Establishment of tree seedlings in the understory of restoration plantations: natural regeneration and enrichment plantings

Little is known about the potential of restoration plantations to provide appropriate understory conditions to support the establishment of seeds arriving from neighboring native forests. In this article, we investigated how seedling establishment is affected in the understory of restoration sites of different ages and assessed some of the potential environmental factors controlling this ecological process. We first compared the density and richness of native tree seedlings among 10‐, 22‐, and 55‐year‐old restoration plantations within the Atlantic Forest region of southeastern Brazil. Then, we undertook a seed addition experiment in each study site, during the wet season, and compared seedling emergence, survival, and biomass on local versus old‐growth forest soil (transferred from a reference ecosystem), in order to test whether local substrate could hamper seedling establishment. As expected, the oldest restoration site had higher density and richness of spontaneously regenerating seedlings. However, seedling establishment was less successful both in the oldest restoration planting and using substrate transferred from a reference ecosystem, where emergence and survival were lower, but surviving seedlings grew better. We attribute these results to lower light availability for seedlings in the understory of the oldest site and speculate that higher incidence of pathogens on old‐growth forest soil may have increased seedling mortality. We conclude that the understory of young restoration plantations provides suitable microsite conditions at the early establishment phases for the spontaneous regeneration or enrichment planting of native trees.     

Seed survival on experimental dishes in a central European old‐growth mixed‐species forest – effects of predator guilds,tree masting and small mammal population dynamics

Predation of tree seeds can be a major factor structuring plant communities. We present a three year study on tree seed survival on experimental dishes in an old‐growth forest in central Europe in Austria. We addressed species specific, spatial and temporal aspects of post‐dispersal seed predation. Seeds of Norway spruce Picea abies, European beech Fagus sylvatica, and silver fir Abies alba were exposed on dishes in different types of exclosures which allowed access only to specific guilds of seed predators. Removal experiments were carried out in two old‐growth forests and a managed forest (macro‐sites), including micro‐sites with and without cover of ground vegetation. We conducted the experiment in three consecutive years with a mast year of beech and spruce before the first year of the study. The seed removal experiments were combined with live trapping of small mammals being potential seed predators. Our experiments showed a distinctly different impact of different predator guilds on seed survival on the dishes with highest removal rates of seeds from dishes accessible for small mammals. We observed differing preferences of small mammals for the different tree species. Seed survival in different macro‐ and micro‐habitats were highly variable with lower seed survival in old growth forests. In contrast to our assumption, and in contrast to the satiation hypothesis which assumes higher seed survival in and directly after mast years, seed survival was lower in the year following the mast year of beech when a population peak of small mammals occurred and higher in intermast periods when subsequently small mammal population crashed. This suggests a higher importance of sporadic masting shortly after mast years in intermast periods for establishment of forest trees provided that pollination efficiency is high enough in such years. Combined with the high seed mortality observed after the mast year, this corroborates the important role of seed predation for forest dynamics. An altered synchrony or asynchrony of masting of different tree species and changed masting frequencies through climate change may thus lead to strong and non‐linear effects on forest dynamics.     

Ecological consequences of a massive flowering event of bamboo (Chusquea culeou) in a temperate forest of Patagonia,Argentina

Question: What changes occur as a consequence of the massive flowering and senescence of the dominant understory species of bamboo, Chusquea culeou (E. Desvaux)? In this study, we documented some of the ecological consequences of this rare event that occurred in 2001, the previous flowering having occurred more than 60 years ago. Location: Nothofagus temperate forest, Patagonia, Argentina. Methods: We assessed changes in environmental variables and bamboo biomass post‐flowering in an old‐growth southern beech forest. In addition, we monitored the demography of emergent Ch. culeou seedling and Nothofagus nervosa saplings, comparing non‐flowered (live understory) and flowered (senescent understory) patches within the forest matrix. Results: Bamboo flowering dramatically increased light availability in the forest understory but, surprisingly, other environmental changes were not observed. Bamboo seedlings emerged in both patch types, and experienced gradual but modest mortality through time. Bamboo dieback promoted higher survivorship and an increment in biomass, height, number of leaves and buds in the saplings of Nothofagus nervosa. Conclusion: The high density of bamboo seedlings 5 years after the flowering event and the independence of emergence from environmental variables suggest that understory regeneration is a gradual process that is not strongly regulated by initial seedling density or resource limitation. In contrast, microenvironmental conditions created after the flowering event significantly increased Nothofagus sapling growth and survival. These results suggest that overstory forest regeneration could be enhanced in this temperate forest in the first years after this infrequent bamboo flowering event.     

Establishment of Fagus sylvatica and Fraxinus excelsior in an old-growth beech forest

Abstract. Distribution of tree seedlings, forest architecture, light conditions, ground vegetation and humus conditions were studied in a 45 m × 100 m area including multiple gaps in an old-growth beech forest. Gaps were created after some beech trees had been felled in severe storms in February 1990. A group of adult ash trees is found near the study site. The data were analyzed by Correspondence Analysis. Young seedlings (< 4 yr), of both Fraxinus (a sun species) and Fagus (a shade species), were most abundant under the crown of beech trees in semi-shade conditions, and where beech litter did not accumulate. Differences in the dissemination of Fraxinus and Fagus explained differences in the establishment of the two species. In contrast, older seedlings of beech established before the storms were more numerous in the gaps, suggesting a change in the ecological requirements of beech seedlings in the course of time.     

Multiple stages of tree seedling recruitment are altered in tropical forests degraded by selective logging

Tropical forest degradation is a global environmental issue. In degraded forests, seedling recruitment of canopy trees is vital for forest regeneration and recovery. We investigated how selective logging, a pervasive driver of tropical forest degradation, impacts canopy tree seedling recruitment, focusing on an endemic dipterocarp Dryobalanops lanceolata in Sabah, Borneo. During a mast‐fruiting event in intensively logged and nearby unlogged forest, we examined four stages of the seedling recruitment process: seed production, seed predation, and negative density‐dependent germination and seedling survival. Our results suggest that each stage of the seedling recruitment process is altered in logged forest. The seed crop of D. lanceolata trees in logged forest was one‐third smaller than that produced by trees in unlogged forest. The functional role of vertebrates in seed predation increased in logged forest while that of non‐vertebrates declined. Seeds in logged forest were less likely to germinate than those in unlogged forest. Germination increased with local‐scale conspecific seed density in unlogged forest, but seedling survival tended to decline. However, both germination and seedling survival increased with local‐scale conspecific seed density in logged forest. Notably, seed crop size, germination, and seedling survival tended to increase for larger trees in both unlogged and logged forests, suggesting that sustainable timber extraction and silvicultural practices designed to minimize damage to the residual stand are important to prevent seedling recruitment failure. Overall, these impacts sustained by several aspects of seedling recruitment in a mast‐fruiting year suggest that intensive selective logging may affect long‐term population dynamics of D. lanceolata. It is necessary to establish if other dipterocarp species, many of which are threatened by the timber trade, are similarly affected in tropical forests degraded by intensive selective logging.     

Disentangling the effects of facilitation on restoration of the Atlantic Forest

Facilitation is an important ecological mechanism with potential applications to forest restoration. We hypothesized that different facilitation treatments, distance from the forest edge and time since initiation of the experiment would affect forest restoration on abandoned pastures. Seed and seedling abundance, species richness and composition were recorded monthly during two years under isolated trees, bird perches and in open pasture. Seed arrival and seedling establishment were measured at 10 m and 300 m from the forest edge. We sampled a total of 131,826 seeds from 115 species and 487 seedlings from 46 species. Isolated trees and bird perches increased re-establishment of forest species; however, species richness was higher under isolated trees. Overall, abundance and richness of seeds and seedlings differed between sampling years, but was unaffected by distance from the forest edge. On the other hand, species composition of seeds and seedlings differed among facilitation treatments, distance from the forest edge and between years. Seedling establishment success rate was larger in large-seeded species than medium- and small-seeded species. Our results suggest that isolated trees enhance forest re-establishment, while bird perches provide a complementary effort to restore tree abundance in abandoned pastures. However, the importance of seed arrival facilitation shifts toward establishment facilitation over time. Arriving species may vary depending on the distance from the forest edge and disperser attractors. Efforts to restore tropical forests on abandoned pastures should take into account a combination of both restoration strategies, effects of time and proximity to forest edge to maximize regeneration.     

Gruppo VI: Bloenergetlca e Fotosintesi

Abstract

Understanding forest stand structural change is crucial for forest ecosystem research and management. The aim of our research was to identify structural and compositional changes and possible natural and anthropogenic causes of these changes in an old‐growth silver fir‐beech forest in the Dinaric mountains of Slovenia. Data on stand structure of the Rajhenavski Rog (51.14 ha) forest reserve originated from traditional methods, including five successive full callipering (1957–2007), two regeneration inventories and analysis of developmental phases (1984, 2007). A drastic decrease in the silver fir proportion in the last 50 years was probably caused by two indirect anthropogenic factors: polluted air and heavy browsing damage of silver fir regeneration. Nevertheless, other factors including reciprocal replacement of fir and beech and climate change also influenced the species composition, as similar processes were documented in the past with less air pollution. Furthermore, the alternation of fir and beech often developed asynchronously, regardless of ungulate density. Although the vertical profile of the old‐growth forest was opened due to the loss of fir trees during past decades, the establishment of more light‐demanding species was uncommon, primarily because the growing stock and forest climate were sustained. The results suggest a high stability of mixed Dinaric old‐growth forest ecosystems.     

Short‐term changes affecting regeneration of Fagus crenata after the simultaneous death of Sasa kurilensis

Abstract. Question: The aim of the present study is to determine whether seed/seedling predation will increase and Fagus survival will decline with the recovery of the Sasa cover. Methods: We examined Fagus crenata regeneration for seven years in an old‐growth Fagus‐Sasa forest near Lake Towada, northern Japan, by examining the effects of simultaneous death of Sasa, tree canopy gap formation, mast seeding of Fagus and seed and seedling predation by rodents on the survival of Fagus seeds and current year seedlings. We established four types of sites differing in forest canopy (closed or gap) and Sasa status (dead or alive) following the simultaneous flowering and death of Sasa kurilensis (dwarf bamboo) in 1995. Results: Fallen Fagus seed was abundant in 1997 and 2000 (mast years). In sites with alive Sasa, survival from the first growing season was low due to high seed and seedling predation. In sites with dead Sasa, seed survival under the canopy was high for both mast years, but in gaps it varied between years. Seedling survival was highest in canopy gaps with dead Sasa (gap‐dead) in 1998, because of higher light levels and lower predation by rodents. However, seedling survival in these plots was low in 2001, apparently because rapid Sasa recovery favoured rodent predation. In both mast years, Sasa die‐back had significant positive effects on seed and seedling survival under closed canopies because the seedlings there were more successful in escaping predation. Conclusion: The change in successful sites for the early stage of regeneration of Fagus appears to reflect the combined effects of canopy gap, seed/seedling predation and revegetation of Sasa.     

Fallow deer impacts on Wakatipu beech forest

This study uses data from forty-nine 20 m × 20 m permanent plots measured in 1976, 1982, 1989 and 1997-2002 in Wakatipu Forest, western Otago. We relate changes in red (Nothofagus fusca), silver (Nothofagus menziesii) and mountain beech (Nothofagus solandri var. cliffortioides) forest vegetation to the presence of fallow deer (Dama dama). Vegetation composition is likely to have been altered prior to plot establishment, and results show that there was little change in vegetation composition during the study. There are some signs of fallow deer reducing silver beech seedling abundance at some plots, but most beech stands appear to be in the late phase of development and undergoing self thinning, so are probably not immediately vulnerable to suppression of canopy regeneration. In the future, widespread disturbance is likely to release plants in the understorey from competition for light and nutrients, and at that stage fallow deer browsing of beech species may be able to alter successional pathways more than has occurred over the past two decades. Seedling and sapling density of the palatable Griselinia littoralis (broadleaf) was probably insufficient to ensure successful regeneration. Comparisons of seedling densities with other mixed beech forests throughout New Zealand suggest that fallow deer density will need to be nearly zero in Wakatipu Forest before regeneration of all palatable subcanopy hardwood species is assured.     

Multivariate path analysis of the relationships between seedling regeneration and environmental factors beneath a dwarf bamboo understory

Seedling emergence and establishment are fragile processes that determine the direction and structure of forest succession and regeneration. However, seedling emergence and establishment are easily affected by biotic and abiotic (environmental) factors. A dense and expanding understory of dwarf bamboo is one such important factor that can seriously hinder the seedling regeneration. We conducted a field experiment to investigate the emergence and establishment of canopy tree seedlings under artificially controlled densities of dwarf bamboo. We found that understory dwarf bamboo obstructed seedling emergence but reduced the death of seedlings. Although understory dwarf bamboo reduced the median retention time of seedlings, dense bamboo increased the mean survival time of seedlings. Our results suggest that understory dwarf bamboo has multiple selectivities for tree seedling emergence and establishment: high‐density dwarf bamboo was beneficial to evergreen species but lower‐density of bamboo was conducive to the survival of deciduous species, it means the dwarf bamboo potentially alters successional trajectories of forest communities. Path analysis revealed that the most important factors affecting tree seedling emergence and death were the abundance of seeds in the seed bank and the density of emerged seedlings, and that the soil temperature promoted seedling emergence but increased seedling death, the thickness of litter limited seedling emergence, and the leaf area index of the bamboo canopy limited seedling death. The present study suggests that dwarf bamboo can directly alter the microenvironment, significantly reducing light levels and soil temperature but increasing the thickness of litter and soil humus, thereby indirectly impacting the regeneration of tree seedlings. Our results indicate that various factors affected seedling emergence, and there were complex indirect relationships among these factors. In general, biological factors had a stronger influence on tree seedling regeneration than environmental factors.     

Seedling establishment of deciduous trees in various topographic positions

Abstract. We investigated the effect of topography‐related environmental factors (i.e., ground‐surface stability and soil moisture) on seedling establishment of 8 deciduous tree species. A field experiment was carried out using canopy species, which were classified into 3 groups based on the spatial distribution of adult trees (ridge, slope and valley). Demographic parameters were compared among species during the early stage of seedling establishment among 3 topographic positions in combination with gap and canopy conditions. The percentage of emerging seedlings tended to be lower on the ridge irrespective of the adult topographical distribution patterns. There was no clear trend in seedling emergence among the species groups classified by their spatial distribution. Seedling survival during 2 growing seasons was significantly different among species, topographic positions and light conditions. On the ridge, seedlings of the species dominating ridge tops had greater survival than those of other species, probably due to differences in demand for soil moisture. On the slope, frequent physical damage caused by surface material movement was observed and some species showed greater adaptability to the disturbed slope habitat. Survival of all seedlings was highest in the valley plots. Light conditions were the critical factor for seedling survival in some species. The results of this study suggest that topography creates diverse habitats for the establishment of tree seedlings. In addition to soil moisture, surface material movement may be a significant factor affecting seedling establishment.     

The potential of small exclosures in assisting regeneration of coffee shade trees in South‐Western Ethiopian coffee forests

Ethiopian Afromontane moist forests where coffee grows as understorey shrub are traditionally managed by the local communities for coffee production through thinning of the shade tree canopy and slashing of competing undergrowth. This management practice has a negative impact on the coffee shrubs, because the removal of shade tree saplings and seedlings reduces the succession potential of the shade tree canopy, which threatens the very existence of the shade coffee production system. We assessed the functionality of small exclosures to initiate coffee shade tree canopy restoration through natural regeneration. Our results show that small exclosures have a strong restoration potential for the coffee shade trees preferred by farmers (Albizia schimperiana, A. gummifera and Millettia ferruginea), as evidenced from their seedling abundance, survival and growth. The regeneration of late‐successional tree species of the moist Afromontane forest was not successful in the small exclosures, most probably due to the low abundance or absence of adult trees as seed sources for regeneration. Therefore, temporary establishment of small exclosures in degraded coffee forest fragments where shade trees are getting old or dying is recommended for sustainable shade coffee production.     

Conspecific density dependence and community structure: Insights from 11 years of monitoring in an old‐growth temperate forest in Northeast China

Forest community structure may be influenced by seedling density dependence, however, the effect is loosely coupled with population dynamics and diversity in the short term. In the long term the strength of conspecific density dependence may fluctuate over time because of seedling abundance, yet few long‐term studies exist. Based on 11 years of seedling census data and tree census data from a 25‐ha temperate forest plot in Northeast China, we used generalized linear mixed models to test the relative effects of local neighborhood density and abiotic factors on seedling density and seedling survival. Spatial point pattern analysis was used to determine if spatial patterns of saplings and juveniles, in relation to conspecific adults, were in accordance with patterns uncovered by conspecific negative density dependence at the seedling stage. Our long‐term results showed that seedling density was mainly positively affected by conspecific density, suggesting dispersal limitation of seedling development. The probability of seedling survival significantly decreased over 1 year with increasing conspecific density, indicating conspecific negative density dependence in seedling establishment. Although there was variation in conspecific negative density dependence at the seedling stage among species and across years, a dispersed pattern of conspecific saplings relative to conspecific adults at the local scale (<10 m) was observed in four of the 11 species examined. Overall, sapling spatial patterns were consistent with the impacts of conspecific density on seedling dynamics, which suggests that conspecific negative density dependence is persistent over the long term. From the long‐term perspective, conspecific density dependence is an important driver of species coexistence in temperate forests.     

Gnawing damage by rodents to the seedlings ofFagus crenata andQuercus mongolica var.grosseserrata in a temperateSasa grassland-deciduous forest series in southwestern Japan

The effects of dwarf bamboo,Sasa, cover on the initial morrality of hardwood seedlings were investigated by transplanting 1-year-old beech (Fagus crenata) and current-year oak (Quercus mongolica var.grosseserrata) seedling to three different stands; old-growth beech and secondary oak forests withSasa undergrowth, and aSasa grassland in a grassland-forest series near the top of Mt Jippo, southwestern Japan. The most frequent cause of seedling morrality was gnawing of the stems by rodents. In the beech forest, the gnawing was more likely to occur underSasa cover, suggesting that it provides a good habitat for rodents on the beech forest floor. TheSasa under growth may thus play an imporrant role in regeneration of beech forest. In the oak floor, mortality of both species was low and only a little gnawing occurred during a year. However, no natural oak seedling were found in the forest even after a mast year. This may be because most of the acorns disappeated before establishment. The early-stage demography of hardwood seedling as oak may thus play an imporrant role in regeneration of oak forest. In theSasa grassland where the seed supply is small, almost all of the seedlings died fromo gnawing regardless of the presence ofSasa cover. These factors prevent the recruitment of a sizable seedling bank. Rodents may thus play an imporrant role in maintenance of theSasa grassland.     

Natural regeneration of selected timber species in the Republic of Congo

Natural regeneration of timber species is critical to the sustainable management of tropical forests. To understand what determines regeneration success of timber species in the Congo Basin, we evaluated whether seedling recruitment rates differed between forest logged 30 years previously and unlogged forest and determined the environmental factors that influence seedling density, growth and survival. We monitored the fate of 2186 seedlings of seven timber species within 462, 25‐m² plots located along 21 transects. We characterized seedling plots by light availability, soil nutrient availability and pH, and abundance of mammalian herbivores and then used linear and generalized linear mixed models to evaluate the variables that influenced seedling density, growth and survival. Light availability and canopy openness were 18% and 81% higher in logged than unlogged forest, and concentration of soil nutrients varied between sites. Seedling density was 32% higher in unlogged than logged forest. Taking all species together, seedling survival was positively correlated with calcium and negatively with magnesium and available phosphorus. Rates of seedling growth increased with available light. Taken separately, seedlings of the selected timber species responded differently to abiotic and biotic factors, demonstrating species‐specific regeneration requirements.     

Driving factors and temporal fluctuation of Collembola communities and reproductive mode across forest types and regions

Despite the major role of Collembola in forest soil animal food webs, ecological and evolutionary determinants of their community composition are not well understood. We investigated abundance, community structure, life forms, and reproductive mode of Collembola in four different forest types (coniferous, young managed beech, old managed beech, and unmanaged beech forests) representing different management intensities. Forest types were replicated within three regions across Germany: the Schorfheide‐Chorin, the Hainich, and the Swabian Alb, differing in geology, altitude, and climate. To account for temporal variation, samples were taken twice with an interval of 3 years. To identify driving factors of Collembola community structure, we applied structural equation modeling, including an index of forest management intensity, abiotic and biotic factors such as pH, C‐to‐N ratio of leaf litter, microbial biomass, and fungal‐to‐bacterial ratio. Collembola abundance, biomass, and community composition differed markedly between years, with most pronounced differences in the Schorfheide, the region with the harshest climatic conditions. There, temporal fluctuations of parthenogenetic Collembola were significantly higher than in the other regions. In the year with the more favorable conditions, parthenogenetic species flourished, with their abundance depending mainly on abiotic, density‐independent factors. This is in line with the “Structured Resource Theory of Sexual Reproduction,” stating that parthenogenetic species are favored if density‐independent factors, such as desiccation, frost or flooding, prevail. In contrast, sexual species in the same year were mainly influenced by resource quality‐related factors such as the fungal‐to‐bacterial ratio and the C‐to‐N ratio of leaf litter. The influence of forest management intensity on abundances was low, indicating that disturbance through forest management plays a minor role. Accordingly, differences in community composition were more pronounced between regions than between different forest types, pointing to the importance of regional factors.     

Succession and Micro-elevation Effects on Seedling Establishment of Calophyllum brasiliense Camb. (Clusiaceae) in an Amazonian River Meander Forest1

I investigated the effects of successional stage and micro‐elevation on seedling establishment of Calophyllum brasiliense (Clusiaceae), a common canopy tree of seasonally flooded lowland forest along the Manú River meander zone in southeastern Peru. To compare seedling establishment between microhabitat types, I planted C. brasiliense seeds in a fully crossed experimental design of three successional stages (early, mid, and mature) and two micro‐elevations (levees and backwaters). Seedling establishment success in this study was affected by both successional stage and micro‐elevation, but micro‐elevation was most important in mid‐successional habitats. In general, seedlings in early succession experienced better conditions than in mature forest; light levels were higher, herbivory lower, and seedling growth higher. In mid‐successional forest, micro‐elevation determined habitat quality; backwaters had higher light levels, lower herbivory, and higher seedling growth and survival than levees. Mid‐successional backwaters were similar in quality to early successional forest for seedling establishment, while levees in that same successional stage were the poorest microhabitats for establishment. Although mid‐successional backwaters are similar to early succession for seedling establishment, in the long run, seedlings that establish in mid‐succession have a lower chance of reaching reproductive size before their habitat ages to mature forest than members of their cohort that established in early succession. I hypothesize that successful recruitment for C. brasiliense in the Manú River meander system requires dispersal to early successional habitat.     

Long‐term forest structure and regeneration after wildfire in Russian Karelia

We examined forest structure and regeneration in a 350‐ha forest dominated by Pinus sylvestris 31 yr after a wildfire in the Vienansalo wilderness, Russian Karelia. In most parts of the area, the 1969 fire was not stand replacing but had left larger trees alive so that the area generally remained forest covered. In some localities, however, all trees apparently died and distinct gaps were formed, suggesting that the fire severity varied considerably, contributing to increased variation in stand structure. Living and dead wood volumes were similar, 112 and 96 m³.ha^‐1, respectively. The tree species proportions of dead vs living wood indicated that prior to fire disturbance Picea was more common in the area. Regeneration was abundant (saplings, ca. 14 000 ind.ha^‐1, height 20 ‐200 cm) and tree seedling recruitment had occurred over a long period of time. Regeneration density was highest on the mesic Vaccinium‐Myrtillus forest site type, decreasing towards nutrient‐poor site types. The most common regeneration microsites were level ground (56% of saplings), immediate surroundings of decayed wood (23%) and depressions (11%). The high proportion of saplings on level ground suggests that after the fire regeneration conditions have been favourable across the whole forest floor. Nevertheless, the areas in the vicinity of decayed wood have been particularly important microsites for seedling establishment. The results provide an example of the effects of wildfire on forest structure in a natural Pinus sylvestris dominated forest, demonstrating the non stand replacing character of fire, high variability in stand structure and the abundance of post‐fire regeneration.     

Variation of plant diversity in a temperate unmanaged forest in northern Spain: behind the environmental and spatial explanation

The persistence of seedlings in the forest understorey is of major importance for the maintenance and regeneration of canopy trees in several forested ecosystems. In the present study, we examine the small-scale spatial pattern of a mixed beech and oak seedling–sapling bank in two areas of an unmanaged temperate deciduous forest with different environmental conditions. We used environmental, biotic and spatial variables to establish the main factors that explain the spatial pattern of these seedling–sapling banks at different scales. The stand structure in both areas was similar, but while in plot A beech dominated the canopy, plot B was dominated by oaks. In both areas, established beech individuals showed a clear reverse J-shaped distribution, whereas established oaks showed a unimodal distribution with only a few young individuals. Seedlings of beech and oak were distributed in aggregates, whereas beech saplings had a random distribution. At broader scales, the abundance of seedlings and saplings is affected by the environment as well as by inter-species competition, while at finer scales the spatial pattern is mainly influenced by stochastic processes, probably related to seed predation and establishment. The structure of the seedling–sapling bank indicates an advantage of beech over oak as far as regeneration is concerned. Beech seedlings and saplings tolerate the stress induced by the canopy and the understorey and persist for many years, while oak seedlings decline in a few years. Therefore, if current conditions persist, after canopy opening beech seedlings and saplings can grow rapidly into the canopy and the stands will move towards beech dominance.     

Tree seedling dynamics in two abandoned tropical dry forests of differing successional status in Yucatán,Mexico: a field experiment with N and P fertilization

The effects of nitrogen (N) and phosphorus (P) fertilization on seedling survival, recruitment, species richness and diversity in two abandoned tropical dry forests (10-yr old, young forest, and c. 60-yr old, old forest) in Yucatán, Mexico, were studied over two years. The seedling dynamics in the control plots were found to be highly seasonal with highest recruitment and lowest death rates during the rainy season. A low percentage of seedlings were resprouts; this important mechanism for forest regeneration had a higher-than-expected survival when compared to seedlings regenerated from seeds. Nutrient addition had significant effects on seedling dynamics in both of the forest regeneration stages. In the young forest, N fertilization facilitated the increase of seedling density. In the old forest, the addition of P decreased seedling diversity, while it increased the recruitment of only a few species. In both forests, P fertilization increased the survival time of seedlings when interacting with light availability and bulk density on the topsoil layer. Results suggest that low nutrient availability combined with low light availability constrain forest succession in Yucatán, Mexico. This revised version was published online in August 2006 with corrections to the Cover Date.