部分菊科入侵种的有性繁殖特征与入侵性的关系
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摘要
生物入侵是自然生态系统面临的最重要的全球性问题之一,它不仅是导致生物多样性丧失的主要原因之一,而且威胁着全球的生态环境和经济发展。
菊科是我国种子植物中最大的科之一,同时也是我国入侵种最多,危害最为严重的科。本文经过实地调研和全面查阅文献,共整理出入侵我国的菊科种58种,其中12个种已被列为2007年新公布的中华人民共和国进境植物检疫性有害生物名录。
本文对中国12个菊科入侵种的繁育系统、3个入侵种的有性繁殖特征与入侵性的关系、10个入侵种的瘦果的特性对入侵性的贡献进行了研究。目的是阐明部分菊科入侵种的繁殖特征,为菊科入侵种的管理提供有益的基础信息。
Baker认为,具有单亲繁殖特征的种在长距离扩散后,比那些依赖于双亲繁殖或传粉者的种更容易成功建群,这个论述被称为Baker定律。有关Baker定律在外来入侵种中的适用性一直存在着争议。本文随机选择了12个菊科入侵种,即紫茎泽兰(Ageratina adenophorum或Eupatorium adenophora)、胜红蓟(Agratum conyzoides).钻形紫菀(Aster subulatus)、三叶鬼针草(Bidens pilosa)、野塘蒿(Conyza bonariensis)、小蓬草(C.canadensis)、苏门白酒草(C.sumatrensis)、线叶金鸡菊(Coreopsis lanceolata)、一年蓬(Erigeron annuus)、加拿大一枝黄花(Solidago canadensis)、续断菊(Sonchus asper)和苦苣菜(S.oleraceus),对它们的繁育系统进行了研究。结果表明,其中的10个种(占83.3%)具有自交亲和和自交结实特性。结实率的统计分析和花粉一柱头的亲和性试验的综合分析表明,这12个入侵种中,2个种即紫茎泽兰和一年蓬(占16.7%)为无融合生殖,3个种即胜红蓟、小蓬草和苏门白酒草(占25%)为自体受精,5个种即野塘蒿、钻形紫菀、三叶鬼针草、续断菊和苦苣菜(占41.7%)为部分自体受精,2个多年生菊科入侵种即线叶金鸡菊和加拿大一枝黄花(占16.7%)为自交不亲和种。线叶金鸡菊和加拿大一枝黄花虽然自交不亲和,但它们可通过发达的根状茎进行繁殖。
以上结果表明,(1)Baker定律适用于本研究中的所有12个菊科入侵种,它们或者进行营养繁殖、或者通过形成不需要异株配子或传粉者的种子进行繁殖。(2)12个种中的10个种(占83.3%)在套袋的花序中结出了可育瘦果,并且其结实率在40%以上,表明通过自体受精、自动自交和无融合生殖产生种子的自交结实方式在这些菊科种的成功入侵中起重要作用。(3)对这12个入侵种的繁育系统与其入侵性相关性的分析表明,繁育系统与入侵性的强弱和危害程度方面存在着极高的相关性：繁育系统为无融合生殖或自体受精的几个种在相关文献中大多被列为入侵性强的种,如紫茎泽兰、一年蓬、胜红蓟、小蓬草和苏门白酒草,而繁育系统为部分自体受精或自交不亲和的几个一年生种,如野塘蒿、钻形紫菀、三叶鬼针草、续断菊、苦苣菜被列为入侵性一般的种。(4)对本研究的结果和已发表的资料的分析表明,12个一年生菊科入侵种中,11个具有两性花的种均可自交亲和或无融合生殖,具备自交结实的特性。12个多年生菊科入侵种中,除3个无融合生殖种外,其余的9个种显示出自交不亲和或不完全自交不亲和的特性。因而,自交结实在植物入侵中起重要作用的观点可能更多地与一年生菊科入侵种相关。
为进一步了解菊科入侵种的繁殖生物学特性在其成功入侵中的作用,对具有不同花序类型、开花时期不同、主要依靠种子进行繁殖的苏门白酒草、续断菊和三叶鬼针草的繁殖特性进行了深入研究,内容包括头状花序和小花的生物学特征和开花动态、花粉/胚珠(P/O)比、花粉活力和柱头可授性、交配系统等以及不同交配方式所产生的种子的萌发率,并对这些繁殖特征与入侵性的关系进行了分析。结果表明：(1)3个入侵种均为自交与异交混合型交配系统,具有同花序内自交结实的能力和灵活的交配机制,这些繁育系统的特征有利于其成功入侵。(2)它们所具有的持续结实或大结实量、高结实率、以及种子的高萌发率和扩散能力等特性在与当地种的竞争和促进入侵方面有重要作用。(3)苏门白酒草和续断菊均有多种昆虫访花,昆虫的访花活动为这些同花序自交亲和的入侵种提供了受精的机会,可避免缺乏专化传粉者、从而降低交配成功率的缺陷。此外,还对测定花粉活力时应当注意的问题和整个头状花序中雄蕊先熟现象进行了探讨。
本论文还对部分菊科入侵种的种子的扩散特性和萌发特性进行了研究,主要包括测量不同种种子的物理特性和冠毛特性以及在静止空气中的沉降速度和扩散潜力。另外,在室内和野外埋藏下种子萌发情况进行了观察和比较分析,对这些入侵种的萌发能力和幼苗建群特性进行了研究。结果表明,10个被研究的菊科入侵种种子的沉降速度从快到慢依次为：胜红蓟、牛膝菊(Galinsoga parviflora)、一年蓬、紫茎泽兰、钻形紫菀、续断菊、苦苣菜、野塘蒿、小蓬草和苏门白酒草,表明其被动传播的能力由弱变强。沉降速度与其种子的千粒重(带冠毛)和冠毛性状中的任一单独性状都不是简单的一一对应关系,而是各种性状综合作用的结果。相关性分析表明,种子的传播能力与种子的千粒重有明显的正相关(相关系数=+0.4767,p=0.0038),与冠毛长度、冠毛数量和冠毛上的倒钩对数均成显著负相关。光照培养箱中的萌发实验的结果表明,在25℃、12h光照的情况下,除钻形紫菀和一年蓬以外,被研究的其它8个种的终萌发率的超过了50%,而且都显示出早期快速萌发的特点。对10个种的累积萌发趋势进行了曲线回归分析,对应的最佳模型均为三次曲线方程。不同埋藏深度下种子萌发实验的结果表明,8个菊科入侵种的种子在表土的萌发率均最高,随着覆土厚度的增加,种子的萌发率逐渐降低,当覆土厚度超过3cm以上时,所有种子均不能萌发；种子在不同的埋藏深度的萌发情况与种子的大小和萌发能力,较大的或萌发能力强的种子破土能力相对较强。上述种子传播能力和种子萌发特性等的研究结果,可为有的放矢地采取有效措施进行防除和控制其蔓延提供理论依据。
Biological invasion is one of the most important problems that the global natural ecosystem faces. It not only causes the biodiversity decrease, but also harms the global ecological environment and the economic development.
Asteraceae (Compositae) is not only one of the biggest families in angiosperm, but also the most harmful family with greatest number of invasive species in China. Referred to the related documents and wide spot investigation, a list of58Asteraceae invasive species in China was recorded in this dissertation. Among them,12species were listed as the quarantine pests that forbidden import into China in2007.
This dissertation focuses on the study of breeding system of12Asteraceae invasive species in China. On this basis, the detailed study is made on the correlation between reproductive traits of3species and invasiveness, and on the contribution of the characteristics of seeds in10Asteraceae invasive species to invasiveness. The main objective of this reseach is to elucidate the reproductive traits of Asteraceae and provide the useful information on the management of Asteraceae invasive species.
According to Baker's Law, species with uniparental reproduction have a higher chance of establishment after long-distance dispersal than species relying on other mates and pollinators. But there has always been the disagreement on the applicability of Baker's law to invasive plants. We randomly select12Asteraceae invasive species to study their breeding system, which are Ageratina adenophora (Eupatorium adenophorum), Ageratum conyzoides, Aster subulatus, Bidens pilosa, Conyza bonariensis, C. canadensis, C. sumatrensis, Coreopsis lanceolata, Erigeron annuus, Solidago canadensis, Sonchus asper and S. oleraceus. In total, ten species of them (83.3%) set viable seeds in capitula enclosed by paper bags and show the self-compatibility and autonomous self-fertilization. Integrated experiments of the breeding system and pollen-pistil compatibility reveal that two species (Ageratina adenophora and Erigeron annuus) are apomixis (16.7%), three species (Ageratum conyzoides, Conyza canadensis, C. sumatrensis) are autogamy (25%), five species (Aster subulatus, Bidens pilosa, Conyza bonariensis, Sonchus asper and S. oleraceus) are partial autogamy and self-compatibility (41.7%) and other two species (Coreopsis lanceolata and Solidago canadensis) are self-incompatibility (16.7%). Although the latter two species are self-incompatible, they can vegetatively reproduce by robust rhizome.
Our results demostrated that:(1) Baker's law is propitious to explain the invasiveness of all12Asteraceae species studied in terms of vegetative reproduction and not requiring a mate or pollinator for seed production;(2)Ten of the12species of Asteraceae (83.3%) were capable of seed production when pollinators were excluded with paper bags, and their seed-set percentages were more than40%, suggesting that the autonomous seed production through autogamy, autonomous self-fertilization and apomixis play an important role in plant invasiveness;(3) In our study,12invasive species demonstrated the high correlation between breeding system and invasiveness and harmfulness. A few species whose breeding system are apomixis or autogamy are species of severe invasiveness, were listed as noxious or regional noxious weeds in the related literature, such as Ageratina adenophora, Erigeron annuus, Ageratum conyzoides, Conyza canadensis and C. sumatrensis. A few annual species whose breeding system were partial autogamy and self-compatibility are general invasive species or weeds, such as Aster subulatus, Bidens pilosa, Conyza bonariensis, Sonchus asper, S. oleraceus;(4) It was recently reported that autonomous seed production plays an important role in plant invasiveness. Our result combining with survey of published literature about the correlation of life form with autonomous seed production indicated that eleven of the12annual Asteraceae invasive species were hermaphroditic. They are capable of autonomous seed production as a consequence of self-compatibility or apomictic. Among the12perennial species of Asteraceae with successful invasion, nine of them showed self-incompatibility or partial self-incompatibility. Thus, autonomous seed production playing an important role in plant invasiveness may relate more to annual Asteraceae invasive species and less to perennial Asteraceae species.
A total of49Asteraceae species were reported as invasive species in China, but few studies on reproductive biology have been reported. In order to understand the role of sexual reproductive traits in successful invasion in Asteraceae invasive species, we studied the reproductive traits in3Asteraceae invasive species (Conyza sumatrensis, Sonchus asper and Bidens pilosa), in terms of the biology and phenology of capitula and florets, pollen/ovule (P/O) ratio, pollen viability, stigma receptivity, mating system, flower visitors, association of these traits with invasiveness. The result demonstrated that these species are self-compatible and have a mixed mating regime of self-and cross-pollination, both are propitious to invasion. Apomixis was not discovered in decapitated inflorescence. The reproductive traits of autonomous seed production, versatile mating system of self-and cross-pollination and generalized pollination system may contribute to successful invasion. The invasiveness is increased by the high and rapid production, high percentage and rapid germination rate and high dispersal capabilities of achenes (seeds).
Understanding of seed biology of invasive plants will be helpful to predicting the trend of invasiveness and evaluating the strategy on sustainable management. Seed dispersal by wind is the prevailing dispersal pattern in Asteraceae invasive plants. In this study, the dispersal and germination characteristics of the seeds with pappus in10Asteraceae invasive plants are determined. The physical character, settlement velocity in the still air and the dispersal potential of these seeds are compared. Also, the germination of seeds which are buried in soil at different depths, and the character of seedling establishment in these invasive plants were studied. The results showed that, the settlement velocity of seeds of ten Asteraceae invasive species ranks from Ageratum conyzoides, Galinsoga parviflora, Erigeron annuus, Ageratina adenophora, Aster subulatus, Sonchus asper, Sonchus oleraceus, Conyza bonariensis and Conyza canadensis to Conyza sumatrensis. The settlement velocity of the seed in the still air is interactively affected by1000-seed weight and the pappus character. The result of correlation analysis demonstrated that, the dispersal ability of seeds is positively related to1000-seed weight (R=0.4767, P=0.0038), whereas is negatively related to the length and number of pappus, and the number of barbs on pappus. The result of germination in incubator indicated that, except Aster subulatus and Erigeron annuus, the germination of the other8seed species was all above50%at25℃and under12hours' illumination, and the germination was rapid at early stage after sowing. The equations of cumulative germination of the10seed species between the timing of incubation are fitted to cubic curve. The germination of the10seed species was the highest at the surface layer soil, and decreased with the increasing depths of burial. When the depth of burial was above3cm, all seeds could not germinate. The results of dispersal ability and germination character mentioned above may provide theoretical basis for the efficient control of these invasive plants.
菊科是我国种子植物中最大的科之一,同时也是我国入侵种最多,危害最为严重的科。本文经过实地调研和全面查阅文献,共整理出入侵我国的菊科种58种,其中12个种已被列为2007年新公布的中华人民共和国进境植物检疫性有害生物名录。
本文对中国12个菊科入侵种的繁育系统、3个入侵种的有性繁殖特征与入侵性的关系、10个入侵种的瘦果的特性对入侵性的贡献进行了研究。目的是阐明部分菊科入侵种的繁殖特征,为菊科入侵种的管理提供有益的基础信息。
Baker认为,具有单亲繁殖特征的种在长距离扩散后,比那些依赖于双亲繁殖或传粉者的种更容易成功建群,这个论述被称为Baker定律。有关Baker定律在外来入侵种中的适用性一直存在着争议。本文随机选择了12个菊科入侵种,即紫茎泽兰(Ageratina adenophorum或Eupatorium adenophora)、胜红蓟(Agratum conyzoides).钻形紫菀(Aster subulatus)、三叶鬼针草(Bidens pilosa)、野塘蒿(Conyza bonariensis)、小蓬草(C.canadensis)、苏门白酒草(C.sumatrensis)、线叶金鸡菊(Coreopsis lanceolata)、一年蓬(Erigeron annuus)、加拿大一枝黄花(Solidago canadensis)、续断菊(Sonchus asper)和苦苣菜(S.oleraceus),对它们的繁育系统进行了研究。结果表明,其中的10个种(占83.3%)具有自交亲和和自交结实特性。结实率的统计分析和花粉一柱头的亲和性试验的综合分析表明,这12个入侵种中,2个种即紫茎泽兰和一年蓬(占16.7%)为无融合生殖,3个种即胜红蓟、小蓬草和苏门白酒草(占25%)为自体受精,5个种即野塘蒿、钻形紫菀、三叶鬼针草、续断菊和苦苣菜(占41.7%)为部分自体受精,2个多年生菊科入侵种即线叶金鸡菊和加拿大一枝黄花(占16.7%)为自交不亲和种。线叶金鸡菊和加拿大一枝黄花虽然自交不亲和,但它们可通过发达的根状茎进行繁殖。
以上结果表明,(1)Baker定律适用于本研究中的所有12个菊科入侵种,它们或者进行营养繁殖、或者通过形成不需要异株配子或传粉者的种子进行繁殖。(2)12个种中的10个种(占83.3%)在套袋的花序中结出了可育瘦果,并且其结实率在40%以上,表明通过自体受精、自动自交和无融合生殖产生种子的自交结实方式在这些菊科种的成功入侵中起重要作用。(3)对这12个入侵种的繁育系统与其入侵性相关性的分析表明,繁育系统与入侵性的强弱和危害程度方面存在着极高的相关性：繁育系统为无融合生殖或自体受精的几个种在相关文献中大多被列为入侵性强的种,如紫茎泽兰、一年蓬、胜红蓟、小蓬草和苏门白酒草,而繁育系统为部分自体受精或自交不亲和的几个一年生种,如野塘蒿、钻形紫菀、三叶鬼针草、续断菊、苦苣菜被列为入侵性一般的种。(4)对本研究的结果和已发表的资料的分析表明,12个一年生菊科入侵种中,11个具有两性花的种均可自交亲和或无融合生殖,具备自交结实的特性。12个多年生菊科入侵种中,除3个无融合生殖种外,其余的9个种显示出自交不亲和或不完全自交不亲和的特性。因而,自交结实在植物入侵中起重要作用的观点可能更多地与一年生菊科入侵种相关。
为进一步了解菊科入侵种的繁殖生物学特性在其成功入侵中的作用,对具有不同花序类型、开花时期不同、主要依靠种子进行繁殖的苏门白酒草、续断菊和三叶鬼针草的繁殖特性进行了深入研究,内容包括头状花序和小花的生物学特征和开花动态、花粉/胚珠(P/O)比、花粉活力和柱头可授性、交配系统等以及不同交配方式所产生的种子的萌发率,并对这些繁殖特征与入侵性的关系进行了分析。结果表明：(1)3个入侵种均为自交与异交混合型交配系统,具有同花序内自交结实的能力和灵活的交配机制,这些繁育系统的特征有利于其成功入侵。(2)它们所具有的持续结实或大结实量、高结实率、以及种子的高萌发率和扩散能力等特性在与当地种的竞争和促进入侵方面有重要作用。(3)苏门白酒草和续断菊均有多种昆虫访花,昆虫的访花活动为这些同花序自交亲和的入侵种提供了受精的机会,可避免缺乏专化传粉者、从而降低交配成功率的缺陷。此外,还对测定花粉活力时应当注意的问题和整个头状花序中雄蕊先熟现象进行了探讨。
本论文还对部分菊科入侵种的种子的扩散特性和萌发特性进行了研究,主要包括测量不同种种子的物理特性和冠毛特性以及在静止空气中的沉降速度和扩散潜力。另外,在室内和野外埋藏下种子萌发情况进行了观察和比较分析,对这些入侵种的萌发能力和幼苗建群特性进行了研究。结果表明,10个被研究的菊科入侵种种子的沉降速度从快到慢依次为：胜红蓟、牛膝菊(Galinsoga parviflora)、一年蓬、紫茎泽兰、钻形紫菀、续断菊、苦苣菜、野塘蒿、小蓬草和苏门白酒草,表明其被动传播的能力由弱变强。沉降速度与其种子的千粒重(带冠毛)和冠毛性状中的任一单独性状都不是简单的一一对应关系,而是各种性状综合作用的结果。相关性分析表明,种子的传播能力与种子的千粒重有明显的正相关(相关系数=+0.4767,p=0.0038),与冠毛长度、冠毛数量和冠毛上的倒钩对数均成显著负相关。光照培养箱中的萌发实验的结果表明,在25℃、12h光照的情况下,除钻形紫菀和一年蓬以外,被研究的其它8个种的终萌发率的超过了50%,而且都显示出早期快速萌发的特点。对10个种的累积萌发趋势进行了曲线回归分析,对应的最佳模型均为三次曲线方程。不同埋藏深度下种子萌发实验的结果表明,8个菊科入侵种的种子在表土的萌发率均最高,随着覆土厚度的增加,种子的萌发率逐渐降低,当覆土厚度超过3cm以上时,所有种子均不能萌发；种子在不同的埋藏深度的萌发情况与种子的大小和萌发能力,较大的或萌发能力强的种子破土能力相对较强。上述种子传播能力和种子萌发特性等的研究结果,可为有的放矢地采取有效措施进行防除和控制其蔓延提供理论依据。
Biological invasion is one of the most important problems that the global natural ecosystem faces. It not only causes the biodiversity decrease, but also harms the global ecological environment and the economic development.
Asteraceae (Compositae) is not only one of the biggest families in angiosperm, but also the most harmful family with greatest number of invasive species in China. Referred to the related documents and wide spot investigation, a list of58Asteraceae invasive species in China was recorded in this dissertation. Among them,12species were listed as the quarantine pests that forbidden import into China in2007.
This dissertation focuses on the study of breeding system of12Asteraceae invasive species in China. On this basis, the detailed study is made on the correlation between reproductive traits of3species and invasiveness, and on the contribution of the characteristics of seeds in10Asteraceae invasive species to invasiveness. The main objective of this reseach is to elucidate the reproductive traits of Asteraceae and provide the useful information on the management of Asteraceae invasive species.
According to Baker's Law, species with uniparental reproduction have a higher chance of establishment after long-distance dispersal than species relying on other mates and pollinators. But there has always been the disagreement on the applicability of Baker's law to invasive plants. We randomly select12Asteraceae invasive species to study their breeding system, which are Ageratina adenophora (Eupatorium adenophorum), Ageratum conyzoides, Aster subulatus, Bidens pilosa, Conyza bonariensis, C. canadensis, C. sumatrensis, Coreopsis lanceolata, Erigeron annuus, Solidago canadensis, Sonchus asper and S. oleraceus. In total, ten species of them (83.3%) set viable seeds in capitula enclosed by paper bags and show the self-compatibility and autonomous self-fertilization. Integrated experiments of the breeding system and pollen-pistil compatibility reveal that two species (Ageratina adenophora and Erigeron annuus) are apomixis (16.7%), three species (Ageratum conyzoides, Conyza canadensis, C. sumatrensis) are autogamy (25%), five species (Aster subulatus, Bidens pilosa, Conyza bonariensis, Sonchus asper and S. oleraceus) are partial autogamy and self-compatibility (41.7%) and other two species (Coreopsis lanceolata and Solidago canadensis) are self-incompatibility (16.7%). Although the latter two species are self-incompatible, they can vegetatively reproduce by robust rhizome.
Our results demostrated that:(1) Baker's law is propitious to explain the invasiveness of all12Asteraceae species studied in terms of vegetative reproduction and not requiring a mate or pollinator for seed production;(2)Ten of the12species of Asteraceae (83.3%) were capable of seed production when pollinators were excluded with paper bags, and their seed-set percentages were more than40%, suggesting that the autonomous seed production through autogamy, autonomous self-fertilization and apomixis play an important role in plant invasiveness;(3) In our study,12invasive species demonstrated the high correlation between breeding system and invasiveness and harmfulness. A few species whose breeding system are apomixis or autogamy are species of severe invasiveness, were listed as noxious or regional noxious weeds in the related literature, such as Ageratina adenophora, Erigeron annuus, Ageratum conyzoides, Conyza canadensis and C. sumatrensis. A few annual species whose breeding system were partial autogamy and self-compatibility are general invasive species or weeds, such as Aster subulatus, Bidens pilosa, Conyza bonariensis, Sonchus asper, S. oleraceus;(4) It was recently reported that autonomous seed production plays an important role in plant invasiveness. Our result combining with survey of published literature about the correlation of life form with autonomous seed production indicated that eleven of the12annual Asteraceae invasive species were hermaphroditic. They are capable of autonomous seed production as a consequence of self-compatibility or apomictic. Among the12perennial species of Asteraceae with successful invasion, nine of them showed self-incompatibility or partial self-incompatibility. Thus, autonomous seed production playing an important role in plant invasiveness may relate more to annual Asteraceae invasive species and less to perennial Asteraceae species.
A total of49Asteraceae species were reported as invasive species in China, but few studies on reproductive biology have been reported. In order to understand the role of sexual reproductive traits in successful invasion in Asteraceae invasive species, we studied the reproductive traits in3Asteraceae invasive species (Conyza sumatrensis, Sonchus asper and Bidens pilosa), in terms of the biology and phenology of capitula and florets, pollen/ovule (P/O) ratio, pollen viability, stigma receptivity, mating system, flower visitors, association of these traits with invasiveness. The result demonstrated that these species are self-compatible and have a mixed mating regime of self-and cross-pollination, both are propitious to invasion. Apomixis was not discovered in decapitated inflorescence. The reproductive traits of autonomous seed production, versatile mating system of self-and cross-pollination and generalized pollination system may contribute to successful invasion. The invasiveness is increased by the high and rapid production, high percentage and rapid germination rate and high dispersal capabilities of achenes (seeds).
Understanding of seed biology of invasive plants will be helpful to predicting the trend of invasiveness and evaluating the strategy on sustainable management. Seed dispersal by wind is the prevailing dispersal pattern in Asteraceae invasive plants. In this study, the dispersal and germination characteristics of the seeds with pappus in10Asteraceae invasive plants are determined. The physical character, settlement velocity in the still air and the dispersal potential of these seeds are compared. Also, the germination of seeds which are buried in soil at different depths, and the character of seedling establishment in these invasive plants were studied. The results showed that, the settlement velocity of seeds of ten Asteraceae invasive species ranks from Ageratum conyzoides, Galinsoga parviflora, Erigeron annuus, Ageratina adenophora, Aster subulatus, Sonchus asper, Sonchus oleraceus, Conyza bonariensis and Conyza canadensis to Conyza sumatrensis. The settlement velocity of the seed in the still air is interactively affected by1000-seed weight and the pappus character. The result of correlation analysis demonstrated that, the dispersal ability of seeds is positively related to1000-seed weight (R=0.4767, P=0.0038), whereas is negatively related to the length and number of pappus, and the number of barbs on pappus. The result of germination in incubator indicated that, except Aster subulatus and Erigeron annuus, the germination of the other8seed species was all above50%at25℃and under12hours' illumination, and the germination was rapid at early stage after sowing. The equations of cumulative germination of the10seed species between the timing of incubation are fitted to cubic curve. The germination of the10seed species was the highest at the surface layer soil, and decreased with the increasing depths of burial. When the depth of burial was above3cm, all seeds could not germinate. The results of dispersal ability and germination character mentioned above may provide theoretical basis for the efficient control of these invasive plants.
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