普洱季风常绿阔叶林次生演替中木本植物幼苗更新特征

以时空替代的方法,将针阔混交林、季风常绿阔叶林的次生林与成熟林等3个处于同一空间下的群落作为次生演替进程中的3个阶段,研究云南普洱地区次生演替过程中的木本植物幼苗更新特征,分析了次生演替过程中木本植物幼苗的物种组成、密度、高度级及与环境因子的相关性。结果表明:在8个共144 m2的幼苗样地中调查木本植物幼苗101种2014株,其中乔木幼苗是主要组成。随着次生演替的进行,木本植物幼苗、乔木与藤本幼苗密度逐渐增加,灌木幼苗密度无显著变化;藤本植物幼苗的物种丰富度随着次生演替进行而增加,乔木与灌木幼苗则无显著变化,成熟季风常绿阔叶林中木本植物幼苗Shannon-Wiener指数要显著小于针阔混交林与次生季风常绿阔叶林。次生与成熟季风常绿阔叶林木本植物幼苗多度随高度级增加而减少,针阔混交林则呈现偏峰曲线,幼苗密度均集中分布在高度20 cm以内,3个群落演替阶段木本植物幼苗物种丰富度随高度级增加呈现偏锋曲线。相似性系数反映出乔木和藤本幼苗的更新来源与群落的物种组成存在着紧密的联系。乔木幼苗密度分布与样地坡度之间存在着显著的负相关,灌木幼苗密度与土壤pH值之间存在着显著正相关。     

哀牢山湿性常绿阔叶林及不同类型次生植被的幼苗更新特征

采用"空间代替时间"的方法,对云南哀牢山徐家坝地区湿性常绿阔叶林及不同类型次生植被中木本植物幼苗的种类组成、数量及更新方式等进行了调查.结果表明:在所有调查样地中,共记录到木本植物幼苗46种,其中乔木31种、灌木15种.荒草地、滇山杨林、栎类萌生林、原生林4种不同植被类型的幼苗密度分别是(421.9±59.5)、(34.4±8.1)、(31.6±7.2)、(18.5±6.6)株·25 m-2.随着演替的进行,灌木的比例逐渐减小,乔木的比例则逐渐增加.在森林恢复演替过程中,处于演替初期的荒草地灌木萌生更新占优势,到了演替后期则以乔木萌生幼苗更新为主.随着群落演替进程的增加,幼苗密度逐渐减少,幼苗生活型和木本植物的幼苗更新方式均发生变化.     

滇中次生常绿阔叶林优势树种的空间格局

采用Ripley的点格局分析方法对滇中次生常绿阔叶林中优势种滇青冈(Cyclobalanopsis glaucoides)和滇油杉(Keteleeria evelyniana)的分布格局以及不同径级分株之间的相互关系进行了分析.结果表明:(1)总体来看,两个优势物种各径级株数分布较均匀,二者的增长处于稳定期.(2)两个优势物种在总体上及不同径级阶段主要呈聚集分布.随径级的增加,滇青冈种群的聚集程度逐渐降低,而滇油杉种群的聚集程度呈现降低-增加-降低的趋势.两个优势物种的幼树、中树和大树主要呈空间正相关或无空间关联性.(3)两优势种群不同径级之间在不同尺度下基本上无关联性,这可能是二者的生存策略存在较大差异而造成的.研究表明,在滇中森林恢复过程中,应结合植物种群的密度控制及种间相互作用来构建群落的结构.     

云南松天然林区植物群落结构的空间动态研究

于2005年4月采用中点四分法对云南省禄丰县一平浪林场云南松天然林区植物群落结构及空间动态进行了实地调查研究.结果表明:从山坡上部到下部,物种的组成越来越丰富,其中的优势种群云南松属于衰退型、滇油杉属于增长型、黄毛青冈属于稳定型.根据更新层的重要值来判断,山坡上部将演替成为以滇油杉为主的单优群落;山坡中部将演替成为滇油杉和黄毛青冈为主的针叶、阔叶混交林;山坡下部将演替成为以黄毛青冈为主的常绿阔叶林.研究还表明,在林冠层中,除了滇油杉在山坡上部为聚集分布外,其他树种的分布格局均呈随机分布;在更新层中,优势种类均为聚集分布.     

吉林蛟河不同演替阶段针阔混交林乔木幼苗数量组成及其时间动态

为了解蛟河地区木本植物幼苗的数量特征及其年际动态, 以及影响幼苗数量的生态学过程, 该文以不同演替阶段针阔混交林监测样地为基础, 在样地内共设置了451个种子雨-幼苗观测样站.通过2012-2014年连续3年的幼苗监测调查, 对木本植物幼苗物种组成,数量特征,高度级结构及其年际动态进行了分析, 并用多元回归的方法分析了幼苗数量与林分类型,同种成体胸高断面积之和及林冠开阔度的关系.结果表明: (1)林下幼苗层主要木本植物的组成在不同演替阶段林分中差异不大, 并与样地内主要树种组成保持着一定相似性, 但幼苗重要值排序在年际间和不同演替阶段稍有波动.(2) 3个样地内幼苗个体数量和物种数都主要集中在5-20 cm高度级之间, 之后随高度级增加而逐渐减少, 但幼苗个体数量减少更快.表明与种间竞争相比, 物种内部的相互作用是导致幼苗死亡的主要原因, 证实了负密度制约效应在幼苗更新过程中的作用.(3) 5个主要树种的幼苗数量都与同种成体胸高断面积之和呈显著的正相关关系, 还呈现出明显的生境偏好, 表明扩散限制和生态位过程也会影响幼苗的数量和分布.该文间接证实了负密度制约效应,扩散限制和生态位过程对温带地区不同演替阶段森林群落中幼苗数量组成的影响.     

吉林蛟河不同演替阶段针阔混交林木本植物幼苗空间分布与年际动态

幼苗的空间分布对群落更新具有重要意义。于2011年夏季在蛟河地区3个不同演替阶段针阔混交林样地内共设置了451个种子雨-幼苗观测样站,并于2012—2014年对幼苗样方内胸径1 cm的木本植物幼苗进行了连续3a的调查。对木本植物幼苗的数量分布以及水曲柳(Fraxinus mandshurica)、五角枫(Acer mono)、杉松(Abies holophylla)和东北枫(Acer mandshuricum)4个主要树种幼苗与大树的空间关系及其年际变化进行了分析,并用Syrjala检验分析了样站尺度上幼苗密度、丰富度空间分布在年际间的差异。结果显示:(1)木本植物幼苗的数量分布在不同群落和年际之间都表现出明显的差异,样站尺度上3个样地内的幼苗密度存在着较大的空间变异性,而幼苗丰富度的空间异质性较低。表明幼苗分布的空间异质性对幼苗密度有着重要影响。(2)样站尺度上幼苗数量和物种数的空间分布在不同年份之间是存在差异的。在一定程度上证实了种子产量、扩散方式和群落组成对幼苗空间分布的影响。(3))从死亡幼苗与大树空间关系来看,4个主要树种的死亡幼苗与成树表现出相似的分布格局,这既表明成体植株的空间分布特征也能够影响幼苗的空间分布格局,也从侧面说明了负密度制约效应对幼苗空间分布的影响。     

滇青冈幼苗的光合和生长对不同生长光强的适应

研究了3种光强(全日照的80%、20%和2%)下生长的滇青冈幼苗的光合-光强响应和生长状况。结果表明:3种光强生境滇青冈幼苗光饱和点和最大净光合速率没有显著差异,20%和2%生长光强下光补偿点显著低于80%生长光强。3种光强生境幼苗的光合饱和点在400~450μmol·m-2·s-1左右,光补偿点也很低(6~10μmol·m-2·s-1),表现出阴生植物的特征。但滇青冈幼苗较强的耐荫能力,是以牺牲生长为代价的。在2%的弱光生境中滇青冈幼苗比叶重显著降低,单位面积和单位干重叶绿素含量均显著增多,具有利用较强辐射的潜力,一旦林窗出现就可以利用增加的辐射提高光合速率,及时进入快速高生长。     

秦岭山地油松群落更新特征及影响因子

采用样方法,对秦岭山地油松次生林群落更新特性和相关环境因子进行了研究.结果表明:油松次生林更新层乔木树种共36种,占总种数的51.4%;优势种有短柄枹栎、锐齿栎和榛子等;幼苗库丰富,高度级及龄级较小;更新方式以实生为主,该森林群落处于演替中期、林分密度对林下乔木幼苗、幼树数量影响极显著(P<0.01),当林分密度从580株.hm-2增加到1500株.hm-2时,林下更新的幼苗、幼树密度呈增加趋势,随着林分密度的继续增加,其密度逐渐减少;坡向对幼苗、幼树密度的影响显著(P<0.05),当坡向由南偏西10°到南偏西40°,林下幼树密度逐渐减少,至阴坡(北偏东10°)幼树数量又逐渐增加.不同坡位林分更新特征各异,平坡林分中幼苗、幼树密度均较大;从坡下向上,幼苗密度逐渐减少,而幼树密度逐渐增大;海拔从1159 m增至1449 m,幼树密度逐渐增加,至1658 m时,幼树密度逐渐减小,而幼苗密度则一直呈现增加趋势.处于阴坡的中密度林分更新效果最佳.合理调整林分密度是加快林分更新的有效途径之一.     

岷江干旱河谷黄蔷薇和川滇蔷薇更新能力及其限制因素

植物种子、种子库和幼苗库的研究是种群更新和植被恢复的重要内容.研究了岷江干旱河谷两种乡土灌木(黄蔷薇和川滇蔷薇)的种子、种子库和幼苗库特征.黄蔷薇和川滇蔷薇种子产量较高, 分别为(627±216)和(7832±4347)果/丛,但种子质量较差,饱满率分别为(49.62±1.03)%和(73.83±3.42)%,活力为(32.25±0.67)%和(55.38±2.5)%.黄蔷薇种子产量和活力低于川滇蔷薇,但其种子重量和休眠程度却远高于川滇蔷薇.两种蔷薇土壤种子库的水平和垂直分布格局相似:在水平分布上,离植株基部越近,种子密度越高;在垂直分布上,种子主要分布在地表层.黄蔷薇和川滇蔷薇种子库密度均较高,分别为(560±90)粒m2和(1955±235)粒/m2,但活力种子数量较少,分别为(26±4)粒/m2和(152±18)粒/m2, 处于休眠状态的种子,黄蔷薇(60.6%)较川滇蔷薇(18.4%)高,可萌发的种子,两者分别为(10±1)粒/m2和(124±14)粒/m2.黄蔷薇(65.3%)的种子库中被啮齿动物取食的比例较川滇蔷薇(0.5%)高.两种蔷薇的幼苗密度差异不大,黄薇幼苗密度((4.28±0.49)株/m2)略小于川滇蔷薇((5.24±1.28)株/m2).幼苗组成的总体趋势是高度级低的幼苗所占的比例相对较高,且Ⅰ级幼苗数显著高于较其他级幼苗数.研究表明,两种蔷薇种子存在休眠,能形成持久种子库,种子特征对种子库和幼苗库具有深刻影响.两种蔷薇更新状况不佳,比较而言,黄蔷薇更新较川滇蔷薇好.黄蔷薇更新的主要限制因素是动物对种子的破坏,而川滇蔷薇更新的主要限制因素是幼苗存活定居.     

长白山次生针阔混交林群落结构特征及群落动态

长白山次生针阔混交林是东北地区典型的次生林类型之一,处于向地带性顶级植被阔叶红松林方向演替的重要阶段。全面分析长白山次生针阔混交林群落动态,能为东北地区森林保护、恢复和可持续经营提供理论支撑。基于长白山地区5.2 hm~2次生针阔混交林动态监测样地,以2005和2017年样地内所有胸径(DBH)≥5cm的木本植物为研究对象,分析了12年间群落的物种组成、物种多样性、胸径结构和密度结构等非空间结构,以及林木空间分布格局、树种混交和林木竞争等空间结构的变化。结果表明:群落个体数量由6614株降至5884株,2017年调查到木本植物13科25属42种,较2005年增加2属2种,群落物种组成和多样性变化较小。平均胸径由14.4 cm增加至15.5 cm;径级结构在两次调查中均呈倒"J"型,虽然群落保持稳定的更新态势,但是各径级的个体数量显著下降。胸径变异系数和基尼系数分别由0.77和0.37增至0.79和0.38,群落复杂程度提高。群落株数密度由1272株/hm~2降至1132株/hm~2,林分密度由33.12 m~2/hm~2增至34.74m~(2/)hm~2。群落空间结构在两次调查中没有变化,群落水平分布格局为随机分布(■=0.51),大小分化程度为中庸分布(■=0.50),混交度为强度混交(■=0.76),空间结构稳定。研究还比较了样地和20m×20m样方尺度上的群落结构变化趋势,发现大部分指标在两个尺度上变化一致。综上所述,本研究中的长白山次生针阔混交林的群落非空间结构呈现正向演替趋势,而空间结构保持相对稳定,群落整体在12年内并未发生本质性改变。在未来经营中,可适当伐除假色槭和色木槭等林下层树种,为红松、蒙古栎和紫椴等顶级树种的生长和更新提供空间。     

滇青冈和高山栲种子昆虫寄生及其对种群更新的影响

对昆明西山地区滇青冈(Cyclobalanopsis glaucoides)和高山栲(Castanopsis delavayi)种子(坚果)虫蛀率在两个年份、整个落种期和山坡不同部位的变化情况进行研究,结果表明：① 寄生于滇青冈种子内的象甲为青冈象(Curculio megadens)和Curculio sp.,寄生于高山栲种子内的象甲是Curculio sp.和剪枝象(Cyllorhynchites sp.);② 滇青冈样地的虫蛀率为2484%~2760%,样地间无显著差异,两年间的平均虫蛀率亦无显著差异;高山栲样地间虫蛀率为1794%~2655%,有显著差异性,两年间的平均虫蛀率也有显著差异。③ 高山栲的下落种子虫蛀率随时间逐步下降,9月和10月中下旬出现两个虫蛀率的高峰期;高山栲种子内寄生的昆虫落地后即入土化蛹,滇青冈落地种子内的寄生昆虫在初期会滞留一段时间;④ 虫蛀率与林下实生苗数量无显著相关性,实生苗多集中在大树周围以及草本层植物较丰富的小生境。     

松嫩平原碱化草甸恢复演替系列的群落种子流分析

对松嫩平原碱化草甸恢复演替系列4个群落优势种种子源库、土壤种子库及幼苗库进行了比较,建立了种子流模型.结果表明:演替初期虎尾草群落优势种种子源库密度最大,为(446182±180455)粒.m-2,分别是演替中期星星草群落的7.2倍、演替后期星星草 羊草群落的11.4倍、演替顶极羊草群落的164.8倍.土壤种子库和幼苗库密度均以虎尾草群落最大,分别为(63650±14541)粒.m-2和(39160±15192)株.m-2,羊草群落最小,分别为(14310±7686)粒.m-2和(790±745)株.m-2,大体呈随着恢复演替进程而递减的趋势.全体演替系列群落输出的实生苗均以虎尾草为主,占79.8%～100%.在种子流中,优势种的种子源库向土壤种子库输入率在10%～35%之间,输出率差异则较大,虎尾草群落高达62.3%,但星星草群落和羊草群落均没有优势种的幼苗输出.     

秦岭山地锐齿栎次生林幼苗更新特征

分析了秦岭山地锐齿栎次生林群落乔木及木本幼苗物种特征值、更新生态位宽度及不同影响因子下（包括林分密度、坡向、海拔等）乔木更新动态。结果表明,乔木层物种有25种,锐齿栎占有明显地优势,重要值为149.18%。其次为华山松(Pinus armandii)和千斤榆(Carpinus cordata)等;更新层共有木本植物41种,其中乔木种有28种,占所有木本植物总数量的68%,优势种有青蛙皮槭(Acer grosseri)、木姜子(Litsea pungens)等。林下有丰富的幼苗库,径级及龄级比较小。更新方式以实生为主;分析了10种更新优势乔木种群的生态位宽度,同种幼树种群生态位宽度均大于幼苗种群生态位宽度。幼苗种群生态位宽度从大到小依次为榛子(Corylus heterophylla)、锐齿栎、青蛙皮槭等。幼树种群生态位宽度从大到小依次为榛子、青蛙皮槭、锐齿栎等;林分密度对林下乔木幼苗和幼树数量影响不同。当林分密度（株?hm-2）从720增加到1460时,幼苗密度逐渐增大。随后随着林分密度的增加,又呈现出减少的趋势。而幼树密度随着林分密度增加一直呈现出减少的趋势。坡向对锐齿栎次生林林下乔木幼苗和幼树的影响各异,阳坡（南偏西3）均利于幼苗和幼树的更新。南偏西45林分内,幼苗密度急剧减少。随着坡向转为阴坡,幼苗密度又逐渐增加。而幼树密度逐渐减少。当海拔从1083 m增加到1547 m时,幼树密度逐渐增加。随后,随着海拔增加到1882 m,幼树密度则逐渐减小。而幼苗密度随着海拔的增高一直呈现减小的趋势。     

Hierarchical neighbor effects on mycorrhizal community structure and function

Theory predicts that neighboring communities can shape one another's composition and function, for example, through the exchange of member species. However, empirical tests of the directionality and strength of these effects are rare. We determined the effects of neighboring communities on one another through experimental manipulation of a plant‐fungal model system. We first established distinct ectomycorrhizal fungal communities on Douglas‐fir seedlings that were initially grown in three soil environments. We then transplanted seedlings and mycorrhizal communities in a fully factorial experiment designed to quantify the direction and strength of neighbor effects by focusing on changes in fungal community species composition and implications for seedling growth (a proxy for community function). We found that neighbor effects on the composition and function of adjacent communities follow a dominance hierarchy. Specifically, mycorrhizal communities established from soils collected in Douglas‐fir plantations were both the least sensitive to neighbor effects, and exerted the strongest influence on their neighbors by driving convergence in neighbor community composition and increasing neighbor seedling vigor. These results demonstrate that asymmetric neighbor effects mediated by ecological history can determine both community composition and function.     

Comparison of fungal communities associated with spruce seedling roots and bryophyte carpets on logs in an old-growth subalpine coniferous forest in Japan

Fungal communities associated with plant tissues were compared between two bryophyte species dominating decaying logs (Scapania bolanderi and Pleurozium schreberi), and roots of spruce seedlings growing on the bryophytes and in the ground soil, to evaluate the contribution of fungal communities to seedling regeneration. Using high-throughput DNA sequencing, a total of 1233 fungal operational taxonomic units (OTUs) were detected. Saprotrophic Ascomycota were dominant in bryophytes, whereas ectomycorrhizal (ECM) Basidiomycota were dominant in spruce roots. Fungal communities were significantly different between the two bryophyte species. In addition, fungal communities of spruce seedlings were significantly affected by the substrates on which they were growing. Some ECM fungi were detected from both of the bryophytes and the spruce seedlings growing on them; however, the dominant OTU identities differed between the two bryophyte systems. The possible effects of functional differences between dominant fungal OTUs on spruce seedling regeneration are discussed.     

Mycorrhizal networks affect ectomycorrhizal fungal community similarity between conspecific trees and seedlings

Ectomycorrhizal (EM) networks (MN) are thought to be an important mode of EM fungal colonization of coniferous seedlings. How MNs affect EM communities on seedlings, and how this varies with biotic and abiotic factors, is integral to understanding their importance in seedling establishment. We examined EM fungal community similarity between mature trees and conspecific interior Douglas-fir (Pseudotsuga menziesii var. glauca) seedlings in two experiments where seed and nursery-grown seedlings originating from different locations were planted at various distances from trees along a climatic gradient. At harvest, trees shared 60% of their fungal taxa in common with outplanted seedlings and 77% with germinants, indicating potential for seedlings to join the network of residual trees. In both experiments, community similarity between trees and seedlings increased with drought. However, community similarity was lower among nursery seedlings growing at 2.5 m from trees when they were able to form an MN, suggesting MNs reduced seedling EM fungal richness. For field germinants, MNs resulted in lower community similarity in the driest climates. Distance from trees affected community similarity of nursery seedlings to trees, but there was no interaction of provenance with MNs in their effect on similarity in either nursery seedlings or field germinants as hypothesized. We conclude that MNs of trees influence EM colonization patterns of seedlings, and the strength of these effects increases with climatic drought.     

Seedling regeneration,environment and management in a semi‐deciduous African tropical rain forest

Questions: How is seedling regeneration of woody species of semi‐deciduous rain forests affected by (a) historical management for combinations of logging, arboricide treatment or no treatment, (b) forest community type and (c) environmental gradients of topography, light and soil nutrients? Location: Budongo Forest Reserve, Uganda. Methods: Seedling regeneration patterns of trees and shrubs in relation to environmental factors and historical management types were studied using 32 0.5‐ha plots laid out in transects along a topographic gradient. We compared seedling species diversity, composition and distribution patterns along topographic gradients and within types of historical management regimes and forest communities to test whether environmental factors contributed to differences in species composition of seedlings. Results: A total of 85 624 woody seedlings representing 237 species and 46 families were recorded in this rain forest. Cynometra alexandri C.H. Wright and Lasiodiscus mildbraedii Engl. had high seedling densities and were widely distributed throughout the plots. The most species‐rich families were Euphorbiaceae, Fabaceae, Rubiaceae, Meliaceae, Moraceae and Rutaceae. Only total seedling density was significantly different between sites with different historical management, with densities highest in logged, intermediate in logged/arboricided and lowest in the nature reserve. Forest communities differed significantly in terms of seedling diversity and density. Seedling composition differed significantly between transects and forest communities, but not between topographic positions or historical management types. Both Chao‐Jaccard and Chao‐Sørensen abundance‐based similarity estimators were relatively high in the plot, forest community and in terms of historical management levels, corroborating the lack of significant differences in species richness within these groups. The measured environmental variables explained 59.4% of variance in seedling species distributions, with the three most important being soil organic matter, total soil titanium and leaf area index (LAI). Total seedling density was positively correlated with LAI. Differences in diversity of >2.0 cm dbh plants (juveniles and adults) also explained variations in seedling species diversity. Conclusions: The seedling bank is the major route for regeneration in this semi‐deciduous tropical rain forest, with the wide distribution of many species suggesting that these species regenerate continuously. Seedling diversity, density and distribution are largely a function of adult diversity, historical management type and environmental gradients in factors such as soil nutrient content and LAI. The species richness of seedlings was higher in soils both rich in titanium and with low exchangeable cations, as well as in logged areas that were more open and had a low LAI.     

昆明西山滇青冈林内滇青冈种子库动态的研究

通过对昆明西山滇青冈林内滇青冈种子库的跟踪取样调查和种子埋藏试验,对滇青冈种子库的动态进行了研究。昆虫在种子成熟前侵入种子,经种子雨进入种子库时已有71．8％的种子失去萌发能力。种子雨输入种子库的绝大部分种子停留在表面种子库,其中48．55％的种子被虫害,25．36％被某些非生物或生物搬运,17．39％的腐烂,8．7％的被动物当场取食,没有种子萌发,影响种子库动态的各种因子的作用大小在时间上是变化。被搬运的种子中,有4．9％的由表面种子库转移到埋藏加。土层是滇青冈种子的安全生境,土壤种子库的存在时间超过250天。埋入土壤的试验种子一直处于静止状态,到6月雨季后有80％种子萌发,20％的腐烂。萌发种子数是当年产种子的0．26％。滇青冈林内的滇青冈种子库是季节性的,种子库对种群个体的补充作用是有限的。     

濒危植物秦岭冷杉地理分布和生物生态学特性研究

秦岭冷杉(Abies chensiensis)为中国特有三级保护植物,仅岛屿化分布于陕西、甘肃、四川、河南的秦巴山地和湖北神农架中高山地区,分布海拔1300—2300m。为了对秦岭冷杉种群进行深入研究和有效保护,通过样地调查和相关资料分析,对其地理分布、群落特性、种群特性进行了系统研究。秦岭冷杉群落包含种子植物40科63属102种,可划分成11个地理区系成分,其中温带分布型属占84．21％,热带分布型属占15．79％,温带分布是群落的基本特性。秦岭山区的秦岭冷杉群落可分为5个群丛,基本特点为：生境阴湿;群落郁闭度大,结构复杂;种群个体年龄差异大;天然更新主要发生在保护较好的阴坡林窗地段。秦岭冷杉苗期生长缓慢,在30—70年龄阶段,高生长旺盛;在40—100年龄阶段,胸径生长较快。秦岭冷杉开始结实年龄在30—40年龄阶段,结实间隔期为3—5年。秦岭山区成熟个体平均产球果8个／株,饱满种子56粒／球果。种群更新的关键问题是种子产量低,林下幼苗稀疏,更新缺乏后继资源。种子向幼苗转化率低是生活史中的脆弱环节。在未来经营保护中,应以就地保护为主,通过抚育管理,促进天然更新;丰年应适时采种,就地育苗,扩大人工种群。