Mediterranean temporary ponds. A disappearing ecosystem

Mediterranean temporary ponds (MTPs) is a priority habitat according to the Natura 2000 network of the European Union (Natura code 3170, Habitats Directive 92/43/EEC) and is located in various Mediterranean countries. Priority habitats are those habitat-types or elements with a unique or important significance to a diverse group of species. They consist of a unique flora composition, succession stage and/or structural factor. Increased urbanisation and agriculture in addition to climate change, in the Mediterranean region, has led to the extinction of a very large number of temporary ponds. This paper is a review of the value and the numerous threats, concerning MTPs. Protection and conservation of the ponds is very important and appropriate managerial measures should be immediately taken, to save such a vulnerable ecosystem.     

Impact of ploughing on soil seed bank dynamics in temporary pools

We examined the impact of ploughing on soil seed banks of plant communities living in temporary marshes located in agricultural fields. The quantity, quality and vertical distribution of seeds were quantified under ploughed or unploughed treatment at community level. We also focussed on a typical semi-aquatic species, Damasonium alisma, to investigate the impact of ploughing at population level. We used two complementary techniques to study seed banks: hand sorting and seedling emergence. We found that species richness of seeds, seed abundance and germination ability were strongly affected by ploughing at community level. Concerning D. alisma, most of the seeds (56%) were stored in the two deepest soil layers among the four considered in ploughed pools. Moreover, the germination rate was higher for buried seeds (84%) than for seeds collected at the surface (33.6%). These patterns were almost inverted in unploughed pools. Our results agree with the temporal storage effect generally suggested to describe the seed bank property of plant communities. But in addition, we showed that ploughing induces a spatial storage effect in accumulating species and individuals in the seed banks that favourably influence community dynamics. We conclude that, in contrast to what is usually thought, ploughing disturbance can be of benefit for such ephemeral wetland vegetation.     

Biotic and abiotic parameters that distinguish types of temporary ponds in a Portuguese Mediterranean ecosystem

Temporary ponds are seasonal wetland habitats subjected to extreme and unstable ecological conditions. Some are classified as priority habitats for conservation by the European Union Habitats Directive. Our study area was the coastal plain of southwest Portugal, which spans across 100 km north to south and hosts a large number of temporary ponds as a consequence of climatic and edaphic characteristics. Field sampling of floristic and edaphic data was carried out in 24 temporary ponds every spring between 2005 and 2008. We recorded a total of 174 plant species identified within visually homogeneous plots. We included the data in a geographic information system and classified ponds according to their floristic composition, using a biotic regionalization analysis based on species presence/absence, which is a practical and unambiguous criterion. We found three significantly different groups of ponds which corresponded to an eco-physiognomic pond typology: Mediterranean temporary ponds, marshlands, and disturbed ponds. For the first two pond types we defined characteristic or indicator plant species. We searched also for relationships between pond type and a series of large-scale climatic, geographic, and geological variables, as well as local-scale physical and chemical properties of the soil. Pond type was distinguished by a complex combination of some of these variables, including environmental energy, soil texture, nitrogen content of the soil and pH. A practical way of discriminating between different types of ponds is important so that management and conservation measures can be defined accordingly.     

Soil seed bank and community dynamics in an annual-dominated Mediterranean salt-marsh

Abstract. The seed bank in the soil and litter of an annual grassland in the upper salt marsh of the Guadalquivir delta (SW Spain) contained on average 49110 seeds/m²; 29 species were represented. Field censuses of the grassland, in two consecutive springs, yielded 7 410 and 9 520 plants/m² respectively. A combined Correspondence Analysis of the seed bank samples and field censuses showed differences in grassland composition between wet and dry years. The environmental changes induced different expressions of the soil seed bank, mediated by demographic processes. The effect of salinity (25 dS/m of NaCl) was studied on intact soil blocks transplanted to a greenhouse. Salinity did not affect the emergence of seedlings, but reduced the growth of the dominant species, Trisetaria panicea. The importance of the role of seed banks in community dynamics and their use for conservation management are discussed.     

围封沙质草甸土壤种子库与幼苗库的特征

采用野外定点取样和萌发实验相结合的方法,研究了科尔沁沙地围封沙质草甸土壤种子库和幼苗库的特征.结果表明:土壤种子库总密度为13854±1651有效种子数/m2,而幼苗库的总密度为808±164株/m2,仅为土壤种子库总密度的6%,土壤种子库与幼苗库总密度之间存在显著的差异 F=36.0,n=219,P<0.001 .土壤种子库与幼苗库的植物种数分别为37和44,其中一、二年生草本植物分别占种子库与幼苗库植物种数的62%和50%,而多年生草本植物所占的比例分别为24%和41%,灌木所占的比例很小,分别为14%和9%.土壤种子库与幼苗库种类组成上的相似性系数为0.667,表现出较小的异质性.土壤种子库密度与幼苗库密度之间存在显著的相关性,其间关系可用三次曲线来描述.     

西双版纳热带森林土壤种子库的季节变化

通过萌发实验法对西双版纳地区的一类热带季节雨林 (番龙眼、千果榄仁 )和 2类次生林(白背桐林、中平树林 )的土壤种子库的季节变化进行了探讨。结果表明 :该地区的土壤种子库动态具有明显的季节性。季节雨林的土壤种子库储量相对稳定 ,土壤上层 ( 0～ 2 cm )的种子储量在雨季末期较大。2类次生林土壤种子库的变化则相反 ,土壤种子库中的种子种类在旱季末期较雨季末期多 ,土壤上层的种子储量在旱季末期较大。各样地均有一些种类只出现在旱季末期或雨季末期。种子在土壤种子库的动态与植物的繁殖物候和所处的环境紧密相关 ,不同种类植物的土壤种子库由于植物本身的生物学特性、传播方式和所处环境的影响而表现出不同的动态模式     

Seasonal fires shape the germinable soil seed bank community in eastern Mediterranean woodlands

季节性火烧塑造了东地中海地区林地的可萌发种子库群落本研究探索了在典型东地中海林地中,季节性火烧是否以及如何与微生境类型(即黄连木属(Pistacia)灌木下、岩蔷薇属(Cistus)灌木下或林冠间隙)相互作用,从而影响可萌发种子库(germinable soil seed bank, GSSB)群落的组成。我们进行了一项野外实验,其中包括预定的春秋火烧,然后利用从火烧地块和相邻未火烧对照地收集的土壤样本对种子萌发的模式进行定量。研究结果表明,秋烧期间,土壤温度比春烧的要高,而春烧期间土壤温度则比秋烧的变化更大。火烧导致可萌发种子库整体的密度、丰富性和多样性降低。秋烧地块内黄连木属和岩蔷薇属灌木的可萌发种子库丰富性和多样性显著降低,这些模式主要表现在一年生植物中。在火烧样地收集的样本中,低矮灌木的可萌发种子库密度更高,在黄连木属和岩蔷薇属灌木下采集的样品中,这种模式更为明显。在季节性火烧与出现的独特物种会共同导致可萌发种子库的群落组成发生显著的变化。这些结果表明,季节性火烧与空间异质性互相作用,主要通过对一年生植物和低矮灌木的可萌发密度的差异效应来影响可萌发种子库群落的组成。这些研究发现意味着,过去几十年东地中海盆地的季节性火烧频率上升,可能转化为进化生态选择压力的变化,并对这一独特生态系统内的植物产生影响。     

Aerial seed bank dynamics and seedling emergence patterns in two annual Mediterranean Asteraceae

Question: We explored the functional significance of seasonal aerial seed banks in two coexisting, heterocarpic annual Asteraceae with dormant (Chrysanthemum coronarium) and non‐dormant (Anacyclus radiatus) achenes. We hypothesised that the plant achene pool is a significant component of total seed reserves, and that within‐season seedling emergence timing is shaped by achene release patterns. Location: SW Spain. Methods: In an observational study, we established temporal achene release patterns. We also quantified the aerial and soil achene pools throughout the release season, and assessed seedling emergence timing. Sowing experiments were used to explore the influence of achene release dynamics on emergence timing, and to establish achene morph‐specific patterns of between‐year distribution of germination. Results: Achene release extended from late spring to late autumn (Chrysanthemum), or from early autumn to early winter (Anacyclus). Within species, achene morphs differed in release timing. Only in Chrysanthemum, a small achene fraction seemed to persist in the soil, and between‐year germination distribution differed among morphs. In coexisting populations, the Anacyclus plant achene pool was an order of magnitude higher than the soil pool throughout the release season, whereas in Chrysanthemum both pools were of the same magnitude during autumn. Within‐year seedling emergence was significantly staggered beneath parent plants compared with the pattern resulting solely from the germination response in soil, with the exception of Chrysanthemum in one of the two study years. Conclusions: Results suggest that seasonal aerial seed banks are effective within‐season, risk‐reducing traits in ruderal Mediterranean habitats characteristic of the study species.     

Submerged macrophyte seed bank in a Mediterranean temporary marsh: abundance and relationship with established vegetation

The abundance and composition of the submerged macrophyte seed bank in the Doñana marsh (southwestern Spain) was evaluated to assess its relationship with the overlying vegetation. The results obtained demonstrate the existence of a dense seed-bank, both in terms of the number of seeds and their biomass, which represented about 10% (5% for angiosperms and >20% for Charophyta) of the total plant biomass, which ensures the maintenance of the annual submerged macrophyte populations of the seasonally inundated Donñana marsh. Seed bank and established vegetation were coupled, as reflected in the existence of significant correlations between their structure and abundance. This coupling was lacking for Charophyta, whose oospores are widespread and abundant across the marsh, even at locations where they are absent, or rare, in the established vegetation. These differences between the seed bank of annual angiosperms and Charophyta appear to reflect, in part, structural differences between angiosperm seeds and Charophyta oospores, with important ecological consequences. An important aspect of these differences is the allocation of Charophyta reproductive effort to many small propagules lacking embryo storage, compared to those of angiosperms, which ensures their efficient dispersal and numerical abundance in the seed bank.     

Demography and role of the seed bank in Mediterranean and desert populations of wild barley

We compared the demography of two populations of wild barley, Hordeum spontaneum, that occupy environments of different aridity (desert and Mediterranean) and represent distinct parts of the species' distribution (periphery and core) in Israel. Our demographic study included a survey of survival and fecundity of adults in quadrate plots marked at two sites in natural vegetation (one site per population), and the creation of experimental seed banks examined over three years. We measured variability in population growth rate and investigated the role of the seed bank in the demography of two populations. The latter employed construction of two-stage, three age-class transition matrices and usage of elasticity analysis. At both sites, population dynamics appear to be transient with high fluctuations of population growth rate due to variability in annual rainfall and, most likely, intra- and inter-specific density dependence. The importance of the seed bank differed among two populations. Seed dormancy had almost no demographic effect on the Mediterranean population, while it was of paramount importance in the desert population. The two populations also differed in seed and seedling survival and per adult fecundity, presumably due to the difference between two sites in (i) relative impact of seed predation, (ii) effects of density dependence on population demography, and (iii) lower vigor of seeds of desert vs. Mediterranean origin. Our study highlights the importance of life history adaptations that may evolve under specific selective forces in different parts of a species' range, which, in certain circumstances, may be critical for the persistence of a species.Wir verglichen die Demographie von zwei Populationen Wilder Gerste, Hordeum spontaneum, die Habitate unterschiedlicher Trockenheit (Wüste und mediterran) besetzen und die unterschiedliche Bereiche der Artenverbreitung (Peripherie und Kern) in Israel repräsentieren. Unsere demographischen Untersuchungen bestanden in einer Erfassung der Überlebensrate und der Fruchtbarkeit ausgewachsener Pflanzen in quadratischen Versuchsfeldern, die in zwei Untersuchungsflächen in der natürlichen Vegetation (eine Untersuchungsfläche pro Population) markiert wurden und in der Herstellung einer experimentellen Samenbank, die über drei Jahre beobachtet wurde. Wir erfassten die Variabilität der Populationswachstumsrate und untersuchten die Rolle der Samenbank in der Demographie der beiden Populationen. Letzteres erforderte die Konstruktion von Zwei-Stadien, Drei-Altersklassen-Transitions-Matrizen und die Verwendung einer Elastizitäts-Analyse. Die Populationsdynamiken scheinen in beiden Untersuchungsflächen kurzlebig zu sein, mit einer hohen Fluktuationsrate des Populationswachstums aufgrund der Variabilität des jährlichen Niederschlags und am wahrscheinlichsten aufgrund der intra- und interspezifischen Dichteabhängigkeit. Die Bedeutung der Samenbank unterschied sich zwischen den beiden Populationen. Die Samendormanz hatte bei der mediterranen Populationen nahezu keinen demographischen Effekt, während sie bei der Wüstenpopulation von herausragender Bedeutung war. Die beiden Populationen unterschieden sich auch in der Überlebensrate der Samen und Keimlinge und der Pro-Pflanzen-Fruchtbarkeit, vermutlich aufgrund der Unterschiede der beiden Untersuchungsflächen in (i) der relativen Wichtigkeit der Samenprädation, (ii) den Effekten der Dichteabhängigkeit der Populationsdemographie und (iii) der geringen Vitalität der Samen der Wüsten-Herkunft im Gegensatz zur mediterranen Herkunft. Unsere Studie hebt die Bedeutung einer Anpassung im Lebenszyklus hervor, die bei spezifischen selektiven Faktoren in den unterschiedlichen Teilen eines Verbreitungsgebietes einer Art evolvieren können und die unter bestimmten Umständen wichtig für den Bestand einer Art sein können.     

Factors influencing above‐ground and soil seed bank vegetation diversity at different scales in a quasi‐Mediterranean ecosystem

Questions

Are factors influencing plant diversity in a fire‐prone Mediterranean ecosystem of southeast Australia scale‐dependent?

Location

Heathy woodland, Otways region, Victoria, southeast Australia

Methods

We measured patterns of above‐ground and soil seed bank vegetation diversity and associated them with climatic, biotic, edaphic, topographic, spatial and disturbance factors at multiple scales (macro to micro) using linear mixed effect and generalized dissimilarity modelling.

Results

At the macro‐scale, we found species richness above‐ground best described by climatic factors and in the soil seed bank by disturbance factors. At the micro‐scale we found species richness best described above‐ground and in the soil seed bank by disturbance factors, in particular time‐since‐last‐fire. We found variance in macro‐scale β‐diversity (species turnover) best explained above‐ground by climatic and disturbance factors and in the soil seed bank by climatic and biotic factors.

Conclusions

Regional climatic gradients interact with edaphic factors and fire disturbance history at small spatial scales to influence species richness and turnover in the studied ecosystem. Current fire management regimes need to incorporate key climatic–disturbance–diversity interactions to maintain floristic diversity in the studied system.

     

Effect of the filling season on aquatic plants in Mediterranean temporary ponds

Aims Pond environmental conditions may differ among years with regards to the season in which ponds begin to fill. We experimentally evaluated how seedling emergence, plant growth and phenology differed among years in which filling occurred in winter, autumn or spring.Methods We collected sediments from a natural temporary pond and located them in aquariums. They were placed in a climatic chamber that simulated annual variation in field environmental temperatures and light conditions. Aquariums were assigned to one of three treatments, which differed in the date on which they were filled with water (autumn, winter and spring). We counted the number of seedlings of different species emerged and recorded data about the presence of flowers, seeds or spores every week. The experiment was finished in June, when we harvested the plants and estimated their biomass.Important findings In most species, seedling emergences were primarily related to time after filling, and thus synchronized their life cycles with the unpredictably timed wet phase of the ponds. Autumn filling resulted in the highest numbers of seeds/spores. However, winter filling promoted plant growth the most. In the spring filling treatment, more terrestrial plant seedlings emerged and fewer seeds/spores were produced. When ponds are flooded earlier, plants may produce a higher number of propagules. However, in years when inundation is delayed to spring and hydroperiods are short, seedling emergence deplete the seed bank and there is little to no seed production, while terrestrial monocots are able to colonize pond basin.     

Smoke interacts with fire history to stimulate soil seed bank germination in Mediterranean woodlands

Aims Fire has important consequences on vegetation dynamics. In fire-prone areas, natural selection favors plant species, characterized by a large soil seed bank, and that their germination is stimulated by fire. Although seed germination stimulated by fire heat is common in the eastern Mediterranean Basin, only little is known about germination stimulation by smoke. We examined the interactive effect of aerosol smoke and fire history on the germinable soil seed bank (GSSB) community in eastern Mediterranean woodlands.     

Restoration of high species density in calcareous grassland: the role of seed rain and soil seed bank

Abstract. A small-scale restoration experiment concerning a species-rich calcareous grassland was conducted over a 7-yr period. 20 yr of abandonment (1971-1990) resulted in both a strong reduction of the number of species and an absolute dominance of the shrub Crataegus monogyna and the grass Brachypodium pinnatum - which contributed 98% of the above-ground herb phytomass. The plot (3 m x 6 m) was cleared in September 1990 and thereafter yearly mown in autumn. Small soil seed bank samples were taken immediately after clearing and again in the autumn of 1996. The inner part of the plot showed a reduced soil seed bank compared to the plot underneath the vegetation outside the plot in 1990. Six years later an increased number of seeds in the soil in this inside area could be recorded mainly in the upper soil layer, although the amount was only 40% compared to the outside or control area. Ca. 20% of the species present in the restored vegetation were recorded in seed traps during the main seed dispersal period in 1997. Restoration of high species richness in the plot was a fast process during the first few years: from seven species in 1990 towards 39 three years later. This was significantly faster than the restoration process of the calcareous grassland at the same slope 20 yr earlier. This was probably due to the increased size of the community species pool at the site as a result of appropriate management during the last decades. Relative species number increase was highest at the largest scale during this experiment, viz. at the total plot area. The success of restoration of species-rich, dry grassland vegetation largely depends on the interaction between several conditions, e.g. seed availability, either originating from the seed rain or soil seed bank, favourable germination conditions and a successful development to a mature, seed producing individual. Germination and establishment strongly depend on suitable vegetation structure as a result of sustainable management.     

铜尾矿废弃地与相邻生境土壤种子库特征的比较

土壤种子库是植被群落的重要组成部分,其特征决定了群落的演替方向。对铜陵杨山冲尾矿库6个植被群落及相邻两处生境的土壤种子库进行了研究。结果表明,尾矿内各植被群落种子库的种子密度间存在较大差异,变化范围为57—593粒/m2,平均种子密度为370粒/m2,尾矿坝体和周围山体种子库种子密度分别为999和121粒/m2,各样地土壤中种子均主要集中在0—5 cm范围内。尾矿种子库与坝体种子库间的相似性指数大于其与周围山体种子库的相似性指数,但尾矿内各植被群落种子库间的相似性较差,相似性指数的变化范围为0.308—0.636。可见,群落微生境对土壤种子库的种子密度和物种组成产生了较大的影响。尾矿种子库种类与地上植被的平均相似指数为0.355,低于坝体种子库种类与地上植被相似性系数。3处样地土壤种子库的物种多样性、丰富度和均匀度指数均表现为坝体>尾矿>山体。尾矿种子库在农田土壤中萌发并正常生长的幼苗为16科44属53种,而在尾矿基质中仅为9科36属45种,均以禾本科、菊科和豆科植物萌发和生长状况较好,这3科构成了尾矿地上植被的主要类型。尾矿种子库在尾矿基质中萌发成活率为62.2%—91.2%,约为坝体和山体土壤种子库在尾矿基质中萌发成活率的2倍。说明,组成尾矿种子库的种子大部分已经适应了尾矿的极端恶劣环境或者其本身就是耐性极强的植物类型。     

The impact of vertic soil cracks on submerged macrophyte diaspore bank depth distribution in Mediterranean temporary wetlands

In temporary wetlands, a majority of diaspores are located in the upper layers of the soil. In general, it is assumed that the vertical movement of diaspores is caused by animal disturbance, and that reduction in this activity concentrates the diaspores in the topsoil. The occurrence of seasonal cracks and the dispersion of topsoil during dry periods are processes that have been underestimated in diaspore bank studies. Our results indicated that diaspore size can partially explain the density of diaspores of submerged macrophytes observed at four discrete soil depths in Doñana National Park temporary marshes (SW Spain). The top layers of the soil exhibit the largest number of species. Spore density of Riella spp., a liverwort, increases with depth, creating an important reserve of viable diaspores in deep soil. Downward movement of diaspores in soil cracks during dry periods, and the extended longevity of Riella diaspores together explain this inverse depth distribution.     

The impact of vertic soil cracks on submerged macrophyte diaspore bank depth distribution in Mediterranean temporary wetlands

In temporary wetlands, a majority of diaspores are located in the upper layers of the soil. In general, it is assumed that the vertical movement of diaspores is caused by animal disturbance, and that reduction in this activity concentrates the diaspores in the topsoil. The occurrence of seasonal cracks and the dispersion of topsoil during dry periods are processes that have been underestimated in diaspore bank studies. Our results indicated that diaspore size can partially explain the density of diaspores of submerged macrophytes observed at four discrete soil depths in Doñana National Park temporary marshes (SW Spain). The top layers of the soil exhibit the largest number of species. Spore density of Riella spp., a liverwort, increases with depth, creating an important reserve of viable diaspores in deep soil. Downward movement of diaspores in soil cracks during dry periods, and the extended longevity of Riella diaspores together explain this inverse depth distribution.     

Postfire seed bank dynamics in semiarid grasslands

We evaluated the combined effects of fire after drought on the seed bank composition and its role in the postfire recovery of NW Patagonia grasslands. During three years, we monitored the seed bank and the aboveground vegetation. Species were arranged in functional groups and Detrended Correspondence Analysis was used to separate sites according to species and functional groups. Similarity between aboveground vegetation and seed bank was calculated with SØrensen Index. In the first year, the seed density was similar in the control and burned sites and was lower than following years in all the sites. The species that survived the high temperatures were all annuals with the exception of the perennial species Fabiana imbricata and Rumex acetosella. In the second postfire year, the diversity and seed density increased due to the contribution of fugitive species (rare in the community) and exotic annual species. Seed bank of perennial species was the most affected by fire and just recovered in the third year. Drought did not affect the similarity between the seed bank and vegetation. Fire had low impact on the total seed bank, probably due to the heat buffering nature of the soil, whereas drought reduced significantly seed bank size and richness. Seed bank contributes to grassland richness maintenance.