Patterns of tree replacement: canopy effects on understory pattern in hemlock-northern hardwood forests

The effect of canopy trees on understory seedling and sapling distribution is examined in near-climax hemlock-northern hardwood forests in order to predict tree replacement patterns and assess compositional stability. Canopy trees and saplings were mapped in 65 0.1-ha plots in 16 tracts of old-growth forests dominated by Tsuga canadensis, Acer saccharum, Fagus grandifolia, Tilia americana, and Betula lutea in the northeastern United States. Seedlings were tallied in sub-plots. Canopy influence on individual saplings and sub-plots was calculated, using several indices for canopy species individually and in total. For each species sapling and seedling distributions were compared to those distributions expected if saplings were located independently of canopy influence. Non-random distributions indicated that sapling and seedling establishment or mortality were related to the species of nearby canopy trees. Hemlock canopy trees discriminate against beech and maple saplings while sugar maple canopy favors beech saplings relative to other species. Basswood canopy discourages growth of saplings of other species, but produces basal sprouts. Yellow birch saplings were rarely seen beneath intact canopy. Since trees in these forests are usually replaced by suppressed seedlings or saplings, canopy-understory interactions should influence replacement probabilities and, ultimately, stand composition. I suggest that hemlock and basswood tend to be self-replacing, maple and beech tend to replace each other, and birch survives as a fugitive by occupying occasional suitable gaps. This suggests that these species may co-exist within stands for long periods with little likelihood of successional elimination of any species. There is some suggestion of geographical variation in these patterns.     

Seedling and Sapling Dynamics of Treefall Pits in Puerto Rico1

Seedling and sapling dynamics in a Puerto Rican rain forest were compared between forest understory and soil pits created by the uprooting of 27 trees during Hurricane Hugo. Soil N and P, organic matter, and soil moisture were lower and bulk densities were higher in the disturbed mineral soils of the pits than in undisturbed forest soils ten months after the hurricane. No differences in N and P levels were found in pit or forest soils under two trees with N–fixing symbionts (Inga laurina and Ormosia krugii) compared to soils under a tree species without N–fixing sym–bionts (Casearia arborea), but other soil variables (Al, Fe, K) did vary by tree species. Forest plots had greater species richness of seedlings (<10 cm tall) and saplings (10–100 cm tall) than plots in the soil pits (and greater sapling densities), but seedling densities were similar between plot types. Species richness and seedling densities did not vary among plots associated with the three tree species, but some saplings were more abundant under trees of the same species. Pit size did not affect species richness or seedling and sapling densities. Recruitment of young Cecropia schreberiana trees (>5 m tall) 45 months after the hurricane was entirely from the soil pits, with no tree recruitment from forest plots. Larger soil pits had more tree recruitment than smaller pits. Defoliation of the forest by the hurricane created a large but temporary increase in light availability. Recruitment of C. schreberiana to the canopy occurred in gaps created by the treefall pits that had lower soil nutrients but provided a longer–term increase in light availability. Treefall pits also significantly altered the recruitment and mortality of many understory species in the Puerto Rican rain forest but did not alter species richness.     

Environmental Correlates of Tree and Seedling–Sapling Distributions in a Mexican Tropical Dry Forest

Bray and Curtis ordination was used to explore which environmental variables explained importance values and the presence–absence of tropical tree seedlings, saplings and adults in La Escondida-La Cabaña, Sierra de Manantlán, Jalisco, Mexico. The diameters of trees ≥2.5 cm DBH and the presence and height of seedlings and saplings were measured in nine 0.1 ha sites. Four matrices including presence–absence data and importance value indices for trees and seedlings and saplings were analyzed through Bray and Curtis ordination. The matrices were based on density, frequency, and dominance of adult trees as well as seedlings and saplings. The environmental matrix consisted of 18 variables, including elevation, slope, canopy gaps, disturbance, and soil variables. We recorded 63 tree species and 38 seedling and sapling species in the nine sites. The ordination explained 70.9% of the variation in importance value data for trees and 62.6% for seedlings and saplings. The variation explained in presence–absence data for trees was 67.1 and 77.4% for seedlings and saplings. The variance in the ordination axes of seedlings and sapling presence–absence data was poorly explained by the number of gaps in the tree, shrub, or herb layer, suggesting little light specialization by seedlings and saplings. Habitat specialization for soil nutrients appears to be important in explaining the presence–absence of seedlings and saplings. Seedling and sapling specialization along different soil microsites could promote species coexistence in this forest, while heterogeneity in light conditions may instead determine differences in growth and, thus, importance value of trees. We hypothesize that in tropical dry forest in Jalisco, Mexico, a habitat specialization for soil resources is likely more importance at early stages in tree life histories than in later life history.     

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．     

Altitudinal variations in seedling and sapling density and age structure of timberline tree species in the Sergyemla Mountains, southeast Tibet

The growth situation of seedlings of treeline species can be used as an indicator for the upper or lower shift of treeline under global climate change, since any advance of a treeline would require seedling survival above the current treeline elevation. We investigated seedling and sapling density of two dominant conifers Abies georgei var. smithii and Sabina saltuaria at lower, middle and upper altitude of two contrasting slopes in Sergyemla Mountain, southeast Tibet. Seedling and sapling age structure of A. georgei var. smithii was analyzed further. Seedling density of both species, as well as sapling density of S. saltuaria, showed a declining trend as elevation increased. Sapling density of A. georgei var. smithii was somewhat higher at middle altitude than other positions. The frequency of old-aged A. georgei var. smithii seedlings/saplings (age >10 year) was the lowest at the highest altitude and tended to increase with the declining elevation. The results indicate that the higher density of seedling and sapling in high-altitude might be attributed to better soil moisture availability and solar insolation than in the mid- and low-altitudes during the growth season. Yet confined by other environmental factors, A. georgei var. smithii seedlings in high-altitude suffer from lower survival rate and lower possibility to grow tall compared with those in mid- and low-altitudes.     

Altitudinal variations in seedling and sapling density and age structure of timberline tree species in the Sergyemla Mountains, southeast Tibet

The growth situation of seedlings of treeline species can be used as an indicator for the upper or lower shift of treeline under global climate change, since any advance of a treeline would require seedling survival above the current treeline elevation. We investigated seedling and sapling density of two dominant conifers Abies georgei var. smithii and Sabina saltuaria at lower, middle and upper altitude of two contrasting slopes in Sergyemla Mountain, southeast Tibet. Seedling and sapling age structure of A. georgei var. smithii was analyzed further. Seedling density of both species, as well as sapling density of S. saltuaria, showed a declining trend as elevation increased. Sapling density of A. georgei var. smithii was somewhat higher at middle altitude than other positions. The frequency of old-aged A. georgei var. smithii seedlings/saplings (age >10 year) was the lowest at the highest altitude and tended to increase with the declining elevation. The results indicate that the higher density of seedling and sapling in high-altitude might be attributed to better soil moisture availability and solar insolation than in the mid- and low-altitudes during the growth season. Yet confined by other environmental factors, A. georgei var. smithii seedlings in high-altitude suffer from lower survival rate and lower possibility to grow tall compared with those in mid- and low-altitudes.     

Marginal holly (Ilex aquifolium L.) populations in Mediterranean central Spain are constrained by a low-seedling recruitment

Seedling density and survival was studied in holly woodland located in the mountains of the Sistema Central, Madrid (Spain) in 1998. Ten permanent quadrats of 50×50 cm were randomly placed in each of the following habitats: under closed holly canopy, in canopy gaps, at the edge of the holly woodland, in the adjacent open grassland, and under fleshy- and dry-fruited shrubs located in this grassland. Emerged seedlings were periodically monitored and their survival was checked during the following 2 years. Additionally, a set of 20 seedlings were planted in each of the cited habitats and their survival was followed in parallel with the naturally emerged seedlings. The influence of light on holly survival was tested by experimental planting of holly seedlings under six different controlled light intensities.Seedling emergence was strongly habitat dependent, with the highest seedling densities in the closed holly woodland and in canopy gaps, and a complete absence in the grassland. Survival to the first year was nil. No naturally emerged seedling reached 1 year of age, independent of the habitat studied. Only one of the planted seedlings in the field survived to the first year. Despite this, mean seedling lifespan was different between habitats, with canopy gaps, woodland edge, and under fleshy-fruited shrubs being the habitats with highest average lifespan. Seedling response to the distinct light intensities is largely similar to that seen in the different field habitats, but other factors in addition to light, mainly summer drought and disturbances are proposed to explain the 100% mortality observed.Seedling incorporation is the recruitment dynamic bottleneck of this holly population, being more severe here than in other places studied further north. This may explain the increasing rarity of the species in the southern half of the Iberian Peninsula, where Ilex is close to its southern limit of its distribution.     

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.     

Seedling recruitment ofChamaecyparis obtusa andSciadopitys verticillata in different microenvironments in an old-growthSciadopitys verticillata forest

Seedling recruitment ofChamecyparis obtusa andSciadopitys verticillata in different microenvironments in an old-growthS. verticillata forest was studied for five years. Exposed mineral soil beneath a gap, which was newly created by a typhoon, supported the best seedling emergence forC. obtusa. S. verticillata seemed to require no specific microenvironment for its seedling emergence. Mortality of seedlings was higher for both species beneath a closed canopy, where all new seedlings ofC. obtusa died within a year after their emergence. Beneath the gap, mortality of seedlings was generally lower for both species, regardless of the presence of litter on the ground. The occurrence of saplings was more frequent forC. obtusa beneath the gaps and forS. verticillata beneath a closed canopy. Difference of seedling recruitment behaviour could maintain the coexistence ofC. obtusa andS. verticillata in the canopy layer of this forest in a non-equilibrium state caused by occasional disturbance.     

Facilitation and interference underlying the association between the woody invaders Pyracantha angustifolia and Ligustrum lucidum

Questions: 1. Is there any post‐dispersal positive effect of the exotic shrub Pyracantha angustifolia on the success of Ligustrum lucidum seedlings, as compared to the effect of the native Condalia montana or the open herbaceous patches between shrubs? 2. Is the possible facilitation by Pyracantha and/or Condalia related to differential emergence, growth, or survival of Ligustrum seedlings under their canopies? Location: Córdoba, central Argentina. Methods: We designed three treatments, in which ten mature individuals of Pyracantha, ten of the dominant native shrub Condalia montana, and ten patches without shrub cover were involved. In each treatment we planted seeds and saplings of Ligustrum collected from nearby natural populations. Seedlings emerging from the planted seeds were harvested after one year to measure growth. Survival of the transplanted saplings was recorded every two month during a year. Half of the planted seeds and transplanted saplings were cage‐protected from rodents. Results: Ligustrum seedling emergence did not differ among treatments while growth was significantly higher in the absence of shrub cover. Sapling survival was significantly higher under the canopy of Pyracantha, intermediate under Condalia, and lowest in the absence of shrub cover. Caging did not affect growth but enhanced seedling emergence and sapling survival. Conclusion: The differential sapling survival in the shrub canopy treatments is consistent with natural sapling distribution. Pyracantha and, less so, Condalia, has a nurse‐plant effect on Ligustrum. This results from contrasting effects of the shrubs on different stages of the life cycle of Ligustrum: no effect on seedling emergence, negative on seedling growth, and positive on sapling survival. This suggests that efforts to control the expansion of Ligustrum over the landscape should tackle Pyracantha as well.     

A test of the escape and colonization hypotheses for zoochorous tree species in a Western Amazonian forest

In order to assess the importance of seed dispersal (escape and colonization hypotheses), I used transplant experiments for seeds and seedlings of 5–11 plant species with fleshy fruits in a lowland tropical forest (Tinigua National Park, Colombia). I controlled seed density, distance to parental tree, and habitat type. I monitored seed removal, seedling survival, and seedling growth during the first year of development for an average of 554 seeds and 169 seedlings for each species. I supplemented the experimental results with measurements of natural recruitment. I found little support for the escape hypothesis during the seed and seedling stages. For six species that showed differences in seed removal associated with distance, five showed highest removal away from, than close to parent trees, suggesting predator satiation. Seedling survival during the first year was not consistently associated with low densities and long distances from parent trees. For the majority of species, seedlings did not survive flooding in low basins, and there was growth advantage for most plant species in canopy gaps. These differences imply advantages for seed dispersal to adequate habitats, as predicted by the colonization hypothesis. In contrast to experiments, strong negative distance-dependent effects were evident when analyzing natural recruitment patterns. The ratio between saplings and seedlings was higher away from parent trees for the species with enough recruitment to be analyzed and this suggests that a negative distance-dependent effect may also occur after seedling establishment. This pattern is suspected for several other species, but an analysis with some of the other most common trees showed a variety of negative, neutral, and positive distance dependent effects. This study emphasizes the importance of long-term studies to asses the role of seed dispersal.     

Demographic implications of life-history stage characteristics in two African acacias at a Makeni savanna plot in Zambia

Aims In spite of the importance of African acacias in vegetation succession and provision of goods and services, little is known about life-history variations within and among species. Much of the work done on African acacias has focused on seed predation and germination and seedling establishment, especially of Acacia tortilis, Acacia nilotica and Acacia karroo. The primary aim of the present work is to investigate differences in the demography of Acacia polyacantha and Acacia sieberiana and the relationship between life-history characteristics and population size. A secondary objective is to assess how fire, an important ecological factor in savanna vegetation, might modify the growth and demographic dynamics of the two acacias.Methods The study was conducted at the Makeni savanna plot in central Zambia, southern Africa. Seedling emergence from both non-scarified and scarified seeds sown at different times in the wet season and the fate and growth of seedlings and saplings were monitored over a period of 4 years. Annual growth of permanently marked sample trees in annually burnt and fire-protected blocks was recorded over a 6-year period (2002–08) in order to assess inter-specific differences and how fire modifies tree growth patterns. Censuses of natural saplings and trees were conducted periodically in sample blocks to determine recruitment into these life-history stages.Important findings Seedling emergence and sapling survival rates were much higher in A. sieberiana than in A. polyacantha. However, both seedling and sapling growth rates were higher in A. polyacantha than in A. sieberiana but tree growth rates were similar in the two species. Under fire protection tree growth was significantly influenced by tree size and year while under annual burning only tree size significantly affected tree growth. The dominance of A. sieberiana over A. polyacantha at the study site was attributed to higher seedling emergence rate, higher sapling survival rate and a large sapling bank that forms a reliable source of tree recruitment. The life-history stage characteristics of A. polyacantha suggest that this is an early successional species.     

福建漳江口红树林区秋茄幼苗生长动态

通过福建漳江口红树林自然保护区内8个样方24个小样方人工种植600个秋茄胚轴,在3a时间内对秋茄胚轴建立、幼苗生长以及环境因子进行定期观测。研究结果表明：林缘空地的秋茄生长状况良好,白骨壤林内最不利于秋茄幼苗的生长。潮位、盐度、底质土壤理化因子不是造成该样地各样方间秋茄幼苗生长差异的主要原因。动物取食、光照状况以及种间竞争是限制秋茄生长的主要环境因子。秋茄胚轴在长根前易于随潮水漂走,底质土壤中自骨壤致密的根系抑制了秋茄胚轴的定植,导致白骨壤林内秋茄幼苗漂走的数量最多。昆虫和螃蟹等动物的取食是导致林内已经固着生长的秋茄幼苗大量死亡的最主要原因,而林外被取食的幼苗个体极少。此后秋茄幼苗能否继续成长,主要取决于幼苗所接受到的光照条件。3a后,在荫蔽的树冠下,秋茄幼苗无法存活;而在林外,秋茄幼苗已经长成幼树。在林外滩涂上迅速生长的互花米草,也将影响秋茄幼苗的更新和生长。     

Invasion by Ligustrum lucidum (Oleaceae) in NW Argentina: early stage characteristics in different habitat types

Currently biological invasions are considered one of the world's most serious conservation problems. Ligustrum lucidum is the most abundant exotic tree in secondary forest patches of montane forests of NW Argentina. We studied the determinants of success of the early stages of its life cycle in distinct habitat types, with the hope of identifying vulnerabilities that could be exploited to control the invasion. Seed arrival, germination, seedling recruitment and survival, and sapling growth were studied in edges, gaps and forest interior. Seed arrival was also assessed under perches and in open fields. Germination was studied in forest and grassland patches. L. lucidum seedling survival and sapling growth were compared with the most abundant native species survival and growth. Seed arrival was strongly seasonal with a peak in mid-August. Seed rain did not differ significantly among habitat types, however there was a tendency for edges to receive more seeds when only dispersed seeds were considered. Perches strongly enhanced seed arrival; more than 40 times the number of seeds were dispersed beneath citrus plants (i.e. perches) than found in paired open areas. In the forest, seeds in gaps and edges had higher germination rates, but there was no difference in seedling survival. Fruits under closed canopy exhibited the lowest germination. Germination and survival were low in open areas. Neither seedling recruitment nor sapling growth differed between gaps and forest interior. L. lucidum saplings grew significantly more than saplings of the most common native species, and also showed higher seedling survival. L. lucidum is a prolific fruit producer, is capable of germinating and surviving in a broad range of forest environments, it is relatively shade tolerant and has higher survival and faster growth rate in comparison to the most common native species. All these characteristics highlight its potency as a successful invader, and point to few vulnerabilities that could be targets of control measures.     

植物种群更新限制——从种子生产到幼树建成

更新限制是指种子由于各种原因,不能够萌发并生长成幼树。它作为解释生物多样性的理论,一直受到国内外群落生态学家关注。从种源限制、传播限制和建成限制3个角度,对更新限制机制研究进展进行了综述。从种源限制而言,时空因素是影响植物种群更新限制的重要因素,因为植物结实量存在明显时空变化,造成植物更新个体出现明显的时空规律。从传播限制而言,传播数量、距离和食果动物行为均限制植物种群更新。数量上,缺乏有限传播者势必减少传播数量,但如果种子拥有较高质量,则能逃脱数量限制;距离上,植物更新个体显示出明显的Janzen-Connell格局,但传播距离趋向稳定,形成植物种群的进化稳定对策;食果动物行为上,不同传播者对更新贡献存在差异,捕食者直接降低更新,融入两类动物行为的模型更能反映食果动物对更新的限制。从建成限制而言,环境因子制约植物生长。小尺度下,微生境的好坏对于植物幼苗建成至关重要;大尺度下,植物提供较好的广告效应则能摆脱生境限制。将传播者行为、捕食者行为与幼苗的空间分布格局、种子传播机理模型等结合,建立植物更新限制机理模型应是更新限制未来的研究热点。选择稀有种和古老种为主题的长期更新限制研究,为种群恢复提供指导,也是未来重要研究方向。     

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。     

Initial regenerative processes ofDistylium racemosum andPersea thunbergii in an evergreen broad-leaved forest

We studied the initial regenerative processes ofDistylium racemosum andPersea thunbergii, major canopy species, in an old-growth evergreen broad-leaved forest, Kagoshima, southwestern Japan. Although the two species coexisted, the patterns of seed production, seedling emergence, and sapling survival were different between them. During the observation (1989–1991),P. thunbergii produced large numbers of seeds in alternate years, whileD. racemosum had no mast year. The density of established seedlings was high and increased with mass seed production forP. thunbergii, but was fairly low forD. racemosum. The mortality of seedlings and saplings ofP. thunbergii was higher than that ofD. racemosum. In a closed stand, the sapling bank was maintained by dense seedling supply forP. thunbergii, and by extremely low mortality of individuals germinated before the beginning of observation forD. racemosum. Since saplings ofD. racemosum showed continuous height growth beneath the closed canopy, the possibility of recruitment into the upper layer seems to be high. The strategy ofP. thunbergii might be to wait for suitable conditions favoring sapling growth on various site by means of frequent and large seed productivity and wide dispersal of seeds. Thus both species might be able to coexist through environmental heterogeneity.     

The abundance and growth of Avicennia marina and Rhizophora stylosa in the low shore zone of Darwin Harbour,Northern Territory

Abstract The growth and distribution of the two mangrove seedling species, Avicennia marina (Forsk.) Vierh. and Rhizophora stylosa Griff., were studied in the low-shore zone at three locations along two estuaries of Darwin Harbour to determine how growth varied along the estuary and within the habitats of the low shore zone. The low shore environment was heterogenous for factors such as salinity and light, with substantial variation within and between locations. Seedlings of Avicennia were in higher densities in light gaps than in the adjacent forest, and density in both habitats decreased towards the mouth of the estuary. Growth of Avicennia seedlings, as measured through in situ growth trials and plant harvests, increased towards the mouth of the estuary. Results for Rhizophora seedlings were more complex although some similar trends were evident. This may have been due to the greater early survival of Rhizophora seedlings compared to Avicennia seedlings. The importance of propagule origin was investigated, for Rhizophora, by transplanting propagules between locations along one estuary. Increased seedling survival was observed at the propagule source site; however, this effect was subtle and probably not the most important factor affecting seedling survival at a site.     

Small-scale patterning in the recruitment of forest species during succession in tropical dry forest,Mozambique

Within a few years of field abandonment on the coastal dunes, a considerable number of species of trees and woody lianes of the climax evergreen forest enter the succession. At a relatively large-scale of analysis (a 100 m² plot), there is no evidence of soil development during the early succession, with no accumulation of soil organic matter nor nutrients being detected. At this scale there is, therefore, little support for the facilitation model of succession. However, when smaller scales of analysis are employed (e.g. a plot equivalent to the canopy area of a sapling) a very different picture emerges; most recruitment of forest woody species occurs beneath saplings found in early succession. Nutrient concentrations were higher in soils under sapling canopies than in soils beyond sapling canopies. With increasing sapling size, an increasing number of seedlings of forest woody species were found and there is progressive soil amelioration. Thus, during the early succession, there is considerable small-scale patterning in soil development and recruitment. This pattern is largely established through bird dispersal, with the intensity of recruitment being dependent on the distance to seed sources.     

Patterns of seedling recruitment in West-Mediterranean Quercus ilex forest influenced by canopy development

Abstract. Quercus ilex (holm oak) coppice forests belong to the most representative communities in the western part of the Iberian Peninsula. Due to the high sprouting potential of holm oak after coppicing, the dynamics of recruitment of new individuals in existing populations has been largely overlooked. In these forests, the density of seedlings increases in old stands, as a result of the remarkable increase in the recruitment of 1-yr and 2-yr old seedlings, probably due to better environmental conditions for germination. Older seedlings show a tendency towards increasing survival under some degree of canopy closure, since their density appears to be higher just before complete canopy closure takes place. Nevertheless, the reduced growth of seedlings in older stands due to low light levels causes age distributions of holm oak seedlings to change drastically, with saplings older than 15 yr becoming very unusual in these stands. Therefore, a bottleneck of recruitment appears at this sapling stage and prevents vigorous regeneration just before thinning. Present conditions of genet density and management tend to prevent population renewal. Dynamics of the seedling bank reinforce the role of gap formation through small or large scale perturbations in population turnover.