Woody plant encroachment impacts on soil carbon and microbial processes: results from a hierarchical Bayesian analysis of soil incubation data

Quantitative methods were used to examine soil properties and their spatial heterogeneity in a 0-year fenced mobile dune (MD0), an 11-year fenced mobile dune (MD11) and a 20-year fenced mobile dune (MD20) in Horqin Sandy Land, Northern China. The objective of the study was to assess the effect of vegetation restoration on heterogeneity of soil properties in sand dunes and to provide a concept model to describe the relationship between vegetation succession and spatial heterogeneity variation of soil properties in the dunes. The results showed that the average values of vegetation cover, species number and diversity, soil organic carbon (C), total nitrogen (N), and electrical conductivity (EC) increased with the increase in fenced age of mobile dunes, while soil water content (0–20 cm) showed the reverse trend. Geostatistical analysis revealed that the spatial heterogeneity of soil organic C, total N, EC, very fine sand content, and soil water content (0–20 cm) increased from MD0 to MD11 with succession from sand pioneer plant to shrub species then decreased from MD11 to MD20 due to continuous development of herbaceous plants. Canonical correspondence analysis (CCA) showed that there was a relatively high correspondence between vegetation and soil factors, suggesting that the major gradients relating soil organic C, total N, EC, pH, slope, very fine sand content, and soil water content are the main factors for the distribution of dune plants and account for 68.1% of the species-environment relationship among the three sites. In addition, the distribution of the sand pioneer plant was positively related to the relative height of the sampling site and soil water content, and that of most herbaceous plants were determined by soil organic C, total N, EC, pH, and very fine sand content in mobile dunes. The conceptual model of relationship between vegetation succession and spatial heterogeneity of soil properties in mobile dunes suggests spatial patterns of soil properties are most strongly related to plant-induced heterogeneity in dune ecosystems prone to wind erosion, and conversely, the magnitude and degree of spatial heterogeneity in soil properties can influence the plant distribution pattern and vegetation succession of mobile dunes.     

Tricholoma matsutake in a natural Pinus densiflora forest: correspondence between above- and below-ground genets, association with multiple host trees and alteration of existing ectomycorrhizal communities

Tricholoma matsutake (matsutake) is an ectomycorrhizal (ECM) fungus that produces economically important mushrooms in Japan. Here, we use microsatellite markers to identify genets of matsutake sporocarps and below-ground ECM tips, as well as associated host genotypes of Pinus densiflora. We also studied ECM fungal community structure inside, beneath and outside the matsutake fairy rings, using morphological and internal transcribed spacer (ITS) polymorphism analysis. Based on sporocarp samples, one to four genets were found within each fairy ring, and no genetic differentiation among six sites was detected. Matsutake ECM tips were only found beneath fairy rings and corresponded with the genotypes of the above-ground sporocarps. We detected nine below-ground matsutake genets, all of which colonized multiple pine trees (three to seven trees per genet). The ECM fungal community beneath fairy rings was species-poor and significantly differed from those inside and outside the fairy rings. We conclude that matsutake genets occasionally establish from basidiospores and expand on the root systems of multiple host trees. Although matsutake mycelia suppress other ECM fungi during expansion, most of them may recover after the passage of the fairy rings.     

Fairy rings,associated fungi,and assessment of their distribution across environmental variables using GIS

Fairy rings are conspicuous features of the Laramie Basin of southeastern Wyoming. Ten fairy ring sites chosen for intensive study were located using a small plane and 415 whole and partial fairy rings were georeferenced and ground-truthed. Fairy ring forming fungi were identified using a combination of taxonomy and ITS fungal barcode sequence comparisons. Precipitation, vegetation, soil type, elevation, slope, and slope aspect were queried using GIS. Eight fairy ring forming species were identified from the sites, including three unknown species, and two taxa not previously reported to form fairy rings. Most fairy rings occurred in drier mixed-grass prairie vegetation, whereas a small percentage occurred in mesic shrub dominated riparian vegetation. Most fungal species exhibited preferences for elevation and slope aspect, and rings were scarce on east facing slopes. The identity and autecology of individual fairy ring fungi and importance of environmental variables to fairy ring biology is discussed.     

Forest age correlates with fine-scale spatial structure of Matsutake mycorrhizas

Examining the fine-scale spatial structure of fungal populations can tell us much about how individual species reproduce and disperse throughout natural landscapes. Here we study the fine-scale genetic structure of Tricholoma matsutake, a prized edible and medicinal mushroom, by systematic sampling of mycorrhizas within fairy rings in 50-y-old and old-growth forests in two villages. Using single nucleotide polymorphism DNA markers we show that mycorrhizas in both forest age classes in both villages showed high levels of genotypic diversity, consistent with a reproductive life history predominated by outcrossing via basidiospore dispersal. Both the percentage of polymorphic loci within fairy rings, as well as genotype diversity were higher in old-growth compared to 50-y-old forests. Fifty-year-old forests showed significant spatial autocorrelation between pairs of mycorrhizas up to 42 m, and a pattern consistent isolation-by-distance structure. Spatial patterns in old-growth forests were random. Furthermore, AMOVA analysis indicates that 11 % of molecular variance in 50-y-old forests is partitioned between villages, whereas no significant variance is partitioned between villages in old-growth forests. We conclude that populations of T. matsutake in 50-y-old forests are the result of a founder effect maintained by local inoculation sources. This pattern attenuates as forests age and accumulate inocula from more distance sources. We speculate on how genetic mosaicism within T. matsutake fairy rings may structure populations within a chronosequence. Finally, we discuss how population spatial dynamics and dispersal strategy in T. matsutake contrast with other ectomycorrhizal species.     

Edaphic properties enable facilitative and competitive interactions resulting in fairy circle formation

Millions of generally regularly spaced, roughly circular barren patches called fairy circles occur in a narrow band ca 100 km inland of the south‐west African coast. These generally have conspicuously taller peripheral grasses in a shorter grass matrix. The origins of these fairy circles are controversial, but one possibility is that they are self‐organizing emergent vegetation patterns that are the consequence of interplay between positive (facilitative) and negative (competitive) interactions between grasses. We hypothesized that the coarse textured sand on which fairy circles occur creates a hydraulically and nutritionally connected landscape, in which neighbouring fairy circles competitively influence each other over several metres, while providing opportunity for focusing of resources around the peripheral grasses. To test our hypotheses we conducted three main groups of analyses: 1) we measured grass biomass to assess facilitative and competitive effects of the component grasses; 2) across a region with fairy circles we measured the size and density of fairy circles and correlated that with water infiltration rates into soil; 3) we measured the capacity of soil to conduct water pulses and ¹⁵N tracers. We found evidence of facilitative interactions in the periphery of the fairy circles and competitive suppression of the matrix grass proximal to the periphery. Across the region, fairy circle size was positively correlated with soil infiltration rates and negatively with precipitation. This suggests that fairy circles emerge in soils with high capacity for water flux that enables landscape hydraulic connectivity. Water‐ and ¹⁵N‐pulse experiments showed that edaphic resources were highly mobile, moving up to 7.5 m over a period of 1–3 weeks. We concluded that the evidence is consistent with an emergent vegetation pattern explanation for the origins of fairy circles and that the circles are more closely associated with a highly connective edaphic environment, rather than with particular biota.     

沙鞭(禾本科)种子在沙丘上的萌发策略及幼苗的耐干燥特性

沙鞭是鄂尔多斯高原流动沙丘上分布的多年生禾草。对位于沙层不同深度的种子萌发和幼苗出土进行了测定。结果表明 ,种子的萌发和幼苗的出土随着沙埋深度的增加而减少 ,最高的幼苗出土率来自 0 .5～ 2 .0 cm浅层沙埋的种子 ,并且沙埋的深度影响种子萌发的时间。沙埋越深 ,越多的种子不能萌发并且处于强迫休眠的状态。但是 ,当被埋较深种子上层的沙层被移走使种子位于较浅的沙层时 ,种子能够克服强迫休眠。种子的强迫休眠还能够通过改善沙层中的空气含量来克服。在自然生境中 ,沙丘的流动性能够使萌发后的幼苗整个暴露在空气中遭受干旱胁迫 ,或者萌发后的幼苗在因无后续的降雨而变干。但是 ,沙鞭在幼苗的早期具有忍受干燥 ,在下次水合后能够恢复生长的能力。幼根的长度和干燥时间影响幼苗的生长恢复率     

Spiral and Rotor Patterns Produced by Fairy Ring Fungi

A broad class of soil fungi form the annular patterns known as ‘fairy rings’ and provide one of the only means to observe spatio-temporal dynamics of otherwise cryptic fungal growth processes in natural environments. We present observations of novel spiral and rotor patterns produced by fairy ring fungi and explain these behaviors mathematically by first showing that a well known model of fairy ring fungal growth and the Gray-Scott reaction-diffusion model are mathematically equivalent. We then use bifurcation analysis and numerical simulations to identify the conditions under which spiral waves and rotors can arise. We demonstrate that the region of dimensionless parameter space supporting these more complex dynamics is adjacent to that which produces the more familiar fairy rings, and identify experimental manipulations to test the transitions between these spatial modes. These same manipulations could also feasibly induce fungal colonies to transition from rotor/spiral formation to a set of richer, as yet unobserved, spatial patterns.     

Structure, Pattern and Mechanisms of Formation of Seed Banks in Sand Dune Systems in Northeastern Inner Mongolia, China

A comparison of structure and pattern of the soil seed bank was made between active and stabilized sand dunes in northeastern Inner Mongolia, China. The objective of this paper was to determine the significance of seed bank in vegetation restoration of sand dunes. The results showed that (1) average seed density decreased from stabilized sand dune to interdune lowland of stabilized sand dune, to interdune lowland of active sand dune, and to active sand dune; (2) horizontally, along the transect from interdune lowland to ecotone and to sand dune top, a ‘V’ shaped pattern was presented in the active dune system, and a reverse ‘V’ shaped pattern in the stabilized sand dune system; (3) vertically, the proportion (accounting for the total seeds) of seeds found in 0–20 mm soil profile decreased from stabilized sand dune to interdune lowland of stabilized sand dune, to interdune lowland of active sand dune, and to active sand dune. The same order was also found in 20–50 mm and 50–100 mm soil profiles; (4) the Sokal and Sneath similarity indices in the species-composition between soil seed bank and above-ground vegetation were ranked as: the stabilized sand dune (24%) > the interdune lowland of active sand dune (21%) > the interdune lowland of stabilized sand dune (18%) > the active sand dune (5%); and (5) vegetation restoration of active sand dunes depends on the dispersal of seeds from nearby plant communities on the interdune lowlands. Much effort must be made to preserve the lowlands, as lowlands are the most important seed reservoir in the active sand dune field.     

Fleshy fungi in grass fields. Dependence on fertilization, grass species, and age of field

An experimental field with grass had 48 squares which were treated with combinations of NPK fertilizers. The fungus flora was studied 1978–1980 by numerous yearly observations. A total of 40 species were noted: 36 agarics and 4 gasteromycetes. 24 species were confined to squares where nitrate was applied. 5 of these species were recorded as fairy ring formers. The fairy rings developed exclusively in the +N squares. Some common species grew in both +N and –N squares, while a few species were confined to –N squares. None of the latter group was found in abundance. The squares were divided in parcels sown with 4 species of lawn grasses. Several fungi showed preference for one or two grass species. The field consists of two parts, sown in 1977 and 1972 respectively. The immigration pattern into the new field is described.     

Sand Dune Restoration in North Brittany, France: A 10-Year Monitoring Study

Dunes, which account for 13% of the Ille et Vilaine north Brittany coast, France, were degraded by high tourist pressure, and they were restored from 1988 onward. Ten years after commencing work an assessment of the restoration was made on three dunes: Les Chevrets, L'Anse Du Guesclin, and Le Verger. Annual monitoring of the vegetation and dune morphology provided an opportunity to study the restoration process. The dune front, which is similar to a reference pioneer dune, lies in front of the mobile dune. The highest part of the dune is an advanced stage of mobile dune. The restoration of the fixed rear dune was accomplished neither in terms of vegetation composition nor in terms of species richness. The vegetation parameter study allows differentiation between the dune front, the dune summit, and the rear dune. The variation in species richness and floristic composition from one zone to another can be explained by abiotic factors such as salinity and the accretion of sand. The restoration was satisfactory in terms of the geomorphology. Marram grass is a good tool for restoring the topography, but it will take a very long time to restore the conservation value of the dune.     

Are Namibian “Fairy Circles” the Consequence of Self-Organizing Spatial Vegetation Patterning?

Causes of over-dispersed barren “fairy circles” that are often surrounded by ca. 0.5 m tall peripheral grasses in a matrix of shorter (ca. 0.2 m tall) grasses in Namibian grasslands remain mysterious. It was hypothesized that the fairy circles are the consequence of self-organizing spatial vegetation patterning arising from resource competition and facilitation. We examined the edaphic properties of fairy circles and variation in fairy circle size, density and landscape occupancy (% land surface) with edaphic properties and water availability at a local scale (<50 km) and with climate and vegetation characteristics at a regional scale. Soil moisture in the barren fairy circles declines from the center towards the periphery and is inversely correlated with soil organic carbon, possibly indicating that the peripheral grass roots access soil moisture that persists into the dry season within fairy circles. Fairy circle landscape occupancy is negatively correlated with precipitation and soil [N], consistent with fairy circles being the product of resource-competition. Regional fairy circle presence/absence is highly predictable using an empirical model that includes narrow ranges of vegetation biomass, precipitation and temperature seasonality as predictor variables, indicating that fairy circles are likely a climate-dependent emergent phenomenon. This dependence of fairy circle occurrence on climate explains why fairy circles in some locations may appear and disappear over time. Fairy circles are only over-dispersed at high landscape occupancies, indicating that inter-circle competition may determine their spacing. We conclude that fairy circles are likely to be an emergent arid-grassland phenomenon that forms as a consequence of peripheral grass resource-competition and that the consequent barren circle may provide a resource-reservoir essential for the survival of the larger peripheral grasses and provides a habitat for fossicking fauna.     

不同沙丘生境主要植物比叶面积和叶干物质含量的比较

研究了生长在不同沙丘生境中 (流动沙丘 ,半固定沙丘和固定沙丘 ) 2 0个植物种 (10个 1年生植物种和 10个多年生植物种 )的比叶面积 (SL A)和叶干物质含量 (L DMC)的变化 ,并且分析了各个沙丘生境的土壤养分特征。结果表明 ,各个植物种的平均 SL A和 L DMC在植物种之间差异显著 ;多数在两种或 3种沙丘生境均有分布的植物其 SL A在不同沙丘生境之间差异显著 ,但是仅有 6个植物种的 L DMC在不同沙丘生境之间表现出差异 (p<0 .0 5 )。与许多研究结果类似 ,1年生植物的 SL A显著大于多年生植物的 SL A,而且两者之间 L DMC存在一定的差异。 1年生植物 SL A和 L DMC之间相关性不显著 ,但多年生植物SL A和 L DMC之间呈显著负相关。综合所有 2 0个植物种可以发现 ,SL A增大时 ,L DMC有下降的趋势     

Allozyme Variance and Clonal Diversity in the Rhizomatous Grass Psammochloa villosa(Gramineae)

Psammochloa villosa (Trin.) Bor (Gramineae) is a rhizomatous grass with characteristic sand fixation. Ten enzymes encoded with 15 allozymic loci were used to examine the genetic structures of four natural populations in Inner Mongolia from different habitats. Electrophoretie data indicated that P. viUosa had lower genetic variation (He = 0.104) and lower clonal diversity (D = 0.764) than other similar plants. Genetic compositions of the 4 populations were different. In Shilongmiao mobile dune and Shihuimiao fixed dune there were more genotypes and higher levels of variation than in Shilongndao fixed dune and Shihuimiao semi-mobile dune. Just hke other clonal plants, most genetic variation of P. villosa resided among populations ( Gst = 62,16% ). Most genotypes were local, and there were no spread genotypes. Factors contributed to the genetic structure are herein discussed.     

根茎禾草沙鞭的等位酶变异及克隆多样性

沙鞭（Ｐｓａｍｍｏｃｈｌｏａｖｉｌｏｓａ（Ｔｒｉｎ．）Ｂｏｒ）是禾本科的一种沙生旱生植物,靠根茎进行无性生殖。采用１０种酶系统共１５个基因位点对中国科学院植物研究所内蒙古伊克昭盟鄂尔多斯沙地草地生态研究站的４个天然居群进行了遗传结构和克隆多样性分析。结果表明,和同类植物相比,沙鞭的遗传变异水平和克隆多样性均较低。沙鞭４个居群的遗传组成明显不同。石龙庙流动沙丘和石灰庙固定沙丘的基株数目多,遗传变异水平较高,而石龙庙固定沙丘和石灰庙半流动沙丘的基株数目少,只有１株或２株,且多样性水平极低。环境差异在沙鞭的等位酶变异上未得到反映。和其它克隆植物相似,沙鞭的大部分变异存在于居群间（ＧＳＴ＝６２．１６％）,居群内变异所占比例较少。沙鞭９１．６７％的基因型属地方型,无广布基因型。     

Spiral vegetation patterns in high-altitude wetlands

When plant communities suffer the stress of limited resources, for instance adverse environmental conditions such as extreme aridity, the spatial homogeneity of the biomass is lost and self-organized patterns may arise. Here, we report the observation of spiral-shaped patterns in the biomass of grass (genus deyeuxia), under highland arid conditions in the north of Chile. The spiral arms are a few meters long and a few centimeters wide. These dynamic structures are observed in the grazing area of an herbivore member of the South American camelids, the vicuna, on the border of highland wetlands. These spirals cannot be explained by the well-established mathematical models which describe other vegetation patterns (that emerge from a Turing-type of instability) such as stripes, rings, or fairy circles. We attribute the formation of spirals to the coupling between the growth of vegetation in semiarid regions and the grazing of vicunas. The mathematical analysis of this coupling reveals an excitable behavior, i.e. small perturbations of the equilibrium generate large trajectories before coming back, that is the origin of the spirals.     

新疆准噶尔盆地南缘土壤生物结皮中地衣物种组成和分布

对新疆准噶尔盆地沙漠南缘土壤生物结皮中地衣种类及分布特征进行了初步研究。结果表明,该地区参与土壤生物结皮的有藓生双缘衣（Diploschistes muscorum（Schreb．）Norm．]、拟橙衣（Fulgensia bracteata（Hoffm．）Raws．]、丽石黄衣[Xanthoria elegans（Link）Th．Fr]、准噶尔橙衣（Caloplaca songoricumA．Abbas）和磷网衣[Psora decipiens（Ehrh．）Hoffm．]等13种地衣,分别属于5目10科。对其分布特征进行了分析,发现地衣的分布与苔藓有相关性;地衣的分布受到环境温度、太阳辐射强度和降水量等因素的影响。聚类分析结果表明,24个样方可分为3组,分别分布于沙丘底部、顶部和中部,其中分布于沙丘底部的样方地衣种类的多样性最高。     

Form and function of grass ring patterns in arid grasslands: the role of abiotic controls

Ring-shaped growth patterns commonly occur in resource-limited arid and semi-arid environments. The spatial distribution, geometry, and scale of vegetation growth patterns result from interactions between biotic and abiotic processes, and, in turn, affect the spatial patterns of soil moisture, sediment transport, and nutrient dynamics in aridland ecosystems. Even though grass ring patterns are observed worldwide, a comprehensive understanding of the biotic and abiotic processes that lead to the formation, growth and breakup of these rings is still lacking. Our studies on patterns of infiltration and soil properties of blue grama (Bouteloua gracilis) grass rings in the northern Chihuahuan desert indicate that ring patterns result from the interaction between clonal growth mechanisms and abiotic factors such as hydrological and aeolian processes. These processes result in a negative feedback between sediment deposition and vegetation growth inside the bunch grass, which leads to grass die back at the center of the grass clump. We summarize these interactions in a simple theoretical and conceptual model that integrates key biotic and abiotic processes in ring formation, growth and decline.     

Coastal plant and soil relationships along the southwestern coast of South Korea

We studied how plant species distribution was regulated by the relationships between vegetation and soil factors on the southwestern coast of South Korea. Vegetation was classified using two-way indicator species analysis (TWINSPAN), thereby producing four vegetation groups that were linked to three habitat types. Two ordination techniques —; detrended correspondence analysis (DCA) and canonical correspondence analysis (CCA) —; were applied to examine the relationships between vegetation and 12 edaphic factors, including soil pH, water and osmotic potentials, moisture content, electrical conductivity, Cl and Na⁺ contents, total Kjeldahl nitrogen, and contents of organic matter, sand, silt, and clay. Results were similar for both types of evaluations. According to DCA and CCA, the 23 communities tended to cluster into three types: salt swamp, salt marsh, and sand dune. The first two canonical axes accounted for 14.9% of the community-soil factor relationship among communities. As identified via CCA, the main gradients were soil-water relations and soil texture.     

沙丘微地形生境蚁丘分布特征及其对恢复过程的响应

为了阐明沙丘微地形生境蚁丘特征及分布规律对沙丘固定过程的响应规律,以科尔沁沙地为研究区,选择不同固沙阶段沙丘的4个方位（西北、东南、西南、东北）布设调查样地,测定了不同样地的蚁丘密度、直径和高度,分析了不同样地蚁丘特征及空间分布规律。结果表明：（1）流动沙丘背风向（即东南方位）蚁丘密度显著高于其他方位,但从流动沙丘到半流动沙丘,背风向（即东南方位）蚁丘密度显著低于其他方位;从半固定沙丘开始,半固定和固定沙丘微地形对蚁丘密度分布的影响较小。（2）在流动沙丘、半流动沙丘、半固定沙丘,不同微地形生境中蚁丘直径和高度均呈现出相似的分布规律,即东南方位均小于其他方位。在固定沙丘,蚁丘高度表现为东南方位亦较低,但蚁丘直径表现为西南和东北方位显著小于西北方位。（3）在流动沙丘,西北、西南和东南方位上蚁丘空间分布均表现为均匀分布型,而在半流动、半固定和固定沙丘,4个方位上蚁丘空间分布均表现为聚集分布型。（4）相关性分析表明,不同固沙阶段不同微生境下蚁丘密度、高度、直径与植物密度、多样性间的相关性不同。随着固沙阶段演替,不同微生境下蚁丘特征与植物密度、多样性的相关性增强。研究表明,在科尔沁沙地,随着流动沙丘固定和生境逐渐恢复,沙丘不同空间方位上蚁丘密度分布差异缩小,而蚁丘直径和高度仍存在沙丘微地形间显著差异性,并且蚁丘空间方位上均呈现聚集分布状态。     

Woody-grass ratios in a grassy arid system are limited by multi-causal interactions of abiotic constraint, competition and fire

Predicting changes in vegetation structure in fire-prone arid/semi-arid systems is fraught with uncertainty because the limiting factors to coexistence between grasses and woody plants are unknown. We investigated abiotic and biotic factors influencing boundaries and habitat membership in grassland (Triodia or ‘spinifex’ grassland)-shrubland (Acacia aneura or ‘mulga’ shrubland) mosaics in semi-arid central Australia. We used a field experiment to test for the effects of: (1) topographic relief (dune/swale habitat), (2) adult neighbour removal, and (3) soil type (sand/clay) on seedling survival in three shrub and two grass species in reciprocal field plantings. Our results showed that invasion of the shrubland (swale) by neighbouring grassland species is negated by abiotic limitations but competition limits shrubland invasion of the grassland (dune). All species from both habitats had significantly reduced survival in the grassland (dune) in the presence of the dominant grass (Triodia) regardless of soil type or shade. Further, the removal of the dominant grass allowed the shrubland dominant (A. aneura) to establish outside its usual range. Seedling growth and sexual maturation of the shrubland dominant (A. aneura) was slow, implying that repeated fire creates an immaturity risk for this non-sprouter in flammable grassland. By contrast, rapid growth and seed set in the grassland shrubs (facultative sprouters) provides a solution to fire exposure prior to reproductive onset. In terms of landscape dynamics, we argue that grass competition and fire effects are important constraints on shrubland patch expansion, but that their relative importance will vary spatially throughout the landscape because of spatial and temporal rainfall variability.