寄生植物在入侵植物防治中的作用——以两种菟丝子为例

生物入侵在全世界广泛发生，目前已经受到全球关注。入侵生物对群落生物多样性和生态系统功能造成严重威胁，导致严重的环境问题和惨重的经济损失。薇甘菊(Mikania micrantha)、五爪金龙(Ipomoea cairica)和南美蟛蜞菊(Wedelia trilobata)是我国华南地区危害最严重的三种外来入侵种，其中以薇甘菊危害最严重，是世界十大有害杂草之一。从20世纪80年代发生以来，薇甘菊已在我国广东农林区域造成严重危害。 机械防治、化学防治和传统的生物防治等治理措施，未能有效治理外来入侵种的危害，直到使用本地种菟丝子(Cuscuta spp)的防治策略。与从原产地引进有害生物天敌的传统生物防治方法不同，本地种由于适应当地气候且与其他物种协同进化，因此，对生态环境潜在的负作用小。从2000年，被发现寄生和抑制薇甘菊之后，菟丝子被认为是防治薇甘菊的有效措施。 为了探讨菟丝子寄生对外来入侵种的治理，及对入侵群落的恢复，本研究在内伶仃岛的林地(入侵种群落被引入菟丝子寄生1-4年)，以及东莞、深圳和海丰的干扰样地(入侵种群落被菟丝子自然寄生5年以上)开展野外调查。在每个样地分别设立外来种入侵亚群和菟丝子治理亚群，通过测定群落结构与组成、土壤性质与养分含量，以及外来种和菟丝子的生长与养分含量等参数之后，本研究得出以下主要结论。 (1) 虽然，被寄生的外来入侵种薇甘菊、五爪金龙和南美蟛蜞菊通过调节资源分配以抵御南方菟丝子(Cuscuta australis)的寄生影响，但是，菟丝子寄生导致外来入侵种生物量降低、繁殖能力下降、养分含量降低。虽然，很多寄生植物都是广谱寄生，能同时寄生多种寄主植物，但是，在本研究的被入侵的群落中，菟丝子主要寄生外来入侵植物。尤其是寄生于南美蟛蜞菊和薇甘菊的菟丝子，生长旺盛、繁殖能力强，表现出高度的适应性。因此，菟丝子对外来入侵种(南美蟛蜞菊和薇甘菊)有寄生偏好性，并对本地种的负面影响小。 (2) 通过吸收寄主的养分，田野菟丝子(Cuscuta campestris)有效地抑制了薇甘菊的危害。由于入侵种的凋落物养分含量高且分解效率高，而且，菟丝子能够促进其它凋落物的分解，并使难以被植物吸收的养分转化成易于被吸收利用的状态。因此，菟丝子与薇甘菊的寄生作用导致土壤养分含量的升高。在薇甘菊被抑制之后，本地种利用丰富的土壤养分资源，提高生长适应性，增强抵抗入侵的能力，甚至抵制薇甘菊的再生。 (3) 菟丝子的寄生作用改变了外来寄主与本地非寄主的竞争平衡，促进本地植物的生长与重建。在外来种被抑制之后，本地种的丰度和群落的物种多样性逐渐增加。本地种如：野葛(Pueraria lobata)和芦苇(Phragmites australis)，取代了入侵群落中的入侵种，成了群落的优势种。而其它原先被薇甘菊抑制的本地草本、藤本和灌木，在引入菟丝子防治之后长势较好。群落稳定性与物种多样性密切相关，被治理群落本地种的增加有利于群落的演替与稳定。 (4) 被干扰的生态系统往往更容易被外来种入侵，而外来入侵种又常导致人工干扰生境的严重退化。在人工干扰样地的菟丝子对薇甘菊的抑制效果与在林地的效果一致，导致被寄生的薇甘菊生长衰退、养分竞争能力下降、入侵危害能力降低。而在薇甘菊被菟丝子治理之后，土壤养分资源增加，入侵群落的物种丰度和生物多样性提高。本 地种的重建与本地群落的恢复密切相关，利于本地被治理群落的稳定，促进被干扰植被的恢复。 菟丝子是一种治理外来入侵种危害的有效措施，适用于破碎的生境和被干扰的生态系统，尤其是在采用目前防治措施难以治理的情况下。本研究为本地种防治外来入侵种提供科学依据，且表明以入侵地的本地种治理外来入侵种有可能成为有效且可持续发展的生物防治策略。

东北地区千里光属、华千里光属及狗舌草属分类学研究

本文在前人工作的基础上，对东北产千里光属Senecio植物进行了形态分类、花解剖及花粉形成结构的研究。结果表明：1. 华千里光组Sect. Sinosenecio和狗舌草组sect. Tephroseris在形态和花结构上与千里光属Senecio植物有显著差异，应从千里光属中独立出来，自立为属，即：华千里光属Sinosenecio对狗舌草属Tephroseriso 2. 东北产华千里光属只有一种，即朝鲜千里光Sinosenecio Koreanus (Kom.) B. Nord。3. 东北产狗舌草属分为2组，共5种1变型，其中T. Campestris (Retz.) Reichb.f.spathulatus (Miq.) R. yin et C. y. Li为新组合，并订正了狗舌草T. campestris (Retz.) Reichb的拉丁名。4. 东北产千里光属分为2组，共5种，2变种1变型。5. 狗舌草属和千里光属花粉形态基本相似，都为球形，表面具刺，3孔沟型。花粉形成不能作为划分属的依据。本文还详细描述了每个种的形态学特征，地理分布和文献考证，并附有每个种的叶形。花解剖图及花粉的扫描电镜、光镜照片。还有分属及分种检索表。

岷江干旱河谷三种蔷薇植物繁殖更新研究

繁殖更新是植物生活史的重要阶段，在退化生态系统中，植物繁殖更新能力往往较差，是植被恢复的限制环节，因而也成为恢复研究重点和核心。本研究选择岷江干旱河谷广泛分布的三种蔷薇：多苞蔷薇（R. multibracteata）、黄蔷薇（R. hugonis）和川滇蔷薇（R. soulieana）为研究对象，通过野外调查，在查明其生长、繁殖更新状况的基础上，采用控制和模拟实验，对种子和幼苗阶段进行了深入研究，综合分析更新潜力，并提出相对应的促进更新和植被恢复措施。主要结论如下： 1）三种蔷薇在岷江干旱河谷广泛分布，生长和繁殖状况良好，结实量大。各生长指标：株高、基径和冠幅，繁殖指标：结实数量、重量和单果重量都具有显著的空间差异性。基径对多苞蔷薇结实量影响最大；而冠幅对黄蔷薇结实量影响最大。海拔和纬度是对蔷薇生长和繁殖影响最大的环境因素，随着海拔和纬度的升高，植株生长更高大，结实量增加；坡度和坡向对其生长和繁殖也有一定影响，随着坡度 和坡向增加，蔷薇生长和结实受到抑制。 2）三种蔷薇在岷江干旱河谷更新现状不佳, 但更新潜力大。活力种子比率低，动物取食以及两年生幼苗的大量死亡是蔷薇更新的主要限制因素。多苞蔷薇和黄蔷薇的结实率低，川滇蔷薇较高。三种蔷薇种子产量大，但种子质量较差，更新具有充足的种源。三种蔷薇都能形成持久种子库，种子库中种子总量大，但有效种子少，黄蔷薇被动物啃食的比例很高，多苞蔷薇和川滇蔷薇也有一部分种子受到动物破坏。三种蔷薇幼苗库组成特征表现为，当年生幼苗所占比例很高，年龄较大幼苗所占比例小。 3）三种蔷薇都具有不同程度休眠，未经处理种子的发芽率极低。黄蔷薇休眠程度最深，为深度生理休眠；多苞蔷薇为中度生理休眠；川滇蔷薇为非深度生理休眠。三种蔷薇种子在形态上发育成熟，种皮具有透水性。蔷薇果果肉和瘦果中含有抑制物质，其浸泡液抑制了油菜种子萌发，果肉抑制作用更强，果肉和瘦果浸泡液的抑制程度分别为：川滇蔷薇>黄蔷薇>多苞蔷薇。切割和硫酸腐蚀提高了川滇蔷薇种子的发芽能力，而对多苞蔷薇和黄蔷薇没有影响。完全去除瘦果果皮和种皮提高了多苞蔷薇种子发芽率，但对黄蔷薇没有影响。赤霉素和烟水对蔷薇种子萌发没有促进作用。三种蔷薇打破休眠所需低温层积时间分别为：黄蔷薇>多苞蔷薇>川滇蔷薇。对于多苞蔷薇和川滇蔷薇，层积前对种子进行硫酸腐蚀或暖温层积能缩短低温层积时间，提高发芽率。对于多苞蔷薇，变温层积中暖温层积和低温层积具有一定的负补性，即延长暖温层积可以缩短种子萌发对低温层积的需要。 4）多苞蔷薇种子形态特征和种子休眠与萌发在不同海拔梯度间存在较大差异。种子采集时间、采集季节和干藏影响多苞蔷薇和川滇蔷薇的种子休眠。多苞蔷薇果实大小、种子大小和千粒重、种皮厚度随海拔升高而增加，而种子饱满率和活力随海拔升高而降低，种子休眠程度也随海拔升高而增加。种皮厚度与种子大小、千粒重成正相关关系，硫酸腐蚀后的种子经过不同时间的低温层积后，种子发芽率与种皮厚度、种子大小、千粒重、海拔成正相关关系。2006 年采集川滇蔷薇和多苞蔷薇种子休眠程度较2005 年低。种子休眠随种子年龄增加而减弱。高温和干旱能减轻多苞蔷薇和川滇蔷薇种子休眠。 5）三种蔷薇的生长和生物量积累在干旱胁迫条件下受到抑制，而生物量分配、叶片形态特征和水分利用特征等都发生了变化。三种蔷薇的根、茎、叶各器官生物量以及总生物量等在干旱胁迫下明显减小，叶片脱落数量增加。在干旱胁迫条件下，较多的生物量分配到地下部分，从而这使R/S 明显增加。比叶面积（SLA）和冠层面积比（LAR）对干旱胁迫的反应不敏感，仅有部分物种在干旱胁迫条件下发生了变化，并且其变化特点在不同年龄幼苗之间有一定差异。干旱胁迫对WUE 的影响在不同物种间存在差异。多苞蔷薇和黄蔷薇的WUE 随着干旱胁迫的增加而增大， 而川滇蔷薇的WUE 则随干旱胁迫增加而减小。在干旱胁迫条件下，多苞蔷薇和黄蔷薇叶片脱落量和生物量减小幅度较川滇蔷薇大，表明其抗旱能力较强。在干旱胁迫条件，三种蔷薇两年生幼苗的生物量减小幅度较当年生幼苗小，表明两年生幼苗的抗旱能力更强。 6）两种植被恢复措施中，幼苗移栽比播种具有更好的植被恢复效果。播种后，蔷薇种子的发芽率较高，但出苗率都很低，即使出苗，幼苗也几乎在一月内全部死亡。 三种微生境条件下（灌木、半灌木和裸地），种子出苗和幼苗成活没有差异。移栽幼苗总体死亡率都比较低，小于20％。特别是两年生幼苗死亡率更低，小于2％。移栽后的幼苗生长状况良好，在整个生长季中，各生长指标不断增加。生境对幼苗的存活率没有显著影响，但对于幼苗的生长和生物量积累有一定影响，裸地更有利于幼苗生长和生物量积累。与当年生幼苗相比，两年生幼苗具有更高的成活率。总之，三种蔷薇在干旱河谷分布广泛、生长繁殖状况良好，结实量大，具有丰富种源，繁殖更新潜力大，但繁殖更新状况不佳；种子散布后动物对种子的取食、种子的深度休眠过程、种子出苗以及当年生幼苗的存活和定居是更新的主要限制环节。水分是影响结实、种子休眠解除和萌发，幼苗存活和定居的最主要的限制因素。在植被恢复中，应在种子成熟季节大量采集种子，在室内打破休眠后进行人工播种，培育两年生幼苗，通过幼苗移栽方式进行植被恢复。川滇蔷薇应栽种在相对湿润的过渡区，而多苞蔷薇和黄蔷薇可以应用于核心区植被恢复。 Regeneration is an important phase in plant life cycle. It has been a key component of ecological restoration in degradation ecosystem in which plants commonly has poor regeneration. In this paper, we investigated the natural growth, propagation and regeneration status of native three rose species, Rosa multibracteata, R. hugonis and R. soulieana, and analysis the limitation in seed germination and seedling establishment stages. Advice on facilitating the use of these plants in restoration based on the results has been proposed. The results were as follows: 1) Three rose plants widely distributed in the dry valley of the Minjiang River, and made a good performance in growth and propagation. There were significant spatial differences in each growth parameter, such as ramet height, basal diameter, crown diameter and propagation parameters including hip number of a clump, hip mass of a clump and a hip mass. Basa diameter was the most important growth parameter influencing fruit number for R. multibracteata and crown diameter was for R. hugoni. Altitude and latitude had the greatest effect on the growth and propagation of rose plants among environmental conditions. Each parameter of growth and propagation increased with the increase of altitude and latitude. In addition, the increase of slope and aspect limited the growth and propagation. 2) Three rose plants had poor natural regeneration, but great regeneration potential. Low seed viability, predation and higher mortality of current year old seedlings were the limitation in regeneration. R. multibracteata and R. hugonis had higher fruiting rates than R. souliean. All three plants produced a great number of seeds, while their viability was poor. Three rose plants had persistent seed banks, with high total seed number but very low viable seed density. Predation was most severe in R. hugonis, and it also existed to some degree in R. multibracteata and R. soulieana. The seedling age-structure was characteristic of current-year seedlings predominating and few older seedlings were observed. 3) Three rose seeds were dormant and untreated seeds germinated with very low germination percentages. The rose seeds had morphological mature embryos, and achenes were permeable. Some inhabit substances existed in hips and achenes for the extracts of hips and achenes inhibited germination of Brassica campestris. The inhibition effect of the extracts of three rose hip and achenes was R. soulieana>R. hugonis>R. multibracteata. Mechanical and H2SO4 scarification increased R. soulieana germination but had no effect on germination of R. hugonis and R. multibracteata seeds. Full removal of pericarp and testa improved the germination of R. multibracteata but did not affect R. hugonis germination. GA3 and smoke water had no positive effect on rose seed germination. The periods of cold stratification required to released seed dormancy was R. hugonis > R. soulieana >R. multibracteata. H2SO4 scarification and warm stratification shortened cold stratification to release dormancy for R. soulieana and R. multibracteata. Warm stratification had complementary effect for cold stratification, i.e. the longer warm stratification seeds received, the shorter cold stratification were required to obtain the same germination percentage. Three rose seeds had different kinds of dormancy; R. hugonis has deep physiological dormancy, R. multibracteata with intermediate physiological dormancy and R. souliean non-deep physiological dormancy. 4）The seeds traits and dormancy of R. multibracteata showed significant difference across altitudes. Year and season of seed collection had significant effect on seed dormancy for both R. souliean and R. multibracteata. Hip size, seed size, seed weight, seed coat thickness and seed dormancy level increased with the increase of the altitude. There were positive relations between seed coat thickness with seed size and seed weight. Germination percentage of seeds treated with H2SO4 scarification following different periods of cold stratification showed positive relation with seed coat thickness, seed size, seed weight and altitude. Seeds of R. souliean and R. multibracteata collected in 2006 had low dormancy level than those collected in 2005. Seed dormancy decreased with increasing seeds age. High temperature and drought were associated with low dormancy level. 5) Seedling growth, the total dry mass and their components of seedlings were reduced, while leaf senescence accelerated under drought stress. More biomass allocation to root system resulted in higher R/S ratio under drought. Water-use efficiency (WUE) of R. multibracteata and R. hugonis increased, while it declined for R. soulieana under drought stress. R. soulieana seedlings had poor drought-resistance capacity it had more senescent leaves, and its reduction of biomass was stronger than two other rose plants under drought. The reduction degree of one year old seedlings under drought stress was slighter than that of current year seedlings. Therefore, one year old seedling was more drought-resistent compared to current year seedlings. 6）Planting seedlings may have better effect in comparison with direct seeding. Most seeds germinated after seeding, but seedling emergence was very low. More than 80 % seedlings from direct seeding died within a months after emergence. Seedling emergence and survival rate did not show difference among microhabitats. Mortality rates of seedlings artificially planted in microhabitats were general lower than 20 %, and the mortality rate of one year old seedlings was lower than 2 %. Each grow parameter including plant height, leaf number and branch number continually increased after planting. Microhabitat type had effect on the growth parameter and biomass production, but it did not influence the seedling survival. Bare land tended to facilitate seedling growth. One year old seedlings had higher survival rate than current year seedlings. In conculsion, the three rose had wide distribution in the dry valley of the Minjiang River. They produced many seeds and had tolerance to drought stress to some degree. But they had poor regeneration in habitats may be caused by predation, seed dormancy，and high mortality in current year seedlings. We recommend that rose plants should be utilized in restoration by planting two-year old seedlings in spring. A large quantity of seeds should be collected artificially in autumn, release seed dormancy in room, and then cultivate two-year old seedlings by seeding in particular container. R. soulieana seedling probably be planted in transition area, and R. multibracteata and R. hugonis can be used in core area of the dry valley of the Minjiang River.