Response of Quercus robur L. seedlings to north-south asymmetry of light within gaps in floodplain forests of Slovenia

To examine Quercus robur establishment and growth in low to intermediate light levels, we analysed regeneration in different microsites created by light asymmetry within gaps in two main floodplain forest regions of Slovenia: Dolinsko and Krakovo. Four years after a mast year in 1995, we installed systematic grids of 1×1 m plots in 11 gaps (0.03–0.40 ha) on wet and dry site variants. In 256 plots, seedling species, cover, density, height, height increment, browsing damage, tree architecture, understory vegetation species and cover, and direct and diffuse light were measured. The average seedling density in all gaps was 15/m², but the highest densities were found in gap positions with low diffuse light levels (10–20%). Competing understory vegetation was more abundant in positions with high diffuse light. In gaps on dry site variants a combination of low diffuse and high direct light was favourable for regeneration. The results indicated that Q. robur can successfully establish in gaps. Here, both light components were sources of within gap resource heterogeneity, therefore knowledge of light asymmetry can improve regeneration success.     

雪灾干扰下林窗对木荷幼苗更新的影响

林窗干扰是维持森林生态系统的重要驱动力之一,对种子萌发、幼苗等自然更新过程、森林物种组成和动态、森林生物多样性的维持具有重要作用。本研究以2008年雪灾干扰后的浙江江郎山木荷林为研究对象,对木荷林窗大小结构、幼苗更新、生长等进行调查研究,结果表明:扩展林窗以50 100 m2的林窗个数最多(占总数的45.45%),各等级林窗中以50 100 m2的林窗占总面积比例最大(占总面积的30.31%)。林窗中木荷幼苗的平均高度和地径较对照林分分别高1.44 cm和0.61 mm,幼树在林窗中的平均高度和地径则比对照林分中分别高45.37 cm和5.00 mm且差异显著;林窗大小对木荷幼苗、幼树的高度和地径生长影响显著,中林窗中幼苗的高度和地径均高于小林窗和大林窗中的幼苗且差异均显著(F=4.893,P=0.007;F=5.203,P=0.004;n=357);幼树的地径在不同大小林窗中差异显著(F=3.569,P=0.037;n=43)。林窗幼苗的更新密度随着林窗面积的增大而增大,在林窗面积达到76 m2时,更新密度达到最大值,而后随着林窗面积的增大下降;中林窗和小林窗中更新苗木以低矮植株(1级、2级)为主,面积100 m2大林窗中,木荷幼苗生长较快。与他人研究的森林天然林窗相比,雪灾干扰后改变了林窗的大小分布结构和面积,50 100 m2的林窗比例较大,一定程度上更利于幼苗更新,具有相对较大的林窗幼苗更新密度;不论林窗大小,林窗内的更新幼苗都比林内多,郁闭度较大的林内或大面积的空地上都不利于更新幼苗的生长。因此,从受灾木荷林窗大小结构、幼苗更新、生长等来看,中林窗是幼苗适宜更新的面积,为木荷灾后恢复与重建提供了科学依据。     

Slit-shaped gaps are a successful silvicultural technique to promote Picea abies regeneration in mountain forests of the Swiss Alps

Many mountain forests in the Swiss Alps are dense and overmature. The resulting lack of tree regeneration threatens their future ability to provide products and services for humans, e.g., protecting settlements and infrastructure against avalanches. To promote natural regeneration, slit-shaped gaps have been cut since the 1980s in many of Switzerland's Alpine forests dominated by Norway spruce (Picea abies). However, little is known about the success of this silvicultural technique. We sampled 38 gaps in 2001 and 2006 and analysed the density and vitality of P. abies seedlings in these gaps, and monitored the growth and survival of selected seedlings between the two inventories. The gaps analysed were located in upper montane and subalpine P. abies dominated forests in the Vorderrhein valley in the Grisons. The density and vitality of large (10–129 cm tall) and small (<10 cm tall) P. abies seedlings were assessed in three parallel transects running perpendicular to the longitudinal axis of each gap. The mean density of large seedlings increased significantly between 2001 and 2006 from 0.1 to 0.3 seedlings/m², whereas that of small seedlings stayed constant at 0.7 seedlings/m², even though it shifted locally between inventories. Significantly higher regeneration densities were found for gaps with NE–SW orientation (afternoon sun) and for those located at lower altitudes. While in gaps on North-facing slopes large P. abies seedlings were more frequent on the middle transect, in gaps on South-facing slopes they were denser near the lower gap edge which is usually less exposed to direct radiation. As expected regeneration density was significantly higher within the gaps than just outside underneath the adjacent stands for all P. abies seedlings. Damage caused by browsing did not turn out to be problematic in this study site. From the selected 280 small seedlings monitored in 2001, 53% died until 2006, 33% became large seedlings and 14% remained small (<10 cm). Our results suggest that creating slit-shaped gaps is a successful procedure to promote P. abies regeneration in the upper montane and subalpine belt of the northern intermediate Alps and helps to preserve protection forests and their goods and services for the coming generations.     

Species diversity and regeneration of old-growth seasonally dry Shorea robusta forests following gap formation

Diversity and regeneration of woody species were investigated in two ecological niches viz. gap and intact vegetation in old-growth seasonally dry Shorea robusta (Gaertn. f.) forests in Nepal. We also related varieties of diversity measures and regeneration attributes to gap characteristics. Stem density of tree and shrub components is higher in the gap than in the intact vegetation. Seedling densities of S. robusta and Terminalia alata (B. Heyne ex Roth.) are higher in the gap than in the intact vegetation, while contrary result is observed for T. bellirica (Gaertn. ex Roxb.) and Syzigium cumini (L. Skeels) in term of seedling density. The complement of Simpson index, Evenness index, and species-individual ratio in the seedling layer are lower in the gap than the intact vegetation. Gap size can explain species richness and species establishment rate. Gaps created by multiple tree falls in different years have higher seedling density of S. robusta than gaps created by single and/or multiple tree falls in the same year. In conclusion, gaps maintain species diversity by increasing seedling density, and favor regeneration of Sal forests. In addition to gap size, other gap attributes also affect species diversity and regeneration. Foundation Project: The study was supported by Swedish International Development Cooperation Agency (SIDA).     

Regeneration niches of Norway spruce (<Emphasis Type="Italic">Picea abies</Emphasis> [L.] Karst.) saplings in small canopy gaps in mixed mountain forests of the Bavarian Limestone Alps

Natural regeneration measurements are the main silvicultural objective in overaged protective forests of the Bavarian Limestone Alps. While manifold problems with these stands, especially the impact of browsing, are widely recognised, the regeneration niches of Norway spruce (Picea abies [L.] Karst.) are insufficiently known. The purpose of this study was to determine favourable combinations of site factors for the development of spruce in small, unfenced canopy gaps, located on Aposerido-Fagetum caricetosum albae forest sites. We recorded the occurrence of spruce saplings (as dependent variable) and of six site factors (as independent variables) on 480 0.5 m²-subplots. In addition, we estimated the coverage of six acid adapted plant species to determine correlations with the humus depth. A binary logistic regression analysis was used to predict the probability of the occurrence of a spruce sapling in dependency of the different site factors. Supported by other studies, we assumed that the supply of solar radiation was adequate for the sufficient regeneration of spruce within the canopy gaps. Other site factors significantly determined the regeneration niches of spruce saplings. More spruce saplings were found near hindrances and on rough surfaces than would be expected from a random occurrence of saplings. These microsite types may have characteristics, especially protection against snow gliding that promotes spruce establishment. A calculated “hindrance index”, which accounted for the number, and the distance of surrounding hindrances might be a good specific value to describe the influence of hindrances on steep slopes. The sapling establishment decreased on thin humus layers. Our assumption for the sites was that thick organic layers might represent a good seedbed for spruce. Decayed dead wood was scarce, but was exceedingly favoured by spruce saplings. Results obtained suggest that the natural regeneration establishment of spruce on steep slopes can be successfully influenced by site factors which inhibit the influence of snow gliding. According to a “positive microsite” concept, we recommend for artificial regeneration measurements with spruce, microsites close to hindrances (e.g. stumps, downed trees) and Vaccinium myrtillus as a predictor for thick, acid humus layers.     

Natural regeneration after gap cutting in Scots pine stands in northern Finland

The objective of the study was to examine the success of regeneration in gaps of variable size in pine-dominated stands in terms of seedling density. It was based on an experiment in central Lapland containing circular clear-cut gaps of 20, 40 and 80 m in diameter on typical sub-xeric and xeric pine sites with site preparation (patch scarification). The observation period covered the first five years after cutting. The average number of seedlings that had emerged after gap cutting was about 22 000?ha^?1 for pine and 7 000?ha^?1 for birch. The proportion of regeneration sample plots without any pine seedlings was less than 10%. The seedling density diminished constantly with greater distance from the edge stand but indicating sufficient density up to the largest gap size in the study (diameter 80 m, area ca. 0.5?ha). Site preparation promoted regeneration remarkably, and we conclude that successful regeneration would be achieved in most cases with soil scarification exposing just 10–20% of the soil surface.     

Tree recruitment in gaps of various size,clearcuts and undisturbed mixed forest of interior British Columbia,Canada

Tree seedling recruitment was monitored after various types of logging in mixed conifer and deciduous forests of northern British Columbia, Canada. Predicting tree seedling recruitment after disturbance is fundamental to understanding forest dynamics and succession and is vital for forest management purposes. Seedling recruitment success in multi-species northern latitude forests varied as a function of mature tree canopy cover, gap size and position in a gap. Recruitment was abundant within canopy gaps across a wide range of gap sizes (20–5000 m²), but recruit numbers dropped off rapidly under the closed forest canopy and in the open conditions of clearcuts. Inside canopy gaps, recruitment was similar by gap position in small gaps (<300 m²) but, in these northern latitude forests, exhibited a trend of increasing density from the sunny north to shady south end of larger gaps. This was true for all tree species regardless of their shade tolerance ranking. There was no evidence of gap partitioning by any of the tree species during the regeneration phase suggesting that adaptation to the subtleties of gap size during early recruitment are not well developed in these tree species. Favorable locations for emergence and early establishment of germinants were less favorable for growth and survival of established seedlings, i.e. the regeneration niches in these forests were discordant. Tree abundance and species diversity appears to be controlled more by differentiation among growth and survival niches than by the regeneration niches. From the perspective of forest management, abundant natural regeneration of all the dominant tree species of these mixed-species forests can be obtained after partial cutting.     

林分间隙形成后季节性干旱的娑罗双树原始林物种多样性及苗木再生(英文)

本文调查了尼泊尔季节性干旱的娑罗双树原始林两种生态位(间隙和全郁闭)下树种多样性及苗木再生情况,并对物种多样性指标和苗木再生属性与林分间隙特征进行了相关性分析。结果表明,间隙生态位下乔木和灌木立木度高于全郁闭。间隙生态位下娑罗双树和榄仁树的幼苗密度也高于全郁闭,但是油榄仁和海南蒲桃的比较结果恰好相反。在幼苗层物种多样性指标(complement of Simpson index,Evenness index,and species-individual ratio)在间隙生态位要低于全郁闭条件。间隙大小能够解释物种丰富度和物种建立速率。多年多样伐木形成的林分间隙下娑罗双树幼苗密度要高于同一年内单一或多样伐木。结论:林分间隙的物种多样性通过增加幼苗密度来维持,并且适于娑罗双树再生。此外,林分间隙的其它属性也会影响物种多样性和苗木再生。     

林隙微生境及更新研究进展

系统评述了国内外关于林隙干扰及更新的最新研究进展.林隙是一种经常发生的小规模干扰,是森林群落时空格局变化的驱动力.不同的林隙干扰状况导致了林隙内光照的差异,进一步造成林隙内温度、湿度、土壤理化性质的变化,形成了微生境的时空异质性.林隙干扰通过改变微生境而对幼苗的出土、存活、定居和生长产生重要影响.林隙内种子库的物种丰富度变化很大,是幼苗分布格局的重要影响因子.更新幼苗的存活率和生长量受林隙大小及幼苗在林隙中位置的影响.指出了这一领域今后的研究方向.     

Natural regeneration of Picea abies on small clear-cuts in SE Norway

The objective was to study the influence of vegetation cover, humus depth, microrelief and distance to seed tree edge on natural regeneration of Norway spruce (Picea abies (L.) Karst.) in five small clear-cuts on bilberry woodland in southeastern Norway. The impact of the vegetation was considered at tree different scales: at the growing point, in the nearest square decimetre around the seedlings and at a 1 m² scale. Most of the regeneration (95%) developed in 1996, after a rich seed year in 1995. The microhabitat was of crucial importance to seedling establishment. Litter, Sphagnum spp. and Polytrichum commune were good substrates for establishment, while areas dominated by Deschampsia flexuosa, Dicranum spp. and Pleurozium schreberi had very few seedlings. Seedling survival after five growing seasons was slightly better in litter than in Sphagnum and other mosses, but the differences were not statistically significant. Increasing humus depth had a positive influence on regeneration, probably due to shallow soils at the sites. Even though depressions covered only 4.9% of the ground, 24.1% of the seedlings occurred here. Survival was, however, lower in depressions than in the other microrelief classes. Distance to the seed tree edge had a significant influence on establishment, with more seedlings establishing close to the edge.     

Spatial variation of seed rain and seed banks in gaps of karst forest in the Maolan Nature Reserve, Guizhou Province

Based on an investigation on gaps and non-gap stands of the Maolan National Karst Forest Nature Reserve, Guizhou Province, quantitative characteristics and dynamic changes of seed rain and seed banks in gaps were analyzed. The results show that the total amount of seed rain was 117.4 ± 32.6 seeds/m² during the period of observation. The number of immature seeds was 56.3 ± 10.3 seeds/m², that of mature damaged seeds was 15.7 ± 4.7 seeds/m², and the number of mature germinated seeds was 45.4 ± 8.2 seeds/m². It is suggested that the seed number is rich for gap regeneration. Seed rain in gaps has spatial and temporal heterogeneities which deeply affect regeneration patterns of gap plants. Along a gradient from the gap center to a non-gap stand, seed density in the litter layer, the number of species, and the Shannon-Wiener diversity index were gradually reduced, but these indices increased in the soil. The seed density in the gap center was 2415 ± 639 seeds/m², near the gap center was 2218 ± 421 seeds/m² and at the gap border area 1815 ± 311 seeds/m². This shows that plants in gaps have good latent regeneration potential. In both gaps and non-gap stands, the Jaccard similarity index of seed in litter layer was the largest, second largest at 5–10 cm soil depth, and the least at the 0–5 cm soil layer the index. The Jaccard index between the soil seed bank and the present plant community was large in the litter layer, but decreased with soil depth both in gaps and non-gap stands. The results show that soil seed banks are the main source of gap regeneration in the karst forests of Maolan and contribute significantly to gap regeneration. __________ Translated from Acta Botanica Yunnanica, 2007, 29(3): 327–332 [译自: 云南植物研究]     

Species diversity,regeneration and dominance as influenced by canopy gaps and their characteristics in tropical evergreen forests of Western Ghats,India

Canopy gaps play a significant role in maintaining structure and composition of tropical forests. This study was carried out in tropical evergreen forests of central Western Ghats in India to understand the influence of canopy gap size and the relationship of gap regime attributes to diversity measures and regeneration. The average gap size in the study area was found to be 396 m2 and around half of gaps were 4–8 years old. Gaps created by natural single tree fall were smaller in size but significantly higher in number. Diversity and regeneration of woody species were compared with canopy gaps and intact vegetation. Species richness and diversity was higher in gaps than in intact vegetation. Macaranga peltata, a shade intolerant species dominated gaps while intact vegetation was dominated by shade tolerant Kingiodendron pinnatum.Gap size significantly influenced species diversity and regeneration. Gap area and age were significantly and negatively correlated with diversity measures but positively correlated with regeneration. Among all the attributes of gaps, regeneration was significantly positively correlated with light intensity. Gaps maintained species diversity and favored regeneration of woody species. In addition to gap size and age, other gap ecological attributes also affected species diversity and regeneration.     

Patterns of regeneration and ground vegetation associated with canopy gaps in a managed beechwood in southern England

In autumn 2001, 15 canopy gaps were selected for study in RumerhedgeWood, a semi-natural, mesotrophic beechwood in southern England.The gaps were located in mature, beech-dominated stands, andhad originated from openings created during a thinning in theearly 1980s and wind damage in 1987/1990 and/or the consequentsalvage operations. The extent of each gap and surrounding treeswere mapped. Tree/shrub regeneration, ground vegetation, bareearth, leaf litter and canopy openness (using a canopy-scope)were measured within and around the gaps using a 5 x 5-m gridand placing a 1 x 1-m quadrat at each grid intersection (totalnumber of quadrats = 400). Most of the gaps were <75 m² inarea. The largest was 241 m². They were generally irregularin shape and there was little or no understorey present. Mostsurrounding trees were beech Fagus sylvatica L. Bramble Rubusfruticosus L. formed a moderate to dense ground vegetation belowmost gaps and declined around the edges only once the gap openingwas substantially obscured. Apart from a few larger saplings,most regeneration was small and of beech. Most of the latterappeared to be in their fifth or sixth growing season, were10–35 cm tall, had an erect base and flat top, had increasedby <5 cm in height during 2001 and were not browsed by deer.Their height and growth form was related to (1) their positionwithin gaps, (2) the degree of canopy openness and (3) the coverof ground vegetation. This was translated into the followingzonation—(1) around the centre of larger gaps: canopyopenness increased to >15 per cent; bramble cover was nearcomplete; litter depth was low; many places had no beech seedlings,but some of the few present were among the tallest, most uprightand fastest growing; (2) towards the edges of the large gapsor directly below smaller gaps: canopy openness was about 4–10per cent; bramble cover was slightly less; beech seedlings weremoderately abundant but patchy, generally shorter, more flattopped and slow growing than in the gap centre, albeit somewere still among the tallest, most upright and fastest growing;(3) beyond the edge of the large/medium gaps (with the gap onlypartially visible) or directly below very small gaps: canopyopenness was only about 2–3 per cent; bramble was muchreduced; beech seedlings were at their most abundant but stillpatchy in distribution and even shorter, more flat topped andslower growing than in the above zone; (4) in an outer zonebeyond or almost beyond the sight of the gaps: canopy opennesswas <2 per cent; bramble was weak and sparse; beech seedlingswere mostly at low densities and predominately short, slow growingand flat topped. This ring pattern of beech regeneration appearedto relate mainly to (1) differences in light availability affectingthe survival, growth rate and form of seedlings; (2) competitionfrom bramble and possibly (3) limited dispersal of beech seedinto gaps. Recommendations are given for managers who wish touse natural regeneration to restock beech woodland.     

Canopy gaps and regeneration in old-growth Oriental beech (Fagus orientalis Lipsky) stands, northern Iran

Virgin beech Fagus orientalis forests in northern Iran provide a unique opportunity to study the disturbance regimes of forest ecosystems without human influence. The aim of this research was to describe characteristics of natural canopy gaps and gap area fraction as an environmental influence on the success of beech seedling establishment in mature beech stands. All canopy gaps and related forest parameters were measured within three 25 ha areas within the Gorazbon compartment of the University of Tehran’s Kheyrud Experimental Forest. An average of 3 gaps/ha occurred in the forest and gap sizes ranged from 19 to 1250 m² in size. The most frequent (58%) canopy gaps were <200 m². In total, canopy gaps covered 9.3% of the forest area. Gaps <400 m² in size were irregular in shape, but larger gaps did not differ significantly in shape from a circle. Most gaps (41%) were formed by a single tree-fall event and beech made up 63% of gap makers and 93% of gap fillers. Frequency and diversity of tree seedlings were not significantly correlated with gap size. The minimum gap size that contained at least one beech gap-filling sapling (<1.3 m tall) was 23.7 m². The median gap size containing at least one beech gap-filling sapling was 206 m² and the maximum size was 1808 m². The management implications from our study suggest that the creation of small and medium sized gaps in mixed beech forest should mimic natural disturbance regimes and provide suitable conditions for successful beech regeneration.     

Tree regeneration in gap-understory mosaics in a subtropical Shorea robusta(Sal) forest

Spatial variation in tree-regeneration density is attributed to the specialization of tree species to light availability for germination and growth. Light availability,in turn, varies across the gap-understorey mosaic. Canopy gaps provide an important habitat for the regeneration of tree species that would otherwise be suppressed in the understory. In subtropical forests, there is still a knowledge-gap relating to how canopy disturbances influence tree-regeneration patterns at local scale, and if they disproportionately favor regeneration of certain species. We aim to analyze whether canopy gaps promote tree regeneration, and tree species are specialized to gaps or understory for germination and growth. We sampled vegetation in 128 plots(0.01 ha), equally distributed in gaps and below canopy, in two subtropical Shorea robusta Gaertn.(Sal) forests in Nepal, recording the number of tree seedlings and saplings in each plot. We compared the regeneration density of seedlings and saplings separately between gaps and the understorey. The mean densities of seedlings and saplings were higher in the gaps at both sites;although there was no difference in the seedling density of the majority of the species between the habitats. No species were confined to either gap or understorey at the seedling stage. We conclude that gaps are not critical for the germination of tree species in Sal forests but these are an important habitat for enabling seedlings to survive into saplings. The classification of trees into regeneration guilds mainly based on germination does not apply to the majority of tree species in subtropical Sal forests. Our results reaffirm that gap creation promotes tree regeneration by favouring seedling survival and growth and can influence forest management for conservation, as well as for plantations.     

西藏色季拉山冷杉原始林林隙更新研究

通过对西藏色季拉山急尖长苞冷杉原始林林隙更新调查，研究了林隙大小与更新以及林隙形成年龄与更新的关系，对林隙和非林隙林分内幼苗和幼树的高生长进行了比较。结果表明：林隙大小在100m^2左右，最适合于苗高20cm以下的更新幼苗生存，超过100m^2，更新幼苗随着林隙面积的增大而减少；林隙大小在300m^2左右，最适合冷杉幼树和小径木的生存。林隙形成年龄在30a左右，更新幼苗和幼树的数量最多，而小径木数量有限，随着林隙形成年龄的增大，小径木数量逐渐增多，而幼苗和幼树逐渐减少；林隙内幼苗、幼树和小径木的密度均大于非林隙林分；林隙对3年生以下幼苗的高生长影响不大，对4年生以上幼树的高生长有促进作用，特别是对9年生至16年生幼树高生长的促进作用最明显。     

Regeneration patterns of Eastern white pine (Pinus strobus L.) in hardwood-dominated forests in Virginia, USA

Eastern white pine (Pinus strobus L.) is a moderately shade-tolerant species that co-occurs with hardwood tree species in many forests of the eastern United States, as well as in pure stands. The species is valued for its timber, as well as for wildlife and recreation. Regeneration of this species is somewhat unpredictable and often occurs in patches of similarly-aged cohorts. We described the regeneration patterns of this species and examined their relation to environmental variables within hardwood forests of southwestern Virginia, USA. An average of 5.3 white pine patches per ha were observed in this study. The majority of patches consisted of saplings (85%), with 9% of patches in pole size classes, and 6% in seedling size classes. The average density of patches was 43.5 stems with an average age of 20 years. The size of patches averaged 80.6 m². The total density of seedlings and the number of regeneration patches of all sizes of regeneration (seedlings, saplings, and poles) in plots was related to the surrounding density of large white pine trees (potential seed trees). The density of seedlings or patches was not significantly related to current vegetation cover or soil surface cover variables, but more than half of regeneration patches were located in or adjacent to old canopy gaps, most of which were old logging gaps. While seedling regeneration may occur within the understory of these forests near seed trees, advancement to the sapling and pole stage appears to be associated with canopy gap formation.     

Thirty years of gap dynamics in a central european beech forest reserve

Investigation of gap characteristics and tree regeneration patternsis central for our understanding of forest dynamics. By integratingaerial photograph analyses and ground surveys, we provide astudy of long-term canopy gap dynamics and tree regenerationpatterns in a Hungarian beech forest reserve. We found (1) thatin spite of the overall increase of gap area during the investigated30-years (from 2.5 to 7.7 per cent), total gap area and averagegap size (40–93 m²) were remarkably similar to those foundin other temperate and tropical forests, (2) if the fate ofindividual gaps was followed, two to three times more intensivecanopy dynamics (gap creation, closure and expansion) couldbe recognized than simple change of gap area indicated, and(3) average seedling density was considered to be sufficientfor natural regeneration. However, it was apparent that recentincreased deer browsing had prevented establishment of youngertrees of 1–2 m in height, as taller saplings were recordedonly in old gaps. Our results not only provide useful informationon forest dynamics but can also contribute to understandingthe potential roles that small forest reserves can play in providingessential reference data for nature-based forest managementof this forest type.     

Interacting effects of canopy gap, understory vegetation and leaf litter on tree seedling recruitment and composition in tropical secondary forests

We experimentally investigated interacting effects of canopy gaps, understory vegetation and leaf litter on recruitment and mortality of tree seedlings at the community level in a 20-year-old lowland forest in Costa Rica, and tested several predictions based on results of previous studies. We predicted that experimental canopy gaps would greatly enhance tree seedling recruitment, and that leaf litter removal would further enhance recruitment of small-seeded, shade-intolerant seedlings in gaps. We created a large (320–540 m²) gap in the center of 5 out of 10 40 m × 40 m experimental plots, and applied the following treatments bimonthly over a 14-month-period in a factorial, split–split plot design: clipping of understory vegetation (cut, uncut), and leaf litter manipulations (removal, addition, control). As expected, experimental gaps dramatically increased tree seedling recruitment, but gap effects varied among litter treatments. Litter addition reduced recruitment in gaps, but enhanced recruitment under intact canopy. Species composition of recruits also differed markedly between gap treatments: several small-seeded pioneer and long-lived pioneer species recruited almost exclusively in gaps. In contrast, a few medium-to-large-seeded shade-tolerant species recruited predominantly under intact canopy. Leaf litter represents a major barrier for seedling emergence and establishment of small-seeded, shade-intolerant species, but enhances emergence and establishment of large-seeded, shade-tolerant species, possibly through increased humidity and reduced detection by predators. Periodic clipping of the understory vegetation marginally reduced tree seedling mortality, but only in experimental gaps, where understory vegetation cover was greatly enhanced compared to intact canopy conditions. Successful regeneration of commercially valuable long-lived pioneer trees that dominate the forest canopy may require clear-cutting, as well as weeding and site preparation (litter removal) treatments in felling clearings. Management systems that mimic natural canopy gaps (reduced-impact selective logging) could favor the regeneration of shade-tolerant tree species, potentially accelerating convergence to old-growth forest composition. In contrast, systems that produce large canopy openings (clear-cutting) may re-initiate succession, potentially leading to less diverse but perhaps more easily managed “natural plantations” of long-lived pioneer tree species.     

Gap size effects on above- and below-ground processes in a silver fir stand

Changes in soil properties related to natural regeneration of silver fir (Abies alba, Mill.) in small (185 m²) and medium (410 m²) gaps, in the Calabria Apennines (Southern Italy), were studied 2 years after gap creation. The organic matter within the medium gaps decayed more rapidly than those in the small gaps. Among the microenvironmental variables and soil properties, soil temperature was most strongly positively related to organic matter decomposition rates. Medium gaps had high soil temperature and photosynthetically active radiation (PAR) transmittance, and low soil moisture. Within medium gaps, we found a great amount of phenolic compounds and a low content of organic matter and humic acids. These results indicate that in the medium gaps mineralization of organic matter was more important than humification. In contrast, in small gaps, the great amount of organic matter, the high content of humic acid and the values of humification index suggested that in these gaps the humification process of organic matter prevailed. Within small gaps, we observed more silver fir regeneration than medium gaps. Difference in organic matter content, particularly in the amount of phenolic compounds, may account for differences observed in natural regeneration of silver fir between small and medium gaps.