荒漠草原土壤种子库对草地管理方式的响应

以宁夏中部不同封育年限(封育1a、3a、5a和7a)、不同放牧方式(中等强度自由放牧、中等强度4区轮牧)、补播改良(2006年补播沙打旺Astragalus adsurgens和紫花苜蓿Alfalfa stiva)及未封育等管理方式下的荒漠草原为研究对象,采用空间梯度代替时间梯度法,于2012年3月下旬用样线法采集土壤样品,采用温室萌发法,统计土壤种子库物种组成和种子数,研究了不同草地管理方式下的种子库与地上植被相似性、多样性等特征。结果表明:1)宁夏荒漠草原土壤种子库中共出现14种植物,分属6科14属,除牛枝子(Lespedeza potanimii)为半灌木外其余均为草本植物。多年生植物占土壤种子库植物总数的65.3%,一年生植物占土壤种子库植物总数的35.7%;多年生植物种子数量随封育年限增加而表现为下降趋势,封育1a和未封育草地最多;2)不同管理方式下土壤种子库和地上植被的共有物种数为2—6种,封育时间延长导致土壤种子库物种与地上植被相似性下降,土壤种子库和地上植被共有物种数有减少趋势,自由放牧降低了土壤种子库和地上植被的共有物种数,补播恢复的草地土壤种子库中未出现补播物种;3)种子库物种多样性与管理方式无明显的相关性;种子库物种多样性指数4区轮牧草地最高,封育7a草地最低,物种丰富度指数封育1a草地最高,均匀度指数自由放牧草地最高,补播草地最低。若荒漠草原仅依靠土壤种子库恢复自然植被,则不建议封育时间过久,需间以适当的干扰(放牧);补播外来种未对荒漠草原土壤种子库产生贡献,反而对乡土种产生了一定限制。     

Plant and soil seed bank diversity across a range of ages of Eucalyptus grandis plantations afforested on arable lands

Aims

Plantation forests are often assumed to have reduced biodiversity relative to unmanaged forests. However, existing knowledge is based on studies of rotation-aged tree crops. We investigated how Eucalyptus afforestation of agricultural land affected plant species composition and biodiversity across a range of plantation ages (1–10 years). We also studied whether the soil seed bank could contribute to regeneration of existing vegetation in such plantations.

Methods

We used a chronosequence approach to evaluate plant and seed species composition and diversity in forests and soil seed banks. We also quantified the similarity of seed banks and aboveground vegetation within plantation sites of a given age. Plantation sites were also compared to a nearby, mature pine forest.

Results

Total plant species number, density and diversity in Eucalyptus grandis plantations increased for the first 3 years plantation establishment, then stabilized or decreased for the next 1–2 years and then increase significantly over the following years. Species number and density in soil seed bank increased significantly with plantation age only after an initial 6-year decrease. Shannon–Wiener index of total species diversity did not significantly differ with plantation age. The understory vegetation and soil seed bank were dominated by pioneer species in the first 3 years, but intermediate-successional and shade-tolerant species gradually invaded as plantations developed further. After 7 years, E. grandis plantation understories were composed of mainly shade-tolerant species. Nevertheless, the diversity of the diversity of intermediate-successional in soil seed banks were higher than that of shade-tolerant species in soil seed banks at this age range (7–10 year). Among species successfully germinated from soil seed banks, 48 % were not found in the aboveground plant community. Similarities between the species in the soil seed bank and the aboveground vegetation were low for both plantation and control forests and did not significantly change with plantation ages.

Conclusions

E. grandis likely produces a changing microclimate during plantation development, which in turn drives composition and diversity dynamics in understory vegetation and soil seed banks after the afforestation of agricultural land. The first 4 years after plantation establishment is associated with lower plant and soil seed bank diversity, meriting a greater focus on biodiversity stabilization and possibly longer rotation periods.     

Comparison of aboveground vegetation and soil seed bank composition at sites of different grazing intensity around a savanna-woodland watering point in West Africa

Grazing removes a plant’s aboveground vegetative and reproductive tissues and can modify the soil seed bank, potentially impacting the restoration of preferred species. Knowledge about aboveground vegetation and species composition of soil seed bank and the processes that contribute to vegetation recovery on and surrounding watering points subjected to grazing is lacking. Successful restoration strategies hinge on addressing these knowledge gaps. We assessed the effects of livestock grazing on aboveground vegetation and soil seed bank characteristics along a river bank and surrounding areas subject to different grazing intensities and draw implications for restoration. Plots (50?×?50 m) were established along five transects representing differing levels of grazing intensity. Soil samples were taken from three layers within each plot to determine soil properties and species composition of soil seed bank using the seedling emergence method. Heavy grazing resulted in the disappearance of perennial grasses, a reduction in species diversity and a decrease in soil nutrients with increased soil depth. Overall, the similarity between the extant aboveground vegetation and flora within the soil seed bank was low. The soil seed bank was dominated by herbaceous species and two woody species, suggesting that many woody species are not accumulating in the soil. With increasing soil depth, the seed density and richness declined. Canonical correspondence analyses (CCAs) showed that emerged seedlings from the soil seed bank were significantly influenced by soil carbon, organic matter, total nitrogen, total potassium and soil cation exchange capacity. This finding suggests that current grazing practices have a negative impact on the vegetation surrounding watering points; hence there is a need for improved grazing management strategies and vegetation restoration in these areas. The soil seed bank alone cannot restore degraded river banks; active transfer of propagules from adjacent undisturbed forest areas is essential.     

Soil seed bank and floristic diversity in a forest‐grassland mosaic in southern Spain

Abstract. Soil seed bank and floristic diversity were studied in a forest of Quercus suber, a forest of Quercus canariensis and a grassland, forming a vegetation mosaic in Los Alcornocales Natural Park, southern Spain. The soil seed bank was estimated by the germination technique. In each community patch, diversity, woody species cover and herbaceous species frequency was measured. Three biodiversity components – species richness, endemism and taxonomic singularity – were considered in the vegetation and the seed bank. Forest patches had a soil seed bank of ca. 11 200–14 100 seed.m^?2 and their composition had low resemblance to (epigeal) vegetation. The grassland patch had a more dense seed bank (ca. 31 800 seed.m^?2) and a higher index of similarity with vegetation, compared with the forests nearby. The complete forest diversity was 71–78 species on 0.1 ha, including 12–15 species found only in the seed bank; the grassland species richness was higher (113 species on 0.1 ha). We discuss the role of soil seed banks in the vegetation dynamics and in the complete plant biodiversity of the mosaic landscape studied.     

Short-term effects on sheep pastureland due to grazing abandonment in a Western Mediterranean island ecosystem: A multidisciplinary approach

Changes of pasture communities consequent to management practices resulting from land abandonment considerably affect the structure and function of the ecosystem. This study analyses the consequences of grazing abandonment in terms of plant and soil microbial diversity and fertility, on a Mediterranean upland sheep pasture, over a short period (five years). Grazing was experimentally excluded by fencing ten 10×10 m permanent plots within an area that had supported grazing until 2000, by 0.23 sheep ha^?1. Plant and soil microbial communities and physicochemical parameters were monitored within the fenced and unfenced control plots, during three sampling times from 2000 (before the fencing) to 2005. Grazing cessation notably altered the floral composition, with an average dissimilarity of 96.7% between the vegetation communities, over five years. No significant change occurred in the control plots that were grazed throughout the sampling period. This work highlighted that, over a short term, the structural change in the specific plant composition affected only the grass species, confirming that grazing favours the small-sized species over the annual species. Further, it was evident that species groups of conservational and phytogeographic interest, like the endemic and Mediterranean-Atlantic species, tended to disappear with pasture abandonment and were substituted by more widespread species throughout the Mediterranean or even the world. Pasture abandonment was accompanied by an increase of soil pH and a decrease in soil organic matter and soil nitrogen. The microbial parameters recorded at three different sampling times revealed a substantial effect of the plant community, or the time of grazing abandonment, on soil microbial abundance and diversity. Considerable importance is given to the consequences of pasture abandonment on the conservation of plant and microbial diversity and on soil fertility.     

Influence of Grazing on Soil Seed Banks Determines the Restoration Potential of Aboveground Vegetation in a Semi‐arid Savanna of Ethiopia

Species composition, number of emerging seedlings, species diversity and functional group of the soil seed banks, and the influence of grazing on the similarity between the soil seed banks and aboveground vegetation, were studied in 2008 and 2009 in a semi‐arid savanna of Ethiopia. We tested whether the availability of persistent seeds in the soil could drive the transition from a degraded system under heavy grazing to healthy vegetation with ample perennial grasses. A total of 77 species emerged from the soil seed bank samples: 21 annual grasses, 12 perennial grasses, 4 herbaceous legumes, 39 forbs, and 1 woody species. Perennial grass species dominated the lightly grazed sites, whereas the heavily grazed sites were dominated by annual forbs. Heavy grazing reduced the number of seeds that can germinate in the seed bank. Species richness in the seed bank was, however, not affected by grazing. With increasing soil depth, the seed density and its species richness declined. There was a higher similarity in species composition between the soil seed bank and aboveground vegetation at the lightly grazed sites compared with the heavily grazed sites. The mean similarity between the seed banks and aboveground vegetation was relatively low, indicating the effect of heavy grazing. Moreover, seeds of perennial grasses were less abundant in the soil seed banks under heavy grazing. We concluded that restoration of grass and woody species from the soil seed banks in the heavily grazed areas could not be successful in semi‐arid savannas of Ethiopia.     

轮牧时间对荒漠草原土壤种子库特性的影响

探讨季节性放牧下土壤种子库的特性对荒漠草原合理利用具有重要作用。以宁夏荒漠草原为对象,通过封育(FY)、传统时间轮牧(FG)、延迟开始轮牧(YG)、提前结束轮牧(TG)、延迟开始并提前结束轮牧(YT)和自由放牧(ZY)处理,研究了0～5、5～10、10～15 cm土层土壤种子库物种组成、种子库密度、垂直分布、多样性及其与地上植被的关系,探讨了种子库与土壤环境因子的关系。结果表明: 研究区土壤种子库物种组成共有9个科21个种。土壤种子库物种组成以TG种子库种类最多,达13种,而FY和ZY土壤种子库种类最少,各8种。ZY土壤种子库密度显著高于FY、YG、TG、YT;FY的土壤种子库中多年生植物种子数量最多,达32.0%;ZY种子库中多年生植物种子数最少,仅为12.4%,主要以有性繁殖的一年生杂类草种子为主;土壤种子库主要集中于0～5 cm表层土壤,随着土层深度的增加,土壤种子库大小降低。土壤种子库的优势度和多样性均以FY最高,优势度、多样性和均匀度均以ZY最低。土壤种子库与地上植被的相似性总体较低,相比之下FG及ZY种子库与地上植被相似性较高,FY最低。土壤水分、有机质、全氮、全磷、土壤容重是影响土壤种子库的主要土壤因子。与传统全年自由放牧相比,季节性四区轮牧能使荒漠草原土壤种子库多年生植物种类、密度以及种子库物种丰富度、多样性和均匀度增加,虽作用不及长期封育草地,但对荒漠草原的利用与保护具有重要意义。     

Resilience,persistence and relationship to standing vegetation in soil seed banks of semi‐arid Australian old fields

Questions: Do soil seed banks of semi‐arid grasslands reassemble after abandonment from cultivation? Do seeds of native and exotic species persist in the soil? Does time since abandonment affect compositional similarity between the vegetation and seed bank? Does the seed bank contribute to resilience in the vegetation? Location: Native grasslands in northern Victoria, Australia. Methods: Seed bank sampling was conducted in spring and autumn over 3 yrs, across a 100‐yr chronosequence. Species richness, composition and germinant density were determined using the seedling emergence method. Seed persistence was assessed by comparing seed densities in spring and autumn. Seed bank composition was compared with the vegetation. Results: The spring seed bank was dominated at all stages by sedges and rushes; hence, native species richness and seed density were largely unaffected by abandonment. In autumn, grassland species contributed more to the seed bank, but richness was reduced after abandonment and showed little recovery, although seed density partially recovered. Seed bank composition showed some recovery in both seasons. Most species had low persistence in the soil. Compositional similarity between the vegetation and seed bank was greater in old fields than uncultivated grasslands in spring, but not autumn. Conclusions: Resilience varied among seed bank parameters and seed banks had low functional importance. Patterns in the seed bank followed, rather than caused, those in the vegetation. Thus, vegetation recovery cannot rely on the seed bank and persistent seeds were not the key mechanism of resilience in the vegetation.     

黄土高原半干旱区退化草地恢复与利用过程研究

草地退化表现为土壤和植被遭到彻底破坏,草地演替过程受到强烈抑制.实验采用长期(30年)封禁措施,定位监测退化草地从次生半裸地演变为近似原生植被(进展演替)的变化过程.结果表明,随着封禁时间的变化,退化草地恢复演替经历了4个阶段,群落盖度、植株密度、物种丰富度和多样性指数、地上生物量和地下生物量在草地群落恢复过程中逐渐增加,其特征变化出现的峰值均在封禁第20年(地下生物量峰值在第15年),其中地上生物量最高达520.5 g/m2;直到封禁的第20~25年,以本氏针茅为建群种的草原群落衰败退化现象明显,而大针茅种群密度剧增;在封禁的第26年以上以大针茅为优势的群落生长较为稳定,从目前群落演替进程看,大针茅有替代本氏针茅的趋势.另外,在草原沟道两侧以斑块状聚集分布有中旱生灌木,群落的演替进入了一个新的阶段.随着封禁时间的延续,退化草地从自然封禁恢复的0~26年,通过侵入-竞争-扩散-定居的几个演替阶段,目前形成以大针茅为建群种相对稳定的"亚顶级".虽然草地生物量有一定下降,但草地质量提高,物种多样性丰富,促进草地的进展演替.草地植物群落主要由禾本科、豆科和菊科组成;多年生植物、C3和旱生物种可以作为草地演替过程和植被恢复的指示物种.长期封育对草地物种更新和生态系统稳定性有负面影响,因此,合理的封育时间是草地生态恢复中非常重要的一个因素.本研究提出,在黄土区退化草地封育10~15年后可以开始进行合理的利用,例如通过两年一次刈割和轻度放牧(2只羊/hm2).本研究可为干旱区、半干旱区相似的退化草地恢复提供理论依据.     

Permanent seed banks in chalk grassland under various management regimes: their role in the restoration of species-rich plant communities

A survey was carried out on four stands of a chalk grassland (open, intensively grazed, tall and scrub) in order to monitor the permanent soil seed bank under various management regimes responsible for the decline of the ecological interest of this ecosystem. There was no real similarity between the seed banks and the associated vegetation, except in the intensively grazed stand. When vegetation changes, seed banks appear to be quite stable and are mostly dominated by three species (Centaurium erythraea, Origanum vulgare, Hypericum perforatum). The seed bank seems to be quite useless in order to restore a species-rich community after both grazing intensification or abandonment. Because recolonization processes are the critical factor, the management of species-rich grassland has to be designed with a view to preserving a given plant community as well as to improving dispersion processes from a source to target areas.     

The effects of three years of fencing enclosure on soil seed banks and the relationship with above-ground vegetation of degraded alpine grasslands of the Tibetan plateau

Background and Aims

Fenced enclosures have become an important method for re-establishing degraded grassland on the Tibetan plateau, and examination of soil seed banks may provide useful insights to understanding the effects and mechanisms of fencing enclosure on the restoration.

Methods

An investigation was conducted into the effects of enclosure for 3 years on the soil seed banks of degraded natural and sown grasslands at eight study sites. Species composition, soil seed bank density and the relationships with above-ground vegetation were analysed based on 4800 soil core samples and counting of seeds extracted from soil samples.

Results

After 3 years of fencing enclosure, soil seed banks differed between the different communities across the study sites. Species numbers and seed density in soil seed banks decreased from natural grassland to sown grassland, with most seeds occurring in the upper 5 cm soil layer. In these alpine grasslands, relatively few species produced high numbers of seeds, although their occurrence across the eight study sites was variable. Total vegetation cover increased with enclosure due to the colonization capacity of the vegetation rather than soil seed banks.

Conclusions

This study provided evidence that soil seed banks do not play an important role in the restoration of degraded alpine grassland when using fencing enclosures. Further studies conducted over longer periods are needed to address this subject.     

Relationship of plant diversity with litter and soil available nitrogen in an alpine meadow under a 9-year grazing exclusion

Grazing exclusion is widely used globally to restore degraded grasslands. Plant diversity has important impacts on grassland ecosystem functions, including grassland productivity and carbon storage. In this study, we selected a Kobresia meadow on the Qinghai–Tibetan Plateau to investigate how grazing exclusion affects plant diversity. Inorganic nitrogen (NH₄ ⁺ and NO₃ ^?) was also measured because its availability impacts plant growth. We found that plant diversity in the meadow was significantly lower under grazing exclusion (fenced meadow) for 9 years compared with moderate grazing. Accumulated litter was significantly higher under grazing exclusion (386.41 g m^?2) compared with grazing (58.77 g m^?2). Soil inorganic nitrogen at 0–5 cm depth was significantly higher under grazing exclusion (13.60 × 10^?2 g kg^?1) than under grazing (9.40 × 10^?2 g kg^?1). The composition of the four functional groups (grasses, sedges, legumes, and forbs) might alter in response to significant changes in the amount of litter and soil available nitrogen content under grazing exclusion compared with grazing. However, the enhanced soil available nitrogen content showed weak feedbacks on plant diversity. In conclusion, light limitation induced by increased amounts of litter may be the main factor causing decreased plant diversity in grazing-excluded meadows compared with moderately grazed meadows.     

Environmental and land use determinants of grassland patch diversity in the western and eastern Alps under agro-pastoral abandonment

Agro-pastoral decline in European mountain areas has recently caused changes to traditional landscapes with negative consequences on semi-natural grassland conservation and the associated biodiversity and ecosystem services. In the Italian Alps, grassland patches enclosed in a forest matrix are progressively disappearing. Two alpine valleys (Pesio and Pejo), having similar land-use history, were chosen as representative of management conditions of western and eastern Italian Alps, respectively. This study aims at interpreting the effect of abandonment on grassland patch plant diversity, considering land cover changes of the last 60 years, and assessing the role of ecological, topographic, management and landscape configuration on current grassland species richness. The total area of grassland patches has declined by 54 and 91 % at Pesio and at Pejo, respectively. Actual grassland patch species richness was mostly influenced by ecological factors, such as quantity of light, soil moisture and reaction, then by topographic features, especially slope, and finally by management intensity. Landscape factors exerted a slightly significant effect on plant diversity. In the two valleys, differences on management practices were detected. Even though in the western valley the conservation of several grazing activities contributed to slow down the process of patch reduction, many species-rich grasslands were generally under-grazed. Conversely, in the eastern valley, despite a denser road network, the stronger decline of grassland patch extension was linked to the hay making decline. At the same time, overuse of grassland patches near farms reduced plant species richness. As a conclusion, plant species richness was weakly related to the area of grassland patches and current and historical landscape configuration were of relatively lower importance than ecological, topographic and management factors, when evaluated at patch-level.     

科尔沁沙质草地放牧和围封条件下的土壤种子库

该文研究了科尔沁沙质草地在放牧和围封条件下土壤种子库密度、组成及其与地上植被的关系。结果表明:1)放牧草地植物种数22种,围封草地植物种数30种,围封使土壤种子库植物种数增加了36%;2)放牧草地土壤种子库密度为16 149±1 900有效种子数·m^-2, 围封草地土壤种子库密度为20 657±3 342有效种子数·m^-2,比放牧草地增加了28%。放牧和围封草地种子库组成密度均以一年生植物为主(分别占99%和98%的比例),多年生植物所占的比例很小;3)放牧草地种子库的Shannon-Wiener指数和丰富度指数分别为0.836 3和4.954 9,明显小于围封草地的0.968 2和7.226 0,表明自由放牧导致物种多样性下降;4)放牧和围封草地土壤种子库密度与地上植被密度均存在显著的相关性(p<0.001)。表明了随着土壤种子库密度的增加, 地上植被密度随之增加,放牧草地地上植被密度78%的变异可归结为土壤种子库密度的变异, 而围封草地地上植被密度58%的变异可由土壤种子库密度的变异来解释。     

Seed Bank and Vegetation Development of Sandy Grasslands After Goose Breeding

Four hypotheses were tested using long-term observations of vegetation development (12 years) and present-day seed bank data in a sandy grassland area overgrazed by domestic geese: i) Gap regeneration is crucial in maintaining species richness; thus, closed vegetation of the lower sites prevents continuous establishment of short-lived species. ii) Short-lived, early successional species comprise most of the seed banks and late successional perennials have at most sparse seed banks. iii) Composition of seed banks is more similar to pioneer vegetation than to later successional stages. iv) The similarity is higher between vegetation and seed banks in the upper-positioned plots than in the closed, lower-positioned ones. Two sites, located in the upper part of dune slopes, and another two, positioned on the lower part, were studied. In each site five 2?×?2 m permanent plots were surveyed between 1991 and 2002. Percentage cover was estimated three times a year. In the last study year, soil seed banks were sampled. Two vertical segments (0–5, 5–10 cm) were separately analyzed. The seedling emergence method was applied on concentrated samples. We found that the vegetation developed from open, annual dominated weedy assemblages to grasslands dominated by perennial graminoids. In the lower-positioned sites perennial clonal grasses (Cynodon dactylon, Poa angustifolia and P. pratensis) formed more closed vegetation, which was accompanied by lower species richness compared to the upper-positioned sites. Seed density varied between 10,300 and 40,900 seeds/m². Significantly higher seed densities were found in upper sites than in the lower ones. Annuals and short-lived perennial dicots comprised most of the seed bank. The dominant perennial graminoids also built up dense seed banks. We found a low to medium similarity between vegetation and the seed bank; similarity was the highest with the vegetation of the 1994–1998 period. In the upper sites the similarity between seed bank and the vegetation of the last studied years was also high. The vertical position had a significant effect on regeneration after overgrazing. The large cover of grasses in lower sites decreased species richness and it also decreased the seed density preventing the seed bank formation of annuals and short-lived perennials. Here, further management practices are needed to increase the species richness.     

Biodiversity of semi‐arid Mediterranean grasslands: Impact of grazing and afforestation

Question: What is the impact of grazing and/or afforestation on grassland diversity, species composition and cover parameters? Location: Semi‐arid Mediterranean grasslands of Jordan. Methods: Vegetation, litter, bare soil and rock cover were compared among four management types – free grazing and protected from grazing with three levels of tree cover. Species composition, plant cover, species richness and evenness were used to evaluate differences in vegetation among management types. Species composition differences among management types were also investigated. Results: Semi‐arid Mediterranean grasslands harbour appreciable levels of plant biodiversity. Grazing did not affect plant diversity, indicating the high resilience against and adaptation to grazing; however,grazing affected species composition and cover parameters. Afforestation seems to protect soil through higher litter cover but its impact on plant biodiversity was negative and markedly affected species composition. Conclusions: Neither protection from grazing or massive afforestation alone are sufficient for conserving biodiversity in this system. A management model is suggested where the landscape should be maintained as a mosaic of four management types: complete protection from grazing, grazing rotation, planting sparse trees in eroded areas and revegetating degraded areas using native, herbaceous and grazing tolerant species.     

The contribution of rewetting to vegetation restoration of degraded peat meadows

Question: What is the contribution of a rise in groundwater level to vegetation restoration of degraded peat meadows compared to abandonment only? Location: Abandoned peat meadows in the central part of The Netherlands. Methods: Comparison of species composition and species abundance of vegetation and seed banks of reference and rewetted peat meadows, using plant trait and seed bank analysis. Results: Vegetation of rewetted meadows shared on average only 27% of their species with the reference meadow, while this was 50% on average for species in the seed bank. Rewetted meadows had a lower total number of species and a lower number of wet grassland and fen species present in the vegetation, but had higher species richness per m², although evenness was not affected. Rewetting increased the dominance of species of fertile and near neutral habitats, but did not result in an increase of species of wet or waterlogged habitats. Re-wetted meadows were dominated by species relying mainly on vegetative reproduction and species with a low average seed longevity compared to the reference meadow. Conclusion: Rewetting was not effective as a restoration measure to increase plant species diversity or the number of wet grassland and fen species in the vegetation. If no additional restoration management is applied, the seed bank will be depleted of seeds of species of wet grassland or fen habitats, further reducing the chances of successful vegetation restoration.     

Variation in small-scale spatial heterogeneity of soil properties and vegetation with different land use in semiarid grassland ecosystem

Soil properties (i.e. soil organic carbon, SOC; soil organic nitrogen, SON; and soil C/N ratio) and vegetation in a semiarid grassland of Inner Mongolia, northern China, were studied with the method of geostatistical analysis. We examined the spatial heterogeneity of soil and plants, and possible impacts of land use on their heterogeneity and on the relationship between soil resources and plant richness. Land use affected small scale spatial heterogeneity in plants and soil. SOC, SON and C/N ratio displayed autocorrelation over a range of ～2 m under most circumstances on sites where livestock grazing had been excluded. The uncontrolled grazing site (UG, i.e. unregulated grazing by excessive livestock) displayed an increased range of spatial autocorrelation and the total amount of variability in soil nitrogen over the other land use types. Plant life forms and plant species exhibited spatial autocorrelation over a range of about 2 m on the grazing exclusion (GE) and mowed (MW) sites, while pattern of spatial autocorrelation for several less common species on the UG site were difficult to predict. Plant species richness was positively related with spatial heterogeneity of SOC, SON and C/N on both GE and MW sites, and with only SOC heterogeneity on the UG site. These suggest that spatial soil heterogeneity plays an active role in maintaining plant species richness. However, we call for caution in generalization of the control of spatial soil heterogeneity over plant richness when multiple modes of disturbances are present, as we found in this study that higher total amount of variation in soil nitrogen and C/N ratio on the over-grazed UG site did not lead to increased plant species richness, and that land use had apparent effects on the patterns of spatial heterogeneity in both vegetation and soil.     

Different responses of plants and herbivore insects to a gradient of vegetation height: an indicator of the vertebrate grazing intensity and successional age

Empirical studies have suggested that species richness of grassland insects usually decreases under grazing management. By contrast, grazing has been shown to increase the species density and richness of vascular plants, especially on productive soils. In order to test the suggested differences in response to management between plants and insects, we simultaneously studied species richness of vascular plants and their insect herbivores, butterflies and moths, in 68 semi-natural grasslands with varying grazing intensity and frequency in SW Finland. Species richness of plants and insects was for the first time related to a quantitative measure of disturbance intensity and successional age, mean vegetation height, by using generalized additive models (GAM). The effects of soil nutrients on vegetation height were accounted for by using phosphorus concentration as a productivity surrogate.
The results showed that species richness of butterflies and moths peaked in taller vegetation compared with vascular plants, corresponding to a lower disturbance intensity and increasing time since abandonment. These patterns are discussed in the light of two hypotheses, the "structural diversity hypothesis" and "dynamic equilibrium model" of Huston, both suggesting a weaker disturbance tolerance of insects compared with plants. Butterflies and moths which are specialists in their larval host-plant use (monophagous and oligophagous species) preferred lower vegetation (higher disturbances) compared with generalists (polyphagous species), as predicted by Huston's model. This difference indicates a stronger relationship with plant species richness for specialist than for generalist butterflies and moths. Our results support the application of regionally and temporally varying grazing intensities in grassland conservation management.     

Earthworms promote greater richness and abundance in the emergence of plant species across a grassland-forest ecotone

Aims Chalk grasslands are subject to vegetation dynamics that range from species-rich open grasslands to tall and encroached grasslands, and woods and forests. In grasslands, earthworms impact plant communities and ecosystem functioning through the modification of soil physical, chemical and microbiological properties, but also through their selective ingestion and vertical transportation of seeds from the soil seed bank. Laboratory experiments showed that seed–earthworm interactions are species specific, but little is known on the impact of seed–earthworm interactions in the field. The overall aim of this study was to better understand seed–earthworm interactions and their impact on the plant community. First we analyzed the composition of seedlings emerging from casts after earthworm ingestion. Then we compared seedling composition in casts to the plant composition of emerging seedlings from the soil and of the aboveground vegetation along four stages of the secondary succession of chalk grasslands.Methods Four stages of the secondary succession of a chalk grassland—from open sward to woods—were sampled in Upper Normandy, France, in February 2010. Within each successional stage (×3 replicates), we sampled the standing vegetation, soil seed bank at three soil depths (0–2, 2–5 and 5–10cm) and earthworm surface casts along transects. Soil and cast samples were water sieved before samples were spread onto trays and placed into a greenhouse. Emerging seedlings were counted and identified. Effect of successional stage and origin of samples on mean and variability of abundance and species richness of seedlings emerging from casts and soil seed banks were analyzed. Plant compositions were compared between all sample types. We used generalized mixed-effect models and a distance-based redundancy multivariate analysis.Important findings Seedling abundance was always higher in earthworm casts than in the soil seed bank and increased up to 5-fold, 4-fold and 3.5-fold, respectively, in the tall grassland, woods and encroached grassland compared to the soil surface layer. Species richness was also higher in earthworm casts than in the soil seed bank in all successional stages, with a 4-fold increase in the encroached grassland. The plant composition of the standing vegetation was more similar to that of seedlings from casts than to that of seedlings from the soil seed bank. Seedlings diversity emerging from casts in the tall and encroached grasslands tended toward the diversity found in woods. Our results indicate that earthworms may promote the emergence of seedlings. We also suggest that the loss of some plant species in the seed bank and the tall grass vegetation in intermediary successional stages modify the local conditions and prevent the further establishment of early-successional plant species.