明显。
2基于核基因标记nrDNA ITS和单拷贝核基因Tpi的分析
ITS和Tpi标记分别检测到33/31种单倍型。在属水平上,两个标记均表现出较高的苷酸多态性和单倍型多态性(Tpi：h=0.92,π=0.02；ITS：h=0.96,π=0.04)。中国和日本群体nrDNA ITS和Tpi无共享单倍型。ITS单倍型序列系统发育分析支持蛛网萼两个变种的分类学地位。
3微卫星多态性和群体遗传结构
开发了12对蛛网萼特异的微卫星引物,将具有较好多态性的7对引物用于群体遗传学分析。共检测到220个等位基因,中国群体的平均等位基因数(NA=66)高于日本(NA=43)。微卫星标记显示出较高的属水平遗传多样性(HT=0.89)。STRUCTURE分析表明,K=7时,中国群体分成5组,日本群体出现遗传混合现象。K=2时,中国的C7群体与日本群体表现出同源性。
4基于多个信息位点的隔离分化遗传学分析
结合cpDNA,ITS,Tpi和SSR数据的IMa分析表明,蛛网萼中国和日本群体的分化时间为0.89Myr。与日本群体(NA=24.45×104)相比较而言,中国群体(NA=46.02×104)具有更高的有效群体大小。
5生态位模型
全部54个蛛网萼分布记录被用于生态位模拟分析。基于生态位模型的分析支持了基于分子遗传标记分析提出的“蛛网萼群体分布区在末次冰期时收缩至避难所,冰后期发生扩张”的假说。变量贡献分析提出蛛网萼的生境与夏季的降雨量关系密切,这一结果为野外生境调查结论提供了进一步佐证。
综上所述,中国和日本的蛛网萼变种分化早于末次冰期,分化以后分别孑遗于中国东海两侧。两个变种的分化很可能是受第四纪早期地质气候变化的影响而导致的异域物种形成。第四纪末次冰期时中国东海路桥的出现并未给蛛网萼中国和日本群体的迁移和基因交流提供便利。不完全谱系分选是导致蛛网萼群体(特别是中国群体)目前遗传格局的主要原因。日本群体存在南、北两个避难所,两个避难所的群体后期扩张的过程中,在本州中部发生了二次接触。同时,本研究还确定了蛛网萼的多个特殊进化单元,并提出了相关保护策略。
Platycrater arguta Sieb. et Zucc (Hydrangeaceae) is an East Asia endemic genus disjuncted distribution in East China and South Japan. It rarely distributed in Zhejiang, Fujian and Jiangxi province in China, and Central Honshu, Kyushu and Shikoku in Japan. The two varieties within the genus, Platycrater arguta var. sinensis H. Hara and Platycrater arguta var. arguta, distributed in China and Japan respectively. After a preliminary survey of specimen, field sampling and habitat observation across the known distribution plots, population genetics study of14Platycrater arguta populations (including314individuals) were carried out using multiple molecular markers (cpDNA psbA-trnH and trnD-trnE regions, nrDNA ITS, single-copy nuclear gene Tpi and seven nSSRs). We integrated the genetic diversity and population structure of Platycrater arguta with the addition of Ecological Niche Modeling analysis to interpret allopatric speciation evolution of Platycrater arguta and population demographic history after speciation. ESUs and ex-situ conservation strategies was also discussed. Major results are shown below:
1. Chloroplast DNA analysis
High level of genetic diversity (RT=4.02, hτ=0.88, πT=0.00475) and significant population genetic structure (NST=0.857> GST=0.787) was observed within Platycrater arguta populations. Strong genetic differentiation (FST=0.915) of the two Platycrater arguta varieties results from geographic isolation. Japan haplotypes are paraphyletic nested by China haplotypes. TCS haplotype analysis shows no shared haplotypes between China and Japan populations. A paraphyletic relationship between China and Japan haplotypes were observed from the phylogenetic analysis. MDA analysis of China and Japan population did not refuse the expansion model (P>0.05).The divergence time of cpDNA lineages are earlier than the Last Glacial Maximum (0.89Ma). Significant expansion was observed in Japan population after the divergence of the two varieties.
2. Nuclear marker analysis based on nrDNA ITS and single-copy nuclear gene Tpi
In total,33/31haplotypes were detected in the ITS and Tpi marker respectively. High level of nucleotide diversity and haplotype diversity were observed at the genus level (Tpi:h=0.92, rc=0.02; ITS:h=0.96, rc=0.04). No shared haplotypes were found for both nuclear sequence markers. ITS phylogenetic analysis supports the status of the two varieties.
3. Microsatellite marker diversity and population genetic structure
Twelve Platycrater arguta specific microsatellite markers were developed using two different strategies. Seven of them, with higher genetic diversity, were used in the population scanning. In total,220alleles were detected. China population (Na=66) has a higher average allele than Japan (Na=43). High level of genetic diversity was also observed in nSSRs data (Hr=0.89). STRUCTURE analysis demonstrate that when K=7, China populations were grouped into five clusters, while genetic admixture pattern were observed in Japan populations; when K=7, C7and Japan populations are homology.
4. Isolation with Migration analysis based on multiple markers
IMa analysis based on cpDNA, ITS, Tpi and SSR markers shows the divergence time of the two varieties are0.89Ma. Compare to Japan population (NA=24.45×104), a higher effective population size were observed in China population (NA=46.02×104).
5. Ecological Niche Modeling
All54distribution records of Platycrater were included in the ENM analysis. Ecological Niche Modeling simulated the potential distribution area of Platycrater arguta in present day, LGM and LIG. The simulations well support the scenario that Platycrater arguta populations contracted and survived in the refuges during the LGM and then expanded after the glaciation. Variable contribution analyses show that BIO18(Precipitation warmest quarter) has the highest relative contribution of the19environmental variables to the maxent model under both two LGM models (CCSM:36.9%; MIROC:48.2%). This result is consistent with our field observation.
In conclusion, the two varieties of Platycrater in East China and South Japan divergence earlier than LGM and survived on the East and South side of East China Sea. Allopatric speciation induced by climate change in early Quaternary isolated the Platycrater arguta ancestral population. The occurrence of East China Sea landbridge during the LGM did not facilitate bidirectional migration of Platycrater arguta after their divergence. Incomplete lineage sorting may play an important role during the evolution of Platycrater (especially for Platycrater arguta var. sinensis). Two refuges were detected in South Japan in the North and South region respectively. Each refuge possesses its own genetic components, a second contact happened in the central region of Japan when the two refuge population expansion during the post glacial period. At the same time, multiple evolutionary units were identified based on our study; related conservation strategies were also discussed.
1基于叶绿体DNA片段的分析
蛛网萼群体具有相对较高的遗传多样性(RT=4.02,hT=0.88,πT=0.00475)和明显的谱系地理结构(NST=0.857>GST=0.787)。两个变种分化明显(FST=0.915),这可能是地理隔离造成的。TCS单倍型谱系关系分析表明,中国和日本群体相对独立,无共享单倍型。系统发育分析表明中国和日本单倍型构成一个并系类群。中国和日本群体的失陪分布分析不拒绝扩张模型(P>0.05)。中国和日本群体的cpDNA单倍型分化时间(0.89Ma)。分化以后,日本群体的扩张比中国群体明显。
2基于核基因标记nrDNA ITS和单拷贝核基因Tpi的分析
ITS和Tpi标记分别检测到33/31种单倍型。在属水平上,两个标记均表现出较高的苷酸多态性和单倍型多态性(Tpi：h=0.92,π=0.02；ITS：h=0.96,π=0.04)。中国和日本群体nrDNA ITS和Tpi无共享单倍型。ITS单倍型序列系统发育分析支持蛛网萼两个变种的分类学地位。
3微卫星多态性和群体遗传结构
开发了12对蛛网萼特异的微卫星引物,将具有较好多态性的7对引物用于群体遗传学分析。共检测到220个等位基因,中国群体的平均等位基因数(NA=66)高于日本(NA=43)。微卫星标记显示出较高的属水平遗传多样性(HT=0.89)。STRUCTURE分析表明,K=7时,中国群体分成5组,日本群体出现遗传混合现象。K=2时,中国的C7群体与日本群体表现出同源性。
4基于多个信息位点的隔离分化遗传学分析
结合cpDNA,ITS,Tpi和SSR数据的IMa分析表明,蛛网萼中国和日本群体的分化时间为0.89Myr。与日本群体(NA=24.45×104)相比较而言,中国群体(NA=46.02×104)具有更高的有效群体大小。
5生态位模型
全部54个蛛网萼分布记录被用于生态位模拟分析。基于生态位模型的分析支持了基于分子遗传标记分析提出的“蛛网萼群体分布区在末次冰期时收缩至避难所,冰后期发生扩张”的假说。变量贡献分析提出蛛网萼的生境与夏季的降雨量关系密切,这一结果为野外生境调查结论提供了进一步佐证。
综上所述,中国和日本的蛛网萼变种分化早于末次冰期,分化以后分别孑遗于中国东海两侧。两个变种的分化很可能是受第四纪早期地质气候变化的影响而导致的异域物种形成。第四纪末次冰期时中国东海路桥的出现并未给蛛网萼中国和日本群体的迁移和基因交流提供便利。不完全谱系分选是导致蛛网萼群体(特别是中国群体)目前遗传格局的主要原因。日本群体存在南、北两个避难所,两个避难所的群体后期扩张的过程中,在本州中部发生了二次接触。同时,本研究还确定了蛛网萼的多个特殊进化单元,并提出了相关保护策略。
Platycrater arguta Sieb. et Zucc (Hydrangeaceae) is an East Asia endemic genus disjuncted distribution in East China and South Japan. It rarely distributed in Zhejiang, Fujian and Jiangxi province in China, and Central Honshu, Kyushu and Shikoku in Japan. The two varieties within the genus, Platycrater arguta var. sinensis H. Hara and Platycrater arguta var. arguta, distributed in China and Japan respectively. After a preliminary survey of specimen, field sampling and habitat observation across the known distribution plots, population genetics study of14Platycrater arguta populations (including314individuals) were carried out using multiple molecular markers (cpDNA psbA-trnH and trnD-trnE regions, nrDNA ITS, single-copy nuclear gene Tpi and seven nSSRs). We integrated the genetic diversity and population structure of Platycrater arguta with the addition of Ecological Niche Modeling analysis to interpret allopatric speciation evolution of Platycrater arguta and population demographic history after speciation. ESUs and ex-situ conservation strategies was also discussed. Major results are shown below:
1. Chloroplast DNA analysis
High level of genetic diversity (RT=4.02, hτ=0.88, πT=0.00475) and significant population genetic structure (NST=0.857> GST=0.787) was observed within Platycrater arguta populations. Strong genetic differentiation (FST=0.915) of the two Platycrater arguta varieties results from geographic isolation. Japan haplotypes are paraphyletic nested by China haplotypes. TCS haplotype analysis shows no shared haplotypes between China and Japan populations. A paraphyletic relationship between China and Japan haplotypes were observed from the phylogenetic analysis. MDA analysis of China and Japan population did not refuse the expansion model (P>0.05).The divergence time of cpDNA lineages are earlier than the Last Glacial Maximum (0.89Ma). Significant expansion was observed in Japan population after the divergence of the two varieties.
2. Nuclear marker analysis based on nrDNA ITS and single-copy nuclear gene Tpi
In total,33/31haplotypes were detected in the ITS and Tpi marker respectively. High level of nucleotide diversity and haplotype diversity were observed at the genus level (Tpi:h=0.92, rc=0.02; ITS:h=0.96, rc=0.04). No shared haplotypes were found for both nuclear sequence markers. ITS phylogenetic analysis supports the status of the two varieties.
3. Microsatellite marker diversity and population genetic structure
Twelve Platycrater arguta specific microsatellite markers were developed using two different strategies. Seven of them, with higher genetic diversity, were used in the population scanning. In total,220alleles were detected. China population (Na=66) has a higher average allele than Japan (Na=43). High level of genetic diversity was also observed in nSSRs data (Hr=0.89). STRUCTURE analysis demonstrate that when K=7, China populations were grouped into five clusters, while genetic admixture pattern were observed in Japan populations; when K=7, C7and Japan populations are homology.
4. Isolation with Migration analysis based on multiple markers
IMa analysis based on cpDNA, ITS, Tpi and SSR markers shows the divergence time of the two varieties are0.89Ma. Compare to Japan population (NA=24.45×104), a higher effective population size were observed in China population (NA=46.02×104).
5. Ecological Niche Modeling
All54distribution records of Platycrater were included in the ENM analysis. Ecological Niche Modeling simulated the potential distribution area of Platycrater arguta in present day, LGM and LIG. The simulations well support the scenario that Platycrater arguta populations contracted and survived in the refuges during the LGM and then expanded after the glaciation. Variable contribution analyses show that BIO18(Precipitation warmest quarter) has the highest relative contribution of the19environmental variables to the maxent model under both two LGM models (CCSM:36.9%; MIROC:48.2%). This result is consistent with our field observation.
In conclusion, the two varieties of Platycrater in East China and South Japan divergence earlier than LGM and survived on the East and South side of East China Sea. Allopatric speciation induced by climate change in early Quaternary isolated the Platycrater arguta ancestral population. The occurrence of East China Sea landbridge during the LGM did not facilitate bidirectional migration of Platycrater arguta after their divergence. Incomplete lineage sorting may play an important role during the evolution of Platycrater (especially for Platycrater arguta var. sinensis). Two refuges were detected in South Japan in the North and South region respectively. Each refuge possesses its own genetic components, a second contact happened in the central region of Japan when the two refuge population expansion during the post glacial period. At the same time, multiple evolutionary units were identified based on our study; related conservation strategies were also discussed.
引文
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