Temporal Asynchrony in Soil Nutrient Dynamics and Plant Production in a Semiarid Ecosystem

A central goal of ecosystem ecology is to understand how the cycling of nutrients and the growth of organisms are linked. Ecologists have repeatedly observed that nutrient mineralization and plant production are closely coupled in time in many terrestrial ecosystems. Typically, mineralization rates of limiting nutrients, particularly of nitrogen, during the growing season determine nutrient availability while pools of mineral nutrients remain low and relatively constant. Although several previous reports suggest nitrogen mineralization has the potential to vary seasonally and out of phase with plant production, such a phenomenon has been poorly documented. Here we report results from a semiarid savanna ecosystem characterized by distinct temporal asynchrony in rates of soil nitrogen cycling and plant production. Periods of positive plant growth following the onset of rains coincide with periods of low N turnover rates, whereas higher rates occur late in the wet season following plant senescence and throughout dry seasons. Plant uptake from the substantial mineral N pool present early in the growing season is sufficient to explain most of the N allocation to aboveground plant biomass during the growing season, even in the absence of any wet-season mineralization. The mineral N pool is subsequently recharged by late wet- and dry-season mineralization, plus urine inputs at sites with high levels of ungulate activity. These findings suggest fundamental changes in the quality of substrates available to decomposers over a seasonal cycle, with significant implications for the partitioning of limiting nutrients by plant species, the seasonal pattern of nutrient limitations of aboveground production, and the effective use of N fertilizers in semiarid ecosystems.     

放牧强度对草甸草原生产力和多样性的影响

采用比较样地法研究了不同放牧强度对羊草草甸草原群落多样性和生产力的影响。结果表明:α多样性指数随放牧强度的增加表现为先增加后降低的趋势,其值在轻度放牧区最大,支持了"中度干扰理论";β多样性指数随放牧强度的增加而变大;不同放牧强度的草地植物种类相似性变差、草地地上净初级生产力与Alatalo均匀度指数具有显著的线性相关性(P0.05);放牧和气候对草地地上净初级生产力均有较大的影响,但二者的互作效应不显著(P0.05);不放牧不能持续维持草地的健康状况,但健康的草地状况可以有效地缓冲放牧和气候干扰,并能够维持较高的草地生产力和生物多样性。     

放牧对草原土壤的影响

介绍了放牧对草原土壤物理性质 (容重、渗透率 )、化学性质 (有机质、N素 )和微生物的影响。由于草原土壤系统本身的复杂性、滞后性和弹性 ,放牧对土壤性质的影响不尽相同。一般而言 ,随放牧强度的增大 ,动物践踏作用的增强 ,土壤孔隙分布的空间格局发生变化 ,土壤的总孔隙减少 ,特别是大孔隙 (>5 0μm)和较大中等孔隙 (9～ 5 0μm)减少 ,使土壤容重增加 ,土壤的渗透阻力加大 ,土壤的保水和持水能力下降。但在有机质含量很低的沙质土壤中 ,超载过牧 ,造成有机质含量降低 ,土壤的团粒结构减少 ,稳定性团聚体减少 ,土壤结构遭到破坏 ,使得土壤容重反而降低。土壤有机质和放牧之间存在复杂的相互关系 ,土壤有机质对放牧的响应受多种因素的影响 ,这些因素包括植被和土壤的初始状况 ;环境因素 ,特别是水分和温度 ;放牧历史 (强度、频率、持续时间和动物类型 )。同时 ,土壤有机质含量低的土壤比含量高的土壤更易受放牧的影响 ,而使有机质发生变化。土壤微生物量碳是最具活性的土壤碳库 ,对环境的变化敏感 ,能较早地指示生态系统功能的变化。当考虑时间尺度时 ,高强度放牧对土壤肥力有负面的影响 ,短期内 ,由于加速了养分的循环效率 ,产生有利的影响 ,但长期无管理的超载放牧必然造成系统物质 (资源 )输入和输     

Oak tree and grazing impacts on soil properties and nutrients in a California oak woodland

There is great interest in understandinghow rangeland management practices affectthe long-term sustainability of California oakwoodland ecosystems through their influence onnutrient cycling. This study examines the effects ofoak trees and low to moderate intensity grazing onsoil properties and nutrient pools in a blue oak (Quercus douglasii H.&A.) woodland in the SierraNevada foothills of northern California. Fourcombinations of vegetation and management wereinvestigated: oak with grazing, oak without grazing,open grasslands with grazing, and open grasslandswithout grazing. Results indicate that oak treescreate islands of enhanced fertility through organicmatter incorporation and nutrient cycling. Comparedto adjacent grasslands, soils beneath the oak canopyhave a lower bulk density, higher pH, and greaterconcentrations of organic carbon, nitrogen, total andavailable P, and exchangeable Ca, Mg, and K,especially in the upper soil horizons (0–35 cm). Incontrast, the light grazing utilized at this site hadminimal effects on soil properties which included anincrease in the bulk density of the surface horizonand an increase in available P throughout the entiresoil profile. While low to moderate intensity grazinghas little effect at this study site, there could bemuch larger impacts under the more intensive grazingpractices utilized on many rangelands. The lack ofoak regeneration and oak tree removal to enhanceforage production may eventually lead to large lossesof nutrients and soil fertility from these ecosystems.Results of this study have important implications forpredicting how management practices may potentiallyaffect oak regeneration, water quality, and ecosystemsustainability.     

Impact of grazing on germination trait selection in an alpine grassland on the Tibet Plateau

放牧对青藏高原高寒草地种子萌发性状选择的影响 以前的研究表明放牧能够引起草地生物与非生物环境的显著变化,但这种变化影响草地群落对种子萌发特征的选择机制尚不清楚。因此,我们旨在回答：放牧是否对草地群落中萌发特征的组成和多样性产生显著影响。我们在实验室检测了研究草地群落内主要物种种子的萌发特性,并比较了这些植物在放牧和非放牧草地上的表现。在此基础上,比较了放牧草地和非放牧草地的各萌发性状的群落加权平均值和萌发性状多样性,从而了解放牧草地和非放牧草地是否存在不同的萌发性状结构。研究结果表明,在物种水平上,放牧和非放牧草地各物种的多度变化与物种的萌发性状无显著关系。但在群落水平上,与非放牧草地相比,放牧草地的物种普遍具有较高的种子萌发率;放牧草地种子萌发对变温的正响应显著大于非放牧草地,而且放牧草地种子萌发温度生态位宽度小于非放牧草地。与非放牧草地相比,放牧草地种子萌发性状多样性增加,萌发性状均匀度降低。放牧可以改变微生境,从而通过环境过滤改变草地群落对萌发性状的选择,导致草地群落的萌发性状于放牧前后在群落水平上存 在显著差异。     

不同放牧制度下呼伦湖流域草原植被冠层截留

冠层截留是降雨过程中的水量分配和流域水平衡的一个重要组成部分,通过水浸泡法和降雨模拟实验研究呼伦湖流域草原3种放牧制度下(休牧、轮牧、自由放牧(超载放牧))植被冠层截留量的变化规律,并利用遥感解译植被归一化指数(NDVI),确定3种放牧制度下草原面积,估算呼伦湖流域草原降雨截留量。研究表明:在休牧、轮牧、自由放牧3种制度下,水浸泡法测定的截留量分别是0.468、0.320、0.271 mm。降雨模拟实验法测得的结果分别是0.957、0.613、0.431 mm。休牧、轮牧、自由放牧草场叶面积指数、盖度、容重、生物量等指标差异显著(P0.05),且单株植被高度、鲜重对截留量影响显著呈线性正相关关系。呼伦湖流域草原一次降雨量为大于等于30 mm全流域降雨,其植被截留量为6.462×106m3。     

Tracking nutrients in space and time: Interactions between grazing lawns and drought drive abundances of tallgrass prairie grasshoppers

1. We contrast the response of arthropod abundance and composition to bison grazing lawns during a drought and non‐drought year, with an emphasis on acridid grasshoppers, an important grassland herbivore.
2. Grazing lawns are grassland areas where regular grazing by mammalian herbivores creates patches of short‐statured, high nutrient vegetation. Grazing lawns are predictable microsites that modify microclimate, plant structure, community composition, and nutrient availability, with likely repercussions for arthropod communities.
3. One year of our study occurred during an extreme drought. Drought mimics some of the effects of mammalian grazers: decreasing above‐ground plant biomass while increasing plant foliar percentage nitrogen.
4. We sampled arthropods and nutrient availability on and nearby (“off”) 10 bison‐grazed grazing lawns in a tallgrass prairie in NE Kansas. Total grasshopper abundance was higher on grazing lawns and the magnitude of this difference increased in the wetter year of 2019 compared to 2018, when drought led to high grass foliar nitrogen concentrations on and off grazing lawns. Mixed‐feeding grasshopper abundances were consistently higher on grazing lawns while grass‐feeder and forb‐feeder abundances were higher on lawns only in 2019, the wetter year. In contrast, the abundance of other arthropods (e.g., Hemiptera, Hymenoptera, and Araneae) did not differ on and off lawns, but increased overall in 2019, relative to the drought of 2018.
5. Understanding these local scale patterns of abundances and community composition improves predictability of arthropod responses to ongoing habitat change.

     

不同放牧强度对赛罕乌拉草原蜘蛛多样性的影响

蜘蛛作为草原生态系统中的主要消费者, 对维系草原生物多样性和生态系统功能具有重大意义。放牧是人类利用草原最普遍的方式, 了解放牧对蜘蛛多样性的影响具有重要生态学意义。本研究调查了内蒙古赛罕乌拉草原上5个不同放牧强度样地中的蜘蛛多样性, 通过单因素方差分析(one-way analysis of variance)比较各样地中的蜘蛛多样性, 非度量多维标度分析(non-metric multidimensional scale, NMDS)和相似性分析(analysis of similarities, ANOSIM)比较各样地间的蜘蛛物种组成相似性, 再结合相关性分析探讨了植被高度对蜘蛛多样性的影响。结果表明: 重度放牧强度样地的蜘蛛多样性显著低于其他未放牧及轻度放牧样地; 具体到常见科上, 放牧强度对织网型的园蛛物种数和个体数影响显著, 而对游猎型的狼蛛、跳蛛却不明显; 织网型蜘蛛主要受植被结构影响, 而游猎型蜘蛛更可能受潜在猎物可得性的影响。NMDS分析表明不同放牧强度下, 蜘蛛类群的物种组成呈现明显的梯度变化, 放牧强度越低, 物种组成和未放牧样地越相近。相关性分析表明草原植被高度与蜘蛛多样性总体上呈正相关关系, 即植被高度越高, 蜘蛛多样性越高。其中依靠植物构建蛛网的园蛛科和在植物上层伏击猎物的蟹蛛科、逍遥蛛科等与植被高度显著相关。这说明植物资源及其空间异质性可能对草原蜘蛛多样性起着主导作用。因此, 降低放牧强度有助于保护草原蜘蛛群落的多样性, 特别有利于织网型蜘蛛。     

Impact of grazing around a watering point on soil status of a semi-arid rangeland in Ethiopia

The influence of long‐term livestock grazing on the soil status of a semi‐arid rangeland was studied along a grazing gradient from a watering point in southern Afar Region of Ethiopia for two seasons. The soil samples were analysed for organic carbon (OC), total nitrogen (N), available phosphorus (P), exchangeable cations (Ca²⁺, Mg²⁺, K⁺ and Na⁺), acidity (pH), cation exchange capacity, and particle size distribution (clay, silt and sand). No significant differences (P > 0.05) were observed for particle size distribution, OC, N, P and K contents along the grazing gradient further than 1500 m from the watering point. Ca²⁺ and Mg²⁺ were found to be dominant cations particularly in the severely degraded area contributing about 74% to the exchangeable cations. The study has also recognized high concentrations of Na⁺ and more than 11% exchangeable sodium percentage. Total exchangeable bases significantly declined (P < 0.05) along the grazing gradient and were indicative of the exchange complex of the soil being saturated with Ca²⁺ and Mg²⁺ ions. Thus this study concluded that there are soil differences in the grazing gradient caused by impact of grazing, particularly in the severely degraded area.     

荒漠草原不同放牧强度背景下添加氮水对凋落物分解的影响

凋落物分解是连接生态系统地上、地下过程的重要环节,决定了生态系统养分循环速率,但到目前为止对凋落物分解在荒漠草地生态系统受放牧以及外源资源补给影响的研究较少。本研究通过对不同放牧强度(对照、轻牧、中牧和重牧)短花针茅草原群落进行添加氮素(10.0 g N m~(-2) a~(-1))和增水(108 mm/a)处理,探讨群落水平凋落物分解速率的变化。研究结果显示,过去不同强度放牧历史对群落凋落物分解影响极显著(P0.0001)。凋落物前期分解(135 d)过程中,凋落物初始C∶N比与凋落物分解速率常数呈显著负相关关系,表明凋落物可降解性在凋落物前期分解中起主要作用。轻度放牧影响下凋落物分解速度最快,这与该条件下凋落物C∶N比显著低于其他放牧强度下的有关,说明适度放牧不仅有利于群落维持,也在一定程度上有利于生态系统养分循环。当凋落物分解更长时间(870 d)后,对照区凋落物分解速率显著低于放牧处理样地,但凋落物初始C∶N比对凋落物分解速率没有显著影响。进一步分析显示,不同放牧强度背景下长期凋落物分解速率与分解环境的土壤微生物多样性成正相关关系,与群落盖度呈极显著(P0.001)负相关关系。添加氮素显著(P0.05)降低凋落物分解速度,但对凋落物氮含量无显著影响。生长季加水未影响凋落物质量及凋落物分解速度。研究结果表明,凋落物前期分解受凋落物质量影响,但较长时间凋落物分解则与分解过程中接受到的太阳辐射量有关。     

The impact of large herbivores on floral composition and vegetation structure in the Naukluft Mountains, Namibia

Floral composition and vegetation structure were investigated on the Naukluft Mountain Plateau to determine (i) whether or not grazing has had an impact on the flora of the Naukluft Mountains during the last two decades and (ii) if so, whether vegetation recovery is influenced by the nature of the rainy season. Vegetation subjected to grazing to large herbivores and vegetation excluded from grazing for 18 and 19 years was compared. Based on classification (TWINSPAN) and ordination (detrended and canonical correspondence analysis) of 20, 10 × 10 m stands within and outside exclosures in two seasons, a clear difference in floristic composition between grazed and not grazed areas was indicated. Although neither the height of the vegetation, not the growth form spectrum differed significantly between grazed and not grazed sites, leaf-succulents were much more abundant when grazing was excluded. This suggests that changes in the vegetation due to grazing pressure have occurred in the last two decades, but the degree of grazing impact is not severe and may be part of the natural savanna ecosystem of the Naukluft Mountains. Whether this trend also occurs in other vegetation types in the Naukluft Mountains remains to the investigated. This revised version was published online in November 2006 with corrections to the Cover Date.     

Ecology of eutrophic waterbodies in a coastal grazing marsh

The ecology of a mainly interconnected, complex system of smallwater-bodies on a coastal grazing marsh in the Greater Thames estuary,north-east of London, England, is described. Sixteen sites were sampled on aregular basis over a two-year period while 37 sites were sampled once in Mayto develop a site classification. Most sites were brackish and alkaline but,following re-wetting after drying out, the pH in some water-bodies decreasedsharply (minimum pH 2.7). All sites had high concentrations of phosphorusbut there were large differences; some, with good growths of submergedaquatic macrophytes, having lower phosphorus than other sites, which weredevoid of macrophytes. Internal loading with phosphorus was important.Nitrate levels were low. Waters were mostly turbid, Secchi depths sometimes falling as low as 2cm, while chlorophyll a was very high, indicating extreme hypertrophy.Diatoms and euglenoids were characteristic members of the phytoplankton;cyanobacterial blooms did not occur. The zooplankton was dominated bycopepods, while Cladocera were much less numerous, largely due to predationby shrimps in waterbodies without macrophytes. The classification of siteswas influenced primarily by water depth, some waterbodies drying out, whilephosphorus and turbidity were also important in differentiating sites. Animal guano is the likely cause of the hypertrophic state but livestockgrazing is an essential management tool for grazing marshes. The dredging ofwaterbodies (with removal of sediments from the site) and the instigation ofa flushing regime, requiring a new system of sluices, should reduce nutrientloading and assist the establishment of submerged macrophytes, providingrefuges for cladocerans from predation.     

Intensified grazing affects endemic plant and gastropod diversity in alpine grasslands of the Southern Carpathian mountains (Romania)

Alpine grasslands in the Southern Carpathian Mts, Romania, harbour an extraordinarily high diversity of plants and invertebrates, including Carpathic endemics. In the past decades, intensive sheep grazing has caused a dramatic decrease in biodiversity and even led to eroded soils at many places in the Carpathians. Because of limited food resources, sheep are increasingly forced to graze on steep slopes, which were formerly not grazed by livestock and are considered as local biodiversity hotspots. We examined species richness, abundance and number of endemic vascular plants and terrestrial gastropods on steep slopes that were either grazed by sheep or ungrazed by livestock in two areas of the Southern Carpathians. On calcareous soils in the Bucegi Mts, a total of 177 vascular plant and 19 gastropod species were recorded. Twelve plant species (6.8%) and three gastropod species (15.8%) were endemic to the Carpathians. Grazed sites had lower plant and gastropod species richness than ungrazed sites. Furthermore, grazed sites harboured fewer gastropod species endemic to the Carpathians than ungrazed sites. On acid soils in the Fagaras Mts, a total of 96 vascular plant and nine gastropod species were found. In this mountain area, however, grazed and ungrazed sites did not differ in species richness, abundance and number of endemic plant and gastropod species. Our findings confirm the high biodiversity of grasslands on steep slopes in the Southern Carpathian Mts and caution against increasing grazing pressure in these refuges for relic plants and gastropods as well as for other invertebrates.     

沙质草地植物功能性状对放牧、增水、氮添加及其耦合效应的响应机制

人类活动和气候变化对陆地生态系统结构和功能的影响日益明显。在中国半干旱草原区,植物功能性状对这些变化的响应机制仍不是很清楚。以科尔沁沙质草地植物群落功能性状（CWM_trait）、一年生和多年生植物平均功能性状为研究对象,开展非生长季放牧、增水、氮添加及其耦合效应的模拟控制试验,于2016年8月中旬在沙质草地开展了植物功能性状的调查及测定,主要包括植物高度、比叶面积（specific leaf area,SLA）、叶片干物质含量（leaf dry matter content,LDMC）、叶片氮含量（leaf nitrogen content,LNC）和叶片碳同位素（leaf carbon isotopes,δ¹³C）。结果表明：氮添加显著提高了CWM_height和CWM_LNC（P<0.05）,降低了CWM_LDMC（P<0.05）,同时放牧和增水的耦合效应对CWM_SLA具有显著影响（P<0.05）;放牧显著降低了一年生植物的平均高度（P<0.05）,氮添加显著提高了一年生植物的平均高度、LNC和δ¹³C （P<0.05）;氮添加显著提高了多年生植物的平均SLA、LNC和δ¹³C （P<0.05）,同时放牧和增水的耦合效应对多年生植物的平均SLA和LDMC具有显著影响（P<0.05）。主成分分析表明,放牧促进了植物群落和多年生植物SLA与LDMC、LNC和δ¹³C之间的负相关关系,增水促进了植物群落和多年生植物高度与SLA之间的正相关关系以及LDMC、LNC和δ¹³C之间的正相关关系,说明多年生植物在调控沙质草地植物群落响应外界干扰时具有较强的主导性作用。非生长季放牧、生长季禁牧有利于沙质草地的可持续发展和管理,增水能够减缓放牧压力对于草地植物的影响,氮添加有利于植物同化δ¹³C,并提高植物叶片的水分利用效率。     

Conundrums in mixed woody–herbaceous plant systems

Aims To identify approaches to improve our understanding of, and predictive capability for, mixed tree–grass systems. Elucidation of the interactions, dynamics and determinants, and identification of robust generalizations that can be broadly applied to tree–grass systems would benefit ecological theory, modelling and land management. Methods A series of workshops brought together scientific expertise to review theory, data availability, modelling approaches and key questions. Location Ecosystems characterized by mixtures of herbaceous and woody plant life‐forms, often termed ‘savannas’, range from open grasslands with few woody plants, to woodlands or forests with a grass layer. These ecosystems represent a substantial portion of the terrestrial biosphere, an important wildlife habitat, and a major resource for provision of livestock, fuel wood and other products. Results Although many concepts and principles developed for grassland and forest systems are relevant to these dual life‐form communities, the novel, complex, nonlinear behaviour of mixed tree–grass systems cannot be accounted for by simply studying or modelling woody and herbaceous components independently. A more robust understanding requires addressing three fundamental conundrums: (1) The ‘treeness’ conundrum. What controls the relative abundance of woody and herbaceous plants for a given set of conditions at given site? (2) The coexistence conundrum. How do the life‐forms interact with each other? Is a given woody–herbaceous ratio dynamically stable and persistent under a particular set of conditions? (3) The net primary productivity (NPP) conundrum. How does NPP of the woody vegetation, the herbaceous vegetation, and the total ecosystem (woody + herbaceous) change with changes in the tree–grass ratio? Tests of the theory and conceptual models of determinants of mixed woody–herbaceous systems have been largely site‐ or region‐specific and have seldom been broadly or quantitatively evaluated. Cross‐site syntheses based on data and modelling are required to address the conundrums and identify emerging patterns, yet, there are very few data sets for which either biomass or NPP have been quantified for both the woody and the herbaceous components of tree–grass systems. Furthermore, there are few cross‐site comparisons spanning the diverse array of woody–herbaceous mixtures. Hence, initial synthesis studies should focus on compiling and standardizing a global data base which could be (1) explored to ascertain if robust generalizations and consistent patterns exist; and (2) used to evaluate the performance of savanna simulation models over a range of woody–herbaceous mixtures. Savanna structure and productivity are the result of complex and dynamic interactions between climate, soils and disturbances, notably fire and herbivory. Such factors are difficult to isolate or experimentally manipulate in order to evaluate their impacts at spatial and temporal scales appropriate for assessing ecosystem dynamics. These factors can, however, be evaluated with simulation models. Existing savanna models vary markedly with respect to their conceptual approach, their data requirements and the extent to which they incorporate mechanistic processes. Model intercomparisons can elucidate those approaches most suitable for various research questions and management applications. Conclusion Theoretical and conceptual advances could be achieved by considering a broad continuum of grass–shrub–tree combinations using data meta‐analysis techniques and modelling.     

放牧对草原植物功能性状影响研究进展

植物功能性状的表达和植被环境适应性相关,植物功能性状之间的权衡变化体现了植物在放牧胁迫下资源的重新整合和获取.本文总结了放牧干扰下植物功能性状表达的差异性,着重将放牧干扰与植物功能性状相结合,介绍了植物功能性状的变异来源是植物遗传特征与环境过滤相互协调的结果,归纳了放牧对植物营养性状、繁殖性状的影响,以及植物可以通过调...