干旱持续时间对内蒙古草原中坚韧胶衣和分指地卷固氮活性的影响

坚韧胶衣Collema tenax和分指地卷Peltigera didactyla是分布于我国北方干旱和半干旱草原中的两种固氮地衣,对生态系统氮素循环具有重要影响,其生理活性对土壤水分条件比较敏感。目前,干旱持续时间对该区域中地衣固氮活性的影响尚不清楚。本文调查了一个克氏针茅Stipa krylovii草原生态系统中坚韧胶衣和分指地卷的盖度,采用乙炔还原法研究了不同干旱持续时间(分别为0、1、2、4、8和16d)对两种地衣固氮活性的影响。结果表明,坚韧胶衣盖度(3.26±1.24%)显著高于分指地卷(0.02±0.07%)(p<0.01);在绝大多数干旱持续时间处理下,两种地衣之间的固氮活性差异并不显著,且均随干旱持续时间延长而呈S形下降。不超过2d的干旱时间并不引起地衣固氮活性的显著变化,4d的干旱时间导致坚韧胶衣和分指地卷固氮活性分别下降了34%和54%,干旱持续8d导致两种地衣固氮活性降幅达90%,16d的干旱使二者降幅均超过99%。表明干旱对坚韧胶衣和分指地卷固氮活性的抑制作用随干旱持续时间的延长而增强。干旱对地衣固氮作用的负效应可能与地衣体内光合产物和能量水平、复水至表达固氮活性之间的延迟时间和达到最高固氮活性的延迟时间有关。     

不同水分条件下坚韧胶衣固氮活性对冻融的响应

坚韧胶衣Collema tenax是干旱和半干旱生态系统中生物土壤结皮的重要组分,其固氮作用对生态系统氮素循环具有重要影响。该地衣生长于温度和水分条件变化剧烈的土壤表面,但在我国北方其固氮活性对冻融的响应尚不清楚。采用乙炔还原法研究了坚韧胶衣固氮活性在不同水分条件下(湿冻组:地衣体含水量200%干重;干冻组:地衣体含水量20%干重)和4次冻融处理之间的差异。坚韧胶衣固氮活性为2,371.0-8,701.8nmolC2H4/m2·h,湿冻组固氮活性低于干冻组,对照组固氮活性最高,湿冻组和干冻组的固氮活性(对照的百分比)均与冻融次数呈显著的线性负相关关系(干冻组:R2=0.916,p0.001;湿冻组:R2=0.965,p0.001),但湿冻组的斜率绝对值(25.05)明显高于干冻组斜率绝对值(7.60)。结果表明低温对坚韧胶衣固氮活性具有显著抑制作用,干燥条件下坚韧胶衣固氮活性对于低温胁迫的敏感性比地衣体湿润条件下更低,这种较低的敏感性可能是该地衣对我国北方干燥和冷热变化剧烈的气候特点的生理适应策略。     

十二种地卷目地衣固氮活性的种间差异

地卷目地衣是对许多生态系统的氮素循环具有重要影响的固氮类群,但其固氮活性的种间差异研究甚少。将不同产地和不同馆藏时间的12种地卷目地衣于相同条件下培养,即：亚石胶衣Collema subflaccidum、亚黑胶衣C. subnigrescens、坚韧胶衣C. tenax、裸果猫耳衣Leptogium hildenbrandii、猫耳衣L. menziesii、柄猫耳衣L. pedicellatum、土星猫耳衣L. saturninum、黑猫耳衣L. trichophorum、犬地卷Peltigera canina、分指地卷P. didactyla、裂芽地卷P. praetextata和地卷P. rufescens。以乙炔还原法（acetylene reduction assay）于培养1d、10d和15d测定其固氮活性,结果显示所有地衣的固氮活性在培养10d与15d之间差异不显著,表明经过至多10d的培养后地衣固氮活性即可完全恢复;培养10d和15d的固氮活性平均值表现出显著的种间差异（P＜0.001）：黑猫耳衣Leptogium trichophorum固氮活性最高[（4.532±0.368）μmol C2H4/gdw·h],约为犬地卷Peltigera canina的2倍[（2.349±0.223）μmol C2H4/gdw·h];其他10种地衣固氮活性大致相近。显示地衣固氮作用的可塑性较大,是地衣具有较强适应能力的又一例证。     

内蒙古草原放牧恢复过程地衣生物量分布及其影响因素的研究

1　引　　言在干旱半干旱草原生态系统中,地衣是植物群落中的一个重要组成部分.地衣是真菌和具有光合功能的藻类的共生体,共生光合生物通过光合作用为自身和共生真菌提供碳水化合物,共生真菌为共生光合生物提供物理保护、水和矿物元素[9] .地衣芽枝可通过截获风蚀物质而增加土壤表面粗糙度,有利于积蓄降水和养分[1] ,地衣的共生真菌菌丝可以通过分泌粘质物而改善土壤团粒结构[3 ] .另外,地衣还可有效地分解矿物元素,同时还有较强的固氮功能[9,17] ,地衣不仅影响土壤的理化性状而且对维管植物也有潜在的影响.因此,地衣在草原生态系统中可促进…     

荒漠地衣结皮研究进展

微型生物结皮在干旱、半干旱荒漠地区广泛分布,对维持荒漠生态系统的稳定性具有重要作用。地衣结皮是微型生物结皮的重要组成部分及主要类型之一,在固沙、固碳和固氮能力方面独具优势。本文从区域尺度和局部尺度综述了国内外荒漠地衣分布、群落组成及其影响因素,从微尺度探讨了荒漠地衣在形成过程中可能相关的生物因子的作用与功能。虽已有研究发现气候类型、降水量、土壤理化性质、微地形和温度会对荒漠地衣生长型、种类、丰度及盖度产生影响,来自地衣体、地衣结皮和地衣结皮土壤中的生物因素与维持地衣正常生命活动及分布之间存在相关关系,但上述结论尚具有一定的局限性,主要表现在区域尺度仍缺乏专门对荒漠地衣的研究,微尺度缺乏对地衣相关生物功能的实验性探索及验证。上述局限在一定程度上限制了通过人工手段大规模应用荒漠地衣结皮的研究。本文基于国内外研究进展及存在问题,对荒漠地衣结皮应用进行了展望,以期为人工构建荒漠地衣结皮和干旱、半干旱荒漠治理及生态恢复提供新的思路和参考。     

古尔班通古特沙漠生物结皮固氮活性

利用乙炔还原法对新疆古尔班通古特沙漠不同类型的生物结皮(藻结皮、地衣结皮、苔藓结皮)的固氮活性(nmolC2H4m-2h-1)进行了定量研究。结果表明,不仅采样时段和结皮类型对生物结皮的固氮活性具有显著的影响(p<0.05),二者的交互效应同样对生物结皮的固氮活性具有极显著的影响(p<0.01)。各类型生物结皮固氮活性变化趋势表现为:3～5月间,藻结皮(2.26×103)>地衣结皮(6.54×102)>苔藓结皮(6.38×102)。6～10月份,各类型生物结皮的固氮能力显著提高(p<0.05),藻结皮的固氮活性最高(9.81×103),依次为地衣结皮9.06×103、苔藓结皮2.03×103。11月～翌年2月间,月均温都低于0℃,抑制了生物结皮的固氮活性,藻结皮、苔藓结皮的固氮活性降幅极显著(p<0.01),分别低达4.18×102、5.43×102,地衣结皮降低至2.78×103。生物结皮成为古尔班通古特沙漠除豆科植物外重要的氮源,为该沙漠1年生浅根系草本植物的种子萌发与植物体的生长提供丰富的有机质源,从而有利于这些植物种群的繁衍与更新,并与之共同促进对沙面的固定。     

腾格里沙漠东南缘生物土壤结皮的固氮潜力

以腾格里沙漠东南缘广泛分布的3类典型生物土壤结皮(藻类结皮、地衣结皮和藓类结皮)为研究对象,在野外环境下连续一年(2010年6月至2011年5月)测定了不同类型结皮的固氮潜力、季节变化,及其对环境因子的响应特征.结果表明：整个试验期间,藻类、地衣和藓类结皮的固氮活性分别为14～133、20～101和4～28 mol·m^-2·h^-1,差异显著,这种差异主要是由结皮种类组成的差异所致.3类结皮固氮活性对环境因子的响应特征基本一致.3类结皮固氮活性与降水量关系不显著,但与试验前3天小于3 mm的降水量均呈显著的正相关关系,说明该地区3类结皮在3 mm降水条件下即可达到最大固氮速率.3类结皮固氮活性与试验期温度均呈显著的二次函数关系,随气温升高均呈先上升后下降的趋势,藻类和地衣结皮的固氮活性在超过30 ℃后即迅速下降,而藓类结皮的固氮活性在超过25 ℃后即开始下降,说明不同类型结皮具有不同的固氮适宜温度区间.3类结皮固氮活性的季节变化均表现为秋季>春季>夏季>冬季,夏季高温和冬季低温抑制了结皮固氮酶活性,而春末秋初适宜的水热条件促进了其固氮活性的提高,结皮固氮活性的季节变化主要受水热因子的共同调控.温带荒漠区生物土壤结皮在湿润条件下全年均具有固氮能力,环境因子对其氮固定的控制作用层次分明,水分是影响其固氮速率和持续时间的关键因子,在水分和碳源充足的条件下,温度则是制约其固氮能力的主要因子.     

火烧对古尔班通古特沙漠土壤养分和土壤酶活性的影响

以古尔班通古特沙漠2016年6月意外火烧事件为背景,对比分析火烧和未火烧样地区不同土壤深度化学性质、土壤酶活性的变化特征,为全面评估火烧对温带荒漠生态系统的影响提供土壤学依据。结果表明:火烧样地和对照样地土壤养分含量和酶活性总体上均表现为上层土壤(0—5 cm)大于下层土壤(5—10 cm),仅土壤氧化酶(过氧化氢酶)活性表现为下层土壤大于上层土壤。同时,火烧和土壤深度存在交互效应,火烧对土壤特性的影响受土壤深度的限制。土壤化学性质受火烧的影响主要表现在0—5 cm土壤层,尤其是速效养分在火烧之后有显著增加趋势。火烧对5—10 cm土壤层土壤化学性质无显著影响。火烧后,硝态氮(NO3-N)含量显著下降,铵态氮(NH4-N)含量上升。土壤水解酶、氧化酶活性在火烧之后降低,具体表现为,蔗糖酶、脲酶、β-葡萄糖苷酶活性在0—5 cm和5—10 cm土壤层均极显著下降,而碱性磷酸酶活性仅在0—5 cm土壤层显著下降。过氧化氢酶活性则在5—10 cm土壤层活性显著下降。表明上层土壤水解酶活性对火烧干扰较为敏感,氧化酶活性在火烧干扰下相对稳定。从各土层土壤酶活性的变化特征来看,火烧对水解酶活性的影响随土壤深度增加而降低,而氧化酶则呈现相反的趋势。总体而言,火烧显著提高了古尔班通古特沙漠土壤速效养分的含量,降低了土壤酶活性,不同土壤酶对火烧响应的敏感程度不同。为深入评估火烧干扰对温带荒漠生态系统的影响提供一定理论依据。     

沙坡头地区生物土壤结皮的固氮活性及其对水热因子的响应

氮是除水分之外影响干旱区生态系统生物活性的关键因子。生物土壤结皮是干旱半干旱荒漠地表景观的重要组成部分, 也是荒漠生态系统氮素的主要贡献者。通过野外调查采样, 利用开顶式生长室, 模拟不同降水梯度, 采用乙炔还原法连续测定了沙坡头地区典型生物土壤结皮(藻类结皮、地衣结皮和藓类结皮)在其主要固氮活跃期(6-10月, 湿润期)的固氮活性, 及其对水热因子的响应特征。结果表明, 试验期三类生物土壤结皮的固氮活性介于2.5 × 10³-6.2 × 10⁴ nmol C₂H₄·m^-2·h^-1之间, 其中藻类结皮的最高(平均达2.8 × 10⁴ nmol C₂H₄·m^-2·h^-1), 地衣结皮的次之(2.4 × 10⁴ nmol C₂H₄ ·m^-2·h^-1), 藓类结皮的最低(1.4 × 10⁴ nmol C₂H₄·m^-2·h^-1), 差异显著(p < 0.001)。在模拟降水3 mm时, 三类结皮均可达到最大固氮速率, 当发生> 3 mm的降水事件时, 它们的固氮速率无显著增加; 不同结皮的固氮活性与温度均呈显著的负相关关系(r_藻类结皮 = -0.711, r_地衣结皮 = -0.732, r_藓类结皮 = -0.755, p < 0.001), 藻类和藓类结皮的固氮活性的最适温度区间为25-30 ℃, 地衣结皮为20-30 ℃。三类结皮之间的这种固氮差异主要归因于结皮组成生物体即隐花植物的差异, 藻类结皮主要成分为大量的蓝细菌和一些绿藻, 地衣结皮也由大量的固氮藻和真菌共生形成, 而藓类结皮的主要组成部分苔藓植物并不具有固氮作用, 其微弱的固氮量是结皮中混生的少量蓝细菌或地衣所致。     

刈割和氮添加对松嫩平原羊草草原碳固持的影响

最近30年来,碳固持作为陆地生态系统的重要生态功能受到前所未有的关注。草原是全球面积最大的陆地生态系统类型,割草和放牧是中国草原的最主要利用方式,同时氮沉降是草原生态系统面临的重要全球变化因子,然而它们对草原生态系统碳固持的影响尚未有一致的结论。本研究在松嫩平原羊草草原通过同化箱法观测并对比了刈割和模拟氮沉降条件下生态系统碳通量的变化。结果表明:无论从季节动态还是日动态来看,氮添加处理的净生态系统气体交换量(NEE)、总生态系统呼吸(TER)和总生态系统生产力(GEP)总体上均高于对照,而刈割处理的NEE、TER和GEP均低于对照;就土壤呼吸而言,各处理间无显著差异;整个生长季对照、刈割和氮添加处理累计碳释放量分别为107.8、285.2和102.9 g·m~(-2)·a~(-1);由此看来,整个实验区域是碳源,氮素添加可以在一定程度上减弱草原的碳源作用,且有向碳汇转变的趋势,而高频次和高强度刈割加重了草原的碳源作用。建议在生产实践中对草原实行轻度放牧和施肥管理,既能加强草原生态系统的生产功能,又能增加以碳固持为主的生态功能,从而实现草原生态系统的可持续利用。     

Litter dynamics in post-fire successional forests of Quercus ilex

Forest floor regeneration after fire in a holm oak forest was studied in a chronosequence in Catalonia (NE Spain). The sampled area represented situations of high fire disturbance because of the frequency of wildfires and the high impact of erosive processes. Forest floor bulk samples from the L, F and H layers were taken in 5 burned plots, aging from 0 to 35 years. They were analyzed for dry standing weight, and organic carbon and N content.Plant cover reached almost 100% two years after the fire. During the first 20 years, shrubs and herbs had the highest surface covering, and thereafter, holm oaks became dominant.Holm oak litterfall stabilized by the fourth year after burning.The major shrub species which were present in the chronosequence had two different patterns with respect to nitrogen: 1) a low litter C/N ratio (i.e. nitrogen fixing species); 2) a high litter C/N ratio.The litter decay coefficients were relatively high, allowing for a rapid structuration and formation of the L and F layers after the fire. Indeed, 95% of the maximum steady standing weight accumulates in 8–9 years. The pattern of cover in the different strata influenced the lateral distribution and variable accumulation of the forest floor layers and determined different turnover rates.     

Microbial community shifts influence patterns in tropical forest nitrogen fixation

The role of biodiversity in ecosystem function receives substantial attention, yet despite the diversity and functional relevance of microorganisms, relationships between microbial community structure and ecosystem processes remain largely unknown. We used tropical rain forest fertilization plots to directly compare the relative abundance, composition and diversity of free-living nitrogen (N)-fixer communities to in situ leaf litter N fixation rates. N fixation rates varied greatly within the landscape, and ‘hotspots’ of high N fixation activity were observed in both control and phosphorus (P)-fertilized plots. Compared with zones of average activity, the N fixation ‘hotspots’ in unfertilized plots were characterized by marked differences in N-fixer community composition and had substantially higher overall diversity. P additions increased the efficiency of N-fixer communities, resulting in elevated rates of fixation per nifH gene. Furthermore, P fertilization increased N fixation rates and N-fixer abundance, eliminated a highly novel group of N-fixers, and increased N-fixer diversity. Yet the relationships between diversity and function were not simple, and coupling rate measurements to indicators of community structure revealed a biological dynamism not apparent from process measurements alone. Taken together, these data suggest that the rain forest litter layer maintains high N fixation rates and unique N-fixing organisms and that, as observed in plant community ecology, structural shifts in N-fixing communities may partially explain significant differences in system-scale N fixation rates.     

Variation in Aboveground Cover Influences Soil Nitrogen Availability at Fine Spatial Scales Following Severe Fire in Subalpine Conifer Forests

Following fire, fine-scale variation in early successional vegetation and soil nutrients may influence development of ecosystem structure and function. We studied conifer forests burned by stand-replacing wildfire in Greater Yellowstone (Wyoming, USA) to address two questions: (1) How do the variability and spatial structure of aboveground cover and soil nitrogen availability change during the first 4 years following stand-replacing fire? (2) At fine scales (2–20 m), are postfire soil inorganic N pools and fluxes related to aboveground cover? Aboveground cover, soil N pools, and annual net N transformations were measured from 2001 to 2004 using a spatially explicit sampling design in four 0.25-ha plots that burned during summer 2000. Within-stand variability (coefficient of variation) in postfire live vegetative cover declined with time since fire, whereas variability in bare mineral soil, charred litter and fresh litter was greatest 2-3 years postfire. The soil nitrate pool was more variable than the soil ammonium pool, but annual net nitrification was less variable than annual net N mineralization. Spatial structure (quantified by semivariograms) was observed in some aboveground cover variables (for example, graminoids and fresh litter), but there was little spatial structure in soil N variables and no obvious congruence in spatial scales of autocorrelation for soil N and aboveground cover. Significant Spearman correlations (at the sample point) indicated that aboveground cover and soil N were coupled following severe fire, and the dominant influence was from aboveground cover to soil N, rather than from soil N to vegetation. Initial patterns of fire severity and re-vegetation contributed to fine-scale heterogeneity in soil N availability for at least 4 years after severe fire.     

Effects of experimental season of prescribed fire and nutrient addition on structure and function of previously grazed grassland

Aims Understanding the drivers of grassland structure and function following livestock removal will inform grassland restoration and management. Here, we investigated the effects of fire and nutrient addition on structure and function in a subtropical semi-native grassland recently released from grazing in south-central Florida. We examined responses of soil nutrients, plant tissue nutrients, biomass of live, standing dead and litter, and plant species composition to experimental annual prescribed fire applied during different seasons (wet season vs. dry season), and nutrient additions (N, P and N + P) over 9 years.Methods Experimental plots were set up in a randomized block split-plot design, with season of prescribed fire as the main treatment and nutrient addition as the subplot treatment. Species cover data were collected annually from 2002 to 2011 and plant tissue and plant biomass data were collected in 2002–2006 and 2011. Soil nutrients were analyzed in 2004, 2006 and 2011.Important findings Soil total phosphorus (P) levels increased substantially with P addition but were not influenced by prescribed fire. Addition of P and N led to increased P and N concentrations in live plant tissues, but prescribed fire reduced N in live tissue. Levels of tissue N were higher in all plots at the beginning of the experiment, an effect that was likely due to grazing activity prior to removal of livestock. Plant tissue N steadily declined over time in all plots, with annually burned plots declining faster than unburned plots. Prescribed fire was an important driver of standing dead and litter biomass and was important for maintaining grass biomass and percent cover. Nutrient addition was also important: the addition of both N and P was associated with greater live biomass and woody forbs. Removal of grazing, lack of prescribed fire, and addition of N + P led to a reduction of grass biomass and a large increase in biomass of a woody forb. Annual prescribed fire promoted N loss from the system by reducing standing dead and litter, but maintained desirable biomass of grasses.     

Annual burning of a tallgrass prairie inhibits C and N cycling in soil,increasing recalcitrant pyrogenic organic matter storage while reducing N availability

Grassland ecosystems store an estimated 30% of the world's total soil C and are frequently disturbed by wildfires or fire management. Aboveground litter decomposition is one of the main processes that form soil organic matter (SOM). However, during a fire biomass is removed or partially combusted and litter inputs to the soil are substituted with inputs of pyrogenic organic matter (py‐OM). Py‐OM accounts for a more recalcitrant plant input to SOM than fresh litter, and the historical frequency of burning may alter C and N retention of both fresh litter and py‐OM inputs to the soil. We compared the fate of these two forms of plant material by incubating ¹³C‐ and ¹⁵N‐labeled Andropogon gerardii litter and py‐OM at both an annually burned and an infrequently burned tallgrass prairie site for 11 months. We traced litter and py‐OM C and N into uncomplexed and organo‐mineral SOM fractions and CO₂ fluxes and determined how fire history affects the fate of these two forms of aboveground biomass. Evidence from CO₂ fluxes and SOM C:N ratios indicates that the litter was microbially transformed during decomposition while, besides an initial labile fraction, py‐OM added to SOM largely untransformed by soil microbes. Additionally, at the N‐limited annually burned site, litter N was tightly conserved. Together, these results demonstrate how, although py‐OM may contribute to C and N sequestration in the soil due to its resistance to microbial degradation, a long history of annual removal of fresh litter and input of py‐OM infers N limitation due to the inhibition of microbial decomposition of aboveground plant inputs to the soil. These results provide new insight into how fire may impact plant inputs to the soil, and the effects of py‐OM on SOM formation and ecosystem C and N cycling.     

Photosynthesis of the cyanobacterial soil-crust lichen Collema tenax from arid lands in southern Utah, USA: role of water content on light and temperature responses of CO2 exchange

1. The gelatinous cyanobacterial Collema tenax is a dominant lichen of biotic soil crusts in the western United States. In laboratory experiments, we studied CO₂ exchange of this species as dependent on water content (WC), light and temperature. Results are compared with performance of green-algal lichens of the same site investigated earlier.
2. As compared with published data, photosynthetic capacity of C. tenax is higher than that of other cyanobacterial and green-algal soil-crust species studied. At all temperatures and photon flux densities of ecological relevance, net photosynthesis (NP) shows a strong depression at high degrees of hydration; maximal apparent quantum-use efficiency of CO₂ fixation is also reduced. Water requirements (moisture compensation point, WC for maximal NP) are higher than that of the green-algal lichens. Collema tenax exhibits extreme 'sun plant' features and is adapted to high thallus temperatures.
3. Erratic rain showers are the main source of moisture for soil crusts on the Colorado Plateau, quickly saturating the lichens with liquid water. High water-holding capacity of C. tenax ensures extended phases of favourable hydration at conditions of high light and temperature after the rain for substantial photosynthetic production. Under such conditions the cyanobacterial lichen appears superior over its green-algal competitors, which seem better adapted to habitats with high air humidity, dew or fog as prevailing source of moisture.     

Cumulative effects of fire on a tussock pampa grassland

Abstract. In order to test for cumulative effects of fire on Paspalum quadrifarium‐dominated grasslands (‘pajonal’), we analysed the impact of single and repeated fires on the community structure and post‐fire recovery of canopy after a final, simultaneous fire event. Nine plots were defined within a homogeneous pajonal stand, and treatments of low (LF), medium (MF) and high frequency (HF) of fire were defined by the application of one, two or four cold‐season burns, respectively, along a 6‐yr period. Both burned and unburned plots were exposed to grazing by cattle during the summer following the first and the third years of that period. High cattle preference for burned sites conditioned fire temperature and vegetation responses to the following burning events. Cumulative effects between successive burning events were observed for the cover of basal area of the dominant and other sprouting species, the cover and thickness of the litter layer, the seed bank size of the principal recruiter species, and the floristic composition. While light interception by the canopy was positively related to fire frequency during the early growth season, further growth of P. quadrifarium determined a greater light interception in LF than in MF and HF. These patterns of light interception were associated with a faster occupation of the inter‐tussock areas by opportunistic species in plots subjected to frequent fires (HF and MF) than in plots with low fire frequency (LF), and a more lasting regrowth of P. quadrifarium in the LF plot than in the HF ones. High fire frequencies reduced the dominance of P. quadrifarium. Percent of species classified as subordinated graminoids or forbs did not vary among treatments. However, the abundance of different forb species was differentially favoured by contrasting frequencies of fire, describing some coarse relationships between their specific responses and their dispersal strategies.     

Barriers to shrub reestablishment following fire in the seasonal submontane zone of Hawai'i

Introduced grass species have invaded extensive areas of Hawaii Volcanoes National Park and increased the size and frequency of fire. Following fire, grass cover is enhanced while native shrub cover is reduced; the reduction in most shrubs persists for at least 20 years even in the absence of fire. Shrub seedlings were planted in burned and unburned plots with and without grass cover. Biomass of 14 month old shrub seedlings was generally highest in recently burned/grass removed plots, intermediate in old burn/grass removed plots, and lowest in unburned/grass removed plots. In contrast, shrub biomass in plots with grass cover was low and did not differ significantly among burn treatments. Light competition is likely to be responsible for differences in shrub growth rates; grass cover reduced light to 1–10% of background levels. In addition, pool sizes of available soil N were highest in recently burned, intermediate in old burn, and lowest in unburned areas.