Seed banks and seed population dynamics of halophytes

In this review I will describe the importance of seed banks and thepopulation dynamics of seeds on the distribution of species in salinehabitats. The main questions being examined in this review include: 1.Does the seed bank represent the flora of the entire salinity gradient or isit restricted to the species in each zonal community? 2. Is the size andspecies composition of the persistent seed bank regulated by the degree ofsalt stress in habitats along an environmental gradient? 3. Does thepopulation dynamics of seeds influence the temporal and spatial distributionof plant species in saline habitats? Seed banks may be transient orpersistent depending upon the physiological responses of species and thesoil environment in which the seeds are found. The formation of zonalcommunities in salt marsh environments is affected by changes in soilsalinity and flooding along an elevational gradient. Population dynamics ofseeds have been found to determine the spatial and temporal distributionof species along salinity gradients. The flora and relative density of speciesof zonal communities are significantly dependent upon the stress toleranceof species at different stages of development and the presence of transientor persistent seed banks. The occurrence of a seed bank is related to thesalinity tolerance of species at the germination stage of development, aseeds ability to tolerate hypersaline conditions and flooding, and whetheror not species are able to maintain a persistent seed bank until hypersalineconditions are alleviated.

     

淡水湿地种子库的小尺度空间格局

以长江中下游淡水湿地湖里沼泽为对象,研究了种子库在小尺度空间范围的水平分布格局。在沼泽中选择地表植被分布比较均一的16m×4m大小的方形区域,按1m间隔获取64个内径7·8cm、深5cm的圆柱状土样,通过幼苗萌发法鉴定每个土样的种子库组成。采用2个格局指数(离散系数和Lloyd平均拥挤指数)以及Moran空间自相关系数分析了种子库中优势种的空间分布格局。结果显示种子库由17个物种组成,多年生的锐棱荸荠(Eleocharisacutangula)和龙师草(E·tetraqueter)是种子库中密度最大的物种。两个格局指数显示7个分布频率大于10%的物种的种子全部为聚集分布。MoransI统计分析显示其中只有3个物种为显著的正的空间自相关,表明这3个物种的种子斑块大小超过了目前的两个取样单位之间的距离(1m),而其它4个物种的种子斑块则小于这个尺度。     

Transferring sediment containing intact seed banks: a method for studying plant community ecology

The experimental study of the ecology of natural plant communities is necessary to demonstrate the effects of environmental factors on plant growth. It allows one to make predictions of the likely effects of environmental changes on plant communities. Manipulating an entire ecosystem is complicated, and is rarely attempted. An alternative method is applied here for the submerged plant communities of the seasonally-flooded oligohaline marshes of the Camargue. It consists of collecting undisturbed sediment samples with their contained intact seed bank, and submitting them to different salinities. The study of total biomass and of the biomass of the more frequent species was carried out using parametric tests. The precision of the results depended on the frequency of the species in the replicates. When the frequency was low, non-parametric statistics were necessary. The method seems to be particularly suitable for the study of communities of annual species in aquatic environments.

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Résumé L'étude expérimentale de l'écologie des communautés naturelles de plantes est nécessaire pour démontrer les relations de cause à effet entre les facteurs environnementaux et l'abondance des plantes. Elle l'est également pour faire des prédictions réalistes sur les effets de modifications envisagées de l'environnement sur des communautés de plantes. La manipulation de l'écosystème complet est complexe, ce qui rend son utilisation rare. Une méthode alternative a été utilisée pour l'étude de la composition spécifique des communautés de plantes submergées des marais temporaires de Camargue. Elle consiste à prélever des échantillons non perturbés de sédiment, avec leur stock de semences intact, et à les soumettre à des conditions de salinité. L'étude de la biomasse totale et de la biomasse des espèces les plus fréquentes est faite au moyen de test paramétriques. La précision attendue des résultats dépend de la fréquence de l'espèce dans les réplicats. Lorsque la fréquence n'est pas élevée l'emploi de statistiques non paramétriques est indispensable. La méthode se révéle particulièrement bien adaptée à l'étude des communautés d'annuelles en milieu aquatique.

     

Wetland restoration by natural succession in abandoned pastures with a degraded soil seed bank

Natural restoration of historical wetland plant communities in fallow fields with a degraded seed bank has been assumed to be possible only if source populations of the target species are present adjacent to the abandoned fields and a high density of suitable microsites is available. However, few studies have monitored both factors simultaneously and verified this assumption. We hypothesized that plant communities that are similar to historical wetlands, including back marshes, back swamps, and bogs, will reestablish in abandoned pasturelands in cases when (1) gaps for new recruitment emerge, followed by the decline of pastures; and (2) seeds with longevity are supplied from the surrounding remnant plant communities of wetlands. We conducted a survey of vegetation and microsites in pastures, abandoned pastures, and reference wetlands followed by structural equation modeling to verify our hypothesis for the natural restoration of Phragmites australis–Phalaris arundinacea and Alnus japonica–Spiraea salicifolia communities. These communities represent historical back marshes and back swamps along a river. However, our hypothesis was not verified for the natural restoration of Vaccinium oxycoccos–Sphagnum spp. communities, which represent plant communities in historical bogs grown on acidic peat that are maintained by rainfall and fog. Our findings partly support our hypothesis that decline in pastures creates gaps and that cumulative seed dispersal from nearby remnant wetlands allows the original wetland plant communities to regenerate. Further case studies are needed to determine how the natural restoration of bog plant communities occurs.     

Restoration of salt marshes in the Netherlands

The conquest of land from the sea has been a long tradition in the Netherlands. When salt marshes were high enough, they were embanked when it was economically feasible, and transformed into intensively exploited agricultural land. This resulted in the transformation of halophytic communities to glycophytic communities. Often as an alternative, a low levee, a summerdike was built, which greatly reduced the flooding frequency of the landward summerpolder, hence creating a sedimentation deficit therein. Such summerpolders now cover 1200 ha in the Netherlands, 2100 ha in NW-Germany and small areas in England. Due to continuous embankments, the present salt-marsh area is relatively small with respect to the tidal basins. Discussions have been started how to increase the salt-marsh area. Two options will be discussed, firstly de-embankment of summerpolders and maintenance of the protective seawall, secondly increase of the effects of saline seepage behind the seawall by top soil removal. Both options include the restoration of salt-marsh communities (target communities) in intensively agriculturally exploited sites that have been salt marshes before. From the few examples abroad and experiments it is discussed (1) to which extent the sedimentation deficit in summerpolders could be compensated for, (2) if the soil seed bank is likely to contribute to re-establishment of salt-marsh communities, (3) if the dispersal of propagules of halophytic plants will be possible by hydrochory when the summerdike is breached, (4) to what extent is dispersal by endozoochory through waterfowl important in case re-establishment in a saline seepage area behind the seawall without open connection to the sea is envisaged. Two case studies of de-embanked summerpolders in the Netherlands revealed that the sedimentation deficit can be counteracted by rapid sedimentation, provided enough transport is possible from the foreshore. Dispersal by incoming tidal water from the nearby salt-marsh source area into the target area is possible for many salt-marsh plant species. The rate of success seems to depend on the relative position of source area and target area. A case study in a saline seepage area after top soil removal in the Netherlands, showed that the number of viable seeds dispersed by droppings from waterfowl is limited. Hence the possibilities for restoration of inland halophytic plant communities seem much lower than after de-embankment of summerpolders.     

杂草种子传播研究进展

种子传播将母株生殖周期的末端与它们后代种群的建立连结了起来,广泛认为,其对植被结构具有深刻的影响。种子传播的整个过程称为种子传播循环。研究表明,杂草种子传播的因子多种多样,包括仅依赖自身来完成的主动传播,以及依赖风、水、动物、人类等外界媒介的被动传播。其中,人类传播杂草种子是影响最广泛的一种,对现代植物的分布格局产生了深刻的影响。杂草种子的传播,对杂草种子库的数量和空间动态影响很大。研究种子传播的主要方法有荧光染料标记法、放射性同位素标记法、稳定同位素分析、分子遗传标记等。结合近几年国内外的研究进展,作者就杂草种子传播对种子库数量和空间动态影响的精确直接研究、杂草种子传播的过程及传播后的命运、杂草种子适应传播的机理、生态控草措施研究、外来杂草入侵蔓延与其种子传播的关系等方面提出了展望。     

Soil seed bank and driftline composition along a successional gradient on a temperate salt marsh

Abstract. This study focuses on the relationship between vegetation succession and soil seed bank composition on the Schiermonnikoog (The Netherlands) salt marsh over 100 yr. The importance of driftline material in seed dispersal and the relationship with succession is also investigated. The results indicate that the majority of species have a transient or short‐term seed persistent bank. Seeds of most species are able to float over the salt marsh and become concentrated in the driftline higher up the marsh. After plants have established a seed bank forms, which disappears when vegetation is replaced by later‐successional species. Exceptions are Spergularia mar‐itima, which is still present in the seed bank of late successional stages, and Juncus gerardi and Glaux maritima, which appear in the seed bank of early successional stages, but are absent in the vegetation. Based on the results of this study constraints and possibilities for salt‐marsh restoration by de‐embankment are discussed.     

Long-distance seed dispersal in plant populations

Long-distance seed dispersal influences many key aspects of the biology of plants, including spread of invasive species, metapopulation dynamics, and diversity and dynamics in plant communities. However, because long-distance seed dispersal is inherently hard to measure, there are few data sets that characterize the tails of seed dispersal curves. This paper is structured around two lines of argument. First, we argue that long-distance seed dispersal is of critical importance and, hence, that we must collect better data from the tails of seed dispersal curves. To make the case for the importance of long-distance seed dispersal, we review existing data and models of long-distance seed dispersal, focusing on situations in which seeds that travel long distances have a critical impact (colonization of islands, Holocene migrations, response to global change, metapopulation biology). Second, we argue that genetic methods provide a broadly applicable way to monitor long-distance seed dispersal; to place this argument in context, we review genetic estimates of plant migration rates. At present, several promising genetic approaches for estimating long-distance seed dispersal are under active development, including assignment methods, likelihood methods, genealogical methods, and genealogical/demographic methods. We close the paper by discussing important but as yet largely unexplored areas for future research.     

The role of avian ‘seed predators’ as seed dispersers

Seed dispersal is a central process in plant ecology with consequences for species composition and habitat structure. Some bird species are known to disperse the seeds they ingest, whereas others, termed ‘seed predators’, digest them and apparently play no part in dispersal, but it is not clear if these are discrete strategies or simply the ends of a continuum. We assessed dispersal effectiveness by combining analysis of faecal samples and bird density. The droppings of seed dispersers contained more entire seeds than those of typical seed predators, but over a quarter of the droppings of seed predators contained whole seeds. This effect was further magnified when bird density was taken into account, and was driven largely by one frequent interaction: the Chaffinch Fringilla coelebs, a typical seed predator and the most abundant bird species in the area and dispersed seeds of Leycesteria formosa, a non‐native plant with berry‐like fruits. These results suggest the existence of a continuum between seed predators and seed dispersers.     

Relationships between seed germinability of Spergularia marina (Caryophyllaceae) and the formation of zonal communities in an inland salt marsh

BACKGROUND AND AIMS: The formation of zonal communities may be attributed to differences in germination across the community and to timing of germination of seeds present in the seed bank. Our goals were two-fold: (1) to assess the annual germination pattern of Spergularia marina; and (2) to determine whether germination of S. marina differed across zonal communities. METHODS: Fresh seeds were buried in an experimental garden in polyester bags. Bags were harvested monthly for 1 year and exposed to differing 12 h/12 h temperature regimes (5/15 degrees C, 5/25 degrees C, 15/25 degrees C and 20/35 degrees C) with a 12 h dark/12 h light photoperiod. Replicate seeds were exposed to 24 h dark. Seeds were also placed in different zonal communities to assess germinability in the field. KEY RESULTS: Spergularia marina has a primary physiological dormancy. Conditional dormancy occurs from December to May and non-dormancy from June to November. Field germination initiates in the spring when temperatures are cool and salinity is low due to flooding, and ceases in the summer when temperatures exceed germination requirements. Spergularia marina has a light requirement for germination. CONCLUSIONS: If seeds become buried in the field or are light inhibited by Phragmites australis, they will remain dormant until they receive an adequate amount of light for germination. Since S. marina can germinate across all zones in a salt-marsh community, the formation of zonal communities is not determined at the germination stage, but at some later stage of development.     

Overseas seed dispersal by migratory birds

Long-distance dispersal (LDD) promotes the colonization of isolated and remote habitats, and thus it has been proposed as a mechanism for explaining the distributions of many species. Birds are key LDD vectors for many sessile organisms such as plants, yet LDD beyond local and regional scales has never been directly observed nor quantified. By sampling birds caught while in migratory flight by GPS-tracked wild falcons, we show that migratory birds transport seeds over hundreds of kilometres and mediate dispersal from mainland to oceanic islands. Up to 1.2% of birds that reached a small island of the Canary Archipelago (Alegranza) during their migration from Europe to Sub-Saharan Africa carried seeds in their guts. The billions of birds making seasonal migrations each year may then transport millions of seeds. None of the plant species transported by the birds occurs in Alegranza and most do not occur on nearby Canary Islands, providing a direct example of the importance of environmental filters in hampering successful colonization by immigrant species. The constant propagule pressure generated by these LDD events might, nevertheless, explain the colonization of some islands. Hence, migratory birds can mediate rapid range expansion or shifts of many plant taxa and determine their distribution.     

A seed predator drives the evolution of a seed dispersal mutualism

Although antagonists are hypothesized to impede the evolution of mutualisms, they may simultaneously exert selection favouring the evolution of alternative mutualistic interactions. We found that increases in limber pine (Pinus flexilis) seed defences arising from selection exerted by a pre-dispersal seed predator (red squirrel Tamiasciurus hudsonicus) reduced the efficacy of limber pine's primary seed disperser (Clark's nutcracker Nucifraga columbiana) while enhancing seed dispersal by ground-foraging scatter-hoarding rodents (Peromyscus). Thus, there is a shift from relying on primary seed dispersal by birds in areas without red squirrels, to an increasing reliance on secondary seed dispersal by scatter-hoarding rodents in areas with red squirrels. Seed predators can therefore drive the evolution of seed defences, which in turn favour alternative seed dispersal mutualisms that lead to major changes in the mode of seed dispersal. Given that adaptive evolution in response to antagonists frequently impedes one kind of mutualistic interaction, the evolution of alternative mutualistic interactions may be a common by-product.