贵州雷公山秃杉林不同林冠环境下箭竹分株种群结构特征

通过对贵州雷公山秃杉林的4种林冠环境,即林下(FU)、中林窗(MG)、大林窗(LG)和林缘旷地(FEW)内的箭竹的分株种群结构（包括株高、基径、生物量、叶和分株数等）和年龄结构进行较系统对比研究。结果显示:（1)从林下→中林窗→大林窗,箭竹分株种群的株高(h)、基径（bd)与生物量显著增加。（2)在4种林窗环境中,箭竹各构件的生物量的百分比发生相应变化,枝和叶片生物量的百分比沿林下→中林窗→大林窗→林缘旷地的顺序显著减小,在林缘旷地,地下茎、粗根（d＞0.5 mm)和细根（d≤0.5 mm)生物量的百分比显著高于其他3种林冠环境。（3)箭竹分株单位叶面积叶重沿林下→中林窗→大林窗→林缘旷地的顺序显著增大,单叶生物量与叶面积以中林窗最大,林缘旷地次之,二者与大林窗或林下差异显著;大林窗的单株叶片数显著高于其余3种林冠环境。（4)在4种林冠环境中,箭竹分株种群死亡率以林下最低,但各种群平均年龄间无显著差异。研究表明,箭竹分株种群对林冠环境变化的响应主要体现在形态与生物量的分配上,而非种群的年龄。     

卧龙亚高山暗针叶林不同林冠环境下华西箭竹分株种群结构特征

详细调查了林下、中林窗、大林窗和林缘旷地等4种亚高山暗针叶林林冠环境下的华西箭竹（Fargesia nitida）分株种群,对其表现结构（株高、基径和生物量）和年龄结构进行了较系统的对比研究.主要研究结果如下：（1）分株水平上,华西箭竹表现结构在4种林冠环境下均有极显著差异（p＜0.01）,且随林冠郁闭度的减小,分株个体的高度、基径和生物量有递增的趋势（林下＜中林窗＜大林窗）;（2）分株各构件在总生物量中所占比例随林冠环境变化而改变：除林缘旷地外,叶的生物量百分比与林冠郁闭度呈正相关.林缘旷地中,地下茎和根系的生物量百分比均高于其余3种环境;（3）不同林冠环境下分株单位叶面积叶重存在显著差异（p＜0.05）,且随林冠郁闭度的减小而增大.单叶生物量和单叶面积均以中林窗最大,林缘旷地次之,二者与大林窗或林下均有显著差异（p＜0.05）.华西箭竹的单株叶片数量以大林窗最大,与其余3种环境有极显著差异（p＜0.01）;（4）分株种群的死亡率以林下最低（P＜0.01）,但各种群平均年龄间无显著差异（p＞0.05）.可见,华西箭竹分株种群对林冠环境变化的反应主要体现在形态和生物量分配上,而非种群的年龄上.     

卧龙亚高山暗针叶林不同林冠环境下华西箭竹的克隆生长

该文调查了林下、中林窗、大林窗和林缘旷地等4种亚高山暗针叶林林冠环境下的华西箭竹(Fargesia nitida)分株种群,对其当年生立竹和母株的特征进行了比较研究,并探讨了母株年龄及大小对克隆生长的影响。主要研究结果如下:1)华西箭竹当年生立竹的分株密度以林下种群的最低,从林下→中林窗→大林窗,随林冠郁闭度的减小,华西箭竹基株当年生立竹的分株数逐渐增大。2)4种林冠环境下,当年生立竹和母株的分株高度、基径和生物量均有显著差异,且随林冠郁闭度的减小有递增的趋势(林下<中林窗<大林窗)。3)各林冠环境均以二龄母株产生的当年生立竹数量最大,不同林冠环境之间母株的平均年龄,以及处于同一林冠环境的各龄级母株产生的当年生立竹平均分株数和平均基径均无显著差异。4)不同林冠环境下,华西箭竹当年生立竹基径与一级母株、二级母株基径均呈正相关关系,但当年生立竹基径随一级母株增加的速度快于随二级母株的增加。5)当年生立竹的基径与地下茎直径呈显著的正相关;母株的基径与当年生立竹的地下茎直径呈正相关关系,而与其地下茎长度相关性不明显。     

亚高山暗针叶林不同林冠环境下华西箭竹的克隆可塑性

以亚高山暗针叶林3种林冠环境中以及暗针叶林林缘的华西箭竹（Fargesia nitida）为对象,对其无性系数量特征、无性系根茎特征、分株生物量以及分株形态特征进行了对比研究。结果表明：（1）林冠环境的差异导致了不同种群的基株密度和每基株分株数的显著差异,但林冠环境差异不影响分株密度。林冠郁闭度愈大,每基株分株数愈少,分株分布愈均匀。（2）不同林冠环境间。分株生物量、分株构件生物量和分株构件的生物量分配百分率均有显著差异。开敞的林冠环境有利于华西箭竹的生长和生物量积累。（3）随着林冠郁闭度的增加,华西箭竹通过增大分枝角度、叶生物量分配百分率、比叶面积和叶面积率以提高光能利用效率,有效适应弱光环境。（4）隔离者长度、隔离者直径和分枝强度在林缘和林窗环境中要显著大于林内环境;同级隔离者分枝角度随林冠郁闭度的增加而最大,其值在林下显著大于林窗和林缘,而异级隔离者分枝角度的变化则正好相反。研究表明,华西箭竹种群在不同的林冠环境中发生了明显的可塑性变化,这些可塑性变化是种群对林冠郁闭度差异的适应性反应的结果,有利于增强种群对环境中有限光资源的利用。     

常绿阔叶林林冠环境对栲幼苗建成的影响

植物幼苗建成阶段是决定种群自然更新的关键生活史阶段。研究林冠环境对常绿阔叶林优势种幼苗建成阶段的影响对该类森林的恢复和管理具有重要意义。2014-2016年, 该研究在重庆市缙云山国家级自然保护区的常绿阔叶林的不同林冠环境(大林窗: >150 m ², 中林窗: 100-150 m ², 小林窗: 50-100 m ², 对照: 林下)下进行栲(Castanopsis fargesii)种子野外播种实验, 并对栲幼苗命运和生长情况进行了3年的持续监测。结果表明: (1)栲幼苗出土时间从7月持续到12月, 出苗时间较长, 大林窗对幼苗出土具有延迟作用; (2)栲种子野外平均萌发率为(62.8 ± 2.0)%, 第3个生长季(2016年)末幼苗平均存活率为(65.1 ± 2.2)%, 枯萎是栲幼苗死亡的主要原因; (3)林冠环境对栲种子萌发率及第1个生长季(2014年)末的幼苗存活率无显著影响, 对第2个(2015年)和第3个生长季末的幼苗存活率具有显著影响; (4)林冠环境在第1个生长季对幼苗生长无明显影响, 但在第2个和第3个生长季具有显著影响, 大、中林窗中幼苗总生物量、株高、基径、根长和叶片数显著高于林下, 比叶面积显著低于林下; (5) 3个生长季内, 4类林冠条件下栲幼苗的叶质量比和茎质量比升高, 根质量比和根冠比降低, 并且从第2个生长季开始大林窗中栲幼苗的叶质量比显著高于林下, 根质量比和根冠比显著低于林下。栲幼苗早期的存活和生长依赖种子储存的能量, 受林冠条件影响较弱, 后期则依赖光合作用, 受林冠条件影响较强, 从整个幼苗建成过程看, 大、中林窗更有利于栲幼苗定居。     

缺苞箭竹密度对其生物量分配格局的影响

研究了一个生长季节内缺苞箭竹(Fargesiadenudata)紫果云杉(Piceapurpurea)原始林下不同密度缺苞箭竹群落的生物量及其分配格局.结果表明,缺苞箭竹群落生物量、净生产量、平均单株生物量、地上部分生物量、地下部分生物量随密度的增加而增大,而缺苞箭竹地上部分净增长率却随密度的增加而降低.在一个生长季节内,缺苞箭竹地上部分与地下部分生长相关性随密度的增加而增大.除指数生长期(7、8月)外,缺苞箭竹地上部分/地下部分生物量比在生长季节内随密度增加而增大,但在缺苞箭竹生长的指数生长期,中等密度有较大的地上部分/地下部分生物量比.缺苞箭竹生物量在各器官的分配取决于密度和生长时期,密度对缺苞箭竹的生物量分配格局有显著影响.     

格氏栲天然林林窗植物物种多样性与系统发育多样性

林窗环境异质性导致群落物种多样性与系统发育多样性(phylogenetic diversity, PD)存在差异, 研究不同大小的林窗中群落的物种多样性与系统发育多样性有助于揭示林下生物多样性的形成及维持机制。本文以格氏栲(Castanopsis kawakamii)天然林为研究对象, 通过Pearson相关性分析与广义线性模型探讨了林窗内物种多样性与系统发育多样性间的相互关系及其环境影响因素。结果表明: (1)大林窗(面积 > 200 m²)植物种类及多度均高于中林窗(50 m² ≤ 面积 < 100 m²)、小林窗(30 m² ≤ 面积 < 50 m²)和非林窗(面积 = 100 m²)。大林窗群落系统发育结构趋于发散, 中、小林窗和非林窗群落系统发育结构受到生境过滤和竞争排斥综合作用。(2)群落系统发育多样性指数与物种丰富度(species richness, SR)、Margalef丰富度指数和Shannon-Wiener指数均呈显著正相关, 这与林窗内稀有种种类组成多于优势种有关。(3)林窗面积对物种多样性存在显著正效应; 土壤全氮含量对系统发育多样性和系统发育结构存在显著正效应。林窗形成提高了格氏栲天然林群落物种多样性和系统发育多样性, 林窗面积与土壤全氮共同驱动了格氏栲天然林林窗物种多样性和系统发育多样性的变化。     

常绿阔叶林林冠环境对栲幼苗建成的影响

植物幼苗建成阶段是决定种群自然更新的关键生活史阶段。研究林冠环境对常绿阔叶林优势种幼苗建成阶段的影响对该类森林的恢复和管理具有重要意义。2014–2016年,该研究在重庆市缙云山国家级自然保护区的常绿阔叶林的不同林冠环境(大林窗:150 m~2,中林窗:100–150 m~2,小林窗:50–100 m~2,对照:林下)下进行栲(Castanopsis fargesii)种子野外播种实验,并对栲幼苗命运和生长情况进行了3年的持续监测。结果表明:(1)栲幼苗出土时间从7月持续到12月,出苗时间较长,大林窗对幼苗出土具有延迟作用;(2)栲种子野外平均萌发率为(62.8±2.0)%,第3个生长季(2016年)末幼苗平均存活率为(65.1±2.2)%,枯萎是栲幼苗死亡的主要原因;(3)林冠环境对栲种子萌发率及第1个生长季(2014年)末的幼苗存活率无显著影响,对第2个(2015年)和第3个生长季末的幼苗存活率具有显著影响;(4)林冠环境在第1个生长季对幼苗生长无明显影响,但在第2个和第3个生长季具有显著影响,大、中林窗中幼苗总生物量、株高、基径、根长和叶片数显著高于林下,比叶面积显著低于林下;(5) 3个生长季内, 4类林冠条件下栲幼苗的叶质量比和茎质量比升高,根质量比和根冠比降低,并且从第2个生长季开始大林窗中栲幼苗的叶质量比显著高于林下,根质量比和根冠比显著低于林下。栲幼苗早期的存活和生长依赖种子储存的能量,受林冠条件影响较弱,后期则依赖光合作用,受林冠条件影响较强,从整个幼苗建成过程看,大、中林窗更有利于栲幼苗定居。     

华北落叶松人工林林隙大小对其更新的影响

对关帝山华北落叶松人工林20个林隙的幼苗(高度<1 m)、幼树(高度≥1 m,胸径<5 cm)进行调查，分析4个面积林隙等级(<60 m²、60～120 m²、120～180 m²、≥180 m²)下华北落叶松幼苗和幼树的更新密度、生长指标和空间分布。结果表明：在不同面积的林隙等级中，幼苗、幼树的生长指标(基径、高度)和幼苗更新密度均在小林隙(14～60 m²)下最好，幼树更新密度在中林隙(60～120 m²)下最大，且同等级面积林隙下幼树密度均大于幼苗密度。华北落叶松幼苗、幼树在小林隙和中林隙下更新状况良好，大林隙(120～180 m²)和特大林隙(≥180 m²)则不利于华北落叶松更新。华北落叶松幼苗、幼树主要分布在林冠投影区域以及林冠空隙区域的边缘。通过补种或间伐等人工干扰手段将林隙面积控制在14～120 m²,可以促进华北落叶松的更新。     

模拟采食干扰下克隆整合对两种箭竹分株种群更新的影响

动物对植物的采食会刺激植物进行补偿性更新生长, 克隆整合效应能够通过分株之间的物质传输增强克隆植物的这种补偿生长。现今对克隆整合效应在箭竹(Fargesia)补偿更新中的作用仍未得到全面认识。2011年10月到2012年11月, 设立了糙花箭竹(Fargesia scabrida)和缺苞箭竹(F. denudata)各40个样方, 分别进行不剪除样方内分株和剪除样方内分株数量的25%、50%、75%四种模拟采食干扰处理, 并将样方四周的根状茎切断或保持连接。从2012年6月起观测并统计了箭竹分株种群的累积出笋率、总出笋率、补充率, 以及新生分株的株高、基径和单株生物量。结果表明: (1)在不剪除分株的样方, 切断根状茎连接显著增加了糙花箭竹的出笋率和补充率, 但降低了新生分株的株高和单株生物量, 也显著降低了缺苞箭竹的出笋率和补充率; (2)保持根状茎连接时, 25%的剪除强度仅仅降低了糙花箭竹新生分株的单株生物量; 同样在保持根状茎连接的条件下, 25%、50%的剪除强度使缺苞箭竹种群的补充率有所降低, 而切断根状茎后缺苞箭竹在25%的剪除强度下的分株补充率反而升高; (3) 75%的剪除强度并未影响两种箭竹新生分株数量更新, 但造成新生分株质量显著下降; 切断根状茎连接显著降低了糙花箭竹的新生分株的株高和基径, 对缺苞箭竹影响不显著。实验证明克隆整合影响了两种箭竹新生分株的萌发、存活和生长, 但不是两种箭竹进行补偿更新的主要机制, 仅在糙花箭竹分株种群受到重度采食干扰后的更新中才起到明显的促进作用; 两种箭竹均能在50%的剪除强度下通过补偿生长恢复种群的稳定, 75%的剪除强度则会造成箭竹新生分株质量的下降。     

Response of clonal plasticity of Fargesia nitida to different canopy conditions of subalpine coniferous forest

The aim of this study is to explore the effects of canopy conditions on clump and culm numbers, and the morphological plasticity and biomass distribution patterns of the dwarf bamboo species Fargesia nitida. Specifically, we investigated the effects of canopy condi-tions on the growth and morphological characteristics of F. nitida, and the adaptive responses of F. nitida to dif-ferent canopy conditions and its ecological senses. The results indicate that forest canopy had a significant effect on the genet density and culm number per clump, while it did not affect the ramet density. Clumps tended to be few and large in gaps and forest edge plots, and small under forest understory plots. The ramets showed an even distribution under the closed canopy, and clus-ter distribution under gaps and forest edge plots. The forest canopy had a significant effect on both the ramets'biomass and biomass allocation. Favourable light conditions promoted ramet growth and biomass accumulation. Greater amounts of biomass in gaps and forest edge plots were shown by the higher number of culms per clump and the diameter of these culms. Under closed canopy, the bamboos increased their branching angle, leaf biomass allocation, specific leaf area and leaf area ratio to exploit more favourable light conditions in these locations. The spacer length, specific spacer length and spacer branching angles all showed significant differences between gaps and closed canopy conditions. The larger specific spacer length and spacer branching angle were beneficial for bamboo growth, scattering the ramets and exploiting more favourable light conditions. In summary, this study shows that to varying degrees, F nitida exhibits both a wide ecological amplitude and high degree of morphological plasticity in response to differing forest canopy conditions. More-over, the changes in plasticity enable the plants to optimize their light usage efficiency to promote growth and increase access to resources available in heteerogeneous light environoments.     

不同生境下刺五加种群构件生物量结构与生长规律

从构件水平对针阔混交林、蒙古栎林和次生杂木林3个生境的刺五加种群各功能构件的生物量结构及生物量比率与年龄之间的关系进行了定量分析. 结果表明,刺五加种群的个体生长和各构件生物量动态与环境条件关系密切.不同环境条件下,刺五加种群各功能构件生物量平均值具有相同的规律：茎构件>根茎构件>叶构件.3个生境中刺五加整体种群水平具有一定的相似性.在郁闭度为40%的蒙古栎林中,刺五加种群个体生物量和各构件生物量较针阔混交林和次生杂木林中大.不同生境下刺五加种群分株生物量的差异蕴涵着重要的生长调节和物质分配策略.在分株较小的幼龄个体以及郁闭度较大而不利于分株生长的次生杂木林中,分株优先建造叶器官,以保证充分的物质生产. 3个生境刺五加种群的叶、茎与分株的相对增重均具有相同的幂函数异速生长规律.     

紫耳箭竹克隆形态可塑性对典型冠层结构及光环境的响应

在重庆金佛山国家自然保护内,选择了3种典型群落类型(落叶阔叶林、常绿落叶阔叶混交林和常绿阔叶林),使用Hemiview数字植物冠层分析系统量化群落冠层结构和光环境特征,并对林下紫耳箭竹(Fargesia decurvata)的形态可塑性特征进行调查,分析冠层结构和光环境特征改变下紫耳箭竹形态可塑性的差异,并探讨它们之间的相互关系。结果表明:(1)随着落叶阔叶林"常绿落叶阔叶混交林"常绿阔叶林演替的进行,群落的冠层开度降低,叶面积指数增加,平均叶倾角变小,趋于水平化,冠层对光的截获能力提高,林下光照的强度降低(P0.05)。(2)随着光照强度的降低,紫耳箭竹分株矮小化,叶片变窄,生物量积累降低,但通过增大比茎长、叶面积率和比叶面积提高对光的利用效率,并增大分枝角度和比隔长有效适应弱光环境。(3)在光照条件差的常绿阔叶林下,紫耳箭竹降低对地下茎的投资,将较多的生物量用于秆的增高增长和叶片的生长;而在光照条件好的落叶阔叶林环境下,紫耳箭竹降低对枝、叶生物量的分配,则加大对地下茎的投资,可认为是克隆植物对水分资源所表现的一种觅食行为。研究表明,紫耳箭竹种群随着冠层结构的改变发生了明显的可塑性变化,这些可塑性变化是种群对冠层结构和光环境差异的适应性反应的结果,有利于增强种群对异质生境中光资源的获取和利用;群落内部可以通过调控冠层结构的改变协调和控制小径竹种群的发展。     

Response of clonal plasticity of Fargesia nitida to different canopy conditions of subalpine coniferous forest

The aim of this study is to explore the effects of canopy conditions on clump and culm numbers, and the morphological plasticity and biomass distribution patterns of the dwarf bamboo species Fargesia nitida. Specifically, we investigated the effects of canopy conditions on the growth and morphological characteristics of F. nitida, and the adaptive responses of F. nitida to different canopy conditions and its ecological senses. The results indicate that forest canopy had a significant effect on the genet density and culm number per clump, while it did not affect the ramet density. Clumps tended to be few and large in gaps and forest edge plots, and small under forest understory plots. The ramets showed an even distribution under the closed canopy, and cluster distribution under gaps and forest edge plots. The forest canopy had a significant effect on both the ramets’ biomass and biomass allocation. Favourable light conditions promoted ramet growth and biomass accumulation. Greater amounts of biomass in gaps and forest edge plots were shown by the higher number of culms per clump and the diameter of these culms. Under closed canopy, the bamboos increased their branching angle, leaf biomass allocation, specific leaf area and leaf area ratio to exploit more favourable light conditions in these locations. The spacer length, specific spacer length and spacer branching angles all showed significant differences between gaps and closed canopy conditions. The larger specific spacer length and spacer branching angle were beneficial for bamboo growth, scattering the ramets and exploiting more favourable light conditions. In summary, this study shows that to varying degrees, F. nitida exhibits both a wide ecological amplitude and high degree of morphological plasticity in response to differing forest canopy conditions. Moreover, the changes in plasticity enable the plants to optimize their light usage efficiency to promote growth and increase access to resources available in heterogeneous light environments. __________ Translated from Acta Ecologica Sinica, 2006, 26(12): 4019–4026 [译自: 生态学报]     

百里香无性系的克隆生长特征

植物克隆生长及其与生态适应性的关系是当今植物种群生态学研究的热点和前沿课题,但目前小半灌木克隆生长的研究开展不多。百里香(Thymus serpyllum var. asiaticus)是一种具有地面匍匐茎的草本状小半灌木,可在土壤侵蚀剧烈、基岩大面积裸露的砒砂岩区形成百里香单优群落,在维持生态系统稳定方面具有重要的生态学作用。皇甫川流域是砒砂岩大面积分布的典型区域,在这一地区对百里香无性系的克隆生长进行研究,不仅具有重要的学术价值,而且在生态环境建设方面也具有一定的现实意义。在皇甫川流域选择含三级分株的百里香无性系,对其各级分株的总生物量、各构件生物量及数量、各构件生物量占总生物量的百分比及其月变化进行了研究。结果表明: 1)母株与子代相比,在总生物量、构件生物量及数量上占有绝对优势,而且具有体型大、结构复杂的特点; 2)对生物量分配格局的研究显示,母株根的生物量在总生物量中所占的比例最大,其叶所占的比例较低。子代叶的生物量在总生物量中所占的比例最大,其根所占的比例较低;3)不同级别分株在生物量分配上的差异,揭示了相互连接的分株在功能上的差别,母株可能更侧重于养分和水分的吸收,子株则更侧重于光合生产;4)构件枝、茎、花生物量分配比月变化显示,子1代各构件的生长规律与母株的基本一致,子2代与母株和子1代的相比差异较大,分析认为这可能是分株间不同程度的生理整合作用造成的结果。     

The effects of physiological integration on biomass partitioning in plant modules: an experimental study with the stoloniferous herb Glechoma hederacea

Morphological and physiological plasticity are crucial attributes enabling plants to acquire resources from heterogeneous habitats. Although physiological integration can modify biomass partitioning in modules, especially when connected modules experience different conditions, its ecological importance has been largely overlooked. This experiment examined its effects on above- and belowground biomass partitioning by modules in the stoloniferous herb Glechoma hederacea. We studied how biomass allocation to roots by younger ramets was affected by connection to older ramets, and by nutrient conditions. A lower proportion of biomass was allocated to roots by younger ramets growing under low nutrient (LN) conditions when connected to older ramets in high nutrient (HN) conditions than when they were isolated, demonstrating localised modification of biomass partitioning due to physiological integration. The proportion of biomass allocated to roots by younger ramets was also lower when connected to older ramets in HN conditions than when connected to older ramets in LN conditions. Thus, the effect of integration on biomass partitioning depended on the nutrient conditions experienced by connected ramets. Such changes in biomass partitioning would result in more extensive stolon growth, and greater lateral displacement of new ramets. Understanding the ecological implications of phenotypic plasticity in plants will require further examination of the effects of physiological integration when connected modules experience contrasting growing conditions. This study demonstrates that such integration affects the biomass allocation strategy of connected ramets, enhancing resource acquisition in heterogeneous habitats. The widespread success of clonality in many communities is likely to be strongly promoted by this characteristic.     

Effects of Aspect on Clonal Reproduction and Biomass Allocation of Layering Modules of Nitraria tangutorum in Nebkha Dunes

The formation of many nebkha dunes relies on the layering of clonal plants. The microenvironmental conditions of such phytogenic nebkha are heterogeneous depending on the aspect and slope. Exploring the effects of aspect on clonal reproduction and biomass allocation can be useful in understanding the ecological adaptation of species. We hypothesized that on the windward side layering propagation would be promoted, that biomass allocation to leaves and stems of ramets would increase, and that the effects of aspect would be greater in the layering with larger biomass. To test these hypotheses, we surveyed the depth of germination points of axillary buds, the rate of ramet sprouting, the density of adventitious root formation points, and the biomass of modules sprouting from layering located on the NE, SE, SW and NW, aspects of Nitraria tangutorum nebkhas. The windward side was located on the NW and SW aspects. The results indicated that conditions of the NW aspect were more conducive to clonal reproduction and had the highest rate of ramet sprouting and the highest density of adventitious formation points. For the modules sprouting from layering on the SW aspect, biomass allocation to leaves and stems was greatest with biomass allocation to adventitious roots being lowest. This result supported our hypothesis. Contrary to our hypothesis, the effects of aspect were greater in layering of smaller biomass. These results support the hypothesis that aspect does affect layering propagation capacity and biomass allocation in this species. Additionally, clonal reproduction and biomass allocation of modules sprouting from layering with smaller biomass was more affected by aspect. These results suggest that the clonal growth of N. tangutorum responses to the microenvironmental heterogeneity that results from aspect of the nebkha.