沙丘坡面异质性小生境中准噶尔无叶豆对水分条件变化的响应

以多年生克隆植物准噶尔无叶豆(Eremosparton songoricum(Litv.)Vass.)为材料,选择河边(A种群)和沙漠腹地(B种群)两个沙丘,研究从沙丘底部至顶部,沿着水分条件连续变化的梯度,准噶尔无叶豆在分株种群和克隆片段水平的形态变化特征,以期能揭示其在异质性小生境内利用水分资源的对策,并为准噶尔无叶豆的资源保护、培育和利用提供有意义的参考.研究发现:①在分株种群水平,A种群分株高度及地上部生物量显著高于B种群,而B种群地下部(根)的生物量则显著高于A种群;②在克隆片段水平,随着沙丘底部至顶部,A种群与B种群克隆片段高度和地上生物量都减小,而分株密度都增加,但升高或降低的强度不同;A种群根的生物量和长度增加,主要是水平的位于地下0～10 cm层面的直径10mm以下的根长度增加,而B种群根的生物量减小,但长度却在增加,主要是水平的位于地下0～10 cm层面的直径6mm以下的细根长度增加.水平细根的长度增加,更利于无性系进行广泛觅食,同时促进无性系尽快越过不利生境斑块和提高分株在有利生境中的生长概率.结果表明,准噶尔无叶豆对沙丘坡面水分条件连续变化的异质性小生境存在分株种群及克隆片段两个等级的可塑性响应,并通过可塑性变化适应了沙丘坡面水分条件的分异.     

人工固沙区与流沙区准噶尔无叶豆种群数量特征与空间格局对比研究

准噶尔无叶豆(Eremosparton songoricum) 是中亚荒漠特有小半灌木,稀有种,在我国仅片段化分布于新疆古尔班通古特沙漠;它是流动沙丘的先锋物种,也能成功定居于人工固沙区(草方格)。对比研究了人工固沙区及自然流沙区的准噶尔无叶豆群落物种组成、种群密度、高度、盖度、生物量等特征及种群空间分布格局。结果表明:人工固沙区内的准噶尔无叶豆群落物种的科、属、种数均明显多于流沙区,而且2种生境物种相似性指数仅为0.522,表明人工固沙生境显著改变了群落物种组成与结构。除植株密度外,人工固沙区种群的高度,盖度,地上、地下及总生物量密度,单株地上、地下及总生物量等数量特征均显著高于流沙区。2种生境中准噶尔无叶豆种群空间分布的关联维数均接近2.0,体现了较强的个体空间相关性和空间占据能力。2种尺度(1 m和2 m)下的聚集度分析表明,2种生境中种群基本为随机分布。因而,人工固沙生境并未明显改变种群空间分布格局,这可能是种群的固有特性。人工固沙区的土壤有机质、全氮和有效氮含量显著高于流沙区,且与种群数量特征(植株密度除外)呈显著正相关。总之,与流沙区相比,人工固沙使地表得到固定,提高了土壤有机质和氮素含量,明显改变了定居于其中的准噶尔无叶豆种群主要数量特征,增强了种群的生存和适应能力。该结论对荒漠濒危植物的保育具有重要指示意义。     

荒漠绿洲边缘区泡泡刺种群对风沙干扰的响应

研究了位于黑河中游荒漠绿洲外缘 (荒漠与绿洲过渡带 )戈壁与沙漠两种生境下泡泡刺种群对风沙干扰的响应。结果表明 ,因风沙干扰的响应 ,泡泡刺种群以斑块状格局形式存在 ;独立灌丛沙堆的形成途径主要有 4种 :种子发育、根蘖繁殖、较大的沙堆退化 (生境破碎化 )和邻近沙堆的兼并。水分条件是泡泡刺种群生长的重要制约因素 ,长期对有限的水分资源竞争 ,使其大小和空间分布都形成了明显的特征。绿洲与戈壁交错带 (类似戈壁生境 )样地的泡泡刺种群的沙埋深度、种群高度、种群盖度、种群大小、生物量都小于绿洲与沙漠交错带 (类似沙漠生境 )样地的泡泡刺种群的相应的种群特征变量 ,而绿洲与戈壁交错带样地的泡泡刺种群密度却大于绿洲与沙漠交错带样地的相应的种群特征变量。绿洲与戈壁交错带样地的泡泡刺种群密度为33.3株 / m2 ,而绿洲与沙漠交错带样地的泡泡刺种群密度为 2 5 .4株 / m2 。两样地泡泡刺种群的沙埋深度、种群高度、盖度、密度及大小存在着显著差异 ,只有生物量无显著差异。两样地中泡泡刺的沙埋深度对泡泡刺种群特征的影响明显 ,无论戈壁还是沙漠生境 ,沙埋深度与种群高度、种群大小、种群生物量明显正相关。适度沙埋可以促进泡泡刺种群的生长 ,但当沙埋深度超过10 0 cm时 ,又会对泡泡刺种群     

不同生境下细叶鸢尾表型可塑性及生物量分配差异性

细叶鸢尾(Iris tenuifolia Pall.)为鸢尾科鸢尾属多年生草本,是干旱半干旱地区的野生观赏花卉之一。该植物在自然条件下萌发率极低,主要靠多年生根系进行繁殖。为了研究细叶鸢尾在异质性生境中的可塑性及其生态适应性,对3种不同生境(种群1:干旱严重;种群2:较干旱;种群3:相对湿润)中的细叶鸢尾自然分布种群的生物学特性,地上、地下生物量分配等进行比较。研究发现:3个不同生境的种群植株在外部形态上差异明显。种群1的根长与根冠比显著大于种群2和种群3,种群3的株高值较大;种群3地上生物量的比重显著大于种群1和种群2,但地下生物量与地上生物量分配呈反比。结果表明:细叶鸢尾植株具有较强的可塑性,在不同的环境条件下,通过外部形态结构和生物量的配比关系,能够形成有利于自身生长发育的适应对策。     

不同生境下假臭草生长特征分析

通过测定庄稼地、弃耕地、桉树林、公路边4种生境下假臭草叶面积(LA)、比叶面积(SLA)、LDMC和株高等形态特征、植株各构件生物量及生物量分配比和假臭草种群盖度、密度等种群数量性状,研究不同生境类型下假臭草个体的形态特征、生物量及生物量分配和种群生长状况的差异。结果表明:光照和土壤养分对假臭草的形态特征、生物量及生物量分配和种群数量性状的影响显著;低光照、高土壤养分生境下假臭草的LA、SLA、株高较高,假臭草叶片的LDMC较小,植株的花序数较少,生物量主要向叶和茎器官进行分配,根和花果器官的生物量分配则受到限制;高光照、养分贫瘠生境下假臭草的LA、SLA、株高较低,假臭草叶片的LDMC较大,生物量主要向根和花果器官进行分配,种群的盖度、密度、生物量和高度等种群数量性状较低,种群生长状况较差;高光照、肥沃养分生境为假臭草的最适生境,假臭草构件的资源配置较为优化,株高、基径粗、冠宽和花序数等形态指标较大,种群盖度、密度、生物量等种群数量性状较大,种群生长较好。     

种群密度对大果虫实形态特征与异速生长的影响

为揭示种群密度与植物形态特征、器官生物量间的异速生长关系,阐明植物在退化土地恢复过程中的适应策略,以大果虫实为材料,通过异速生长分析方法,研究种群密度对其形态特征、生物量分配与异速生长的影响。结果表明,种群密度对大果虫实的株型构建产生了显著地影响。随着密度增大,大果虫实株高呈减小趋势,其分枝数及分枝长度明显减小。大果虫实各器官生物量随密度增大而显著减小。随密度增大,茎和繁殖器官生物量分配呈减小趋势,根和叶片生物量分配呈增大趋势。这与最优化分配理论中水分、营养物质及光资源受限时的情况一致。密度对大果虫实株高:根生物量间异速生长具有显著影响,且对株高与器官生物量间异速指数和个体大小产生了极显著影响。密度对根:地上部分、叶片:根、繁殖器官:根生物量间的影响属于表观可塑性,而对根:茎、茎:地上部分、叶:其他器官及繁殖器官:其他器官生物量间的影响属于真正可塑性,说明密度改变了大果虫实的株型发育系统,并影响各器官间的异速生长,进而权衡器官生物量分配以完成生活史。     

不同生境下入侵植物胜红蓟种群构件生物量分配特性

为探究入侵植物胜红蓟在种群构件水平上的生长特性,以更好地了解其整体的生存策略和入侵性,通过测定胜红蓟各构件的生物量、各构件生物量相关性、繁殖构件/营养构件比和根冠比,对Ⅰ房前屋后、Ⅱ菜园地、Ⅲ桔园和Ⅳ路边4种不同生境下的胜红蓟种群构件的生物量分配进行了分析研究。结果表明:不同生境下胜红蓟种群各构件的生物量既存在差异,又存在相同的趋势,即:茎生物量叶生物量根生物量花果生物量;生境Ⅱ胜红蓟总生物量、茎、叶和花果生物量均最大,生境Ⅳ的均最小,两者之间均存在显著差异(P0.05);不同生境中胜红蓟种群的根、茎、叶、花果生物量分配与其各构件生物量大小顺序并不一致,前者各生境间的差异小于后者各生境间的差异;4种生境下繁殖构件与营养构件生物量比大小顺序依次为ⅣⅢⅡⅠ,其值分别为0.069、0.061、0.050和0.042,根冠比大小顺序依次为ⅢⅠⅣⅡ,其值分别为0.211、0.179、0.154和0.090;胜红蓟植株株高与各构件及各构件之间存在极显著正相关(P0.01),株高与各构件的相关性中,株高与茎生物量的相关性最强,相关系数为0.525,各构件的相关性中,茎生物量与叶和花果生物量的相关性较强,其相关系数均大于0.900。胜红蓟种群在不同生境下各构件的生物量、繁殖构件/营养构件比和根冠比表现出的差异显示了其较强的表型可塑性和对异质环境的适应性,可能增强了其入侵性。不同生境下胜红蓟种群各构件生物量相同的趋势及生物量分配比间相对较小的差异显示了胜红蓟种群构件生物量分配模式的相对稳定性。胜红蓟种群各构件间的极显著相关性显示了其整体性的生存策略。     

不同光照条件下聚花过路黄的克隆构型和分株种群特征

以都江堰灵岩山常绿阔叶林林下、林缘和林缘旷地3种不同光照环境中匍匐茎草本聚花过路黄(Lysimachla congestiflora)为对象,对其匍匐茎节间长、分枝强度、分枝角度、分株种群密度、分株种群生物量等指标进行了测定和分析．结果表明,聚花过路黄的匍匐茎节间长、分枝强度、分株种群密度和分株种群生物量在3种生境间差异显著．Kruskal Wallis检验表明,匍匐茎节间长度和分枝角度的频次分布在3种生境间差异显著．3种生境中匍匐茎节间长度分布偏斜度(skewness)的大小分别为：林缘旷地>林缘>林下．林缘旷地与林缘和林下生境在分枝角度、分株种群高度和分株种群根冠比差异显著,而且林缘旷地生境中分枝角度分布偏斜度最小．林缘和林下生境在分枝角度、分株种群高度和分株种群根冠比差异不显著．从林缘旷地、林缘到林下,聚花过路黄的克隆构型和分株种群特征发生了相应改变．结合克隆植物对资源的利用对策,讨论了不同生境中聚花过路黄克隆构型和分株种群特征可塑性的生态适应意义．     

荒漠草原异质生境下猪毛蒿种群动态

以荒漠草原猪毛蒿种群为对象,设置风沙土、灰钙土和基岩风化沉积土3个生境下增水30%、减水30%和对照3个处理,编制猪毛蒿种群的动态生命表,绘制种群存活和死亡曲线,分析异质生境下猪毛蒿种群动态规律,研究猪毛蒿年内年际间种群动态特征.结果表明: 猪毛蒿种群存活曲线趋于Deevey-Ⅰ型,即生长前期存活率较高,趋于稳定状态,死亡率基本维持在较低的水平,生育期末种群死亡率则上升较快.各生境下猪毛蒿个体的存活数均呈前期波动后期下降的趋势,风沙土和灰钙土生境下死亡率的波动变幅较大,增减水处理对猪毛蒿种群死亡率的影响无显著差异.土壤类型对猪毛蒿株高、冠幅、密度、盖度和生物量均有显著影响,降水处理对猪毛蒿株高、冠幅和盖度影响显著,对密度和生物量无显著影响,而土壤类型和降水处理的交互作用仅对株高和冠幅有显著影响.基岩风化沉积土生境下猪毛蒿生物量的可塑性指数显著高于风沙土和灰钙土,灰钙土和基岩风化沉积土生境下盖度可塑性显著高于风沙土.增水处理的密度可塑性指数显著高于对照和减水处理.株高和冠幅的可塑性指数较高,说明在不同的生境压力下,猪毛蒿可以通过优先调节这两个参数来应对生境的变化.     

不同居群准噶尔无叶豆果实和种子特性及种子萌发差异

研究准噶尔无叶豆(Eremosparton songoricum) 6个居群间果实和种子特性及种子萌发差异, 以揭示异质生境下准噶尔无叶豆果实和种子的生态适应机制。结果显示: 居群间准噶尔无叶豆的植株距离(F = 2.34, p = 0.03)和植株冠幅(F = 8.49, p < 0.01)存在显著差异, 沙漠北缘E、F居群和沙漠腹地C居群(受人类干扰剧烈)的植株距离和植株冠幅高于沙漠腹地A、B、D居群; 居群间准噶尔无叶豆果实和种子的长度、宽度、厚度、重量存在显著差异, 居群E、F和C的大部分参数显著高于其他沙漠腹地居群; 居群间果实多子性(F = 6.96, p < 0.01)也存在显著差异, 居群C的果实多子性最高((32.50 ± 4.79)%); 萌发结果表明, 居群间新鲜的成熟种子萌发不存在显著差异, 且萌发率都低于15%; 所有居群的大部分种子都存在物理性休眠现象, 人为划破种皮能显著提高种子萌发率, 但在低温(15/5 ℃)条件下, 所有居群的种子萌发率都较低, 说明低温抑制了种子萌发; 经人为划破种皮解除物理休眠后, 种子的休眠没有完全释放, 居群C、E和F (大种子居群)的种子萌发率显著高于居群A、B和D (小种子居群) (F = 30.77, p < 0.01), 说明准噶尔无叶豆种子不仅存在物理性休眠现象, 也可能存在生理休眠现象。不同程度的种子复合休眠可能是准噶尔无叶豆不同居群适应古尔班通古特沙漠的重要生存策略。     

根茎型木本克隆植物准噶尔无叶豆的种群数量动态

 根据根茎型木本克隆植物的特征, 不以种群的分株数量代表种群大小, 而尝试以不同茎级的根茎长度代表种群大小, 运用种群静态生命表、存活曲线、生殖力表和Leslie矩阵模型, 研究了准噶尔无叶豆(Eremosparton songoricum)的两个种群——A种群(46°31.09′ N, 88°33.06′ E, 紧邻乌伦古湖)和B种群(46°28.07′ N, 88°33.07′ E, 位于沙漠腹地)的种群数量动态。结果表明: 种群存活表现为Deevey-I型。A种群在中龄阶段受到的人为干扰较大, 死亡率出现高峰, 种群的净增长率(R₀)、内禀增长率(r_m)和周限增长率(λ)较低, 表现为衰退型种群, Leslie矩阵模型的模拟结果表明, 15 a内种群呈现下降趋势; B种群所受到的压力主要是干旱贫瘠的荒漠环境所导致的系统压力, 种群的R₀、r_m和λ值适中, 表现为缓慢增长型种群, Leslie矩阵模型的模拟结果表明, 15 a内种群呈现先下降、再上升的趋势。此外, 研究结果验证了Leslie矩阵模型可以扩展应用到根茎型木本克隆植物这类特殊生活型植物的种群数量动态研究上。     

水分异质性影响两种根茎型克隆植物赖草和假苇拂子茅的水分存储能力

水分在自然系统中呈异质性分布。有关水分异质性对克隆植物生长、形态和生理影响的研究已有大量的工作, 但是水分异质性对克隆植物存储能力, 尤其是水分存储能力影响的研究却十分缺乏。该文将两种根茎型克隆植物赖草(Leymus secalinus)和假苇拂子茅(Calamagrostis pseudophragmites)进行水分异质性和同质性实验处理, 探讨水分异质性对克隆植物水分存储能力、生长和形态的影响。在异质性水分处理下, 两种克隆植物的间隔子、枝和根的含水量均显著增加。两种克隆植物对水分异质性分布的适应策略有所不同, 赖草通过降低单个克隆分株的生长、提高芽的数量以应对水分异质性, 而假苇拂子茅通过增强整个分株种群的地下部分(根状茎、根和芽)生长来应对水分资源的异质性分布。水分储存能力的增强可以提高克隆植物适应水分异质性的能力。     

Ecological benefits of integration of Calamagrostis epigejos ramets under field conditions

We established two independent experiments to estimate the ecological consequences of artificial severing on ramets of the competitively strong perennial grass Calamagrostis epigejos. We compared the responses of mature ramets of different size growing in different environments. Finally, we tested whether the response of young ramets to the severing depends on the density of surrounding vegetation.Severing decreased biomass and number of rhizomes of young ramets but did not affect their competitive tolerance. It decreased survival of mature ramets (probably due to traumata caused by cutting attached rhizomes) but did not influence total biomass of survived ramets. The response of total biomass of mature ramets to severing changed with size of the ramets. Further, biomass allocation to rhizomes changed differently after ramet severing in the two environments.The results suggest that field grown young ramets of Calamagrostis benefit from physiological integration. In contrast, mature ramets seem to be independent units according to the resource economy. Ecological benefits of integration depended on size of ramet clumps: ramets in clump had larger survival probability than control ramets. They also depended on environmental conditions: severing increased formation of new rhizomes at a sand dune subsrate, but it had a negative effect on plants in the forest experimental site. This intra-specific variation should be taken into account when trying to explain ecological patterns of integration benefits of clonal plants.     

增加水分与养分对克隆植物羊柴自然种群繁殖权衡的影响

 羊柴(Hedysarum laeve)是豆科多年生半灌木,在自然条件下可以同时进行有性繁殖和克隆繁殖。该文在野外条件下研究了不同水平的水分和养 分处理对羊柴种群的繁殖权衡的影响。结果表明,与对照相比,增加一定量的水分处理显著减少了花和荚果的生物量;显著增加了克隆分株枝 的生物量,显著减少了分株根茎的生物量, 但没有影响其它部分的生物量。增加一定量的水分会抑制有性繁殖,改变生物量对克隆繁殖分株各 部分的分配比例。与对照相比,增加一定量的养分能够促进有性繁殖,抑制克隆繁殖。     

CLONAL INTEGRATION ACROSS A SALT GRADIENT BY A NONHALOPHYTE,HYDROCOTYLE BONARIENSIS (APIACEAE)

This study examined the benefits associated with resource sharing among interconnected ramets spanning a soil salinity gradient. Clones of Hydrocotyle bonariensis, a rhizomatous dune perennial, expand into salt marsh communities from surrounding upland dune systems in coastal North Carolina. In rhizome-severing experiments conducted under both field and laboratory conditions, Hydrocotyle was shown to proliferate ramets under saline conditions, provided that these ramets were connected to other ramets growing in nonsaline conditions. Ramets that benefited from resource integration did not appear to be affected by local salt exposure in that these ramets were morphologically similar to those grown under nonsaline conditions. Supporting ramets incurred no net cost in terms of biomass or ramet production, but there was an increased percent allocation to roots and rhizomes. Ramets grown in saline conditions without the benefit of clonal integration showed high mortality and produced little or no net clonal growth. It is likely that the acropetal movement of water allowed Hydrocotyle clones to ameliorate the heterogeneous saline conditions associated with coastal environments.     

The significance of rhizome connection of semi-shrub Hedysarum laeve in an Inner Mongolian dune,China

Hedysarum laeve, a rhizomatous clonal semi-shrub, commonly dominates the inland dunes in semiarid areas of northern China. This species propagates vegetatively by extension of horizontal woody rhizomes resulting in programmed reiteration of apical and/or axillary meristems. In this study, the plants were experimentally manipulated by cutting rhizome connections and ¹⁴C-labelling techniques were employed to investigate the ecological significance of rhizome connections within the H. laeve clone. Severance of rhizome connections had a great effect on the performance of young ramets within a clone. Young ramets severed from their parent ramets experienced a significant reduction both in ramet growth and vegetative propagation, as compared with the intact young ramets. Within clonal fragments, consisting of three interconnected ramets including a mother ramet, a daughter ramet and a granddaughter ramet, ¹⁴C-photosynthates from the fed leaves of mother ramets were acropetally transported to all clonal component parts. The ¹⁴C-photosynthate translocation within the clonal fragment provides evidence that the young ramets were supported by their parent ramets. Our results suggest that the woody rhizome connections among the interconnected ramets are ecologically and strategically important for the species to grow in the sand dune habitat.     

Clonal plant allocation to daughter ramets is a simple function of parent size across species and nutrient levels

The evolution of clonal growth is a widespread phenomenon among plant species, characterized by the production of genetically identical clonal fragments (ramets) via rhizomes or stolons that form an interconnected clonal organism (genet). Clonal plant species are known to differ in their investment into ramet production, and exhibit considerable variation in ramet morphology both within and among species. While patterns of resource allocation are thought to be linked to a number of plant characteristics, many analyses are limited by uncertainty in how clonal plants determine the morphology and resources allocated to new ramets. In this study, we attempted to discern what aspects of parent ramets best predicted resource allocation to new daughter ramets, and the relationship between resource allocation and daughter ramet rhizome morphology. We grew two sedge species, Schoenoplectus tabernaemontani and Eleocharis elliptica, in a greenhouse under two levels of fertilizer addition. By harvesting daughter ramets that had initiated stem production, yet remained aphotosynthetic, we were able to isolate parental investment into non-independent daughter ramets at a point where daughter ramet spacer length became fixed. Our results indicate that parent ramets allocated a non-linear proportion of parent rhizome biomass to the production of daughter ramets. Moreover, this relationship was unaffected by environmental nutrient availability. Daughter ramet biomass, in turn, was strongly correlated with daughter ramet spacer length. These observations shed light on key processes governing clonal growth in plants, and their potential application in unifying allocational and morphological perspectives to explore the fitness implications of variability in clonal growth.     

Life-history monographs of Japanese plants 14: Cardamine leucantha (Tausch) O. E. Schulz (Brassicaceae)

The life-history characteristics of Cardamine leucantha (Tausch) O. E. Schulz (Brassicaceae) are described. The species is an herbaceous perennial that favors open but relatively moist habitats. It is distributed from Kyushu to Hokkaido in Japan but also occurs in Korea, Mongolia, China and the Russian Far East. In southwestern Japan, shoots start sprouting from mid- to late April, reaching approximately 30–70 cm in height, with 5–10 compound leaves. Ramets simultaneously produce one or more stoloniferous rhizomes that elongate until new ramets are formed at the tips. Cardamine leucantha has a pseudo-annual life cycle, in which mother ramets wither at the end of each season and only daughter ramets appear aboveground in the next year. As a result, ramet positions change annually. In a study population, the number of flowers averaged 23.9 ± 21.0 per ramet and fruit set was 44.2 ± 24.8% (10.4 ± 10.1 fruits per ramet). Ramets produced 3.8 ± 2.3 rhizomes that were 22.0 ± 15.6 cm long. The species sometimes forms large populations. A single genet develops into a group of disconnected ramets spreading via clonal growth. Reproductive characteristics (e.g., fruit set and numbers of flowers and rhizomes) vary among populations, resulting in interpopulation differences in genet structure. Because the reference genome became available recently, established molecular tools will be applied effectively for the investigations of C. leucantha as a model clonal plant.     

EFFECTS OF FIRE ON CLONAL GROWTH AND DYNAMICS OF PITYOPSIS GRAMINIFOLIA (ASTERACEAE)

The rhizomatous perennial Pityopsis graminifolia was studied in a Florida sandhill community in an annually burned site, a periodically burned site, and a site that has been protected from fire since 1965. These different fire regimes significantly affected the demography and life histories of both plants and plant parts in this clonal species. Fires resulted in reductions in ramet biomass and height, and an increase in the (root + rhizome)/shoot biomass ratio. Burning also decreased the total number of flower heads and new rhizomes produced per ramet. However, the survivorship of initiated rhizomes was greater in burned sites and resulted in a larger number of established daughter ramets per clone. As a result, in burned sites there was a shift in clone structure toward larger numbers of smaller ramets, but there were no significant reductions in seed or rhizome production on a per genet basis. The results showed that the responses to fire in P. graminifolia are different when measured at the genet vs. ramet level and that the effects of fire on clones can be explained by demographic responses of plant parts. Population regeneration in the study sites was dependent on successful clonal ramet production because no seedling recruitment was observed. This suggests that disturbances other than fire are important for new genet recruitment in these clonal populations.