Interactions between frequency and size of disturbance affect competitive outcomes

Disturbance has many effects on ecological communities, and it is often suggested that disturbance can affect species diversity by altering competitive outcomes. However, disturbance regimes have many distinct aspects that may act, and interact, to influence species diversity. While there are many theoretical models of disturbance-prone communities, few have specifically documented how interactions between different aspects of a disturbance regime change competitive outcomes. Here, we present a model of two plant species subject to disturbance which we then use to examine species coexistence over varying levels of two aspects of disturbance: frequency, and spatial extent (i.e., area disturbed). We show that the competitive outcome is affected differently by changes in each aspect and that the effect of disturbance frequency on species coexistence depends strongly on the spatial extent of the disturbance, and vice versa. We classify the nature of these interactions between disturbance frequency and extent on the basis of the shape of the resulting coexistence regions in a frequency?Cextent parameter plane. Our results illustrate that different types of interaction can result from differences in life-history traits that control species-specific sensitivity to frequency and extent of disturbance. Thus, our analysis shows that the various aspects of disturbance must be carefully considered in concert with the life-history traits of the community members in order to assess the consequences of disturbance.     

塔里木河下游不同地下水位下土壤种子库特征

选取塔里木河下游不同地下水位的25个天然植物群落样地,进行土壤种子库取样,通过萌发试验分析不同地下水位条件下土壤种子库的密度、物种多样性、生活型比例及土壤种子库与植物群落的相似性等特征.结果表明,在干旱地区,地下水位的高低不仅影响植物群落的种类和组成,而且也影响土壤种子库的密度和组成.随着地下水位的逐渐下降,植物群落退化程度的加重也反映到种子库上,表现为种子库密度减少、物种多样性下降、生活型逐渐单一,同时植物群落与土壤种子库的相似性系数也逐渐降低.该研究有助于丰富干旱区土壤种子库的理论和指导塔里木河下游的生态恢复实践.     

Assessing the impact of plant invasions on soil seed bank communities: use of univariate and multivariate statistical approaches

Questions: Are soil seed banks affected by invasions of alien plants? How can we rigorously assess alterations in seed bank communities associated with invasive species and account for the high spatial variability of seed bank data? How do multivariate approaches compare with more traditional approaches based on analysis of variance? Location: Three riparian sites, Ireland. Methods: A protocol based on a combination of multivariate techniques was used to characterize soil seed bank communities associated with the herbaceous invasive species Heracleum mantegazzianum in May and October. Permutational multivariate analysis of variance (PERMANOVA) was used to test the effects of the factors “invasion”, “site”, “plot” and “depth” on the soil seed bank, while multivariate analysis of dispersion (PERMDISP) provided a measure of the variability of seed bank data at different spatial scales. Similarity percentages analysis (SIMPER) was used to identify the species that contributed most to the differences between invaded and uninvaded communities. A comparison between the results of PERMANOVA and ANOVA analyses was also made. Results: The composition of seed bank communities invaded by H. mantegazzianum differed significantly from that of uninvaded seed banks. Invaded seed banks were less diverse and had reduced abundance, and were dominated by only a few species, such as Urtica dioica and Juncus effusus. Such patterns were recorded at each of three depth categories, indicating that invasive plants can affect both the transient and the more persistent component of the soil seed bank. Seed bank variability was significantly higher within uninvaded areas, supporting the notion that invasions tend to lead to more homogeneous communities. Conclusion: The analytical protocol used in this study was effective in quantifying the effect of plant invasions, at different spatial scales, providing a statistically robust analysis of alterations in soil seed bank communities. Compared to ANOVA, this protocol provided more biological information and was more appropriate for analysis of the data. This approach is therefore recommended in soil seed bank and invasion ecological studies.     

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.     

Effects of natural and artificial disturbance on landscape and forest structure in Tiantong National Forest Park,East China

This paper aims to understand the ecological effects of disturbance on broadleaved evergreen forest in East China. We used a manipulative field experiment approximating the common natural and artificial disturbance types in this area to investigate the community physiognomy, floristic composition, and 5-year recovery dynamics of the post-disturbance forest community. The results indicated that the landscape and forest structure have degraded into shrub communities, structure-damaged evergreen broadleaved communities, and so on. The post-disturbance communities presented different means of plant recruitment and vegetation recovery patterns at an early successional stage. The recovery of disturbed forests primarily depended on external seed sources and re-sprouting from stumps, rather than on soil seed banks, as few buried seeds were found. Re-sprouting thus appears to be key in allowing rapid vegetation recovery in evergreen broadleaved forest. Disturbances seem to be one of the most important factors that can contribute to regional species coexistence across temporal and spatial scales in evergreen broadleaved forests.     

植物种群更新的补充限制

补充限制基于生态位理论, 从种子萌发、幼苗存活和生长、繁殖体扩散等生活史阶段的种群统计特征及环境因素(土壤水分、养分、凋落物等)着手, 探讨种群的更新问题。种源限制和微生境限制是补充限制理论研究的核心内容, 但是哪个更为重要并没有统一的结论。种源限制与种子生产、土壤种子库和地下芽库中的繁殖体数量不足有关。其中, 气候的年际波动、土壤种子库寿命和动物的捕食都会影响种子生产在种群更新中的作用; 土壤种子库常被视为种群更新的保险库, 与地上种子雨共同促进种群更新, 但是, 如果土壤里种子具有较高的死亡率和休眠率, 将会降低种子库的作用; 地下芽库及其产生的无性分株对于种群更新的意义更多地体现在干扰后种群更强的恢复能力上。扩散限制是种群更新中的普遍现象, 与种子产量、散布能力、传播媒介、幼苗密度等因素有关。微生境限制主要表现为水分、养分、凋落物等非生物因素以及竞争、捕食等生物因素对种子的活力、萌发性、幼苗的存活力、物质分配等过程的影响, 其重要性随着植物生活史阶段而发生变化。未来需要进行综合的、长期的实验, 并应着重加强种源限制及相关生态过程的进化与生态相结合的机理性研究, 从而更深刻地认识和理解种群更新问题, 建立更为综合、系统的种群更新理论体系。     

The role of the seed bank,seed rain and the timing of disturbance in gap regeneration

Abstract. Question: Does the degree and timing of disturbance contribute significantly to the pattern and process of regeneration in plant communities as a consequence of the availability and number of species of propagules present? Location: Acid grassland at 230 m a.s.l., eastern Scotland, UK. Methods : Plots were surface disturbed or had their soil profile inverted at monthly intervals at 12 dates during a year. Seed bank and seed rain were assessed at each treatment time. The effect of disturbance intensity and timing on the regenerating vegetation was assessed. Results: Removing the seed bank significantly slowed regeneration, as it contributed 43 % of developing cover after one year where it was present. At an individual seed level, seed in the seed rain had a much higher likelihood of contributing to the regenerating vegetation than a seed in the seed bank. Some species showed a reliance on the seed bank for regeneration, and hence there was a significant difference in the vegetation that developed between plots with the seed bank intact and those where it was removed. Winter disturbed plots (little seed rain) had slower rates of re‐vegetation than summer disturbed plots. Timing had little effect on species composition, though a significantly higher cover of perennial forb species developed on the winter disturbed plots. Conclusion: Removing the contribution of the seed bank had a greater effect on the composition of regenerating vegetation than the effect of seasonal variation on the seed rain.     

Response of two oligohaline marsh communities to lethal and nonlethal disturbance

Severity is recognized as an important attribute of disturbance in many plant communities. However, the effects of disturbances of different severity on patterns of regeneration in oligohaline marsh vegetation have not been experimentally examined. In these communities, a critical difference in the effects of disturbance severity may be whether the vegetation dies as a result of the disturbance or is merely damaged and hence capable of resprouting. We described the regeneration of vegetation in two Louisiana marsh community types, one dominated by Sagittaria lancifolia L. and the other by Spartina patens (Ait.) Muhl., following three levels of disturbance: no disturbance, a nonlethal disturbance, and a lethal disturbance. In the nonlethal disturbance, aboveground vegetation was clipped to simulate common disturbances such as fire and herbivory that remove aboveground vegetation but leave rhizomes intact. In the lethal disturbance vegetation was killed using herbicide to simulate disturbances causing plant mortality such as wrack deposition, sedimentation, scouring, and flooding following fire or herbivory. Regeneration was assessed over a 2-year period by measuring plant species richness, relative abundance, relative dominance, cover, and final biomass. To elucidate mechanisms for observed responses of vegetation, the species composition of the seed bank, light penetration, water level, salinity, and soil redox potential were evaluated. Despite differences in the structure of undisturbed vegetation in the two community types, they exhibited the same overall pattern of regeneration. Following nonlethal disturbance, the dominant species resprouted and quickly reestablished the structure of the vegetation. In contrast, recolonization following lethal disturbance occurred primarily via seedling recruitment, which resulted in marked shifts in community structure that persisted throughout the study. While the two communities responded similarly overall to disturbance, the response of individual species was not uniform; abundance, dominance, biomass, or cover increased for some species but decreased for others in response to disturbance. Seed bank species occurred in the vegetation following lethal disturbance in the Spartina community and in both disturbed and undisturbed plots in the Sagittaria community, indicating that the seed bank is a source of propagules for regeneration and maintenance of oligohaline marshes. Of the environmental variables measured, light level was most closely related to the effect of disturbance severity on community structure. Our results suggest that lethal and nonlethal disturbances have differential effects on regeneration of vegetation that can create pattern in oligohaline marshes communities. Received: 29 September 1997 / Accepted: 12 May 1998     

三峡库区消落带水淹初期土壤种子库月份动态

干湿交替往往导致土壤种子库产生剧烈变化,严重影响着地上植被的自然恢复和生长。采用垂直空间梯度代替水淹时间梯度,研究了消落带水淹初期土壤种子库月份动态变化。结果表明：消落带内不同土地环境类型土壤种子库具有不同的月份变化趋势,分别是采伐迹地土壤种子库密度逐渐增加,弃耕地则逐渐降低。随着垂直空间梯度的升高,5、7、9月份消落带土壤种子库密度都表现为逐渐增加,说明水淹时间越长越不利于土壤种子的储存。不同月份共同出现的优势物种包括地果、黄鹌菜、具芒碎米莎草、毛马唐、婆婆纳、匍茎通泉草、鼠曲草、水田碎米荠、雾水葛、细穗腹水草、香附子、一年蓬、知风草、苎麻,这些物种构成了消落带土壤种子库的主要组成部分。各月份土壤种子库的物种生活型非常接近,主要以一年生草本和多年生草本为主,相对缺少乔灌木,且生活型百分比例在不同海拔梯度存在差异。结合土壤种子库物种水分生态功能,将其划分为湿生植物、湿中生植物、中生植物、旱中生植物,发现其主要以中生植物为主,高海拔区段旱中生植物比例相对较高,低海拔区段湿中生植物比例相对较高,而湿生植物比例则变化较小,水淹导致种子库植物呈现向中生和湿中生植物类型的变化趋势。消落带土壤种子库植物具有较高的生物多样性,但不同海拔区段植物变异性较高,没有形成绝对优势的物种,植被群落明显处于演替早期。研究认为,消落带水淹初期,土壤生境、物种组成以及种子储量都发生着剧烈的变化,仅依靠土壤种子库进行消落带植被恢复与重建是不现实的,必须要给予更多的人为干预。     

Effects of disturbance intensity on seasonal dynamics of alpine meadow soil seed banks on the Tibetan Plateau

Background and Aims

Information on soil seed bank processes is crucial for understanding vegetation dynamics. Despite the documented importance of soil seed banks in many ecosystems, their role is not fully understood in some sensitive habitats, such as the alpine meadows of the Tibetan Plateau.

Methods

We studied the seasonal dynamics of the germinable soil seed bank under four disturbance intensities in an alpine meadow on the Tibetan Plateau as well as seed size distribution relative to disturbance intensity. Composition of the seed bank was compared with that of the standing vegetation.

Results

Density of buried seeds increased with disturbance intensity, but species richness and species diversity decreased. Seed density and species richness of the seed bank varied seasonally in all layers (0–2, 2–7, 7–12 cm) and the whole (0–12 cm). The species composition of seed bank was not significantly influenced by season. There was no trend in seed size distribution as disturbance increased. Seasonal seed bank turnover rates increased with increase in disturbance. The result of the NMDS showed that species composition of seed bank and vegetation exhibited a fairly uniform pattern in each season.

Conclusions

Although as a whole the species composition of the vegetation and seed bank showed a relatively low degree of similarity in each season, similarity was highest in the most disturbed habitat. There was no alteration in species composition of seed bank regardless of disturbance intensity, but seed density decreased as disturbance increased. Disturbances in alpine plant communities might increase persistence of regeneration niches. Regeneration from the seed bank together with vegetative reproduction contributed to aboveground vegetation in highly disturbed habitats. Clonal species played an important role in regeneration of vegetation in slightly disturbed areas, where there was little contribution of ruderals from soil seed banks.     

On ecology of post-fire soil fungi: assessing impact of disturbance using species-abundance models as measure of community organization

Mediterranean ecosystems are among those most significantly modified by fires. Such fires lead to evident disturbance of the above- and below-ground ecosystem components, at various spatial and temporal scales, including soil microfungi. The ecological parameters used to measure the effects of disturbance on soil fungal communities include species-abundance distribution patterns, which reflect changes in the relationships between species numbers and their relative abundance, and serve as a critical measure of community organization. Species-abundance distribution patterns were used to assess the disturbance impact of experimental fires on soil fungal communities in Mediterranean maquis (southern Italy) in the short- to mid-term. The trend in the distribution patterns of heat-stimulated and xerotolerant soil fungi over time, their varying responses to low- and high-intensity fire, the efficiency of the soil germplasm bank, and the pivotal role of Neosartorya spp. in post-fire community structure in Mediterranean burned soils may all be used as tools to accurately assess the effects of fire on soil mycobiota.     

Long‐term farming systems and last crop sown shape the species and functional composition of the arable weed seed bank

Questions: The assembly of arable weed communities is the result of local filtering by agricultural management and crop competition. Therefore, soil seed banks can reflect the effects of long‐term cumulative field management and crop sequences on weed communities. Moreover, soil seed banks provide strong estimates of future weed problems but also of potential arable plant diversity and associated ecological functions. For this, we evaluated the effects of different long‐term farming systems under the same crop rotation sequence on the abundance, diversity and community assembly of weed seed bank, as well as on the functional diversity and composition. Location: DOK (biodynamic [D], bioorganic [O], conventional [K]) long‐term trial, Therwil, Switzerland. Methods: The effects of long‐term contrasted farming systems (i.e., biodynamic, organic, conventional, mineral and unfertilised systems) and last crop sown (i.e., wheat and maize) were evaluated on different indicators of species and functional diversity and composition of the weed soil seed bank. Results: The results showed significant influences of 40 years of contrasted farming systems on the diversity and composition of the seed bank, with higher diversities being found in unfertilised and organic farming systems, but also higher abundances than those found under conventional systems. Organic farming also allowed higher functional richness, dispersion and redundancy. Different farming systems triggered shifts in species and functional assemblies. Conclusions: The results highlight the importance of organic management for the maintenance of a diverse arable plant community and its functions. However, such results emphasise the need for appropriate yearly management to reduce the abundance of settled weediness and prevent affecting crop production. The farm management filtered community composition based on functional traits. Although the soil seed bank buffers the long‐term farming and crop sequence, the last crop sown and, thus, the yearly management were important determinants of seed bank composition.     

Disturbance persistence in managed grasslands: shifts in aboveground community structure and the weed seed bank

The length of time and form in which disturbances persist in systems depends on the intensity and frequency of disturbance and on the abilities of resident species to recover from such events. In grazed grasslands, trampling by large mammalian herbivores can periodically facilitate weed establishment by exposing patches of bare ground but whether an intense soil disturbance event results in a temporary increase in weed abundance or a persistent weed problem remains unclear. In May 2002, cattle trampling following heavy rain caused severe damage to nine-month old, rotationally grazed, cool-season pastures (Midwest USA). In September 2002, we compared the aboveground composition of paddocks (i.e., fenced pasture sections) that were heavily disturbed to those that received no damage. Relative to undisturbed paddocks, forage species relative cover was 17% lower in disturbed paddocks, and weed species and bare ground relative cover was 61% and 100% higher, respectively. By September 2004, paddock types did not differ in all aboveground community components. However, the abundance and species richness of weed seeds in the soil seed bank averaged respectively 82% and 30% higher in disturbed paddocks between 2003 and 2004. These findings indicate that a spatially extensive, intense soil disturbance event may soon become undetectable in components of aboveground pasture structure but can persist as an augmented weed seed bank. Because of high weed seed bank longevity, disturbances to formerly disturbed pastures would likely result in higher weed recruitment, with more species represented, than in those which lack previous disturbance. Disturbance history may thus be a useful predictor of weed community composition following subsequent disturbance. Based on empirical data supporting this proposition, we recommend that grassland managers explicitly incorporate disturbance history into dynamic management planning and do not rely exclusively on aboveground characters to evaluate the invasion status or colonization potential of an area by undesirable plants. We emphasize that the ecological legacies of past soil disturbance events cannot only influence the contemporary patterns and processes of grasslands, but importantly, affect their compositional trajectories following subsequent perturbation.     

Soil warming increases plant species richness but decreases germination from the alpine soil seed bank

Global warming is occurring more rapidly above the treeline than at lower elevations and alpine areas are predicted to experience above average warming in the future. Temperature is a primary factor in stimulating seed germination and regulating changes in seed dormancy status. Thus, plant regeneration from seed will be crucial to the persistence, migration and post disturbance recruitment of alpine plants in future climates. Here, we present the first assessment of the impact of soil warming on germination from the persistent alpine soil seed bank. Contrary to expectations, soil warming lead to reduced overall germination from the soil seed bank. However, germination response to soil temperature was species specific such that total species richness actually increased by nine with soil warming. We further explored the system by assessing the prevalence of seed dormancy and germination response to soil disturbance, the frequency of which is predicted to increase under climate change. Seeds of a significant proportion of species demonstrated physiological dormancy mechanisms and germination of several species appeared to be intrinsically linked to soil disturbance. In addition, we found no evidence of subalpine species and little evidence of exotic weed species in the soil, suggesting that the soil seed bank will not facilitate their invasion of the alpine zone. In conclusion, changes in recruitment via the alpine soil seed bank can be expected under climate change, as a result of altered dormancy alleviation and germination cues. Furthermore, the alpine soil seed bank, and the species richness therein, has the potential to help maintain local species diversity, support species range shift and moderate species dominance. Implications for alpine management and areas for further study are also discussed.     

Plant life history stage and nurse age change the development of ecological networks in an arid ecosystem

Understanding how ecological networks are organised over the course of an organism's lifetime is crucial for predicting the dynamics of interacting populations and communities across temporal scales. However, most studies so far considered only one life history stage at a time, such as adult, when studying networks of interacting species. Therefore, knowledge about how multiple life history stages affect the development and stability of plant–plant association networks is lacking. We measured the understory adult plant community and the soil seed bank across a plant age gradient of the nurse shrub Retama sphaerocarpa in an arid ecosystem in Spain. Using a multilayer network approach, we built adult understory–nurse and seed bank–nurse networks and analysed how network nestedness, species’ role, and species specificity varied between them and with nurse plant age. We found that seed bank and adult understory networks changed depending on nurse plant age in two different ways. With increasing nurse plant age, adult understory networks became significantly more nested than seed bank networks. The nested architecture of seed bank networks was therefore a poor predictor of adult understory network nestedness. The contribution and specificity of species to network nestedness increased with increasing nurse plant age more in the adult understory than in seed bank networks, despite high species turnover. Our data show that life history and ontogeny affect the development of plant–plant association networks. Niche construction and environmental filtering along nurse ontogeny seem pivotal mechanisms structuring adult understory networks while the assembly of seed bank networks seems rather stochastic. We highlight the importance of mature plant communities for maintaining rare species populations and supporting the stability of ecological communities through time.     

The role of soil seed banks in the restoration of dry acidic dune grassland after burning of Ulex europaeus scrub

Question: Can the seed bank play a significant role in the restoration of plant communities of dry acidic dune grassland where fire has destroyed Ulex europaeus scrub? Location: Northern French Atlantic coast. Methods: One year after the fire, the seed bank and vegetation were sampled in 1 m × 1 m plots along three transects from the oldest scrub vegetation towards the grassland. Differences in species richness, seed density and contribution of ecological groups in the seed bank and vegetation along the transects were analysed. Results: Seed density and species richness in the seed bank decreased significantly from the grassland towards the centre of the scrub vegetation; 50% of the seed bank consisted of core species of the target plant community, such as Carex arenaria, Aira praecox, Rumex acetosella and Agrostis capillaris. Seeds of these species were also found in the deeper soil layers beneath the oldest scrub vegetation, indicating that they can be considered to be long‐term persistent. Beneath the youngest scrub vegetation, seeds of rare satellite target species also occurred. However, no target species were established on the burned site after one year, resulting in a large discrepancy between seed bank and vegetation. Conclusions: Although the seeds present in the soil indicate that restoration of the acidic grassland based on the seed bank is possible, additional management actions such as mowing and soil disturbance may be necessary to restrict resprouting of Ulex and to stimulate the germination of seeds of target species in the deeper soil layers.     

Seed arrival, ecological filters, and plant species richness: a meta-analysis

Theoretical models predict that effects of dispersal on local biodiversity are influenced by the size and composition of the species pool, as well as ecological filters that limit local species membership. We tested these predictions by conducting a meta-analysis of 28 studies encompassing 62 experiments examining effects of propagule supply (seed arrival) on plant species richness under contrasting intensities of ecological filters (owing to disturbance and resource availability). Seed arrival increased local species richness in a wide range of communities (forest, grassland, montane, savanna, wetland), resulting in a positive mean effect size across experiments. Mean effect size was 70% higher in disturbed relative to undisturbed communities, suggesting that disturbance increases recruitment opportunities for immigrating species. In contrast, effect size was not significantly influenced by nutrient or water availability. Among seed-addition experiments, effect size was positively correlated with species and functional diversity within the pool of added seeds (species evenness and seed-size diversity), primarily in disturbed communities. Our analysis provides experimental support for the general hypothesis that species pools and local environmental heterogeneity interactively structure plant communities. We highlight empirical gaps that can be addressed by future experiments and discuss implications for community assembly, species coexistence, and the maintenance of biodiversity.     

Accumulation of buried seeds and establishment of ruderal therophytic communities in disturbed habitat,central Japan

The establishment and maintenance mechanisms of pioneer communities were investigated in ruderal habitats under two disturbance regimes, frequent and infrequent cutting sites. In the infrequent cutting sites, large perennials dominated through the year and inhibited the invasion of annuals, and the perennial community succeeded to forest stage if the cutting was stopped. In the frequent cutting sites, therophytic communities of winter and summer annuals alternated by season. Fresh seeds of both winter and summer annuals are dormant, but they have different germination times and thus can share the same sites in different seasons. Wind-dispersed biennials and large perennials have nondormant seeds and easily invade the sites; however, they are unable to mature to reproductive phase due to recurrent cuttings. The therophytic species, which can complete their life-cycle in a period between cuttings, accumulate seeds in the soils and are maintained by these buried seeds (seed bank annuals) during recurrent disturbances. The seed bank is compensation for the dispersal inefficiency of seed bank annuals. After abandonment of the frequent cutting sites, the buried seeds of seed bank annuals germinate and become the first-year pioneers. Thus, seed bank annuals are not invasive colonizers but are the remnants of the ruderal weed communities before abandonment.     

漫溢干扰过程中微地形对幼苗定居的影响

通过对比漫溢干扰后边坡、坡中和坡底幼苗的物种组成、密度、幼苗周边的环境条件及土壤种子库的差异来分析漫溢干扰过程中,微地形对幼苗定居的影响.结果表明:(1)微地形对幼苗物种组成和生物多样性指数的影响很大.漫溢干扰后,位于边坡、坡中和坡底幼苗的Simpson多样性指数和Mcintosh均匀度指数在0.01的水平上差异极显著,Margalef丰富度指数在0.05的水平上差异显著.(2)微地形对土壤盐分含量影响很大.方差分析的结果表明,除HCO3-和K+外,边坡、坡中和坡底的pH值、C023-、SO24-、Ca2+和Mg2+各指标在0.01的水平上差异极显著;全盐、总盐、Cl-和Na+各指标均在0.05的水平上差异显著.(3)微地形通过改变幼苗定居的环境条件,进而影响幼苗库的物种组成、密度和生物多样性指数.微地形中,幼苗库的各种生态指标均小于对应的土壤种子库的生态指标,从坡底到坡中到边坡,除单位面积上的植被密度外,土壤种子库和幼苗库的各项生态指标的变化趋势均不同.     

Coexistence of plant species in a biodiversity hotspot is stabilized by competition but not by seed predation

Understanding the mechanisms of species coexistence is a key task for ecology. Recent theory predicts that both competition and predation (which causes apparent competition among prey) can either promote or limit species coexistence. Both mechanisms cause negative interactions between individuals, and each mechanism promotes stable coexistence if it causes negative interactions to be stronger between conspecifics than between heterospecifics. However, the relative importance of competition and predation for coexistence in natural communities is poorly known. Here, we study how competition and apparent competition via pre‐dispersal seed predators affect the long‐term fecundity of Protea shrubs in the fire‐prone Fynbos biome (South Africa). These shrubs store all viable seeds produced since the last fire in fire‐proof cones. Competitive effects on cone number and pre‐dispersal seed predation reduce their fecundity and can thus limit recruitment after the next fire. In 27 communities comprising 49 990 shrubs of 22 Protea species, we measured cone number and per‐cone seed predation rate of 2154 and 1755 focal individuals, respectively. Neighbourhood analyses related these measures to individual‐based community maps. We found that conspecific neighbours had stronger competitive effects on cone number than heterospecific neighbours. In contrast, apparent competition via seed predators was comparable between conspecifics and heterospecifics. This indicates that competition stabilizes coexistence of Protea species, whereas pre‐dispersal seed predation does not. Larger neighbours had stronger competitive effects and neighbours with large seed crops exerted stronger apparent competition. For 97% of the focal plants, competition reduced fecundity more than apparent competition. Our results show that even in communities of closely related and ecologically similar species, intraspecific competition can be stronger than interspecific competition. On the other hand, apparent competition through seed predators need not have such a stabilizing effect. These findings illustrate the potential of ‘community demography’, the demographic study of multiple interacting species, for understanding plant coexistence.