淡水湿地种子库的研究方法、内容与展望

种子库是指存在于土壤表面和土壤中全部存活种子的总和.种子库的研究是深入探讨湿地植被结构、功能与动态等方面的重要基础,同时也是对湿地生物多样性研究的一个重要补充,对于了解植被演替动态、植被更新和受损湿地植被重建与恢复具有指导意义.本文结合国内外淡水湿地种子库研究现状,对湿地种子库的形成、研究方法以及湿地种子库时空格局、种子库与地表植被的关系、水文变化对湿地种子库的影响、湿地种子库在湿地恢复中的应用等主要研究内容进行了系统的总结;同时对湿地种子库的时间动态、种子库的分类、种子库与地表植被的关系、湿地恢复重建及湿地景观营造等研究进行了展望.     

湿地土壤种子库与地上植被相似性关系研究评述

土壤种子库与地上植被的关系是土壤种子库研究的重要组成部分。当前,湿地生态系统面临严重威胁,研究湿地土壤种子库和地上植被关系既可以加强对土壤种子库和植物群落特征的认识,又可以为湿地保护与管理提供理论指导。检索了科学引文索引扩展版(SCIE)数据库中收录的1900—2012年间研究湿地土壤种子库与地上植被关系的文献,通过分析土壤种子库与地上植被的Srensen相似性系数,结果发现:不同湿地类型的土壤种子库和地上植被的相似性存在显著差异,河流湿地中两者的相似性最小;不同植被类型中土壤种子库与地上植被的相似性差异显著:草本群落的相似性大于乔木群落;不同气候带的湿地中两者的相似性也存在显著差异,其中亚热带地区相似性最小。总结了湿地种子库与地上植被相似性关系的时空变化特征。二者的相似性通常随着植物群落的演替而减小,在空间上也随着环境梯度而变化。分析了两者关系的影响因素,如种子传播、环境条件和繁殖策略等。对研究中存在的问题及发展方向提出建议。     

农田开垦对三江平原湿地土壤种子库影响及湿地恢复潜力

种子库是湿地植被恢复的重要途径之一,不同时期的耕作土壤中残留的种子对开垦湿地恢复具有重要的作用.本文采用温室萌发法在两种水分条件下对三江平原天然湿地、不同开垦年限湿地种子库结构和规模进行了研究,以了解不同开垦年限湿地种子库特征及其在湿地植被恢复中的潜力.本次实验共萌发物种50种,随着开垦年限增加,萌发物种逐渐减少,天然湿地、开垦1年、3年、10年、20年的湿地分别为34种、31种、21种、21种和8种,萌发物种数与种子库规模均表现出极显著差异(F1=8.32,F2=5.946,P＜0.001).种子库密度以天然湿地和开垦1年湿地最大,分别为7624粒/m2,9836粒/m2.随着开垦年限增加,种子库规模逐渐减小,开垦3年、10年种子库密度为4336粒/m2,4872粒/m2.开垦20 a后,显著减少为432粒/m2.湿润条件下萌发物种数及种子密度显著高于淹水处理,种子库具有明显的分层现象,0-5 cm土层种子库规模显著高于5-10 cm.小叶章(Calamagrostis angustifolia)作为该地区优势物种,由最初的1192粒/m2,经过20 a开垦后在种子库中消失.研究表明,在一定的开垦年限范围内,开垦湿地土壤中仍然保留大量的湿地物种种子,在湿地恢复中具有重要的作用.     

三江平原恢复湿地土壤种子库特征及其与植被的关系

通过幼苗萌发法和样方调查相结合的方法对三江平原不同演替恢复阶段的种子库特征及其与植被的关系进行了研究。将开垦湿地、不同演替恢复阶段湿地以及天然湿地不同土壤层次(0-5、5-10 cm和根茎)的种子库在两种水分条件下(湿润、淹水10 cm)进行萌发处理。结果表明: 随着演替恢复阶段的进行, 种子库的结构和规模逐渐扩大, 地表群落表现出由旱生物种占优势的群落逐渐演变成以小叶章(Calamagrostis angustifolia)占优势的湿生群落的演替趋势。恢复7年湿地、恢复14年湿地、天然湿地土壤种子库萌发物种数分别为24种、29种、39种, 植被物种数为21种、25种、14种。湿地类型、水分条件和土壤层次均显著影响种子库萌发的物种数及幼苗数(p < 0.01)。种子库具有明显的分层现象, 天然湿地0-5 cm土层种子库种子萌发密度是5-10 cm土层的4倍左右, 而恢复湿地仅1.3倍左右, 且土层间萌发物种相似性系数较低。湿润条件下的萌发物种数显著高于淹水条件, 且两种水分条件下萌发物种的生活型不同。由于恢复时间较短, 不同演替恢复阶段的种子库与植被相似性维持在30%以下。湿地中根茎分蘖出大量的湿地物种, 对于小叶章等优势物种的繁殖具有重要作用。研究表明, 在开垦湿地退耕后的次生演替阶段, 种子库能够保持大量的湿地物种, 通过对湿地种子库与植被的关系研究, 能够为三江平原湿地群落演替与湿地恢复提供策略指导。     

皖江湿地及其开垦为稻田后土壤种子库结构比较

运用镜检法对皖江2处天然湿地(十八索和升金湖)及其开垦的稻田表层土壤(0～10cm)种子库物种进行了鉴定,分析天然湿地开垦为稻田后土壤种子库群落结构及多样性的变化.结果表明:天然湿地种子库密度为83499～109141粒·m2,而开垦为稻田30～50年后种子库密度为9140～47452粒·m2;天然湿地的土壤种子库物种数为16～30种,以蓼科-莎草科或莎草科-蓼科为优势种,开垦为稻田后物种数下降为7～16种,且趋向于以禾本科为单一优势种;此外,与十八索湿地相比,升金湖湿地有较高的土壤种子库物种数、丰富度和多样性,而开垦年限为30年的十八索稻田土壤种子库物种数和丰富度显著高于开垦年限为50年的升金湖稻田.湿地开垦为稻田后,土壤种子库密度和物种数的急剧减少,人为管理生态系统生物多样性受到强烈干扰.因此,湿地保护对于维持长江流域的生物多样性及其流域生态功能具有重要意义.     

莫莫格扁秆藨草恢复湿地土壤种子库对长芒稗入侵的响应

湿地恢复过程中,时常有外来种或本地杂草入侵。土壤种子库作为未来植被的潜在种源,对湿地恢复效果具有重要的指示意义。在莫莫格国家级自然保护区,以恢复白鹤栖息地(扁秆藨草(Scirpus planiculmis)沼泽)为目的,进行了退化湿地的水文恢复;但退化湿地恢复2a后,一年生杂草长芒稗(Echinochloa caudata)在大部分区域成为建群种。以长芒稗入侵湿地和扁秆藨草自然湿地为研究对象,对比分析了长芒稗和扁秆藨草的土壤种子库及生长结实特征。结果表明,在自然湿地扁秆藨草种子库规模是长芒稗的18.42倍,而在恢复湿地长芒稗种子库大小是扁秆藨草的5.04倍。与自然湿地相比,扁秆藨草种子库密度在入侵湿地明显减少,但仍保留了一定量具有活力的种子(664.32±105.98)粒/m~2,这与研究区扁秆藨草较高的种子生产力(9210.4±1513.4)粒/m~2及种子较强的浮力(FP50=39.7d)有关,说明扁秆藨草具备通过种子库或水传播恢复的潜力。长芒稗土壤种子库密度在入侵湿地高达(3345.9±520.3)粒/m~2,明显高于自然湿地种子库规模(P0.01),说明恢复湿地受长芒稗入侵影响严重,这与长芒稗较高的种子生产力(7621.4±376.25)粒/m~2及较弱的种子浮力(FP0=5d)有关,同时也表明长芒稗通过水传播扩散的能力较弱。另外,研究区长芒稗平均高度超过1m,且盖度较大,不仅阻碍扁秆藨草种子的水播,也降低了到达地表的光照水平,从而抑制扁秆藨草更新。因此,在莫莫格受长芒稗入侵湿地,于开花结实前收获长芒稗地上植物体及凋落物应是限制长芒稗扩展、同时促进扁秆藨草恢复的有效措施。     

三江平原沟渠土壤种子库特征及其与地上植被的关系

为了探究三江平原沟渠土壤种子库在湿地植物保护中的作用及其在湿地恢复中的潜力,该文采用幼苗萌发法与样方调查相结合的方法,对三江平原不同开挖年限沟渠的土壤种子库特征及其与地上植被的关系进行了研究。结果表明:沟渠具有较大规模的土壤种子库,边坡种子库显著大于底泥种子库,边坡种子库密度为8 973–25 000 seeds·m–2,底泥种子库密度为506–1 488 seeds·m–2。开挖10年、20年和30年的沟渠土壤种子库共有50种植物萌发,隶属于20科41属。开挖10年、20年和30年的沟渠土壤种子库萌发物种数分别为37种、34种和33种,地上植被物种数分别为25种、33种和22种。土壤种子库和相应地上植被的相似性系数分别为38.7%、35.8%和32.7%。随着植物群落演替的进行,地上植被的Simpson指数、Shannon-Wiener多样性指数和Pielou均匀度指数均逐渐增大。三江平原沟渠土壤种子库和地上植被中保存了大量湿地植物,表明沟渠具有保护植物物种多样性的作用,沟渠土壤种子库具有湿地恢复的潜力。随着沟渠开挖年限增加,沟渠植物群落呈现退化特征,建议对沟渠系统加强管理。     

尕海湖滨湿地种子库初探

在青藏高原湖泊尕海湖滨湿地的高寒草甸中,选取4个样地22个样点研究种子库基本组成及其与地表植被的关系。结果表明,尕海湖滨湿地种子库由13科34属37种湿生植物组成,平均为7400 seeds/m2,种类以一年生草本植物为主;地表植被由17科29属31种湿生植物构成,以多年生草本植物为主,种子库与地表植被组成的差异较为明显,符合高寒草甸的特征。总体而言,种子库垂直分布的各深度间种子密度有明显的相关性,土壤表层的种子库密度显著高于深层,在物种水平上,仅露蕊乌头Aconitum gymnandrum有这一趋势。种子库物种与水位变化研究表明,种子库中物种的生活型随着水位变化而发生变化：随着水位的升高,种子库中的优势种逐渐由以露蕊乌头为指示种的中生植物向以水毛茛Batrachium bungei为指示种的沉水植物过渡。本研究表明,尕海湖滨湿地种子库具有对受损湿地进行修复的潜在可能,同时水位对其分布有一定影响;如果是针对目标植物的选择性修复则要注意该物种是否存在于种子库中。     

黄河三角洲滨海湿地地下水位变化对土壤种子库特征的影响

地下水位是影响湿地生态系统结构和功能的重要环境因子之一，然而其如何影响滨海湿地土壤种子库特征尚不清楚。通过地下水位控制试验，研究黄河三角洲滨海非潮汐湿地地下水位梯度（-20、-60、-100 cm）下，土壤种子库密度、物种多样性等特征，并通过土壤化学指标与对应的土壤种子库特征之间的通径分析，探讨地下水位深度变化对土壤种子库特征的影响机理。研究发现，在该区域秋季采样的土壤种子库中，碱蓬和盐地碱蓬的种子数量最多；地下水位变化对土壤种子库密度无显著影响，但对土壤种子库的物种组成影响较大，随着地下水位深度增加，多年生草本植物种类呈增加趋势。整体上，随着地下水位深度增加，土壤种子库的Margalef丰富度指数和Shannon-Wiener多样性指数随之增大。主成分分析表明，土壤盐是影响黄河三角洲滨海湿地土壤种子库组成特征的主导因素，贡献率为69.482%，土壤养分为次要影响因素，贡献率为23.710%，进一步通径分析表明，土壤水溶性盐总量对种子库物种多样性起到限制作用，同时，有效磷浓度通过水溶性盐总量对Margalef指数和Shannon-Wiener指数表现为间接负作用，总磷浓度对种子库与地上植被相似性起到促进作用。研究表明，黄河三角洲滨海湿地地下水位深度变化，主要通过影响土壤水溶性盐总量，进而影响土壤种子库物种多样性。     

天津滨海典型盐碱湿地土壤种子库特征及CCA分析

选取天津滨海地区典型盐碱湿地,进行土壤种子库采样及萌发试验,同时测定土壤理化性质。采用典范对应分析法(CCA)对土壤种子库和土壤特征因子的关系进行研究。结果表明:4个样地土壤种子库储量较大,但物种组成单一,为滨海地区指示性盐生植物。与人工湿地相比,自然保护湿地具有物种丰富度高,群落结构稳定的优点,但受到干扰最小,种子库密度却最低。种子库种子萌发率曲线呈不规则的抛物形状,萌发数量曲线呈"S"型分布,土壤板结程度对萌发率有较大影响。土壤种子库垂直分层结构明显,随土壤深度加深,种子库密度降低,10～15 cm采样层仍保留大量活性种子。通过CCA分析,发现土壤种子库呈现明显的3个聚类;且土壤种子库受全盐量、有机质、pH影响较大,但土壤通透性这一因素也不容忽视。     

Rapid Seed Bank Development in Restored Tidal Freshwater Wetlands

Studies of seed bank development have rarely been included in evaluations of wetland restoration. We compared the seed bank of a recently restored tidal freshwater marsh in Washington, D.C., Kingman Marsh, with seed banks of another restored site (Kenilworth Marsh) and two reference marshes (Dueling Creek and Patuxent Marsh). The density and richness of emerging seedlings from Kingman Marsh seed bank samples increased from less than 4 seedlings and 2 taxa/90-cm² sample in 2000 (the year of restoration) to more than 130 seedlings and 10 taxa/90-cm² sample in 2003. The most important seed bank taxa at Kingman Marsh included Cyperus spp., Juncus spp., Lindernia dubia , Ludwigia palustris , and the non-native Lythrum salicaria . These taxa are not abundant in most mid-Atlantic tidal freshwater marshes but are almost identical to those described for a created tidal freshwater wetland in New Jersey. Seed banks of both the restored sites contained few seeds of several important species found at the reference sites. Flooding had a significant negative effect on emerging seedling density and taxa density, suggesting that slight decreases in soil elevation in restored wetlands will dramatically decrease recruitment from the seed bank. Because seed banks integrate processes affecting growth and reproduction of standing vegetation, we suggest that seed banks are a useful metric of wetland restoration success and urge that seed bank studies be incorporated into monitoring programs for restored wetlands.     

The potential of soil seed banks of a eucalypt wetland forest to aid restoration

Soil seed banks can play an important role in the regeneration of wetland vegetation. However, their potential role in the restoration of degraded wetland forests is less certain. I surveyed the soil seed bank and extant floras of four sites across a eucalypt wetland forest of variable vegetation condition. At each site, the extant vegetation was surveyed within two 5 × 5 m² quadrats, each from which five composite soil seed bank samples were collected. Across the four sites, 57 (including 18 exotic) species were identified in the extant vegetation, while from the seed bank samples 6379 seedlings emerged from 80 taxa, 33 of which were exotic species. The soil seed bank was dominated by native and exotic monocots, and contained very few seeds of wetland tree or shrub species. Overall, the similarity between the extant and seed bank floras was very low (~24 %). Soil seed banks are likely to be of limited use in the restoration of degraded wetland forests, because the dominant species in such systems—woody and clonal plants—are typically absent from the soil seed bank. Wetland soil seed banks may contribute to the maintenance and diversity of understorey vegetation, however, they may also act as a source of exotic plant invasions, particularly when a wetland is degraded.     

Seed Ecology of the Invasive Tropical Tree <Emphasis Type="Italic">Parkinsonia aculeata</Emphasis>

Parkinsonia aculeata is an invasive tree native to tropical America, but introduced to Australia. Propagation and stand regeneration is mainly by seed. To gain baseline knowledge for management decisions, seed bank dynamics were monitored for two months during the fruit dispersal period at a coastal wetland in Costa Rica (native habitat), and at a coastal wetland and two semi-arid rangeland sites in Northern Queensland, Australia (introduced habitats). Seed bank densities underneath dense, uniform Parkinsonia stands were found to be lowest in the Australian wetland but highest in the Costa Rican wetland. Post-dispersal seed losses were highest in the Australian wetland, primarily due to seed germination and/or death. At the other sites, seed losses were minor during the study period, and predation was the most important cause of losses. At the two rangeland sites bruchid beetles accounted for more than 95% of the seed losses by predation. Total predation was lowest in the Costa Rican wetland. In order to test for intrinsic differences of seed characteristics, germination trials were conducted using both canopy seeds and seeds from the soil seed bank. Dormancy release and germination rate were studied under four temperature treatments. In all populations, dormancy release increased with increasing temperature, but averaged responses were significantly different between Costa Rican and Australian seed populations, and between seeds collected from the soil and from trees. Germination rate of scarified seeds was fastest at 35 °C in all tested seed populations. While high seed germination levels seem to explain low seed bank densities in the Australian wetland, the large seed banks at the rangeland sites reflect the lower incidence of favourable conditions for germination. In the Australian wetland biocontrol with bruchids is unlikely to be successful, while control by conventional methods, such as killing stands by basal bark spraying, seems feasible, due to a lower long-term risk of re-infestation from the soil seed bank. At the rangeland sites conventional control will be difficult and costly. Parkinsonia stands may be better left to their own, while bruchid populations are monitored and management efforts are concentrated on preventing further invasion.     

Restoration and creation of freshwater wetlands using seed banks

The minimum information about a seed bank needed for a wetland restoration or creation project is a species list. There are two basic techniques for determining the composition of seed banks: (1) mechanical separation of seeds from a volume of soil and (2) germination of seeds from a volume of soil under appropriate environmental conditions. The latter method always gives biased results. It is best to collect as many random samples as possible when sampling a wetland seed bank. These can be combined as needed for processing. Field studies in India have demonstrated that vestigial seed banks can be used to re-establish a former vegetation type in a monsoonal wet-land that had become overgrown by a species of grass. In less than a year, 9 of 1 I species in the vestigial seed bank were found growing in areas cleared of the grass. Vestigial seed banks of drained prairie wetlands in the northcentral United States contained a few wetland species after 70 years, although species diversity and seed density declined significantly after 20 to 30 years of drainage and cultivation. In Florida, U.S.A., wetlands have been established in strip-mined areas using donor soils from existing wetlands. Newly established wetlands quickly developed a dense cover of vegetation, although this vegetation often lacked many desirable wetland species. Experimental studies of soil moisture conditions using a seed bank from the Delta Marsh, Canada, demonstrated that soil moisture affected both the total number of seeds, and the relative proportion of seeds of each species that germinated from a seed bank. The density of seedlings of emergent wetland species in the treatments was directly proportional to soil moisture, while that of terrestrial annuals was inversely proportional. Emergent species made up nearly 90% of the seedlings in the wettest treatment and 0% in the driest.From a paper presented at the Third International Wetlands Conference, 19–23 September, 1988, University of Rennes, France.     

Persistence of seed banks in Australian temporary wetlands

1. The ability of seeds to survive periods of drying and wetting that do not lead to seed production will determine the potential species pool for future plant communities of temporary wetlands. I investigated characteristics of the seed banks in sediment from Australian temporary wetlands that might contribute to the ability of aquatic plants to re‐establish after extended drought. 2. Experimental investigation into germination from sediment from six sites in five Australian temporary wetlands, with various water regimes, examined two sources of seed bank depletion: (i) length of time dry (longevity up to 12 years) and (ii) successive annual wetting and germination events (up to seven) with intervening periods dry (leaving a residual seed bank), both without any opportunity for replenishment of the seed bank. 3. These wetlands had species‐rich, long‐lived seed banks that were not exhausted by successive germination events. After three years of dry storage, 90% of the original seed bank species germinated, after six years 75% and after 12 years 20%. After seven successive wetting and drying events without seed bank replenishment, 48% of the original species still germinated. The mean survival time dry for seed bank species, 7.4 years, was longer than the duration of recent droughts. 4. Seed bank composition varied among wetlands and over time; most species did not occur in all wetlands and many occurred in one wetland only. The germination patterns of different species, although differing in detail, tended to be consistent in that all species could survive long dry periods and several wetting and drying events. However, experimental drought significantly diminished species richness and abundance, indicating limits to seed bank persistence. 5. Data from such long‐term studies of seed bank persistence should allow prediction of the species richness and composition of the germinating communities in a wetland whose water regime is intentionally or unintentionally altered. This ability to forecast may become particularly important under climate change and the need to predict future wetland conditions.     

Effects of Water Level on Three Wetlands Soil Seed Banks on the Tibetan Plateau

Background

Although the effect of water level on germination in soil seed banks has been documented in many ecosystems, the mechanism is not fully understood, and to date no empirical studies on this subject exist. Further, no work has been done on the effect of water level on seed banks of drying and saline-alkaline wetlands in alpine areas on the Tibetan Plateau.

Methodology

We examined the effects of water level (0 cm, 5 cm and 10 cm) on seed germination and seedling establishment from soil seed banks at 0–5 cm and 5–10 cm depths in typical, drying, and saline-alkaline wetlands. We also explore the potential role of soil seed bank in restoration of drying and saline-alkaline wetlands.

Principal Findings

Species richness decreased with increase in water level, but there almost no change in seed density. A huge difference exists in species composition of the seed bank among different water levels in all three wetlands, especially between 0 cm and 5 cm and 0 cm and 10 cm. Similarity of species composition between seed bank and plant community was higher in 0 cm water level in drying wetland than in the other two wetlands. The similarity was much higher in 0 cm water level than in 5 cm and 10 cm water levels in all three wetlands. Species composition of the alpine wetland plant community changed significantly after drying and salinization, however, species composition of the seed bank was unchanged regardless of the environment change.

Conclusions/Significance

Water level greatly affects seed bank recruitment and plant community establishment. Further, different water levels in restored habitats are likely to determine its species composition of the plant community. The seed bank is important in restoration of degraded wetlands. Successful restoration of drying and salinization wetlands could depend on the seed bank.     

洪湖湿地退耕初期种子库的季节动态

 为了研究水田退耕初期种子库的特征,在两个不同水位条件下通过幼苗萌发法对洪湖的濒湖退耕田种子库的结构和季节动态进行了研究。结果表明,种子库季节变化明显,种子库的密度于12月达到最大值2.61×105 ind.·m-2,6月时最小,为2.90×104ind.·m-2。而萌发的物种数则相反,6月时最大,12月时最小。Simpson 和Shannon-Wiener多样性指数及Shannon均匀度指数表现出较为一致的季节变化规律,在6月最高,12月和3月最低。该种子库类型属于持续性种子库。种子库与地表植被群落种相似性系数变化较大。0～5 cm和 5～10 cm土层中的种子库萌发的物种数差异不显著,但在数量上有明显分层现象,上层种子库占总数的78%。One-way ANOVA分析表明,萌发时的水位条件可显著影响种子库萌发的种类和数量。结合种子库研究的结果,讨论了种子库在退田还湖后湿地植被恢复过程中的重要性。     

Plant Establishment from the Seed Bank of a Degraded Floodplain Wetland: A Comparison of Two Alternative Management Scenarios

There has been little research examining the soil seed banks of degraded floodplain wetlands and their contribution to wetland rehabilitation in Australia. Our aim was to assess the establishment of plants from the seed bank that may occur following the delivery of an environmental water allocation to Kanyapella Basin, a 2950 ha wetland located on the floodplain of the Goulburn and Murray Rivers in northern Victoria, Australia. Two hypothetical water regimes were investigated (flooded and dry) in a glasshouse experiment, where plants were left to establish from the seed bank over a period of 124 days. Differences in the establishment of plants from the seed bank indicated that the return of a flooding regime is likely to have a significant effect on the composition of the wetland vegetation. Mapping of the distribution of plant species indicated that propagules were highly dispersed across the wetland for the majority of taxa, in contrast to the localised distribution of many of the plant species represented in the extant vegetation. Inundation favoured the establishment of native wetland and floodplain plants, although many areas of Kanyapella Basin that are currently ‘weed-free’ have the potential to become colonised and potentially dominated by introduced plants if the wetland is not managed appropriately. Overall, results supported the aim of management to reestablish a wetting and drying regime through use of an environmental water allocation. This study presents a significant example of the application of seed bank investigations in wetland ecology and management.