The importance of former land use in determining successful re-creation of lowland heath in southern England

In order to reverse losses of lowland heathland substantial areas of former agricultural and forestry land are currently being targeted for restoration. In this study we assessed the success of heathland re-creation on 37 former arable, improved grassland and conifer plantation sites in southern England, UK. The similarity between the vegetation and seed banks of the re-creation sites and undisturbed heathland controls were examined in relation to former land use, soil properties and re-creation management. Former land use was found to be the most important determinant of re-creation success. On former arable, and to a lesser extent improved grassland, antecedent management had caused significant changes to seed bank and soil properties. As a result, regeneration of Calluna heath had been minimal, even on sites where appropriate management (e.g. cropping, addition of pH amendments) had been undertaken and heathland species introduced. In contrast, former plantation seed banks and soils were similar to heathland controls, and, as a result, rapid regeneration of Calluna heath had taken place. It was concluded that conifer removal provides the most practical and cost-effective means of re-creating Calluna heath on former heathland, although further research is required to assess the effect of litter removal and the rate at which heather seed banks decline following conversion to forestry. In contrast, re-creation on former agricultural sites will require effective management to reduce soil pH, fertility and the abundance of competitive species as well as the introduction of heathland propagules. Given limited resources a more realistic objective for these sites is likely to be reversion to an acid grassland or grass-heath.     

The establishment of heathland vegetation on ex-arable land: the response of Calluna vulgaris to soil acidification

The UK Biodiversity Action Plan has identified the creation of lowland heathland as an important objective. Heathland restoration studies have identified soil pH, elevated soil nutrients and large weed seed banks as major problems in the restoration of heathland vegetation on ex-arable land. Heathland vegetation is usually found on nutrient-poor acidic soils. Creating acidic soil conditions on ex-arable sites thus may produce a suitable environment for the establishment of heath vegetation. Soil acidification by the addition of sulphur has been shown to reduce the soil pH and the availability of nutrients in arable soils. A series of experiments was established to investigate the effects of soil acidification using sulphur on the establishment of Calluna vulgaris and the development of weed vegetation. The application of sulphur at 0.24 kg m⁻² to an arable soil was found to increase the survival rate of C. vulgaris cuttings planted in it. The mechanism of increased C. vulgaris survival appeared to be by sulphur application significantly reducing the cover of arable weeds arising from the soil seed bank. Higher rates of sulphur application (0.36 and 0.48 kg m⁻²) resulted in the death of many C. vulgaris plants. However C. vulgaris seedlings were able to establish successfully on these ex-arable soils within 18-24 months following the addition of these levels of sulphur. The application of sulphur appears to offer a practical solution to heathland creation on ex-arable land. However, it may be necessary to provide an interval of between 18 and 24 months between the application of sulphur and the addition of C. vulgaris plants or seeds for the successful establishment of heathland vegetation.     

Effects of experimental restoration on the diaspore bank of an upland moor degraded by Pteridium aquilinum invasion

We assessed the effects of a range of experimental restoration treatments on the diaspore bank (seed and spores) on an upland moor infested by dense Pteridium aquilinum, 9 years after implementation. Experimental treatment included both Pteridium control (cutting, spraying and combinations) and restoration treatments (grazing, seeding); both univariate and multivariate analyses of variance were used to assess significant effects. The seed bank was made up largely of Calluna vulgaris, Juncus effusus and Agrostis capillaris with the majority germinating from the litter (38 per cent) and upper soil layer (53 per cent). Soil depth influenced species composition and density, with greatest numbers near the surface, in keeping with a negative exponential response. In the lowest soil depth only ferns were abundant. A. capillaris seed density was greater in the most disturbed treatment (cut twice yearly with grazing), whereas C. vulgaris was greater in the low‐disturbance treatment (herbicide application only). The diaspore bank composition response identified that certain treatment combinations had a grass‐heath composition in the upper soil layer, which may assist future restoration. There was a significant effect of C. vulgaris seed addition detected in the Pteridium litter layer suggesting that this litter acted as a barrier to seed transfer to the soil. There was little evidence of coupling developing between the diaspore bank and vegetation. The most successful treatment combination was cutting twice per yr with low grazing plus C. vulgaris addition as brash. Where seed is added the Pteridium litter should be disturbed. Copyright © 2007 John Wiley & Sons, Ltd.     

坝上草原退耕地植被不同恢复处理土壤种子库研究

采用野外调查取样和室内实验相结合的方法,研究坝上草原退耕地土壤种子库的物种组成、密度和物种多样性等特征.结果表明:自然恢复、浅耕处理和深耕处理3种不同恢复措施下,土壤种子库物种丰富度(种)大小为浅耕处理(18)>自然恢复(15)>深耕处理(14);自然恢复处理、浅耕处理和深耕处理土壤种子库密度分别为23 949粒·m-2、15746粒·m-2和10600粒·m-2,浅耕处理和深耕处理分别比自然恢复处理减少34.3%和55.7%;不同恢复方式下土壤种子库与地上植被的物种密度的对应关系可用三次曲线来表示.土壤种子库间有较高的相似系数,但随着干扰程度的增加而减少.     

元谋干热河谷山地植被修复区土壤种子库研究

 为了解干热河谷不同植被修复方法的土壤种子库的变化模式,采用对不同处理地段(包括水平阶间坡面、自然坡面、沟底)和对照(放牧地)的土壤种子库组成、大小进行比较研究的方法,研究元谋干热河谷山地植被修复区的土壤种子库。结果表明:1)植被修复区土壤种子库密度、丰富度均大于对照区;2)修复区内第2年土壤种子库密度、丰富度均大于第1年;3)水平阶间坡面和各类型间的土壤种子库密度差异显著;4)种子库中草本植物占很大比例;5)孔颖草和扭黄茅是种子库2大优势种。研究项目所采取植被修复措施在很大程度上改善土壤种子库的数量结构和物种组成。     

黄土丘陵沟壑区红砂灌丛土壤种子库及其自然更新潜力评估

[目的]对黄土丘陵沟壑区红砂灌丛植被土壤种子库的特征及红砂灌丛植被自然更新潜力进行分析和评估,以说明红砂灌丛土壤种子库在植被恢复与重建中的重要作用。[方法]采用土壤种子库"萌发法"(每个地段10个2m×2m样方内分表土层0—2cm和2—5cm土层采集土样)及野外植被调查方法,对黄土丘陵沟壑不同坡位红砂灌丛土壤种子库进行研究。[结果]黄土丘陵沟壑区红砂灌丛植被土壤种子库发芽试验共观察到的4 251株幼苗,分属于9个物种。土壤种子库密度在100~1 000粒/m2,物种数在0.8~1.2种/0.01m2,下坡段的土壤种子库平均种子密度和平均物种数均比上坡段高。3个坡段的2个层次的平均种子密度和平均物种数均随土层加深而减小。3个坡段土壤种子库和地上植被的组成物种多为草本植物和红砂灌丛,地上植被与其土壤种子库的密度及物种数均呈不显著相关;物种组成的Sorensen相似性指数较高,均达到0.60以上,且土壤种子库比地上植被具有更高的物种丰富度。[结论]黄土丘陵沟壑区红砂灌丛具有依靠土壤种子库实现自然更新的潜力,但由于物种组成种类较少,现存灌丛植被一旦遭到破坏,仅靠土壤种子库恢复现存植被是困难的,需要在对自然恢复潜力评估的基础上,积极采取人工保护促进自然恢复的策略。     

风沙黄土区排土场不同恢复类型植物群落与土壤种子库特征

风沙黄土区排土场作为一种人造生态系统,自然条件恶劣,土壤贫瘠,植被恢复困难。为了探明有效的人工促进植被恢复措施,采用植被调查与种子库萌发试验相结合的方法,通过研究不同植被类型地上植物群落与土壤种子库特征及二者的关系,探讨了其植被恢复效益及潜力。结果表明:研究区人工植被恢复下地上植物群落中共47种植物,分属16科40属,土壤种子库共14种植物,分属5科13属,其中均以禾本科、菊科、藜科、豆科占主导地位; 灌木植被的地上植被和土壤种子库的物种多样性均表现为最优; 地上植被与土壤种子库密度的变化范围为88.48～495.47株/m²,74.74～1422.91粒/m²,且均在草地类型下最大。土壤种子库和地上植被的相似性普遍较低,相似性系数仅为0.16～0.23。因此,风沙黄土区排土场的植被恢复与重建需要加强保护与管理,可以考虑构建以草灌配置为主的人工植被恢复模式,保障群落的恢复潜力,并提高群落多样性与稳定性,亦可考虑引入外源种子库提升群落恢复的潜力。     

Soil seed bank compositional change constrains biodiversity in an invaded species-rich woodland

Relationships between plant invasion and the soil seed bank in highly diverse fire adapted mediterranean woodlands are poorly understood, yet critical for that ecosystem’s conservation. Within the biodiversity hotspot of southwest Australia we investigated the composition and diversity of the Banksia woodland soil seed bank in good condition (GC), medium condition (MC) and poor condition invaded by the South African perennial species Ehrharta calycina (PCe) and Pelargonium capitatum (PCp). The investigation assessed three questions: (1) Do soil seed banks of invaded sites have fewer germinants of native species and more germinants of introduced (non-native) species than sites with minimal invasion? (2) Do soil seed banks show shifts in ecological functional types with invasion? (3) Is the soil seed bank of introduced species persistent? Native species germinants, mainly shrubs and perennial herbs, were highest in GC sites and least in poorer condition sites suggesting a reduction in their numbers had occurred over time. Introduced germinants were dominated by perennial and annual grasses, and annual herbs. E. calycina had the greatest seed density (8328 germinants m⁻²). More introduced than native germinants occurred in the litter. Rapid germination of introduced species (30% in week 1) compared to native species (4% in week 1) provides the capacity for their early dominance. A limited native soil seed bank and dominant persistent introduced soil seed bank represent great challenges for the structural and functional conservation and restoration of woodland ecosystems. This study provides key new knowledge, applicable to a wide range of ecosystems, to help formulate conservation protocols to control dominant introduced species and conserve and restore biodiverse-rich woodlands.     

The Vertical Distribution of Soil Seed Bank and Its Restoration Implication in an Active Sand Dune of Northeastern Inner Mongolia,China

Although the functions and characteristics of soil seed banks in topsoil layers have been described for various ecosystems, the spatiotemporal pattern of the seed bank in deep soil and its ecological implications for vegetation restoration of active sand dune have not been fully explored. In 2007 and 2008, seed densities with regard to dune position, soil depth and season were investigated on an active and a stabilized sand dune of northeastern Inner Mongolia, China. Seeds in the 0–10 cm topsoil layer accounted for 60% of total soil seed bank on the stabilized sand dune, while more than 40% of seeds were stored in the 50–100 cm layers on the active sand dune. Seed density declined significantly with soil depth on the stabilized sand dune, but it was relatively constant across the 0–100 cm soil profile on the active sand dune. Seed density fluctuated with soil depth on the active sand dune suggesting that seeds were either relocated upward or downward over time. Seeds of annual non‐psammophytic species accounted for the majority of soil seed bank on the stabilized sand dune, while pioneer psammophytes contributed more to the soil seed bank of the active sand dune. Our data suggest that seeds in the deep soil layers of active sand dunes account for a large proportion of the whole soil seed bank. Because of the effect of wind erosion, seeds in deep soil could be gradually exposed to shallow soil layers and potentially contribute to population recruitment and vegetation restoration on active sand dunes. Copyright © 2015 John Wiley & Sons, Ltd.     

Moderate impacts of plant invasion and management regimes in coastal hind dune seed banks

Plant invaders may directly or indirectly affect ecosystem resilience through their impact on soil seed banks. The invaders, and the application of control measures, change seed bank dynamics by altering the number of seeds entering and leaving the seed bank. We tested the impact of bitou bush (Chrysanthemoides monilifera ssp. rotundata), on the seed bank. We examined seed banks in heavily-invaded, sparsely-invaded and managed dunes, where bitou bush biomass had been controlled. While management of bitou bush may have reduced the density of bitou bush seeds in the soil, it did not reduce the richness of other weed species. Native tree species richness was significantly higher in seed banks of sparsely-invaded than either heavily-invaded or managed sites, perhaps indicating a permanent shift in community structure following invasion. However, remaining indices of native seed bank diversity were similar across all invasion categories, indicating that seed banks of many native species were unaffected by both invasion and management. While examination of seed banks is informative in assessing past and potential community dynamics, low similarity between the standing vegetation and seed bank at all sites indicated that many hind dune species had other storage or regeneration modes and seed banks cannot be relied upon for comprehensive dune restoration.     

Long-term heathland restoration on former grassland: The results of a 17-year experiment

European lowland heaths have declined by up to 80% due to land use change and lack of management. There has been considerable research into the restoration of this threatened habitat. However, long-term outcomes of restoration are poorly understood, especially in situations where past agricultural land use imposes severe constraints on community re-assembly. In 1989 a large-scale experiment was established to examine the effectiveness of five treatments to restore heathland on formerly productive grassland: (i) natural regeneration; (ii) herbicide application to facilitate regeneration; (iii) cultivation and application of seed-rich heathland vegetation; (iv) soil removal and incorporation of heathland topsoil; and (v) heathland translocation. After 17 years the pH of the unamended agricultural soil remained significantly higher than that of the adjacent heathland. All treatments showed different trajectories of vegetation change in the long-term. Natural colonisation by heathland species was slow due to seed limitation, resulting in formation of an acid grassland community. Heathland community assembly was not facilitated by destruction of the initial grassland with herbicide. Incorporation of topsoil had an intermediate effect on pH reduction. This may explain the subsequent failure of the plant community to assemble in the anticipated proportions, and the dominance of leguminous scrub species (Ulex spp.). Turf translocation was effective in reducing pH to the required range and restoring the heathland community in the long-term. However, this technique should only be considered as a means of ‘rescue’ when habitat destruction is otherwise unavoidable. The only practical and sustainable means of increasing heathland extent on former farmland is the application of seed-bearing vegetation cut as part of routine management. However, this technique needs refining in order to establish the full range of characteristic heathland species.     

Soil water depletion and replenishment during first- and early second-rotation Eucalyptus globulus plantations with deep soil profiles

Eucalyptus globulus plantations are thought to use stored soil water when planted on ex-agricultural sites, and we hypothesized that this is likely to affect productivity of 2nd and later rotation plantations because the next rotations have access to less stored soil water. We used a combination of experiments and modeling to understand the impact of E. globulus plantations on soil water stores over the first rotation and early second rotation. The experiments were conducted at 3 contrasting sites in south-western Australia, and modeling was used to extrapolate the results to other climatic zones. Soil water dynamics were assessed to 8 m depth under a range of management options, including spacing and nitrogen addition in the first rotation, and coppice or seedling re-establishment in the 2nd rotation. We found that soil water stores declined over the course of the first rotation at all sites, with some (incomplete) annual replenishment evident at the higher rainfall sites, but less replenishment at depth, especially in the lower rainfall sites. Only the wettest of the 3 sites fully replenished with soil water after harvest of the 1st rotation. Plots with higher stocking rates had higher soil water depletion early in the rotation, although by the end of the first rotation, most treatments had similar soil water deficits of around 800 mm at all of the sites. Of the sites that were responsive to N fertilizer, there was a strong differential in the degree of soil water deficit between N treatments, with N fertilized trees using more of the soil water store each year, but also producing more wood. A process-based plantation growth model, CABALA, was found to be adequate for predicting soil water dynamics under the range of management options that we explored, and we applied it to understanding the potential replenishment of soil under 2nd rotation plantations in a range of climatic zones within the E. globulus estate in south-western Australia. This modeling showed that most sites with soil depths of more than 4 m (i.e., most of the estate) are unlikely to be fully replenished in the 2nd rotation, and that this is likely to have a significant impact on the capacity of sites to achieve similar productivity levels in the second rotation as the first unless the sites are given an opportunity for soil water replenishment between rotations. The results from this study suggest that plantation managers will need to understand soil water dynamics at any given site to be able to predict productivity in 2nd and later rotations, and may need to explore novel management options like fallowing between rotations to allow for soil water replenishment.     

陕北安塞退耕地持久土壤种子库与地上植被的对应关系

通过研究陕北安塞退耕3~31 a的阴阳坡10个样地的持久土壤种子库特征及其与地上植被的关系,分析持久土壤种子库在植被恢复中的作用。结果表明：调查样地持久土壤种子库由15个科36个物种组成,种子密度在1 908~12 617粒/m2之间,1年生物种占80%,随着退耕演替的进行,种子库中1年生物种比例呈下降趋势,多年生物种比例呈上升趋势,地带性物种在持久种子库中的比例也随退耕时间的延长而增加,物种数和种子库密度均随退耕演替先增大后降低;持久土壤种子库与地上植被的相似性较低,Serensen相似性系数为0.21~0.42;通过对适宜物种的确限度及土壤种子库大小的分析,认为该区退耕地的持久土壤种子库具有一定的植被恢复潜力。     

Does the heathland flora in north-western Belgium show an extinction debt?

The large losses of heathland area since the end of the 18th century can be expected to have resulted in the decline or even extinction of many characteristic heathland species. Historical data on plant species distribution patterns can provide valuable information in this context. Therefore, the aims of this research are to study how the loss of heathland area has changed the presence of heathland and forest plant species in north-western Belgium using historical plant distribution data, and to test whether the heathland flora shows an extinction debt. Furthermore, plant traits determining extinction sensitivity are investigated.Our results revealed that, despite the dramatic reduction of heathland area (more than 99% of heathland was destroyed over a 230-year period), the loss of heathland species is relatively limited (11%) and is comparable with that of forest species (11%). Heathland species that have a long-term persistent seed bank or can propagate vegetatively are least sensitive to extinction. For forest species, on the other hand, growth form is the key determinant for extinction sensitivity. Long-lived woody species have a greater chance of persisting.The relatively low extinction numbers probably represent an extinction debt and the full effects of habitat loss may not have been fully manifested yet. Consequently, future extinctions are expected to occur unless environmental conditions are improved. Therefore, heathland restoration and prevention of further heathland area losses is required.     

草炭与珍珠岩对土壤种子库种子萌发特征的影响

[目的]探究基质添加对土壤种子库种子萌发特征的影响,为今后土壤种子库应用于植被恢复工程提供有价值的参考。[方法]借鉴日本植被恢复的先进经验,采用温室萌发法探究有机基质(草炭)、无机基质(珍珠岩)及混合基质(草炭与珍珠岩)添加对土壤种子库种子萌发特征的影响。[结果]草炭与珍珠岩混合基质对土壤种子库密度、物种多样性的促进作用更为明显,当混合基质与表土配比为20%和30%时,Shannon-Wiener多样性指数和种子库密度分别达到最高值;种子库萌发曲线中,混合基质组的萌发效果明显高于空白组和其他组;种子库密度与土壤因子的多元回归分析中,C,N,P,K元素均对种子库密度影响显著,进一步通过CCA排序法,分析发现大部分植物均对土壤含水率和C,N,P,K等元素有一定程度上的要求。[结论]向表土中添加混合基质作为一种人工措施,能有效地促进种子库密度和物种多样性,可为植被恢复工程提供具有实用价值的参考。     

Vegetation Type Affects the Relationship Between Soil Carbon to Nitrogen Ratio and Nitrogen Leaching

Nitrate leaching occurs when the soil's nitrogen immobilisation and plant uptake capacity has been saturated. Several widely-used models of nitrogen saturation incorporate a breakthrough function in which N begins to be leached at C/N values below an upper threshold, and is completely leached at C/N values below a lower threshold. In a survey of deciduous and coniferous woodland, acid grassland and heathland sites for which both C/N and nitrate flux measurements were available, deciduous woodland and acid grassland typically had lower C/N ratios, and began leaching nitrate at a lower C/N ratio, than coniferous woodland and heathland. Least-square fits of nitrate breakthrough functions gave upper thresholds (no nitrate leaching) of 27 mol C mol^?1 N for deciduous woodland and acid grassland and 50 mol C mol^?1 N for coniferous woodland and heathland. Upper thresholds were similar, at 24 and 51 mol C mol^?1 N, respectively, for total inorganic N (NH₄ + NO₃) leaching flux as a proportion of total inorganic N influx. In conifer plantations, stand maturity had a large effect, suggesting that a breakthrough function is unsuitable for modelling systems that are in disequilibrium. However, there was sufficient evidence to suggest that using different breakthrough C/N thresholds for different groups of vegetation would improve predictions of N saturation and leaching at both plot and catchment scales. The difference may be related to the reactivity of soil carbon; soils with a large proportion of recalcitrant carbon are likely to begin leaching nitrate at a higher C/N value than soils with more labile carbon.     

土壤种子库研究的热点问题及发展趋向

土壤种子库是指存在于土壤上层凋落物和土壤中全部存活种子的总和。土壤种子库是种群定居、生存、繁衍和扩散的基础,也是植被潜在更新能力的重要组成部分,在植被的更新、演替和恢复过程中起着重要的作用。本文总结提出了土壤种子库研究的几个热点问题以及发展趋向,包括土壤种子库的研究方法,入侵植物的土壤种子库,种子雨、种子库与地上植被的关系,种子库在退化生态系统植被恢复和生物多样性保护中的作用,土壤侵蚀对土壤种子库的影响,全球变化对土壤种子库的影响。同时对这些领域的研究进展进行了综述,旨在为进一步推动今后土壤种子库的研究工作提供参考。     

Effects of removing the litter and humic layers on heathland establishment following plantation removal

We investigated ways of optimising the establishment of Calluna vulgaris-dominated heathland vegetation following removal of pines from afforested heathland using a randomised block experiment. Removal of the litter layer, or both the litter and humic layers, resulted in a higher initial establishment rate of C. vulgaris compared to in the control plots. The effect of adding heathland clippings suggested that the rate of establishment of C. vulgaris was limited by seed availability in plots from which both the litter and humic layers had been removed, but not in those from which just the litter layer had been removed. Removal of both the litter and humic layers did, though, result in a long-term reduction in the depth of the humic layer, a lower loss-on-ignition and lower level of nitrate and total nitrogen compared to in the control plots. The optimal method for establishing heathland vegetation at a site will depend on: (1) whether there is a viable seedbank of C. vulgaris and other desirable heathland species present; (2) the importance of quickly establishing a high cover of desirable heathland species; (3) the likely long-term effects of soil nitrogen levels on the maintenance of C. vulgaris dominance; (4) the practicality of removing and disposing of the litter and humus layers.     

Habitat selection by the Mediterranean horseshoe bat, Rhinolophus euryale (Chiroptera: Rhinolophidae) in a rural area of southern Italy and implications for conservation

We studied habitat selection by Rhinolophus euryale in a rural area of southern Italy in 1998-2000 by radio-tracking. Two comparisons were carried out, one between habitat occurrence within individual home ranges and within the study area, the other between time spent in each foraging habitat and habitat occurrence within the home range. The first analysis showed that olive groves and conifer plantations were, respectively, the most and the least important habitats. The second analysis highlighted the importance of woodland for R. euryale, while urban sites, open areas and conifer plantations were avoided. We recommend that clearing of continuous, large areas of woodland for tree harvesting should be avoided. Conifers should not be used for reforestation. Urbanisation should be limited in the areas of greatest importance for the species, and linear landscape elements such as tree lines and hedgerows should be maintained.     

Changes in humus properties and collembolan communities following the replanting of beech forests with spruce

Differences of collembolan communities within the organo-mineral A layer were studied in relation to physico-chemical changes in humus at nine sites of beech forests (Fagus sylvatica L.) and first generation spruce stands (Picea abies (L.) Karst.), planted on former beech forest 30 years ago (Central Pyrenees, France). Changes in humus form were caused by the spruce plantation and occurred mainly within the fermentation horizon where acidifying litter accumulation increased the horizon depth. The recent replacement of beech by spruce induced a shift from mull towards moder humus forms, which is explained by the decreasing organic matter turnover rate. A significant decrease in the three exchangeable cations and pH under spruce was also observed. Collembolan species diversity within the A horizon was significantly lower under spruce at this early stage of the silvogenetic cycle. Differences between species composition of Collembola between the two forest stands is related to changes in environmental conditions (e.g. nutrient availability, soil porosity, soil moisture). This study shows how forest management practices are susceptible to modify biological activity within the A horizon under exotic conifer plantations.