Shrub-encroachment induced alterations in input chemistry and soil microbial community affect topsoil organic carbon in an Inner Mongolian grassland

Shrub encroachment frequently occurs in arid and semi-arid grasslands worldwide and affects the regional carbon balance. Many previous studies have revealed the effects of shrub encroachment on bulk carbon content of grasslands, but molecular evidence is surprisingly lacking. In this study, we examined the chemical composition of plant tissues and soil organic carbon (SOC), and soil microbial communities to identify the effects of shrub (Caragana microphylla) encroachment on SOC storage in the top layer (0–10 cm) along a gradient of natural shrub cover in the grasslands of Inner Mongolia. We found that SOC in the shrub patches was derived mainly from leaves, whereas SOC in the grassy matrix was composed of a mixture of fresh root- and leaf-derived compounds. Compared with pure grassland, the SOC decreased by 29% in the shrub-encroached grasslands (SEGs), and this decrease was enhanced by increasing shrub cover. We also found that free lipids and lignin-derived phenols increased while the ratios of ω-C₁₈/∑C₁₈ and suberin/cutin decreased with increasing shrub cover. In addition, the ratios of fungal to bacterial phospholipid fatty acids (PLFAs) and gram-negative to gram-positive bacterial PLFAs decreased with increasing shrub cover. These results indicate that the encroachment of nitrogen-rich legume shrubs can lead to carbon loss by altering the chemical composition of plant inputs as well as the soil microbial community in grassland ecosystems.     

Carabid beetles as indicators for shrub encroachment in dry grasslands

Semi-natural grasslands are key habitats for biodiversity conservation in Central Europe. Shrub encroachment is one of the most threatening drivers of grassland degradation and affects soil properties, microclimate, and vegetation with possible impacts on higher trophic levels. We aimed to analyse the impact of shrub encroachment with broom (Cytisus scoparius) on carabid beetle diversity, species composition, and functional traits. In a field study on dry grasslands on the island of Hiddensee (Germany) we studied 15 sites along a gradient of increasing broom encroachment and classified them into three dry grassland types with low, medium, and high shrub cover. Our results provide evidence that shrub encroachment initially has positive effects on species richness and activity densities of dry grassland carabids. Carabid species composition differed among differently shrub-covered dry grassland types, and sites with low and high shrub cover were each characterised by unique carabid assemblages. The species composition of sites with a medium shrub biomass had a transitional character and contained species which are typical for open dry grassland, but also shared species with sites with a high shrub cover. Among functional trait parameters investigated, especially the body size of carabid beetles was related to environmental parameters associated with shrub encroachment. Body size was positively correlated to shrub biomass and soil humidity, but negatively to temperature. Eurytopy values of carabids were related to high litter cover, i.e. habitat generalist (eurytopic) species mainly occurred in densely shrub-encroached sites. In order to preserve unique carabid assemblages of open dry grasslands with stenotopic and smaller species, it is most important to prevent a shrub encroachment higher than about 60% cover. For management we suggest extensive grazing (by cattle, sheep or horses) to prevent shrub encroachment on dry grasslands. In areas with high shrub cover additionally the use of goats or mechanical removal of shrubs might be necessary.     

Soil fungal composition changes with shrub encroachment in the northern Chihuahuan Desert

Woody species encroachment of grasslands globally causes many socioecological impacts, including loss of grazing pastures and decreased biodiversity. Soil microbial communities may partially regulate the pace of shrub encroachment, as plant-microbial interactions can strongly influence plant success. We measured fungal composition and activity under dominant plant species across a grassland to shrubland transition to determine if shrubs cultivate soil microbial communities as they invade. Specifically, soil microbial communities, abiotic soil properties, and extracellular enzyme activities were quantified for soils under four common Chihuahuan Desert plant species (three grasses, one shrub) in central New Mexico, U.S.A. Extracellular enzyme activity levels were fairly consistent under different plant species across the grassland to shrubland transition. Activity levels of two enzymes (alkaline phosphatase and beta-N-acetyl-glucosaminidase) were lower in the ecotone, presumably because soil organic matter content was also lower in ecotone soils. Community composition of soil fungi mirrored patterns in the plant community, with distinct plant and fungal communities in the shrubland and grassland, while grassland-shrubland ecotone soils hosted a mix of taxa from both habitats. We show that shrubs cultivate a distinct microbial community on the leading edge of the invasion, which may be necessary for shrub colonization, establishment, and persistence.     

Shrub encroachment alters composition and diversity of ant communities in abandoned grasslands of western Carpathians

Invasions of woody species into grasslands abandoned by agriculture are a global phenomenon, but their effects on diversity of other taxa have been rarely investigated across both regional and local scales. We quantified how shrub encroachment affected the activity, composition, and diversity of ant communities in managed and abandoned grasslands in western Carpathians of Central Europe across four regions and four shrub encroachment stages in each region. We surveyed ant communities on 48 sites in total, with each encroachment stage replicated three times in each region and twelve times overall. We used pitfall traps to sample ants over three years (2008, 2009, 2011) and identified 9,254 ant workers belonging to 33 species in total. Although the epigaeic activity and composition of ant communities varied with region, abandoned grasslands supported a greater species richness of ants than managed grasslands regardless of the region, and especially so in more advanced shrub encroachment stages. Since the woody colonization within grasslands was moderate even in the advanced encroachment stages (on average ~40 % of grassland colonized by woody species), it allowed coexistence of forest specialists (e.g. Temnothorax crassispinus) with species typical of open grasslands, thus increasing overall ant diversity. Managed grasslands were not only less species rich compared to abandoned grasslands, but they were characterized by different species (e.g. Lasius niger, Myrmica rugulosa). The differences in ant communities between managed and abandoned grasslands are likely to cause differences in ecological functions mediated by ants (e.g. predation of arthropods or plant seed dispersal).     

Analysis of macrofungal communities reveals a complex reciprocal influence between Mediterranean montane calcareous grassland and surrounding forest habitats

Fungi are essential components of all terrestrial ecosystems. Despite the crucial ecological role of soil fungi in grasslands, knowledge about fungal community diversity and structure in Mediterranean meadow habitats is still fragmentary. We analyzed macrofungal communities in three geographically distinct Mediterranean montane calcareous grasslands and surrounding forests, by means of fruit body surveys. We investigated a number of biotic and abiotic factors influencing the studied fungal communities, including plant species composition. Out of 6365 fruit bodies, a total of 268 species belonging to 84 genera were found. In general, there was a significant correlation between plant species richness and fungal richness. Variation in vegetation and plant community structure accounted for approximately 20% of variance in fungal community structure. Tree and shrub vegetation played a dominant role in shaping the analyzed fungal communities, both in meadows and surrounding forests, with particular influence on ectomycorrhizal, litter, and lignicolous saprotrophic fungi. Fungal biodiversity in the studied meadows was increased by the presence of tree and shrub species from the adjacent forests, but was reduced by the increasing vegetation cover.     

Species richness and composition of shrub-encroached grasslands in relation to environmental factors in northern China

Aims Shrub encroachment has taken place in many of China’s northern grasslands. This study attempts to answer the following questions: which plant communities are present in these shrub-encroached grasslands (SEGs)? What are the species richness and composition of these communities? Which environmental factors determine the spatial distribution thereof?     

Effects of grassland species on decomposition of litter and soil microbial communities

Decomposition of litter is greatly influenced not only by its chemical composition but also by activities of soil decomposers. By using leaf litter from 15 plant species collected from semi-natural and improved grasslands, we examined (1) how interspecific differences in the chemical composition of litter influence the abundance and composition of soil bacterial and fungal communities and (2) how such changes in microbial communities are related to the processes of decomposition. The litter from each species was incubated in soil of a standard composition for 60 days under controlled conditions. After incubation, the structure of bacterial and fungal communities in the soil was examined using phospholipid fatty-acid analysis and denaturing gradient gel electrophoresis. Species from improved grasslands had significantly higher rates of nitrogen mineralization and decomposition than those from semi-natural grasslands because the former were richer in nitrogen. Litter from improved grasslands was also richer in Gram-positive bacteria, whereas that from semi-natural grasslands was richer in actinomycetes and fungi. Nitrogen content of litter also influenced the composition of the fungal community. Changes in the composition of both bacterial and fungal communities were closely related to the rate of litter decomposition. These results suggest that plant species greatly influence litter decomposition not only through influencing the quality of substrate but also through changing the composition of soil microbial communities.     

Soil fungal community changes in response to long-term fire cessation and N fertilization in tallgrass prairie

In grasslands, fire management and fertilization are established drivers of plant community change, but associated soil fungal responses are less well defined. We predicted that soil fungal communities would change seasonally, that decades of fire cessation and nitrogen (N) fertilization would alter fungal distributions, and that plant and fungal community change would be correlated. Surface soils were sampled monthly for 1 y from a 30-y fire by fertilization experiment to evaluate fungal community dynamics and assess correlation with plant community heterogeneity. ITS gene community composition was seasonally stable, excepting increased arbuscular mycorrhizal fungal summer abundance in the burned, fertilized treatment. Long-term treatments affected soil fungal and plant communities, with correlated heterogeneity patterns. Despite woody encroachment in the fire cessation treatment, soil fungal communities did not resemble those of forests. This study provides evidence supporting the strength of feedbacks between fungal and plant community change in response to long-term grassland fire and N management changes.     

Flower visitor communities are similar on remnant and reconstructed tallgrass prairies despite forb community differences

One common goal of habitat restoration and reconstruction is to reinstate the biodiversity found at intact reference sites. However, few researchers have examined whether these practices reinstate communities of flower‐visiting insects. This is unfortunate, as anthropogenically mediated declines in flower visitors, including bees (the primary pollinators for most terrestrial ecosystems), beetles, flies, and butterflies, have been reported worldwide. Biodiversity declines may be especially severe in North America's tallgrass prairie, a once‐vast grassland that has experienced severe destruction and degradation due to agricultural conversion. As such, we assessed the structure of forb and flower‐visiting insect communities as a whole and two subsets of the flower visitor community—bees and phytophagous beetles—across five tallgrass prairie remnants and five reconstructed prairies (former crop fields) in Kansas from 2013 to 2015. Remnant prairies had significantly higher forb diversity and differed significantly in forb composition, compared to reconstructed prairies. Despite the dissimilarities in forb community structure, there were no differences in flower visitor diversity or abundance between remnants and reconstructed prairies. However, when considered separately, bee communities exhibited significantly greater variability in composition on reconstructed prairies, likely due to the abundance of generalist bee species visiting non‐native legumes at two reconstructed prairies. Our work provides evidence that prairie habitat reconstruction is a valuable tool for reestablishing flower‐visiting insect communities and also emphasizes the considerable role that non‐native species may play in structuring grassland plant–bee interactions.     

Life History Traits Reflect Changes in Mediterranean Butterfly Communities Due to Forest Encroachment

The biodiversity of the Southern Balkans, part of the Mediterranean global biodiversity hot-spot, is threatened by land use intensification and abandonment, the latter causing forest encroachment of formerly open habitats. We investigated the impact of forest encroachment on butterfly species richness, community species composition and the representation of life history traits by repeated seasonal visits of 150 one-hectare sites in five separate regions in three countries—Greece, Bulgaria, and the Republic of Macedonia (FYROM—the Former Yugoslav Republic of Macedonia)— 10 replicates for each habitat type of grasslands, open formations and scrub forest within each region. Grasslands and open formations sites hosted in average more species and more red-listed species than scrub forest, while no pattern was found for numbers of Mediterranean species. As shown by ordination analyses, each of the three habitat types hosted distinct butterfly communities, with Mediterranean species inclining either towards grasslands or open formations. Analysing the representation of life history traits revealed that successional development from grasslands and open formations towards scrub forest shifts the community composition towards species overwintering in earlier stages, having fewer generations per year, and inhabiting large European or Eurosiberian (e.g. northern) ranges; it decreases the representation of Mediterranean endemics. The loss of grasslands and semi-open formations due to forest encroachment thus threatens exactly the species that should be the focus of conservation attention in the Mediterranean region, and innovative conservation actions to prevent ongoing forest encroachment are badly needed.     

Legacy effects of afforestation on prairie plant and seed bank communities in a northern Canadian prairie

Afforestation resulting from fire suppression, modified grazing, plantation establishment and climate change poses a threat to northern prairie ecosystems. Trees alter the composition and function of plant and soil fauna communities and can compromise the restoration of afforested prairies. To evaluate the hypothesis that legacies of afforestation persist in restored prairie communities and decrease the potential for restoration, we examined the composition, structure, and diversity of plant and seed bank communities along a 20 year chronosequence of plantation tree removal from a northern fescue prairie in Riding Mountain National Park, Manitoba, Canada. Tree removal increased the abundance of weedy species in the plant and seed bank communities of restored prairies and plant diversity peaked and declined over the 20 year period of passive restoration. As a result, time since tree removal and the encroachment of invasive species were key in explaining the composition of restored prairie communities. Low correlation between the species composition of plant and seed bank communities, including the complete absence of Festuca hallii in restored treatments, demonstrated that legacies of afforestation compromised the potential of seed banks to facilitate prairie restoration. We conclude that tree removal alone is insufficient for the restoration of northern fescue prairies and that, in the absence of active management, the persistence of low-diversity plant and seed bank communities constitutes an important legacy of afforestation and an important barrier to future restoration.     

典型农田退耕后土壤真菌与细菌群落的演替

土壤真菌和细菌作为地下生态系统的重要组成部分,其群落的恢复状况是评价农田退耕还林生态效益的重要指标。以云南省维西县典型退耕还林农田为对象,利用高通量测序等方法比较了不同退耕年限的农田土壤中真菌和细菌群落随植被演替的变化特征。结果发现,农田撂荒后土壤细菌多样性先显著降低后缓慢上升,真菌多样性变化不明显;地上部植被由草本经灌丛再向林地演替的过程中,土壤真菌的群落组成随植被变化呈现明显的改变,主要体现在粪壳菌纲(Sordariomycetes)所占比例的减少(由30%减至10%左右)和伞菌纲(Agaricomycetes)所占比例的增加(由5%以下增至20%以上);而细菌的群落组成无明显变化。聚类分析的结果显示,真菌的群落组成变化与植物群落的演替规律更为同步。不同演替阶段的退耕农田土壤真菌和细菌群落均明显区别于未经扰动的天然林,表明人为扰动对土壤微生物群落的影响可能在较长时间内持续存在。研究揭示了云南典型农田退耕后地下土壤真菌和细菌群落随植被演替的变化特征,为全面评价该地区退耕还林的生态效益提供了数据支撑。     

Covariance between disturbance and soil resources dictates the invasibility of northern fescue prairies

Increasing environmental impacts of exotic organisms have refocused attention on the ability of diverse communities to resist biological invaders. Although resource availability, often related to natural and anthropogenic disturbances, appears central to the invasibility of biological communities, understanding the links between resources, diversity and invasibility is often confounded by the covariance among key variables. To test the hypothesis that community invasibility remains contingent on the type and intensity of disturbance and their impacts on plant community diversity and resource availability, we designed an experiment testing the invasibility of northern fescue prairies by smooth brome (Bromus inermis Leyss.), a Eurasian perennial grass, threatening the structure and function of prairie remnants throughout the Great Plains. Using soil disturbances and herbicide, we imposed treatments manipulating the diversity and resource availability of native prairies. Our observations demonstrate that the vulnerability of native prairies to exotic plant invasions remains contingent on resources. While the establishment of smooth brome seedlings increased with increasing disturbance, its impact depended on the availability of soil nitrogen. As a result, soil burial treatments, simulating disturbance by northern pocket gophers, provided poor recruitment areas for smooth brome, and their low levels of soil moisture and nitrogen, combined with the rapid recovery of the prairie community, compromised seedling establishment. Emphasizing the covariance of diversity and key environmental variables following disturbance, our findings illustrate the importance of disturbance type and intensity on community invasibility. Such a consideration is critical in the conservation and restoration of native prairie remnants throughout the Great Plains.     

Soil C and N responses to woody plant expansion in a mesic grassland

Changes in land management and reductions in fire frequency have contributed to increased cover of woody species in grasslands worldwide. These shifts in plant community composition have the potential to alter ecosystem function, particularly through changes in soil processes and properties. In semi-arid grasslands, the invasion of shrubs and trees is often accompanied by increases in soil resources and more rapid N and C cycling. We assessed the effects of shrub encroachment in a mesic grassland in Kansas (USA) on soil CO₂ flux, extractable inorganic N, and N mineralization beneath shrub communities (Cornus drummondii) and surrounding undisturbed grassland sites. In this study, a shift in plant community composition from grassland to shrubland resulted in a 16% decrease in annual soil CO₂ flux(4.78 kg CO₂ m^–2 year^–1 for shrub dominated sites versus 5.84 kg CO₂ m^–2 year^–1 for grassland sites) with no differences in total soil C or N or inorganic N. There was considerable variability in N mineralization rates within sites, which resulted in no overall difference in cumulative N mineralized during this study (4.09 g N m^–2 for grassland sites and 3.03 g N m^–2 for shrub islands). These results indicate that shrub encroachment into mesic grasslands does not significantly alter N availability (at least initially), but does alter C cycling by decreasing soil CO₂ flux.     

Short-Term Temporal Effects on Community Structure of Lepidoptera in Restored and Remnant Tallgrass Prairies

Understanding the degree to which species assemblages naturally vary over time will be critically important when assessing whether direct management effects or contingency is responsible for species gain or loss. In this study, we tested three predictions related to short‐term variation in prairie moth communities: (1) communities would only exhibit significant temporal variation in newly restored sites (1–3 years old); (2) prairie size and age would positively influence community reassembly, with larger, older restorations sampling a greater proportion of the regional species pool; and (3) older restorations (7–10 years old) would have yet to converge on the community composition of prairie remnants. Moths were sampled from 13 Tallgrass prairie restorations and remnants in central Iowa in 2004–2005. Repeated measures analysis of variance revealed significant effects of sampling year on moth species richness and abundance as well as on the richness of two functional groups, but difference among prairie types was only observed in 2005. Rarefaction analysis revealed that older restorations and prairie remnants supported higher species richness compared to recently planted sites, and nonmetric, multidimensional scaling ordination indicated that restorations older than 7 years were clearly converging on the species composition of remnants. These results suggest that moth communities in restorations and remnants are highly variable in time but that as restorations age, they appear to reaccumulate moth species found in prairie remnants. The long‐term persistence of a particular species assemblage within a given site, however, might be a difficult endpoint to attain in central Iowa prairies because of significant annual variation in species occurrence.     

Responses of soil bacterial and fungal communities to reciprocal transfers of soil between adjacent coniferous forest and meadow vegetation in the Cascade Mountains of Oregon

Little information exists on the responses of soil fungal and bacterial communities in high elevation coniferous forest/open meadow ecosystems of the northwest United States of America to treatments that impact vegetation and soil conditions. An experiment was conducted in which soil cores were reciprocally transplanted between immediately adjacent forests and meadows at two high elevation (∼1,600 m) sites (Carpenter and Lookout) in the H.J. Andrews Experimental Forest located in the Cascade Mountains of Oregon. Half of the cores were placed in PVC pipe (closed) to prevent new root colonization, whereas the other cores were placed in mesh bags (open) to allow recolonization by fine roots. A duplicate set of open and closed soil cores was not transferred between sites and was incubated in place. After 2 year, soil cores were removed and changes in fungal and bacterial biomasses determined using light microscopy, and changes in microbial community composition determined by PLFA analysis, and by length heterogeneity PCR of the internal transcribed spacer region of fungal ribosomal DNA. At both sites soil microbial community structures had responded to treatments after 2 year of incubation. At Carpenter, both fungal and bacterial community structures of forest soil changed significantly in response to transfer from forest to meadow, with the shift in fungal community structure being accompanied by a significant decrease in the PLFA biomarker of fungal biomass,18:2ω6,9. At Lookout, both fungal and bacterial community structures of forest soil changed significantly in response to open versus closed core treatments, with the shift in the fungal community being accompanied by a significant decrease in the 18:2ω6,9 content of closed cores, and the shift in the bacterial community structure being accompanied by a significant increase in bacterial biomass of closed cores. At both sites, fungal community structures of meadow soils changed differently between open and closed cores in response to transfer to forest, and were accompanied by increases in the18:2ω6,9 content of open cores. Although there were no significant treatment effects on the bacterial community structure of meadow soil at either site, bacterial biomass was significantly higher in closed versus open cores regardless of transfer.     

A multifaceted view on the impacts of shrub encroachment

Shrub encroachment, a global phenomenon with management implications, is examined in two papers in the current issue of Applied Vegetation Science. Barbosa da Silva et al. show that encroachment simplifies the herbaceous community, and Pittarello et al. illustrate how pastoral practices can restore encroached grasslands. While detrimental effects of shrub encroachment on grassland vegetation are often reported, we argue for a more holistic view when assessing this land‐cover change.     

Patterns of diversity and adaptation in Glomeromycota from three prairie grasslands

Arbuscular mycorrhizal (AM) fungi are widespread root symbionts that often improve the fitness of their plant hosts. We tested whether local adaptation in mycorrhizal symbioses would shape the community structure of these root symbionts in a way that maximizes their symbiotic functioning. We grew a native prairie grass (Andropogon gerardii) with all possible combinations of soils and AM fungal inocula from three different prairies that varied in soil characteristics and disturbance history (two native prairie remnants and one recently restored). We identified the AM fungi colonizing A. gerardii roots using PCR amplification and cloning of the small subunit rRNA gene. We observed 13 operational taxonomic units (OTUs) belonging to six genera in three families. Taxonomic richness was higher in the restored than the native prairies with one member of the Gigaspora dominating the roots of plants grown with inocula from native prairies. Inoculum source and the soil environment influenced the composition of AM fungi that colonized plant roots. Correspondingly, host plants and AM fungi responded significantly to the soil–inoculum combinations such that home fungi often had the highest fitness and provided the greatest benefit to A. gerardii. Similar patterns were observed within the soil–inoculum combinations originating from two native prairies, where five sequence types of a single Gigaspora OTU were virtually the only root colonizers. Our results indicate that indigenous assemblages of AM fungi were adapted to the local soil environment and that this process occurred both at a community scale and at the scale of fungal sequence types within a dominant OTU.     

草原灌丛化对生态系统多功能性的影响

草原灌丛化现象在干旱半干旱区广泛发生,影响了生态系统的结构、过程和功能。生态系统具有同时提供多种功能的能力,即生态系统多功能性。灌丛化是否会引起草原生态系统多功能性的减少,其内在的作用机制又是什么？这些问题仍有待明晰。理解草原灌丛化对生态系统多功能性的影响,对于促进草原地区"草-畜-人"平衡和实现区域可持续发展至关重要。从响应规律、影响路径和控制因素三个方面总结评述了草原灌丛化对生态系统多功能性影响的研究进展,主要包括:(1)阐明了单一生态系统功能和多种生态系统功能对草原灌丛化的响应特征;(2)从生物路径、非生物路径以及气候变化和人类活动的影响方面探讨了灌丛化对生态系统多功能性的影响路径;(3)从灌丛化物种、灌丛化阶段和草原类型三个方面明晰了草原灌丛化对生态系统多功能性影响的控制因素。在此基础上,针对灌丛化对生态系统多功能性的影响机制,对生产-生态功能权衡的影响等方面对未来研究进行了展望,并面向可持续发展目标探讨了灌丛化生态系统的可持续管理路径。研究可为我国灌丛化草原的恢复和管理提供支撑。     

毛竹种植对土壤细菌和真菌群落结构及多样性的影响

为揭示天然林改为毛竹林过程中土壤微生物变化规律,在浙江省湖州市安吉县和长兴县两地选择不同种植历史的粗放经营毛竹林,分层采集0～20和20～40 cm的混合土壤样品,应用PCR-DGGE技术分析土壤细菌和真菌群落结构及多样性变化.结果表明: 在马尾松林改种毛竹林或毛竹林入侵杂灌阔叶林形成毛竹纯林过程中,土壤细菌和真菌的群落结构均发生明显变化,且细菌结构对毛竹种植的响应更敏感;随着毛竹生长时间的延长,表层土壤细菌群落表现出抵抗干扰、最后向改种毛竹之前状态恢复的趋势.毛竹种植时间、样地和土层均对土壤细菌和真菌多样性产生显著影响,其中样地和土层的影响明显大于种植时间.土壤性质和细菌、真菌结构的冗余分析结果表明,不同地点、不同土层驱动土壤微生物结构随时间变化的主要因子没有一致规律,且第1、2轴对样地变化的解释率大多低于65.0%,说明除本研究分析的5个土壤化学指标外,可能还有其他土壤理化性质共同驱动微生物结构的变化.