Enhancement of late successional plants on ex-arable land by soil inoculations

Restoration of species-rich grasslands on ex-arable land can help the conservation of biodiversity but faces three big challenges: absence of target plant propagules, high residual soil fertility and restoration of soil communities. Seed additions and top soil removal can solve some of these constraints, but restoring beneficial biotic soil conditions remains a challenge. Here we test the hypotheses that inoculation of soil from late secondary succession grasslands in arable receptor soil enhances performance of late successional plants, especially after top soil removal but pending on the added dose. To test this we grew mixtures of late successional plants in arable top (organic) soil or in underlying mineral soil mixed with donor soil in small or large proportions. Donor soils were collected from different grasslands that had been under restoration for 5 to 41 years, or from semi-natural grassland that has not been used intensively. Donor soil addition, especially when collected from older restoration sites, increased plant community biomass without altering its evenness. In contrast, addition of soil from semi-natural grassland promoted plant community evenness, and hence its diversity, but reduced community biomass. Effects of donor soil additions were stronger in mineral than in organic soil and larger with bigger proportions added. The variation in plant community composition was explained best by the abundances of nematodes, ergosterol concentration and soil pH. We show that in controlled conditions inoculation of soil from secondary succession grassland into ex-arable land can strongly promote target plant species, and that the role of soil biota in promoting target plant species is greatest when added after top soil removal. Together our results point out that transplantation of later secondary succession soil can promote grassland restoration on ex-arable land.     

Transfer seeds,hay, or soil blocks? The importance of the completeness of biological inputs to address dispersal and establishment limitations during the restoration of plant assemblages in floodplain grasslands

This study investigated different techniques of grassland restoration to overcome dispersal or establishment limitation, which are key processes influencing early-successional plant community assembly. A fully randomized in situ experiment was set up in a former arable land in a floodplain along the Garonne river (south-western France) to test for the effect of (1) the type and completeness of the biological input (any biological material—seed, hay, and soil—transferred to the site under restoration), (2) soil disturbance by deep tillage, and (3) their interaction on plant community dynamics for 5 years. All inputs influenced the plant community structure and composition. The effects depended on the type of the input but not necessarily in link with its completeness. High density and diversity seed mixture led to high levels of richness and relative abundance of target species like soil blocks, the input considered as the most complete. During the first year, hay transfer mostly influenced community assembly through negative litter effects. Delayed germination of several species contributed to buffer these early effects. This study supports the importance of dispersal limitation during early succession in degraded grassland ecosystems whereas soil disturbance had only subtle effects on the seed bank and standing plant community, indicating that competition and establishment limitation were of secondary importance in our study system. Our results suggest that even low-frequency immigration events (e.g. when a few seeds are transferred within hay) can be of great importance providing that a sufficiently long time period is considered for grassland restoration.     

黄土高原半干旱区不同种植年限紫花苜蓿人工草地土壤微生物和线虫群落特征

建植紫花苜蓿人工草地是黄土高原植被恢复的重要措施之一。土壤微生物和线虫群落特征是评价和调控植被恢复的生态环境效应的重要依据。本研究在宁夏南部山区选取不同种植年限(1、2、6和12年)的紫花苜蓿人工草地为研究样地,以农田和天然草地作为对照,探索黄土高原人工草地植被恢复过程中土壤微生物和线虫群落的演变规律及其影响因素。结果表明: 1)种植苜蓿显著提高了土壤细菌群落的Chao1、ACE和Shannon多样性指数,并在种植苜蓿后第6年达到最高,但在种植6年和12年后真菌群落多样性降低;随着苜蓿种植年限的增加,真菌群落组成从农田逐渐向天然草地方向演变;2)土壤线虫数量与细菌群落多样性的变化趋势相同,在种植苜蓿后第6年出现峰值,该时期线虫群落结构组成与农田较相似,苜蓿12年样地则更接近天然草地;随着苜蓿种植年限的增加,食细菌线虫、植食性线虫比例总体呈上升趋势,食真菌线虫、杂食/捕食线虫比例呈下降趋势,土壤成熟度指数(MI)逐渐减小,植物寄生线虫指数(PPI)和线虫通路指数(NCR)则不断增大;3)在苜蓿人工草地植被恢复过程中,土壤有机碳、全氮和速效磷对土壤微生物群落结构影响较大,并进一步影响线虫群落结构;细菌和真菌群落优势类群和多样性与线虫的不同营养类群及生态指数之间存在密切联系,表明微生物群落结构与多样性对线虫群落具有显著影响;在不同种植年限苜蓿草地中,植物的生物量与多样性的变化可能通过影响土壤微生物与线虫食物资源状况从而引起其群落特征的改变。     

Contrasting success in the restoration of plant and phytophagous beetle assemblages of species-rich mesotrophic grasslands

Over the last 60 years changes to the management of species-rich mesotrophic grasslands have resulted in the large-scale loss and degradation of this habitat across Europe. Restoration of such grasslands on agriculturally improved pastures provides a potentially valuable approach to the conservation of these threatened areas. Over a four-year period a replicated block design was used to test the effects of seed addition (green hay spreading and brush harvest collection) and soil disturbance on the restoration of phytophagous beetle and plant communities. Patterns of increasing restoration success, particularly where hay spreading and soil disturbance were used in combination, were identified for the phytophagous beetles. In the case of the plants, however, initial differences in restoration success in response to these same treatments were not followed by subsequent temporal changes in plant community similarity to target mesotrophic grassland. It is possible that the long-term consequences of the management treatments would not be the establishment of beetle and plant communities characteristic of the targets for restoration. Restoration management to enhance plant establishment using hay spreading and soil disturbance techniques would, however, still increase community similarity in both taxa to that of species-rich mesotrophic grasslands, and so raise their conservation value. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The Restoration of Phytophagous Beetles in Species‐Rich Chalk Grasslands

This study focuses on the restoration of chalk grasslands over a 6‐year period and tests the efficacy of two management practices, hay spreading and soil disturbance, in promoting this process for phytophagous beetles. Restoration success for the beetles, measured as similarity to target species–rich chalk grassland, was not found to be influenced by either management practice. In contrast, restoration success for the plants did increase in response to hay spreading management. Although the presence of suitable host plants was considered to dictate the earliest point at which phytophagous beetles could successfully colonized, few beetle species colonized as soon as their host plants became established. Morphological characteristics and feeding habits of 27 phytophagous beetle species were therefore tested to identify factors that limited their colonization and persistence. The lag time between host plant establishment and colonization was greatest for flightless beetles. Beetles with foliage‐feeding larvae both colonized at slower rates than seed‐, stem‐, or root‐feeding species and persisted within the swards for shorter periods. Although the use of hay spreading may benefit plant communities during chalk grassland restoration, it did not directly benefit phytophagous beetles. Without techniques for overcoming colonization limitation for invertebrate taxa, short‐term success of restoration may be limited to the plants only.     

云南松林次生演替阶段土壤细菌群落的变化

土壤细菌多样性是维持森林生态系统功能的关键因子,森林演替是影响其动态变化的重要因素。研究云南松林不同演替阶段土壤细菌群落结构及其多样性的变化规律,有助于深入理解森林生态系统恢复过程的驱动机制。本研究以云南省永仁县皆伐后形成的针叶林、针阔混交林和常绿阔叶林为对象,基于Illumina Hiseq高通量测序技术,分析森林演替过程中土壤细菌群落组成、结构、多样性及其影响因子的变化。结果表明: 土壤细菌的种群分类单元、Ace指数、Chao1指数和Shannon指数均随着演替进行呈减少趋势,演替早期阶段土壤的细菌总数、菌群丰富度及复杂程度最高。不同演替阶段细菌群落结构存在显著差异,其中,针阔混交林的差异最大,变形菌门和酸杆菌门为各演替序列共有的优势类群,放线菌门、绿弯菌门和Patescibacteria是演替早期的优势类群,且随着演替进行呈现减少趋势;变形菌门和WPS-2相对多度随演替进行呈增加趋势。土壤pH和乔木层物种丰富度是驱动次生演替过程中土壤细菌群落组成变化的关键因子。随着演替的进行,土壤细菌多样性减少,群落组成差异加大。     

The Influence of Soil Inoculum and Nitrogen Availability on Restoration of High-Elevation Steppe Communities Invaded by Bromus tectorum

Cheatgrass ( Bromus tectorum L.) is an exotic annual grass that has invaded approximately 40,000,000 ha of rangelands in the United States, including montane ecosystems that are important habitats for wildlife and livestock. In addition to well-understood mechanisms by which Cheatgrass gains competitive advantage, recent studies have shown that Cheatgrass may also change the associated soil microbial community to impact native perennial plants and promote the persistence of Cheatgrass. Furthermore, reducing plant-available N represents a tool for initiating conditions that accelerate successional change from annual- to perennial-dominated communities. At a montane, mixed shrub–grassland Cheatgrass-dominated site in Colorado, we applied sucrose to reduce available N, and we added soil from a native plant community in order to reestablish the microbial community. This approach tested the idea that intact native soil microbial communities may enhance the beneficial effect of reducing soil N availability in a restoration setting. By the end of the experiment, reduced N availability decreased Cheatgrass by 9.8%, non-native annual/biennial plant cover by 15.0%, and increased relative perennial plant cover by 13.4%; soil inoculation reduced Cheatgrass by 7.6% and increased perennial abundance by 11.3%. Soil inoculum additions and reduced N availability both contributed toward restoring a perennial-dominated community and demonstrates that addition of native soil inoculum may be a useful tool for restoration efforts.     

Artificial Dispersal as a Restoration Tool in Meadows: Sowing or Planting?

Habitat fragmentation and the abandonment of former agricultural practices have led to extremely low dispersal rates for plant species growing in traditionally managed hay meadows in Sweden. Historically, seed dispersal between populations was maintained by hay movement, grazing animals, and farmers sharing their equipment. Because these means of dispersal typically are no longer occurring, artificial dispersal using seeds and plug-plants is tested here as a restoration tool. In this study, we chose two perennial herbs commonly occurring in meadows as test species, viz, Hypochoeris maculata L. (Asteraceae) and Succisa pratensis Moench. (Dipsacaceae). We found that plug-plant transplants were twice as effective as seed sowing for both species. The seed collection site was found to be important for seed-based establishment and survival; consequently, the choice of donor meadow is important when acquiring seeds used for restoration. We also found that survival of plants introduced as seeds was generally lower at sites harboring species favored by nitrogen as well as at sites in later successional phases. Both methods of introducing meadow species worked well, even though long-term establishment may well be more successful with the plug-plant method due to higher plug-plant establishment 2 years after introduction in the field.     

Restoration of upland hay meadows over an 11‐year chronosequence: an evaluation of the success of green hay transfer

Grassland restoration has become a key tool in addressing the drastic losses of seminatural grassland since the mid‐twentieth century. This study examined the restoration by green hay transfer of upland hay meadows, a particularly scarce and vulnerable habitat, over an 11‐year chronosequence. The community composition of 18 restoration meadows was compared with that of donor reference sites in two study areas in the Pennine region of northern England. The study investigated: differences in community composition between donor and restoration meadows; transfer of upland hay meadow target species; and the effect of time and isolation from neighboring meadows on the community composition of the restoration meadows. Results showed that restoration meadows differed from donor meadows in that some target species were easily transferred whilst others were not found in the restoration meadows, or were at low levels of cover. Time had a significant effect on the community composition of the restoration meadows, but the similarity between restoration sites and donor sites did not increase with time, and the effect of isolation was not significant. The study showed that the green hay transfer method increases botanical diversity and is an important first step in meadow restoration. However, further restoration activity, such as seed addition, is likely to be required if restoration sites are to resemble closely the reference donor sites.     

宁南山区不同植被恢复方式下土壤线虫群落特征:形态学鉴定与高通量测序法比较

线虫是土壤食物网的重要组分, 也是土壤健康与生态系统恢复的重要指示生物, 因此准确测定线虫群落特征是发挥其生态指示作用的基础。传统线虫学研究多采用形态学鉴定方法, 但高通量测序技术近年来也逐渐受到重视。然而, 关于这两种方法的对比研究目前仍比较缺乏。本研究同时采用形态学鉴定和高通量测序法, 在黄土高原宁夏南部山区, 对不同植被恢复方式下(农田、自然恢复草地、柠条(Caragana korshinskii)人工林地和苜蓿人工草地)土壤线虫的数量、群落格局和生态指数进行了测定和比较。结果表明: (1)高通量测序仅能提供线虫类群的相对多度, 而形态学鉴定法能够测定土壤线虫的绝对多度, 后者测定结果表明3种植被恢复样地, 特别是自然恢复草地和柠条人工林地, 较农田具有更高的土壤线虫多度; (2)高通量测序法检获的线虫类群数(3纲4目26科42属)高于形态学鉴定法的测定结果(2纲3目18科27属), 但两种方法仅检获15个共有线虫属, 前者检测到的植物寄生线虫属数(22属)较后者(7属)显著增加, 而食细菌线虫和杂食-捕食线虫则相反; (3)在两种方法下, 相比农田, 3种植被恢复样地尤其是自然恢复草地和柠条人工林地, 其食微线虫的相对多度均显著下降, 而植物寄生线虫和杂食-捕食线虫的相对多度大幅上升, 这也导致线虫成熟度指数(MI)和植物寄生线虫指数(PPI)的提高及瓦斯乐卡指数(WI)的显著下降; (4)相比形态学鉴定法, 高通量测序法能检测到更丰富多样的植物寄生线虫, 在该方法下土壤线虫群落的组成、结构和生态指数在植被恢复样地与农田之间的差异也更为显著。综上所述, 采用形态学鉴定和高通量测序法测定的不同植被恢复方式下的线虫群落特征具有显著差异。     

Soil community composition drives aboveground plant–herbivore–parasitoid interactions

Soil organisms can influence higher trophic level aboveground organisms, but only very few studies have considered such effects. We manipulated soil community composition of model grassland ecosystems by introducing nematode communities, microorganisms, neither or both groups. Above ground, aphids ( Rhopalosiphum padi ) and parasitoids ( Aphidius colemani ) were introduced, and we measured individual performance and population dynamics of plants, aphids and parasitoids. In microcosms with nematode inoculations either with or without microorganism inoculation, aphids offspring production was significantly reduced by 31%. Aphid populations on both host plants Agrostis capillaris and Anthoxanthum odoratum were lowest in microcosms with combined nematode and microorganism inoculations. Opposite results were found for parasitoids. While the number of emerged parasitoids did not differ between treatments, parasitoid mortality and the proportion of males were significantly lower in microcosms with nematode and microorganism inoculations. Parasitized aphids were significantly larger in microcosms with nematodes inoculated. Plant biomass did not differ, but in the preferred host plant A. odoratum , foliar phenolic content was reduced in the presence of nematodes, and also the concentration of amino acids in the phloem. This study shows that the composition of the soil community matters for aboveground multitrophic interactions.     

Can Establishment Characteristics Explain the Poor Colonization Success of Late Successional Grassland Species on Ex‐Arable Land?

Abstract Many grassland restoration projects on former arable land face problems because early successional grassland species establish vigorously and persistently but late successional grassland species fail to establish. Differences in establishment characteristics of early and late successional species might provide an explanation for the failure of many late successional species to colonize grasslands on ex‐arable land. I examined whether early and late successional species had different establishment rates in the initial years of a grassland succession, whether a specific establishment stage (seedling emergence, mortality or growth) could be identified as the key process controlling establishment, and what management would enhance the establishment of late successional grassland species. Seeds of three early and three late successional species were sown separately in ex‐arable plots with bare soil, 1‐year‐old vegetation, and 2‐year‐old vegetation. Emergence, mortality, and seedling growth were monitored for 1 year. Early successional species established successfully in the bare soil plots but failed to establish in plots with 1‐ and 2‐year‐old vegetation. Late successional species showed either lower establishment rates in the younger succession stages or decreased establishment with succession that nevertheless resulted in significant establishment in the oldest plots. Seedling emergence proved to be the key factor determining the establishment pattern of early and late successional species. In absolute numbers, emergence of late successional species was, however, similar or higher than that of early successional species, even in the earliest succession stage. The poor establishment of late successional species on former arable land could therefore not be explained solely by differences in establishment characteristics between early and late successional grassland species. Competitive processes between early and late successional species later in the life cycle probably play an important role. The results do point out that establishment of late successional species can be promoted by creating vegetative cover from the start of the restoration effort.     

Responses of Pioneer and Later‐Successional Plant Assemblages to Created Microtopographic Variation and Soil Treatments in Riparian Forest Restoration

Riparian forest restoration generally involves introduction of later‐successional tree species, but poor species suitability to severely altered or degraded site conditions results in high mortality and poor community development. Additionally, while microtopographic heterogeneity plays a crucial role in the development of natural riparian forests, little is known regarding effects of restored or created microtopography on the development of introduced plant communities. The objective of this study was to determine the influence of created microtopography and soil treatments on early development of introduced pioneer and later‐successional plant communities in riparian forest restoration. Ridges, flats, and a mound‐and‐pool complex were created, and pioneer and later‐successional tree assemblages were planted within plots in each of these three microtopographic positions. Straw‐based erosion control mats were placed on half the plots as a source of mulch. After two growing seasons, growth and survival of the pioneer assemblage were equal among microtopographic positions, but survival of the later‐successional assemblage was significantly higher on ridges (59%) than on mounds and pools (22%) and flats (26%). A suitability index indicated that performance of the later‐successional assemblage on ridges was higher than that of the pioneer assemblage for all microtopographic positions. Flood duration explained much of the variation in plant assemblage survival, and erosion control mats had little influence on seedling survival. Restoring microtopographic features has the potential to enhance species survival and promote community development. Microtopographic restoration may be as important in riparian forest restoration as proper species selection and hydrologic reestablishment, especially at severely disturbed sites.     

Plant community composition steers grassland vegetation via soil legacy effects

Soil legacy effects are commonly highlighted as drivers of plant community dynamics and species co‐existence. However, experimental evidence for soil legacy effects of conditioning plant communities on responding plant communities under natural conditions is lacking. We conditioned 192 grassland plots using six different plant communities with different ratios of grasses and forbs and for different durations. Soil microbial legacies were evident for soil fungi, but not for soil bacteria, while soil abiotic parameters did not significantly change in response to conditioning. The soil legacies affected the composition of the succeeding vegetation. Plant communities with different ratios of grasses and forbs left soil legacies that negatively affected succeeding plants of the same functional type. We conclude that fungal‐mediated soil legacy effects play a significant role in vegetation assembly of natural plant communities.     

Temporal variation in plant-soil feedback controls succession

Soil abiotic and biotic factors play key roles in plant community dynamics. However, little is known about how soil biota influence vegetation changes over time. Here, we show that the effects of soil organisms may depend on both the successional development of ecosystems and on the successional position of the plants involved. In model systems of plants and soils from different successional stages, we observed negative plant–soil feedback for early-successional plant species, neutral feedback for mid-successional species, and positive feedback for late-successional species. The negative feedback of early-successional plants was independent of soil origin, while late-successional plants performed best in late- and worst in early-successional soil. Increased performance of the subordinate, late-successional plants resulted in enhanced plant community diversity. Observed feedback effects were more related to soil biota than to abiotic conditions. Our results show that temporal variations in plant–soil interactions profoundly contribute to plant community assemblage and ecosystem development.     

Inoculation with a Native Soil Community Advances Succession in a Grassland Restoration

Restoration on post‐agricultural land may be hindered by the degradation of the soil community, which has been shown to contribute to structuring plant communities and driving succession. Our experiment tested the effect of inoculation with remnant grassland whole soil with or without nurse plants on the survival and growth of uninoculated early and late successional plant species. In 2007 and 2008, we planted uninoculated early, mid, and late successional plant species 0.25–2 m away from a central point of inoculated nurse plants. We found a negative response to inoculation on early successional plants and a positive response to inoculation on mid to late successional plants. This work suggests that the restoration of the soil community is critical to establishing a late successional plant community and that the benefit of inoculated plants can spread to neighbors.     

Evaluation of Initial Restoration Measures during the Restoration of Calcareous Grasslands on Former Arable Fields

In 1993, experiments on the restoration of calcareous grasslands on ex‐arable fields were started in order to provide new habitats for species of a small nature reserve with ancient grasslands north of Munich (Germany). The effects of diaspore transfer by the application of seed‐containing hay on vegetation establishment were studied on restoration fields with and without topsoil removal for 5 years. The aim of the study was to assess plant diversity for the evaluation of restoration success by different methods including determination of species with viable seeds in the hay by germination tests, phenological investigations on hay‐transfer source sites at the time of harvest, and vegetation analyses on the restoration sites. Total seed content of the hay and the number and composition of plant species with viable seeds were affected by the time of harvesting and differed between a site which had been used as arable field until 1959 and ancient grassland sites. Nevertheless, the number of established hay‐transfer species showed only few differences between restoration fields. The proportion of species transferred to restoration fields in relation to the number of species with viable seeds in the hay was between 69 and 89%. Five years after the hay transfer, the proportion of the established species was still between 58 and 76%. Up to now, topsoil removal had no significant effect on the number of established hay‐transfer species. After triple hay application the absolute number of transferred grassland species was higher than on sites with single hay application, but restoration efficiency was lower because many of the species with viable seeds in the hay did not establish. In general, our results showed that the transfer of autochthonous hay is a successful method to overcome dispersal limitation in restoration projects.     

Effects of topsoil removal,seed transfer with plant material and moderate grazing on restoration of riparian fen grasslands

Question: How do moderate grazing, topsoil removal and hay transfer affect species diversity and abundance on a eutrophic fen grassland site? Location: Northern Germany. Method: A three-factorial field experiment with the factors grazing, topsoil removal and hay transfer of diaspore-rich material was established in 2001. Soil nutrients and seed bank were analysed at the beginning of the experiment, species composition and vegetation development was monitored for four years (2002–2005). Results: Topsoil removal had a significant effect on the abundance of different plant species groups: resident vegetation of agricultural grasslands was suppressed, while clonal reed species were facilitated in recolonising the area. The establishment of regionally rare and endangered species of nutrient-poor fens and wet meadows introduced with hay was achieved mainly on plots with topsoil removal, with the exception of Rhinan-thus angustifolius, which also established on plots with intact topsoil. Effects of grazing after four years of experiments were of minor influence on species composition. Conclusion: The establishment of target plant species of nutrient-poor fens is most successful when both an adequate number of viable diaspores and suitable sites for germination and establishment are available. In our experiment this was achieved by the combination of topsoil removal and hay transfer. We recommend this combination, together with continuous management (grazing/cutting), for further restoration in fen grasslands.     

Single introductions of soil biota and plants generate long‐term legacies in soil and plant community assembly

Recent demonstrations of the role of plant–soil biota interactions have challenged the conventional view that vegetation changes are mainly driven by changing abiotic conditions. However, while this concept has been validated under natural conditions, our understanding of the long‐term consequences of plant–soil interactions for above‐belowground community assembly is restricted to mathematical and conceptual model projections. Here, we demonstrate experimentally that one‐time additions of soil biota and plant seeds alter soil‐borne nematode and plant community composition in semi‐natural grassland for 20 years. Over time, aboveground and belowground community composition became increasingly correlated, suggesting an increasing connectedness of soil biota and plants. We conclude that the initial composition of not only plant communities, but also soil communities has a long‐lasting impact on the trajectory of community assembly.     

引入草木樨对半干旱黄土高原区早期植物群落演替和土壤养分的影响

植被演替早期的优势物种是影响演替进程的重要因素。以半干旱黄土高原地区退化耕地植被的自然恢复为对照,于植被恢复的早期引入两年生豆科植物草木樨(Melilotus officinalis L.),在种植密度为11.3kg.hm-2条件下,考察了6年内引入草木樨对退化耕地植被恢复过程中植物群落演替进程的影响。结果表明,与自然恢复植被相比,草木樨的引入不仅可以提高地表植被覆盖度和加速植物群落演替进程,促使一些群落演替后期物种如冷蒿(Artemisia frigida)、阿尔泰狗哇花(Heteropappus altaicus)和短花针茅(Stipa breviflora)等在植被恢复的第3年出现,同时有利于提高植被生产力和稳定性。同自然恢复相比,试验期间引入草木樨后的植被地上平均生物量提高67.90%。同时,植被恢复早期生物多样性与土壤全氮含量呈显著正相关,而草木樨的引入有利于提高土壤全氮含量;在植被恢复初期(2003～2005)引入草木樨的土壤全氮平均含量比自然恢复高7.32%;试验期间土壤速效磷含量与物种丰富度呈显著负相关。因此,在半干旱区黄土高原退化耕地植被恢复早期阶段,两年生豆科植物草木樨的引入和磷肥的施用对于加快植被恢复有着积极作用。