Functional diversity analysis helps to identify filters affecting community assembly after fen restoration by top-soil removal and hay transfer

Top-soil removal followed by species introduction through hay transfer has appeared as a method to restore drained fens. This method addresses abiotic constraints by restoring hydrology and nutrient status, and biotic constraints by removing an unwanted seed bank and counteracting dispersal-limitation. Restoration works by altering environmental filters. Knowledge about the restoration actions effect on functional traits is necessary to understand which types of species may establish. In this study we analyse which factors in top-soil removal followed by hay transfer influence selection and composition of functional traits. Top-soil removal followed by hay transfer from reference sites was conducted at two sites in the Całowanie fen, 33 km SE of Warsaw, Poland. Species and abundance data were recorded for three consecutive years. These data, combined with data on functional traits were used to analyse the effect of the restoration actions on four functional diversity-indices and the community weighted mean of functional traits. Our results reveal a strong habitat filter in the restoration site that follows an elevation gradient. At low elevation this filter selects low values of autochory and specific leaf area and high values of clonal lateral spread, Ellenberg moisture values, and dispersal through hydrochory. The transferred hay differs in trait characteristics compared to the reference site vegetation by having species of higher specific leaf area, lower Ellenberg moisture value and lower dispersal by autochory and hydrochory. The result presented here has three important implications for fen restoration. First, the difference in trait-characteristics between the transferred hay and the reference site it was harvested from limits the restoration potential. Second, since for several fen species important functional traits are filtered along an elevation-gradient, careful planning regarding depth of top-soil removal is needed. Finally the results illustrate how a functional analysis can be used to detect environmental filters acting during ecological restoration.     

Revegetation following soil disturbance in a California meadow: the role of propagule supply

Revegetation following a disturbance event initially should be constrained by the abundance and types of propagules available at the disturbed site. I tested this idea by conducting two experiments in which I created artificial soil disturbances by excavating or burying pre-existing grassland vegetation. In the first experiment, I varied disturbance intensity (depth), to investigate the consequences for revegetation when numbers of surviving propagules (dormant seeds and bulbs) were altered. In the second experiment, I varied the timing of disturbance, to investigate the consequences when disturbed sites experienced differing exposures to seasonal patterns of clonal growth and seed dispersal. I sampled these experiments from 1991 to 1993, and have interpreted their results using measurements of the seed bank, the bulb bank, and the seed rain. In the first (depth) experiment, bulbs declined in abundance with burial depth and were scarcer in deeper excavations. In contrast, numbers of annual graminoids initially showed no trends with respect to disturbance depth. These results reflect the depth distributions of the seed and bulb banks. Since bulbs occur deeply in the soil, progressively deeper disturbances left fewer survivors. Similarly, perennial graminoids could grow through the shallowest burials. In contrast, since the annual-graminoid-dominated seed bank is concentrated near the soil surface, disturbance depth mattered less to these species: any disturbance removing the surface layer was equally destructive. In the second (timing) experiment, more annual graminoids initially occurred in older plots. This result reflects seasonal patterns of seed production: plots exposed to more of the annual-graminoid-dominated seed rain supported higher densities of annual graminoids as a result. In subsequent years, the vegetation of most plots in both experiments was increasingly dominated by annual graminoids, again as a consequence of their great abundance in the seed rain. These results indicate that interactions between soil disturbances and sources of propagules play an important role in controlling early stages of succession in newly created gaps. They also suggest that disturbance may play different roles in communities characterized by species with different reproductive strategies. Understanding sources of colonists will improve our ability to predict the effects of disturbance.     

Seed supply of native and cultivated grasses in pine forests of the southwestern United States and the potential for vegetation recovery following wildfire

In this study we examined seed supply (seed banks and seed rain) and vegetation of seeded cultivated grasses and naturally occurring native grasses following a wildfire in northern New Mexico, USA. We specifically examined density of native and cultivated grass seeds and plants in areas of high, moderate, and low fire severity. We also examined the similarity between density of native and cultivated grass seeds to density of above ground plants. Density of native grass seed per square meter was higher in areas that burned under low fire severity (85.18 ± 44.83) compared to areas of moderate (18.52 ± 11.26) and high (7.41 ± 4.90) fire severity; however, differences were not statistically significant due, in part, to the high error associated with estimates. Density of cultivated grass seed per square meter was higher than that of native grass seed in areas of high (439.60 ± 117.98) and moderate (437.02 ± 146.50) fire severity, areas that were seeded with cultivated grasses after the wildfire for erosion control. Density of seeded grass plants per square meter was also higher than that of native grass plants in areas of high (18.78 ± 4.59 versus 0.33 ± 0.24) and moderate (8.22 ± 1.76 versus 0.22 ± 0.15) fire severity. There was a higher correspondence between the density of cultivated grass seeds and plants (highest value 0.32 ± 0.11) compared to density of native grass seeds and plants (highest value 0.05 ± 0.04). The high density of seeds, plants, and correspondence indicated that seeds from cultivated grasses are more likely to establish as post-fire vegetation than seeds from native grasses. Seeding with cultivated grasses following a wildfire may slow or inhibit recovery of native grasses in the short term. Longer-term implications for site occupancy deserve further study.     

The relationship between productivity and multiple aspects of biodiversity in six grassland communities

Biodiversity is a multifaceted concept but most studies examining the association between the biodiversity of a community and its productivity focus only on species richness. Consequently, studies are needed to examine how other facets of biodiversity vary with productivity if we want to have a better understanding of the distribution of biodiversity across our planet. We evaluated how a number of biodiversity measures (species richness, evenness, dominance, rarity, Simpson’s diversity, and Shannon–Weiner diversity) varied across natural productivity gradients at 6 grassland sites in the continental US. Variation in productivity did not account for a substantial amount of variation in any measure of biodiversity at small spatial scales (≈1 m²) at most sites. When productivity accounted for substantial variation in biodiversity, different measures of biodiversity responded to productivity in different ways. For example, dominance changed in a U-shaped fashion along a productivity gradient whereas richness increased in an asymptotic fashion. Consequently, diversity indices, which account for both species richness and evenness, varied in a hump-shaped fashion along the productivity gradient. Our results highlight that an exclusive focus on the association between species richness and productivity provides an incomplete picture of how a community’s biodiversity is related to its functioning.     

Vertical structure of seed banks and the impact of depth of burial on recruitment in two temporary marshes

The structure of the seed bank (including Chara oospores), in relation to depth within the sediment and disturbance, was studied in two Rhône delta temporary marshes for two years. The seeds of all species were concentrated in the top 2 cm of sediment with very low numbers beeing found below 4 cm. When an exclosure eliminated disturbances of the sediment by animals, the vertical repartition of seeds at site 2 was more pronounced than outside the exclosure.In experiment 1, the emergence capacity of seeds from different depths and buried under layers of sterile equivalent to those in the field was measured. Depending of the species, 22 to 98% of the seeds germinated from unburied seeds in the top 2 cm. Only 1% of the oospores of Chara (from site 2) at 2 to 4 cm depth in the sediment emerged.In experiment 2, surface seed bank samples were placed under 0, 2 or 4 cm sterile sediment depth. The samples contained numerous recent seeds and the emergence percentage reached 41% (for Ruppia maritima). Only the seeds of Zannichellia spp failed to germinate from a depth of 2 cm or more. The emergence percentage from 2 cm depth or more was always lower than at the surface. These experiments showed that both burial and ageing of seeds decrease germination capacity.The majority of the active seeds located at the surface germinate when the marsh is flooded. Seeds located between 2 and 4 cm can be brought back to the surface by disturbances and play the role of a reserve involved in maintenance of populations that go without seed production for one or some years.     

植物群落功能多样性计算方法

以性状为基础的功能多样性指数在预测生态系统功能中起到越来越重要的作用.本文对近年来陆续出现的植物群落功能多样性指数进行综述.依据物种多样性指数的组成,功能多样性指数分为功能丰富度、功能均匀度和功能离散度指数.介绍了这3类指数的计算方法,有助于更好、更准确地理解“生物多样性-环境-生态系统功能”的关系.     

水盐变化对荒漠植物功能多样性与物种多样性关系的影响

研究植物功能多样性与物种多样性关系对土壤水盐环境的响应规律,有助于掌握胁迫环境下植物多样性的维持机理.本研究分析了艾比湖高、低土壤水盐环境下荒漠植物功能多样性、物种多样性差异及二者间关系在不同水盐环境下的变化规律.结果表明:(1)高水盐环境下植物功能离散度显著低于低水盐环境(P<0.05),物种多样性指数Fish-α显著高于低水盐环境;(2)一元与多元回归分析均表明,高水盐环境下物种多样性对功能多样性RaoQ与FRic指数有显著促进作用,低水盐环境下物种多样性对功能多样性和均匀度有显著促进作用,但对功能离散度具有显著抑制作用(P<0.05);(3)物种多样性对功能多样性的影响在低水盐环境下更显著,预测力更强;(4)功能丰富度、离散度与均匀度能较敏感地反映出荒漠植物功能多样性对水盐变化的响应.本研究有助于理解荒漠植物群落构建机制,并为荒漠植物恢复与管理提供理论参考.     

How do depth,duration and frequency of flooding influence the establishment of wetland plant communities?

In many temporary wetlands such as those on the Northern Tablelands of New South Wales Australia, the development of plant communities is largely the result of germination and establishment from a long-lived, dormant seed bank, and vegetative propagules that survive drought. In these wetlands the pattern of plant zonation can differ from year to year and season to season, and depth is not always a good indicator of the plant community composition in different zones. In order to determine which aspects of water regime (depth, duration or frequency of flooding) were important in the development of plant communities an experiment using seed bank material from two wetlands was undertaken over a 16 week period in late spring–early summer 1995–1996. Seed bank samples were exposed to 17 different water-level treatments with different depths, durations and frequencies of flooding. Species richness and biomass of the communities that established from the seed bank were assessed at the end of the experiment and the data were examined to determine which aspects of water regime were important in the development of the different communities. It was found that depth, duration and frequency of inundation influenced plant community composition, but depth was least important, and also that the duration of individual flooding events was important in segregating the plant communities. Species were grouped according to their ability to tolerate or respond to fluctuations in flooding and drying. The highest biomass and species richness developed in pots that were never flooded. Least biomass and species richness developed in pots that were continuously flooded. Short frequent floods promoted high species richness and biomass especially of Amphibious fluctuation-tolerator species and Amphibious fluctuation-responder species that have heterophylly. Terrestrial species were able to establish during dry phases between short floods. Depth was important in determining whether Amphibious fluctuation-tolerator or Amphibious fluctuation-responder species had greater biomass. Longer durations of flooding lowered species richness and the biomass of terrestrial species. Experiments of this kind can assist in predicting vegetation response to water-level variation in natural and modified wetlands.     

坡向因子对黄土高原草地群落功能多样性的影响

研究群落水平上的植物功能性状特征及功能多样性随坡向的变化规律,对认识不同坡向上的植物群落形成过程具有重要意义。以黄土高原不同坡向上的自然草地群落为研究对象,比较研究了植物功能性状(株高和比叶面积)和功能多样性测度指标(功能丰富度、功能均匀度和功能离散度)随坡向的变化规律。研究结果显示:(1)阴坡的株高和比叶面积显著高于其他坡向;(2)一元性状的功能丰富度在不同坡向间均无显著差异;阴坡和半阴坡的多元性状功能丰富度显著高于阳坡;(3)阴坡的株高功能均匀度显著高于半阳坡,而比叶面积功能均匀度在各坡向的差异并不显著;多元性状功能均匀度在不同坡向差异显著,阴坡最高,半阳坡最低;(4)阴坡和半阴坡的株高功能离散度显著高于阳坡和半阳坡,而半阴坡的比叶面积功能离散度显著高于阳坡;半阴坡的多元性状功能离散度Rao指数显著高于阳坡。研究结果暗示了,在阴坡和半阴坡上,植物对群落内的生态位空间和资源利用更充分,种间竞争强度较低,不同物种之间生态位高度分化;而在阳坡和半阳坡上,由于水分等条件的限制植物可占据的生态位空间有限,导致其对占据的生态位空间使用不足,物种间资源竞争较强烈。在群落水平上,研究地区的植物功能性状及功能多样性随坡向的规律性变化,反映了黄土高原植被群落构建过程中坡向因子对功能性状的筛选效应。该研究结果对该区的植被恢复重建的物种选择及植被布局规划具有重要实践意义。