Diversity of soil macro-invertebrates in grasslands under restoration succession

In grasslands under restoration succession, the effect of nutrient impoverishment on the vegetation has been intensively studied. The invertebrate soil animal community has, however, rarely been studied under such conditions. We hypothesised that a plant community in nutrient-poor grasslands supports fewer macro-invertebrate individuals than richer grasslands. We also hypothesised that the composition of the macro-arthropod community is specific for a certain stage of vegetation succession, i.e. nutrient impoverishment. A detailed sampling programme was carried out in four grasslands under restoration succession. The time since last fertilisation differed for these four grasslands (7, 11, 24 and 29 years). The lowest total number of individuals in different taxonomic groups of macro-invertebrates was found in the most impoverished field in the majority of cases. Analysis of the data set for all specimens identified to order or family level with a canonical correspondence analysis resulted in a pattern not related to successional stage. The same holds when the species composition of the ground beetles was analysed separately. Short-term abiotic factors explained a significant part of the variation in both analyses. With the same multivariate analysis, however, the adult weevils, identified to species level, showed a clear relation with nutrient status of the grasslands.     

黄河源区人工草地植被群落和土壤养分变化

[目的]研究黄河源区不同年限人工草地植被群落特征和土壤养分的动态变化,揭示高寒地区人工草地稳定机制与演替规律,为退化高寒草甸(湿地)的近自然恢复和缩短退化草地恢复时间提供理论依据。[方法]选择黄河源区青海省玛沁县3,11,17 a单播垂穗披碱草人工草地,对植被与土壤养分特征进行调查。[结果]随着种植年限增加,人工草地优势种垂穗披碱草盖度降低,植物总盖度、生物结皮盖度、杂类草盖度以及生殖枝数量呈倒"V"型变化,而原生植被莎草科植物盖度、物种多样性逐渐增加,17 a人工草地中莎草科植物的盖度是3,11 a的10倍;人工草地土壤养分中全氮、全钾、速效氮、速效钾以及有机质随年限增加呈现积累趋势,土壤pH值逐渐趋于中性。土壤全氮含量在不同恢复年限之间差异最大,平均准确率降低度为25.71,有机质含量次之,其平均准确率降低度为18.55,而全钾含量及均匀度指数最小,平均准确率降低度均小于5。[结论]高寒地区人工草地群落结构和土壤营养随着建植时间的延长在逐渐恢复,建植17 a的人工草地土壤全氮、有机质含量仅是原生高寒草甸土壤的50%左右,因此,17 a人工草地土壤养分完全恢复还需要较长时间。     

Restoring grassland biodiversity: Sowing low-diversity seed mixtures can lead to rapid favourable changes

We studied the effect of sowing low-diversity seed mixtures (containing 2-3 competitive grass species) on the regeneration of vegetation on croplands previously used as alfalfa fields. In four permanent plots in 10 restored fields (four with alkali and six with loess seed mixture) the cover of flowering plants was recorded and phytomass was measured between 2006 and 2008. We asked three questions: (i) How fast will weedy, short-lived species decrease in abundance during secondary succession enhanced by sowing low-diversity seed mixtures? (ii) Can weeds be suppressed by sowing competitive native grasses, followed up by management by mowing? (iii) Can succession towards the target native grasslands be accelerated by sowing only low-diversity seed mixtures compared to set-aside old field succession? Our results showed that in just two years a vegetation dominated by perennial grasses has developed, which successfully prevented the establishment of weed species. These results suggested that sowing seeds of two or three competitive grass species is an effective tool to eliminate weed domination and to lead towards the restoration of species poor grasslands with grass domination such as alkali grasslands. However, the developed dense perennial grass cover and the accumulated litter may hamper the immigration of specialist species characteristic to reference grasslands. Therefore, the restoration of species-rich grasslands requires the facilitation of the immigration of grassland specialist species by further management (grazing, mowing and/or hay-transport).     

退化演替对高山草地植被和土壤理化特性影响

[目的]分析3种不同草地退化阶段(轻度退化,中度退化和重度退化)草地植被和土壤理化特性的变化规律,为类似区域退化草地植被恢复提供有效途径。[方法]野外植被调查、土壤取样和室内分析。[结果]草地退化不同阶段草地植物群落组成和物种多样性均有差异,退化对草地土壤理化特性有明显影响。重度退化草地土壤容重显著高于轻度退化草地(p0.05)。轻度和中度退化样地0—10cm土壤空隙度显著高于重度退化草地。重度退化草地的土壤有机质、全碳、全钾、全磷和有效钾均明显小于轻度退化草地(p0.05),但土壤pH值和有效氮含量没有显著变化。[结论]高山草地退化演替对该区土壤物理特性具有显著影响。     

Effects of grassland abandonment, restoration and management on butterflies and vascular plants

When semi-natural pastures are abandoned, specialized grassland species are lost as a consequence of succession. As a counter-measure, previously abandoned grasslands may be restored by clearing shrubs and trees and re-introducing grazing livestock. In order to examine the effects of this type of habitat restoration, we compared species richness of plants and of specialized plants thought to be dependent on continuous management and species richness and abundance of butterflies and red-listed butterflies in 12 sets of matched continuously managed, abandoned and restored grassland in southern Sweden. We found no differences in species richness or abundance between the three grassland types. There were, however, some negative effects of abandonment. The number of management-dependent plants decreased with increasing cover of trees and shrubs, and in restored sites species richness of all groups decreased with increasing cover of trees and shrubs before restoration. Also the present management significantly affected both butterflies and plants. Species richness of both groups increased with increasing vegetation height and differed between sites depending on the species of grazers, with negative effects of sheep compared to cattle or horses. Our study indicates that for grassland management to be efficient, the restoration actions should mainly be directed towards sites where the post-abandonment succession has not proceeded too far.     

Increasing dominance of Brachypodium pinnatum (L.) beauv. in chalk grasslands: A threat to a species-rich ecosystem

Chalk grassland is a semi-natural, species-rich vegetation which was once widespread in West Europe. Due to the high species diversity a number of remnants of this vegetation have been set aside as nature reserves and managed by man to prevent natural succession into woodland.

During the last decade an increase of dominance of the grass Brachypodium pinnatum has been observed in chalk grassland within the greater part of the distribution area. Increasing dominance is strongly negatively correlated with the species diversity. Increase of this species is observed in grassland with different management regimes, e.g. sheep grazing, mowing or burning. Brachypodium can gain 80% or more of the relative above-ground phytomass. In such vegetation the species number decreases to less than 50% of the original number within a few years—the low growing forbs especially disappear almost completely. It is argued that the impact of nutrients by air pollution, especially of nitrogen, causes the increasing dominance of Brachypodium. Further research will focus on the effects of this extra nutrient supply.

The aim of the present research is to formulate management strategies in order to prevent degradation of the species diversity in chalk grasslands.     

Management effects on the vegetation and soil seed bank of calcareous grasslands: An 11-year experiment

Calcareous grasslands, most of which are man made and therefore depend on some kind of human interference for their maintenance, are among the most species-rich communities on Earth at a small scale. For many centuries, most of these grasslands have been used as extensive pasture. However, after 1900, and particularly from 1940 onwards, livestock grazing has declined throughout Europe leading to the abandonment of low intensity grasslands over large areas. To conserve the remaining grasslands or to restore recently abandoned grasslands, better insights about the effects of grassland management on above and belowground species diversity are needed. Here, we describe the results of an 11-year experiment to investigate the role of grassland management (grazing, mowing and abandonment) in determining species composition and diversity both in the aboveground vegetation and the seed bank of a calcareous grassland in Belgium. Species diversity declined by about 60% 11 years after abandonment, from 29 species m⁻² to as few as 12 species m⁻². Plots that were grazed remained constant in species richness, whereas mown sites lost about 20% of their original species. Abandoned plots were largely dominated by a few grass species, in particular Festuca rubra. Concomitant with changes in the aboveground vegetation, both the number of species found in the seed bank and seed density (number of seeds m⁻²) had changed significantly 11 years after abandonment. Species diversity and seed density were significantly lower in abandoned plots than in grazed or mown plots. We conclude that abandonment of calcareous grasslands may lead to rapid decline of plant species diversity both in the aboveground vegetation and in the seed bank. As a result, seed banks probably have a limited role to play in the restoration of recently abandoned grasslands.     

青藏高原三江源区不同恢复期高寒草甸土壤线虫群落演变

青藏高原三江源地区的高寒草甸面临着严峻的退化问题，人工建植是三江源地区退化草地的重要修复方式。为探究地下生物对草地人工恢复措施的响应，本研究比较了三江源地区高寒草甸不同恢复期人工草地(建植1、5和10 a)的土壤线虫群落变化。结果显示：与原生植被样地相比，所有恢复期样地的植物地下生物量降低，土壤容重、pH、全磷、全钾和硝态氮含量升高，表明人工草地系统的初级生产力和土壤特性尚未恢复到原生草地状态。不同恢复期样地中土壤线虫的均匀度和多样性指数均显著高于原生植被样地。此外，不同恢复期样地的线虫多度、代谢足迹以及成熟度指数均随恢复年限的增加而增加。相关性分析结果表明，土壤线虫多度与植物地下生物量、土壤有机质、全氮、全磷、矿质氮和速效氮含量显著正相关(P<0.05)，与土壤pH、全钾和容重显著负相关(P<0.05)。尽管三江源区退化草地的人工恢复措施尚未完全恢复牧草生产力至原生植被状态，但土壤生物结构及功能具有改善的趋势，显示该地区退化草地生态系统具有较强的恢复潜力。     

荒漠草原不同植被恢复模式的水土保持生态效益

[目的]以内蒙古四子王旗境内退化荒漠草原为研究区域,研究不同植被恢复模式对植物群落组成、土壤含水量、土壤侵蚀的影响,为退化草原水土流失治理和草原生态建设提供理论依据。[方法]针对退化草原特点、水土流失特征,分析不同植被恢复模式下植物群落特征、土壤水分、地表径流和侵蚀量的变化,运用主成分分析对不同植被恢复模式的水土保持生态效益进行评价。[结果]退化荒漠草原的3种植被恢复模式试验样地的植物种类组成、群落总盖度、地上生物量均高于放牧天然草地,"围栏封育+灌木条带+施肥复壮"、"围栏封育+松土补播+草种包衣"2种植被恢复模式的地上生物量与放牧天然草地之间变化差异达到显著水平(p0.05);3种植被恢复模式的实施有效控制了土壤侵蚀,径流量和土壤侵蚀量均低于天然草地。[结论]"围栏封育+松土补播+草种包衣"植被恢复模式在保持水土资源,增加群落稳定性等方面的生态效益最好。     

黄土区典型退化草地40年封禁恢复过程研究

天然草地退化引起草地植被和表层土壤遭到严重破坏,将使草地植被天然恢复过程受到强烈抑制。为此,试验采用长期(40年)封禁恢复技术,探究了封禁后黄土区退化草地植被变化特征,揭示了退化天然草地封禁恢复后的生态环境效应。结果表明：随着封禁恢复时间的变化,退化草地恢复演替已经历了5个时间段;草地生物量的变化过程已呈现出由先增加到后降低的趋势,尤其是在封禁恢复的第20年生物量达到最高值为520.5 g/m²;直到封禁恢复的第25年出现以本氏针茅为建群种的草原群落,第25年之后优势种大针茅种群密度急剧增加,在封禁的第30～34年以大针茅为群落的优势种生长繁殖较快,从群落恢复进程看大针茅有替代本氏针茅的趋势,但大针茅为有性繁殖,受草地凋落物的影响在群落中持续时间较短;从自然修复的35年后,通过种间竞争—扩散—定居—稳定生长的几个作用机制,又形成了以本氏针茅为建群种的“亚顶级”群落。此外,封禁后的草地物种丰富度和多样性逐渐提高。综上,封禁有利于黄土区草地生态系统恢复,且随着封禁时间的推演,草地生态系统逐渐趋于良性循环。     

黄土丘陵区典型草地演替中植物群落特征与土壤储水量关系

[目的]阐明草地植被演替过程中植被生产力、植物多样性等生态学特征与土壤储水量的关系,为探明黄土高原地区植被恢复的生态环境效应提供一定的科学依据。[方法]采用时空互代的方法对宁夏回族自治区固原市云雾山保护区自然恢复3,8,13,46,66,89a的样地进行取样,分析0—100cm土层土壤储水量的分布及其与地上地下植物生物量、物种多样性的关系。[结果]随着草地演替的进行,植被群落盖度、生物量和物种多样性指数在恢复13a之前显著增加,之后渐趋稳定;土壤含水量逐渐增加,容重逐渐降低。植被群落演替对0—40cm土层土壤储水量没有显著影响,但演替后期对40cm以下土层水分有明显消耗。植被群落生物量及物种多样性指标与表层0—10cm水分呈显著正相关。[结论]草地演替过程中,植被群落生物量和物种多样性的增加与表层土壤储水能力的提升密切相关,但深层根系生物量的增加对下层土壤储水的消耗也逐渐增大。     

Soil degradation and restoration as affected by land use change in the semiarid Bashang area,northern China

The Bashang area, located in the semiarid agropastoral zone, represents a typical degraded ecosystem under intensified human activities. This region has undergone profound land use changes during the past century. Natural grasslands had been progressively cultivated, and subsequently, part of cultivated lands had been abandoned by farmers due to severe desertification. In this study, we collected soil samples (0–20 cm depth) from four farmlands on both flat and gentle slope lands where cultivation had occurred for 0, 8, 30, and 50 years to assess the effects of grassland cultivation on soil degradation. In addition, soil samples were taken and plant species were investigated from eight sites in age sequence of 0- to 50- year-old abandoned field to assess natural restoration process following field abandonment. The results showed that cultivation of grassland result in a significant soil degradation which is manifested by the coarsening in soil texture and the losses in organic C and nutrients. After 50 years of cultivation, soil organic C, total N, and total P concentrations had declined 73–79%, 60–70%, and 67–68% in the 0–20 cm plough layer, respectively. Over half of these losses occurred during the first 8 years of cultivation, and subsequent was slow. After fields were abandoned, vegetation got self-restoration, and plant species composition moved toward that of the natural grassland community with time. Soil organic C, total N, and total P levels gradually improved with increasing years of land abandonment, with a faster restoration rate in the early vegetation recovery stage and a slower rate in the late succession stage. The results suggested that soil degradation may occurred drastically by inappropriate land use and management with a short time, while soil restoration for a degraded ecosystem may take long period of time, especially in this fragile ecosystem. From the perspective of soil resource management and desertification control, conservational tillage and management for farmlands and proper grazing management for recovered grassland should be taken into account in this area.     

宁南黄土丘陵区不同生态恢复模式对土壤养分的影响

[目的]分析植被恢复过程中土壤养分的变化规律,认识和评价植被生态系统功能恢复,促进植被演替和加快生态恢复的人工调控。[方法]通过对宁夏南部山区彭阳县中庄示范区天然草地(封山禁牧)、农耕地(退耕)、人工苜蓿(退耕还林草)3种生态恢复模式进行调查,研究分析不同生态恢复模式对土壤养分的影响。[结果]宁夏黄土丘陵区不同生态恢复模式效果依次为:天然草地农耕地人工苜蓿。随着土层深度的增加,土壤养分含量均呈现降低的趋势。随着植被恢复年限的延伸,土壤养分会逐渐累积而增加。不同恢复年限苜蓿地土壤肥力指数小于农耕地,土壤肥力贫瘠,且随苜蓿种植时间的延长,呈现先增大,再减小的趋势。[结论]在宁夏黄土丘陵区进行植被恢复,能明显提高土壤养分含量,改善土壤肥力状况,但旱作苜蓿粗放经营(只刈割,不培肥),导致土壤综合肥力指数日趋下降。     

不同植被恢复模式下三峡库区万州段消落带土壤养分及其空间分布特征

消落带植被状况与水位消涨是影响土壤养分及其分布的重要因子。本文以三峡库区万州段消落带为研究地点,通过对人工和自然恢复样地沿海拔梯度对土壤进行取样和分析,揭示了不同恢复模式下消落带土壤养分及其空间分布特征。研究结果表明,植被恢复模式对土壤养分有显著影响,人工恢复样地消落带主要土壤养分含量总体高于自然恢复样地,显示人工恢复能有效促进土壤养分在植物群落中的积累;在库区反季节水位消涨的作用下,人工恢复地消落带土壤养分沿海拔梯度呈先增加后减少的空间分布格局,土壤养分含量在消落带中部(海拔165 m)达到最高值,而自然恢复地土壤有机质和全氮的空间分布则随海拔梯度增加而增加,以消落带上部(海拔175 m)的值最高。土壤养分的空间分布格局源自于不同海拔梯度的消落带受水位消涨扰动程度以及植被恢复状况的差异,植被恢复模式对土壤养分的空间分布仍有一定影响。今后应在消落带上部进一步引种适宜的乔灌木物种,不断提高消落带植被对土壤养分的固持能力。     

三江源区草地退化与人工恢复对土壤理化性状的影响

以三江源区不同退化程度的天然草地及不同恢复年限的人工草地为研究对象,系统分析土壤理化性质随取样深度的动态变化。结果表明:(1)人工草地土壤含水量及全氮、全钾、全磷、速效钾、速效磷的含量随恢复年限均表现出先减小后增大的"V"形变化趋势;(2)随着退化程度增加,退化草地土壤中含水量逐渐降低,全氮、全钾、有机质、速效钾含量未表现出逐渐降低趋势,而草地的裸斑面积、土壤及养分的流失量逐渐增加;(3)人工草地及退化草地土壤中全氮、速效钾、有机质含量均随着取样深度增加而逐渐降低,草地恢复年限和退化程度基本不影响养分在土壤中的空间分布;(4)人工草地建设使退化草地有机质、全氮、全钾的含量增加,尤其有利于0-4cm的土壤养分增加。因而,人工草地建设可以作为三江源区退化草地土壤恢复的措施之一。     

Foliar, Physiologial and Growth Responses of Four Maple Species Exposed to Ozone

In order to restore the forest ecosystem in the vicinity of an industrial park, Ulsan, southeastern Korea, which has been heavily acidified by air pollution, a preliminary experiment by applying tolerant plants selected through several procedures, and dolomite and sewage sludge as soil ameliorators was carried out. Furthermore, a restoration based on the results was executed and the effects were evaluated based on the creation of safe sites, where new species can establish: regeneration of the forest with species similar in composition to the natural vegetation of native forests that are distant from the industrial park; increase in species diversity. In a preliminary study, the necessity of soil amelioration was diagnosed. Quercus serrata, Alnus firma and Ligustrum japonicum, which represent for tree, subtree, and shrub layers of vegetation in this region, were used as sample plants. Dolomite, sludge, and a mixture of both materials were applied as soil ameliorators. Bare ground (BG), and two grasslands dominated by forbs (GF) and grass (GG), respectively were designated as experimental plots based on a vegetation map of the corresponding area. BG and GF plots, which have lower organic matter contents, increased the growth of sample plants in response to soil amelioration, whereas that with higher contents, GG plot, did not show this response. The result suggests that necessity of soil amelioration depends on site quality. The effects of soil amelioration depended also on the sample plants. This difference is due to an ecological property of A. firma, which can fix atmospheric nitrogen through a symbiotic relationship with actinomycetic fungi. This result implies that this alder could be used as a substitute for soil ameliorators in restoration plan of this area. The height and standing crop of undergrowth, which forms dense grass mat and thereby impedes establishment of new plants, decreased in the restored stands. Such a decrease in the height and biomass of undergrowth could be recognized as providing safe sites, in which the other plants can invade, by removing the dense carpet formed by Miscanthus sinensis. The results of stand ordination showed a progression of the former bare grounds to either M. sinensis (GG) or Pueraria thunbergiana (GF) stands, suggesting a natural recovery through succession toward the stands dominated by both plants. But the change was not progressed beyond the grassland stage. Active restoration practice, which was carried out by applying tolerant plants, however, led to a change toward species composition similar to the natural vegetation before devastation. Furthermore, restored stands reflected the restoration effect by showing higher diversity than the stands in the degraded state. These results showed that the restorative treatment carried out by introducing tolerant plants functioned toward increasing both biological integrity and ecological stability and thereby could meet the restoration goal.     

Can a mesotrophic grassland community be restored on a post-industrial sandy site with compost made from waste materials?

Restoration of sites degraded by industry to species-rich semi-natural vegetation communities is difficult; it usually involves the addition of soil ameliorants but excessive fertility may favour dominance by competitive species. In a field-experiment we tested the establishment of a biodiverse mesotrophic grassland community using different compost types (comprising of mixtures of waste materials), application rates and seeding (with species in the target community). Compost addition to the alkaline sandy substrate increased soil organic matter, nutrient content and water holding capacity (WHC), whilst decreasing pH. Over the first two growing seasons compost addition, (especially at a higher rate) increased total vegetation cover (from <20% to a maximum of 67%), although the cover of the target community remained below 20%. Seeding with target species greatly increased their establishment on compost-treated plots, demonstrating its value for restoration of mesotrophic grassland communities in such sites lacking a local seed source. Five soil properties accounted for 46% of the variation in target species density: negative correlations with soil pH and %N, and positive correlations with electrical conductivity (EC), %C, and WHC. For this mesotrophic grassland community, high EC and WHC and low pH were most important for forb species and high %C for grasses. Overall, %C was the soil property that best explained variation in the early restoration success of different compost types and application rates; pH and EC were also correlated with the rate of vegetation establishment and available-P was linked to plant community composition. While a longer time period is needed to judge the sustainability of the outcome, this demonstrates the potential to refine compost properties for restoration of biodiversity.     

封育对退化草原植被恢复及土壤理化性质影响的研究进展

封育是退化草原生态系统恢复的有效措施之一。在梳理现有封育研究成果的基础上,归纳封育对植物多样性、生物量、植被演替、土壤物理与化学性状的影响效应,发现:受地理区域、气候条件及封育前退化程度等本底条件的影响,封育条件下植被多样性呈现出显著增加、基本稳定和明显下降3种结果,生物量则表现为围封初期增加,围封时间过长不利于维持较高生物量的变化趋势;封育使得种群拓殖能力与群落资源冗余,土壤种子库结构改善、密度增长,退化草原呈正向演替,并可能出现3种演替模式。封育消除了土壤紧实层,改善了土壤结构与性状,但对土壤养分的影响效应不一致。基于研究现状,提出今后应深入研究的6个内容与方向,对丰富封育条件下退化草原恢复效应具有指导意义。     

Density and success of bird nests relative to grazing on western Montana grasslands

Grassland birds are declining at a faster rate than any other group of North American bird species. Livestock grazing is the primary economic use of grasslands in the western United States, but the effects of this use on distribution and productivity of grassland birds are unclear. We examined nest density and success of ground-nesting birds on grazed and ungrazed grasslands in western Montana. In comparison to grazed plots, ungrazed plots had reduced forb cover, increased litter cover, increased litter depth, and increased visual obstruction readings (VOR) of vegetation. Nest density among 10 of 11 common bird species was most strongly correlated with VOR of plots, and greatest nest density for each species occured where mean VOR of the plot was similar to mean VOR at nests. Additionally, all bird species were relatively consistent in their choice of VOR at nests despite substantial differences in VOR among plots. We suggest that birds selected plots based in part on availability of suitable nest sites and that variation in nest density relative to grazing reflected the effect of grazing on availability of nest sites. Nest success was similar between grazed plots and ungrazed plots for two species but was lower for nests on grazed plots than on ungrazed plots for two other species because of increased rates of predation, trampling, or parasitism by brown-headed cowbirds (Molothrus ater). Other species nested almost exclusively on ungrazed plots (six species) or grazed plots (one species), precluding evaluation of the effects of grazing on nest success. We demonstrate that each species in a diverse suite of ground-nesting birds preferentially used certain habitats for nesting and that grazing altered availability of preferred nesting habitats through changes in vegetation structure and plant species composition. We also show that grazing directly or indirectly predisposed some bird species to increased nesting mortality. Management alternatives that avoid intensive grazing during the breeding season would be expected to benefit many grassland bird species.     

广西喀斯特地区植被演替对土壤质量的影响

以植被演替空间序列代替时间序列的方法,在野外选取广西喀斯特地区4个主要植物群落演替阶段即草地、灌草地、灌丛、乔林的典型样地,对土壤物理、化学和生物学性状进行了比较研究,以探讨植被演替对土壤质量的影响。结果表明:不同的植被演替导致了土壤物理、化学和生物学性质的显著差异。随演替的正向进行,土壤综合肥力指标值(Q I)呈增长趋势,其土壤质量综合指数分别为:草地(0.09)、灌草地(0.16)、灌丛(0.61)、乔林(0.89)。自然植被的正向演替是提高土壤质量的有效途径。导致草地和灌草地土壤质量相对较低的主要因素是植物生产力较低,土壤养分积累较少,土壤有机质等养分含量较低,土壤微生物数量较少,土壤酶活较低。土壤微生物数量结合土壤酶是反映土壤生物学活性和土壤质量的较好指标。