小径竹对亚高山暗针叶林优势 树种林窗更新的影响

在亚高山岷江冷杉林中选取面积≤50m2,50~150m2,>150m2的林窗,每种类型内均包含3种小径竹盖度(≤20%、20%~50%、>50%),共调查林窗9个,并调查包含这3种小径竹盖度的三块林下对照样地,研究了该类森林林窗更新与小径竹生长的关系。结果表明:(1)无论林窗大小,林窗内的更新幼苗数量都比林下的多,林窗更新是岷江冷杉群落更新的主要途径;(2)所选林窗均为发育早期,林窗对更新树种的种类组成和数量的影响主要表现在幼苗上。糙皮桦幼树及幼苗数量随林窗面积的增加而急速增加,它的更新更需要较大的林窗;(3)不同小径竹盖度下幼苗的密度呈现显著性变化,小径竹的生长明显抑制了森林幼苗的更新及填充的进程;(4)华西箭竹的分散程度随林窗面积的增大而降低,而平均高度和基径则有增加的趋势。     

大熊猫栖息地亚高山针叶林结构和动态特征

亚高山针叶林是大熊猫适宜的栖息环境 ,其结构和动态规律严重影响大熊猫的生存和繁衍及其进化潜力的维持 ,是恢复退化大熊猫栖息地的唯一科学依据。通过样方法和中心点四分法调查了大熊猫栖息地 4个亚高山针叶林样地和该样地所在森林的 6 2个林窗 ,研究了该类森林的结构及其更新与大熊猫主食竹生长、林窗干扰的关系 ,以期揭示亚高山针叶林的动态规律 ,为大熊猫栖息地的恢复提供科学依据。研究结果表明 :岷江冷杉 Abies faxoniana和紫果云杉 Picea purpurea是亚高山针叶林的主体 ,其所构成的森林是大熊猫重要的栖息环境。岷江冷杉种群年龄呈现连续分布格局 ,而紫果云杉种群年龄呈间歇性分布 ,其在 2 5 0～ 5 0 0 a之间没有更新。该类森林中普遍存在着林窗干扰和更新现象 ,90 %的林窗下分布有岷江冷杉和紫果云杉更新的幼苗、幼树和大树。但林窗内不同树种更新密度不同 :岷江冷杉更新幼苗、幼树和大树占所有更新树种的 82 .5 % ,而紫果云杉和糙皮桦的更新幼苗、幼树和大树占 11.3%。岷江冷杉和紫果云杉的更新同时受林下大熊猫主食竹生长状况的影响 :主食竹盖度高 ,更新树种幼苗、幼树的密度低 ,即主食竹的生长抑制了岷江冷杉和紫果云杉的更新。     

油松人工林林窗对幼苗天然更新的影响

为了阐明油松人工林间伐产生的林窗对幼苗天然更新的影响,通过典型样地调查,对不同生境的生态因子,以及1 ～ 10年生油松幼苗的年龄结构、高度、基径、针叶长及根、茎、叶生物量进行了调查,并对影响幼苗更新数量的环境因子进行通径分析.结果表明:阴坡与阳坡不同大小林窗下的生态因子存在明显差异.林下缺少3年生以上的幼苗,小林窗下缺少7年生以上的幼苗,大林窗下5、6年生幼苗较少.1 ～10年生油松幼苗可分为1～3年生、4～7年生和8 ～ 10年生3个阶段,1～3年生幼苗的存活是种群能否成功更新的关键.幼苗生长总体表现为阴坡大林窗＞阳坡大林窗＞阳坡小林窗＞阴坡小林窗＞阳坡林下＞阴坡林下.通径分析表明,光照强度对各个阶段的幼苗数量起积极的决定作用,灌木层盖度对4～7年生及8 ～ 10年生幼苗数量均起积极的决定作用,腐殖质干质量对4～7年生幼苗数量起消极作用,但对8～10年生幼苗数量起积极作用.在油松人工林经营中,应进行高强度的间伐,同时清理林下灌木,为油松种群可持续发育提供条件.     

林窗对岷江冷杉更新幼苗根系生物量积累与分配的影响

林窗对更新幼苗生物量积累和分配的影响在种群恢复中具有重要作用,尤其是暗针叶林更为突出。以川西亚高山岷江冷杉原始林林窗、林下的岷江冷杉更新幼苗为对象,研究其粗根和细根生物量的积累与分配过程及其异速生长关系。结果表明：随着岷江冷杉幼苗的生长,粗根、细根生物量均不断积累,但根系生物量中分配到粗根的比例逐渐增加、分配到细根的比例逐渐减少,粗根-细根间生物量的异速生长指数由等速生长转变为异速生长。林窗幼苗的细根生物量、粗根生物量和总根生物量中位数均小于林下幼苗,细根生物量分配比例大于林下幼苗;林窗还显著影响细根、粗根生物量随基径增粗的积累与分配过程,细根生物量分配比例随基径增粗的降幅大于林下幼苗;但林窗对粗根-细根生物量间的异速生长指数影响不显著。说明林窗干扰改变了岷江冷杉更新幼苗地下资源的分配方式,优化了资源配置。     

长苞铁杉幼苗在林窗不同位置的建立

通过2003年12月到2005年1月对福建省天宝岩国家级自然保护区长苞铁杉林内椭圆形林窗(面积118m2)中心、中部、边缘和林下样地进行种子埋藏及幼苗定位观测实验,研究了长苞铁杉幼苗在林窗不同位置中的建立。结果表明:在林窗内不同位置对长苞铁杉幼苗建立有显著影响。林窗中心样地、林窗中部样地、林窗边缘和林下样地内长苞铁杉幼苗发生率分别为10%,10.7%,6%和6%。从林冠下到林窗中心,长苞铁杉种子的幼苗发生率略有增高趋势。在林窗中心样地和林窗中部样地中雨水冲刷是幼苗死亡的最主要原因,而在林窗边缘样地和林下样地中昆虫的取食是幼苗死亡的最主要原因。林窗位置对幼苗的存活率有显著影响,林窗中部样地幼苗存活率最高(11.4%),林窗中心样地幼苗存活率次之(6.7%),而林窗边缘样地和林下样地幼苗则均全部死亡。种子营养消耗完后,在林窗中心、林窗中部、林窗边缘和林下等4个位置样地中,林窗中心样地幼苗平均高度最高。经过一个生长季后,林窗中心样地中幼苗的根生物量、茎生物量和总生物量均略高于林窗中部样地的幼苗,但差异并不显著。林窗中心样地幼苗叶生物量、叶重比、叶/地上等指标显著高于林窗中部样地的幼苗,而茎重比则低于林窗中部样地的幼苗。     

林窗对岷江冷杉幼苗生存过程的影响

岷江冷杉林是我国西南亚高山暗针叶林的主体之一,其健康的种群动态和更新策略在维持长江上游生态安全方面具有重要意义。为了解岷江冷杉更新过程中林窗对幼苗种群动态的影响,通过调查林窗内和林下岷江冷杉更新幼苗的存活情况,采用数量分析方法研究了幼苗的年龄结构和动态特征。结果表明:林窗内和林下岷江冷杉幼苗的年龄结构金字塔均为两头小中间大的纺锤型,存活曲线均为DeeveyⅡ,生命期望以1龄级的最高、林下种群的高于林窗种群。幼苗的死亡率、消失率、累计死亡率和危险率均随龄级的增加而增大,生存率和死亡密度随龄级的增加而减小;岷江冷杉幼苗种群的数量化动态指数Vpi=0.1059,为增长型种群,在外界随机事件干扰时也可持续增长,尤以林窗种群抵御随机干扰、持续增长的特点更为突出;岷江冷杉幼苗种群的周期波动主要受基波控制,林窗种群在9龄级和11龄级处的波动还受谐波的影响。林窗扩大了岷江冷杉更新幼苗的数量规模、提高了种群抵御外界随机干扰的能力。因此,在岷江冷杉林更新管理中,应高度重视林窗的作用。     

卧龙自然保护区亚高山暗针叶林树种更新研究

在卧龙亚高山暗针叶林中选取有代表性的18个林窗斑块以及对照的林下样方进行群落学调查,记录乔木树种和灌木树种的相关数量特征。按照树种在林窗内外的重要值差异结合树种本身生态学特性将群落中出现的乔木层树种划分为先锋组和耐荫组两类生态种组。林窗内外乔木树种的组成明显不同,两类生态种组树种幼苗在林窗与林下环境中的更新表现出差别,这与树种本身的生态学特性以及所处林窗的环境有关,同时亚高山暗针叶林中灌木层优势种小径竹的生长对树种更新方式也产生一定的影响。     

长白山云冷杉针阔混交林不同林隙下幼苗幼树密度及空间分布

2019年8月在云冷杉针阔混交林样地(0.36 hm²),对48个林隙及幼苗(0.2<更新高度RH<1 m)、幼树(RH≥1 m,胸径DBH<5 cm)进行调查,分析林隙大小(<20 m²,小;20～50 m²,中;50～120 m²,大;>120 m²,特大)对林隙内红松、鱼鳞云杉及冷杉幼苗幼树密度和生长指标(高、基径)的短期影响,并采用核密度估计法分析其空间分布规律。结果表明: 云冷杉更新的密度通常随林隙增大而降低,仅对幼树影响显著,小林隙下云冷杉幼树密度分别为0.34和1.74株·m^-2,红松密度不受林隙大小的影响。林隙大小对冷杉幼苗幼树生长指标的影响最大,对红松影响最小,平均最大值多出现在大林隙。红松和云杉幼树的基径和树高最大值均分布在小、中、大林隙东北部,在特大林隙中转移至冠空隙西北部。小林隙有助于幼苗的建立和萌发,可通过择伐冷杉创造小林隙,随后扩大林隙面积(>50 m²)促进幼树生长,需要持续监测来确定林隙大小对森林更新的长期影响。     

林冠环境对亚高山针叶林下缺苞箭竹生物量分配和克隆形态的影响

通过对亚高山针叶林的林下、小林窗(130 m²)、中林窗(300 m²)和大林窗(500 m²) 4种林冠环境中缺苞箭竹(Fargesia denudata)分株种群特征进行调查, 研究其生物量分配格局和克隆形态可塑性。研究结果表明: (1)分株生物量、基径、高度以及分株各构件生物量随林冠郁闭度减小均表现为先增加后减小的趋势, 在小林窗中达到最大值; (2)大林窗中, 根生物量分配和数量显著高于其他林冠环境, 随林冠郁闭度增大, 分株不断增大叶生物量分配、比叶面积和分枝百分比等地上部分投资, 以适应低光环境; (3)比茎长和比地下茎长随林冠郁闭度增大表现为先减小后增加的趋势, 在小林窗值最低, 分枝强度在小林窗和中林窗中显著高于林下和大林窗环境。结果显示, 缺苞箭竹在不同林冠环境中具有生物量分配和克隆形态的可塑性, 以利于种群对光资源的有效利用。小林窗环境是缺苞箭竹较适宜的生境, 生物量积累最多, 长势最好。     

林窗对川西周公山柳杉人工林林下物种多样性的影响

采用典型样地法,以川西周公山柳杉人工林5种不同大小的林窗为研究对象,以林下非林窗为对照,研究了不同大小的林窗对柳杉人工林物种多样性的影响,同时分析了不同梯度林窗下林窗中心、林窗边缘、及林下群落的物种组成、物种多样性的变化情况。结果表明:(1)在所调查的18个样地231个样方中共记录到维管束植物141种,隶属于76科113属;随着林窗面积的增大,群落各层次的物种数呈现出先升高后降低的趋势,灌木层物种数在各林窗梯度上表现为林缘林下林窗中心,草本层物种数在各林窗梯度上表现为林缘林窗中心林下。(2)不同林窗优势种及其重要值不同,即在小林窗内,优势种为柳杉和野桐,其重要值之和高达0.292 3;在大林窗内,杉木及亮叶桦为群落优势物种,群落内出现大量其更新幼苗。(3)不同大小的林窗表现为灌木层物种丰富度指数(D)、Shannon-Wienner指数(H)、和Pielou均匀度指数(Jsw)值在400~450 m2面积的大林窗内达到一个均优水平,草本层物种的多样性在面积为100~150m2的小林窗内达到较高水平;不同梯度的林窗各层次群落D、H值整体表现为林缘林窗中心林下。研究认为:林窗的存在会改变群落物种组成,提高群落物种多样性水平,并且大林窗(400~450m2)更利于柳杉人工林林下树种更新及物种多样性的提高。     

Artificial canopy gaps accelerate restoration within an exotic Pinus radiata plantation

We created small‐scale artificial canopy gaps to accelerate the growth of mature indigenous forest canopy species for restoration of an 18‐year‐old exotic Pinus radiata plantation forest, in the Marlborough Sounds, New Zealand. Small and large circular gaps were formed by felling. Seedlings of two indigenous forest canopy species, Podocarpus totara (Podocarpaceae) and Beilschmiedia tawa (Lauraceae), were planted within artificial gaps and undisturbed plantation canopy. Seedling height growth, mortality, and occurrence of animal browse were monitored at approximately 6‐month intervals over 17 months. Both P. totara and B. tawa differed significantly in height growth and in animal browse occurrence among artificial gap treatments. Growth of the light‐demanding P. totara was better under large canopy gaps, whereas growth of the shade‐tolerant B. tawa increased under gaps of any size but was most consistent under small gaps. For P. totara, any significant restoration benefit of gap formation on height growth was lost when browsed seedlings were taken into account. Animal browse significantly limited B. tawa height growth in large but not in small gaps. Small‐scale canopy gap creation is an effective method of modifying light transmission to the plantation understorey and accelerating seedling growth rates. Canopy gap size can be used to optimize understorey illumination according to species‐specific light requirements. The increased occurrence of animal browse in gaps requires consideration. Artificial canopy gaps within planted monocultures create structural heterogeneity that would otherwise take an extended period of time to develop. These results further support the role of plantations as indigenous forest restoration sites.     

Effects of position, understorey vegetation and coarse woody debris on tree regeneration in two environmentally contrasting forests of north-western Patagonia: a manipulative approach

Aim To investigate the differential effects of position within gaps, coarse woody debris and understorey cover on tree seedling survival in canopy gaps in two old‐growth Nothofagus pumilio (Poepp. & Endl.) Krasser forests and the response of this species to gaps in two forests located at opposite extremes of a steep rainfall gradient. Location Nahuel Huapi National Park, at 41° S in north‐western Patagonia, Argentina. Methods In both study sites, seedlings were transplanted to experimental plots in gaps in three different positions, with two types of substrate (coarse woody debris or forest floor), and with and without removal of understorey vegetation. Survival of seedlings was monitored during two growing seasons. Soil moisture and direct solar radiation were measured once in mid‐summer. Seedling aerial biomass was estimated at the end of the experiment. Results Mid‐summer soil water potential was lowest in the centre of gaps, in plots where the understorey had been removed, and highest at the northern edges of gaps. Direct incoming radiation was highest in gap centres and southern edges, and lowest at northern edges. Seedling mortality was highest in gap centres, in both sites. Coarse woody debris had a positive effect on seedling survival during summer in the mesic forest and during winter in the xeric forest. The removal of understorey cover had negative effects in gap centres during summer. Seedling final aerial biomass was positively affected by understorey removal and by soil substrate in both sites. In the dry forest gaps, seedling growth was highest in northern edges, whereas it was highest in gap centres in the mesic forest. Overall growth was positively related to survival in the xeric forest, and negatively related in the mesic forest. Main conclusions Survival and growth were facilitated by the shade of gap‐surrounding trees only in the xeric forest. Understorey vegetation of both forests facilitated seedling survival in exposed microsites but competed with seedling growth. Nurse logs were an important substrate for seedling establishment in both forests; however, causes of this pattern differed between forests. Water availability positively controls seedling survival and growth in the xeric forest while in the mesic forest, survival and growth are differentially controlled by water and light availability, respectively. These two contrasting old‐growth forests, separated by a relatively short distance along a steep rainfall gradient, had different yet unexpected microenvironmental controls on N. pumilio seedling survival and growth. These results underscore the importance of defining microscale limiting factors of tree recruitment in the context of large‐scale spatial variation in resources.     

东灵山地区辽东栎幼苗的建立和空间分布

研究树木实生苗的建立对森林生态系统的保育和恢复具有重要意义,在北京东灵山地区调查了辽东栎(Quercus liaotungensis)实生和萌生幼苗在几种典型的植被类型中的空间分布以及辽东栎一年生实生幼苗在1个林窗梯度上的建立。选择几种典型的植被类型,研究辽东栎实生幼苗和萌生幼苗在森林中的空间分布。结果显示,在不同植被类型中辽东栎实生和萌生幼苗的密度存在空间差异,实生苗密度随幼苗年龄增大逐渐降低,种群的更新主要依靠萌生苗完成。在1个落叶阔叶林中选择1个林窗,在不同梯度(即林窗中间、林窗和树冠连接处,以及树冠下)上,播种辽东栎坚果,第二年秋季调查实生幼苗的2个生长指标和出苗率。结果显示,林窗对一年生实生幼苗的生长高度有影响。幼苗高度在林窗中比在树冠下大;林窗对辽东栎幼苗最长叶片长度和幼苗的出苗率没有影响。说明辽东栎实生幼苗的良好生长需要阳光比较充足的生境条件。本项研究结果建议,对辽东栎种群的更新,应该适当择伐一些较大个体,使森林形成一些林窗或林中空地,以利于辽东栎实生幼苗的建立,这样才能使辽东栎种群通过实生苗进行更好的自然更新。     

DEMOGRAPHIC ASPECTS OF COEXISTENCE IN ENGELMANN SPRUCE AND SUBALPINE FIR

The mechanisms for the maintenance of coexistence of Engelmann spruce and subalpine fir in subalpine forests of the Colorado Front Range were examined by comparing age, size, and spatial distributions of spruce and fir in two adjacent, previously logged sites of differing moisture availability. Adult tree ages were calculated from stem cores, while seedling ages were calculated from a multiple regression equation based on diameter, height, and number of branch whorls. Tree size was measured by height and diameter; spatial distributions were described by Morisita's index of dispersion. Cumulative age and size distributions were significantly different in the two species, with greater longevity and a larger overall size in spruce than fir. Both species showed a significant linear relationship between size and age, while fir showed a faster height growth rate than spruce. The linear relationship beween age and size was much closer in seedlings than in adults. Seedling spatial distribution was highly clumped in both species, but mature trees showed little or no clumping. Because both species are mainly wind dispersed, the greater clumping in spruce than in fir seedlings suggests that spruce have more specific establishment requirements than fir. Colonization patterns indicated that spruce seedlings were primarily found in forest gaps or associated with fir canopy trees, while fir seedlings were more commonly found in the forest, associated with either spruce or fir canopy trees. Tree density, growth rates, and mortality rates were higher in the wet site, with spruce showing the largest between site differences. These data suggest a new hypothesis for coexistence stating that Engelmann spruce and subalpine fir are maintained as codominants because the greater longevity and size of spruce is balanced by the faster height growth and more flexible seedling establishment requirements of fir.     

岷江冷杉林林窗小气候及其对不同龄级岷江冷杉幼苗生长的影响

对川西亚高山原始岷江冷杉(Abies faxoniana)林林窗内和林冠下小气候及岷江冷杉幼苗生长和生物量进行了两个生长季的连续观测。结果表明:6月林窗内与林冠下太阳辐射的日积累量没有显著性差异,而7~8月的日积累量则有显著性差异。整个生长季节,林窗内太阳辐射的平均积累量为8.10×MJ·m^-2,而林冠下太阳辐射的平均积累量为5.02×MJ·m^-2,两个位点太阳辐射积累量的显著差异主要来自7~8月日积累量的不同;林窗内5和15 cm层土壤的日平均温度比林冠下相应深度分别高2.1和2.7℃,差异显著。林窗内和林冠下3~8年岷江冷杉幼苗高增长率分别为1.2±0.3 cm·a^-1和1.1±0.3 cm·a^-1,差异不显著;9~20年岷江冷杉幼苗高增长率分别为6.2±2.4 cm·a^-1和3.0±0.9 cm·a^-1,差异显著。林窗内岷江冷杉幼苗根、主茎和总生物量与林冠下幼苗根、主茎和总生物量没有显著差异。不同年龄的岷江冷杉幼苗叶和侧枝生物量积累对林窗微环境的响应不同。     

山地落叶阔叶林优势树种米心水青冈幼苗的定居

在郁闭的林冠下、模拟林窗和林间空旷地3种光照环境中,研究了施肥和未施肥的米心水青冈幼苗生存和生长发育过程．结果表明,幼苗在林下的发育受到光照不足的严重抑制,生长在林下的幼苗比生长在林窗和空旷地幼苗死亡快,死亡率高,幼苗在郁闭林下能够生存的时间不超过12周．施肥处理的幼苗比对照的幼苗死亡快,死亡率高．幼苗死亡很可能与土壤中的病原生物有关．虽然由林下光照强度增加到模拟林窗的光照强度后,幼苗发育有明显改善,但幼苗在模拟林窗环境和空旷地的生长没有显著差异．结果说明,米心水青冈新出土的幼苗在郁闭林冠下易受伤害,更新立苗阶段需要有比较好的光照条件,肥沃土壤可能也无助于改善郁闭林冠下幼苗的定居．     

Tree seed germination and seedling establishment in treefall gaps and understorey in a subtropical forest of northeast India

Abstract Seed germination, and survival and growth of seedlings of four dominant tree species, Quercus dealbata, Quercus griffithii, Quercus glauca and Schima khasiana were studied in the treefall gaps and forest understorey of an undisturbed mature-phase humid subtropical broadleaved forest in northeast India. Three important microenvironmental factors namely photosynthetically active radiation (PAR), soil moisture and litter depth, were also measured in the forest understorey and gaps and correlated with seedling mortality. Seed germination of S. khasiana was significantly higher in the treefall gaps than in the understorey; among the tree species studied, it had the highest germination. Quercus seedlings were abundant in the understorey and small gaps, while S. khasiana seedlings were more numerous in the large gaps. The survivorship curves for the seedling populations revealed that the three Quercus species survived better in the understorey, while S. khasiana did so in the gaps. PAR and soil moisture were positively correlated with tree seedling mortality, which occurred mainly during the winter months. The Quercus seedlings grew better in the forest understorey and small gaps and S. khasiana seedlings in the large gaps. The differential performance of the tree seedlings to the conditions prevailing in the understorey and gaps of two sizes indicates that different species were adapted to different light environments depending upon their optimum requirements. This could be an effective mechanism for promoting species coexistence in the forest community.     

林隙与林下环境对锐齿槲栎和米心水青冈种群更新的影响

锐齿槲栎 (Quercusalienavar.acuteserrata ) 和米心水青冈 (Fagusengleriana) 是广泛分布在我国暖温带和亚热带山地中海拔地段的主要建群种, 对水土保持和维持生态系统稳定具重要意义。该文通过在神农架地区的典型样地调查, 分析了这 2个种群在林隙和林下不同光环境中的更新状况和径级结构 ;利用便携式Licor₆ 4 0 0光合仪测量了 2种苗木的光饱和及光诱导曲线, 探讨了光对这 2个树种更新的影响机理。实验结果表明, 在所调查的林地中, 锐齿槲栎和米心水青冈种群都有充足的幼苗库, 锐齿槲栎幼苗以实生为主, 米心水青冈幼苗以萌生为主 ;从幼苗到幼树的转化过程中, 锐齿槲栎的成活率几乎为零, 而米心水青冈的成活率约为 6 %~ 8% ;在两者的混交林中, 米心水青冈种群呈稳定的金字塔形年龄结构, 而锐齿槲栎则是某一生长阶段的种群占据主导地位。光饱和曲线显示, 对 2年生同生群苗木, 在林窗下, 锐齿槲栎的最大净光合速率 (4.6 1μmol·m-2 ·s-1) 比米心水青冈的 (4.16 μmol·m-2 ·s-1) 要高 ;而在林冠下, 米心水青冈的最大净光合速率 (Pn=3.89μmol·m-2 ·s-1) 比锐齿槲栎的最大净光合速率 (Pn=3.6 8μmol·m-2 ·s-1) 要高。无论在林窗下或林冠下, 米心水青冈幼苗的光饱和点比锐齿槲栎的要低。两者混交林下的光诱导曲线显示, 与米心水青冈幼苗相比, 锐齿槲栎幼苗对光的反应速度较慢, 最大净光合速率也较低, 在林冠下表现出较弱的竞争力。资源分配试验证明, 锐齿槲栎在幼苗阶段以地上部分生长为主, 而米心水青冈倾向于把更多的干物质贮藏在根部, 显示出不同的光利用策略。     

Canopy gaps and establishment patterns of spruce (Picea abies (L.) Karst.) in two old-growth coniferous forests in central Sweden

The spatial pattern of seedlings, saplings and canopy trees was studied in two spruce (Picea abies (L.) Karst.) forests in central Sweden. Canopy and forest structure were determined in five 0.25 ha plots. Life stage classes were distinguished on the basis of age and size distributions. Ripley's K-function (1977) was used to analyze the spatial patterns within each class. A random distribution of seedlings gave way to a more aggregated pattern on a small scale during the establishment phase. Saplings and sub-canopy trees were strongly aggregated and canopy trees were again randomly distributed within the plots. The proportion of individuals growing in gaps was used as an index of association between the spatial pattern in saplings and sub-canopy trees and the occurrence of small (50–350 m²) canopy gaps. Under the null hypothesis of independence the expected value of this statistic would equal the canopy gap ratio for the stand. Monte Carlo simulation of this statistic, using fixed sapling positions and randomly repositioned canopy gaps, confirmed the importance of canopy gaps for the final success of establishment of spruce. The association of understorey trees with gaps suggest that small gaps are typically closed by recruitment of new saplings from a sapling bank rather than by the release of larger suppressed trees.     

Tree species distribution in canopy gaps and mature forest in an area of cloud forest of the Ibitipoca Range,south-eastern Brazil

The tree community of both canopy gaps and mature forest was surveyed in a 5 ha plot of cloud forest in the Ibitipoca Range, south-eastern Brazil, aiming at: (a) comparing the tree community structure of canopy gaps with that of three strata of the mature forest, and (b) relating the tree community structure of canopy gaps with environmental and biotic variables. All saplings of canopy trees with 1–5 m of height established in 31 canopy gaps found within the plot were identified and measured. Mature forest trees with dbh 3 cm were sampled in four 40×40 quadrats laid on the four soil sites recognised in the local soil catena. All surveyed trees were identified, measured and distributed into three forest strata: understorey (<5 m of height), sub-canopy (5.1–15 m) and canopy (15.1–30 m). The following variables were obtained for each gap: mode of formation, age, soil site, slope grade, size, canopy openness and abundance of bamboos and lianas. A detrended correspondence analysis indicated that the tree community structure of gaps in all soil sites was more similar to that of the mature forest understorey, suggesting that the bank of immatures plays an important role in rebuilding the forest canopy and that gap phases may be important for understorey shade-tolerant species. There was evidence of gap-dependence for establishment for only one canopy tree species. Both canonical correspondence analysis and correlation analysis demonstrated for a number of tree species that the distribution of their saplings in canopy gaps was significantly correlated with two variables: soil site and canopy openness. The future forest structure at each gap is probably highly influenced by both the present structure of the adjacent mature forest and the gap creation event.