Shade tolerance

Secondary successional vegetation represents one-third of the 7% of the original area covered with tropical dry forest that remains in central Veracruz, Mexico. In this region, fallow periods are short, in general, no longer than 7–10 years, and old secondary successional sites are not available. Therefore we evaluated the potential of very early successional stages of tropical dry forest with different land use histories for recovering the structure and composition of regional forest. We compared five early successional sites (7–72 months) with five nearby forest remnants. Successional sites had reached 38 and 30%, respectively, of the average basal area and density recorded for the forest understories, but only 5 and 10%, respectively, of the basal area and density of forest overstories. A total of 132 tree species were recorded, 45 at successional and 107 at forest sites. Individuals of tree species with animal-dispersed seeds (57%) were significantly better represented than wind-dispersed (22%) and self-dispersing (21%) species in both successional and forest sites. Successional sites had already recruited 10% of the intermediate and shade-tolerant species that grow in forest remnants. However, only 20 species occurred in both early successional and forest sites, several showing resprouting ability. We conclude that the entry of forest species to the successional process at very early stages and the recruitment of individuals from resprouting may facilitate the recovery of the dry forest in Veracruz.

Young forests can be manipulated in diverse ways to enhance their ecological values. We used stem maps from two dense, second-growth stands in western Washington and a spatially explicit light model (tRAYci) to simulate effects of five silvicultural manipulations on diameter distribution, species composition, spatial patterning, and light availability. Each treatment removed 30% of the basal area, but differed in

La luz y el agua son dos de los principales factores que regulan el crecimiento y la supervivencia de las plantas leñosas mediterráneas. El objetivo de este estudio es conocer el nivel de respuesta plástica de plántulas de cuatro especies de Quercus a tratamientos de sombra y sequía. Además queremos evaluar qué variables son más plásticas a uno u otro factor, y probar la hipótesis de que los rasgos fisiológicos son más plásticos que los rasgos estructurales. Se cultivaron en condiciones controladas plántulas de Quercus ilex subsp. ballota, Q. súber, Q. pyrenaica y Q. canariensis en tres tratamientos de luz (100, 27 y 3 % de luz incidente) y con agua abundante. A partir de mediados de primavera, a la mitad de las plántulas se les suprimió el agua, de forma que recibieron dos tratamientos de agua (riego versus no riego) dentro de cada tratamiento de luz. Se analizaron distintos rasgos fisiológicos como la tasa de fotosíntesis y respiración, el punto de compensación y de saturación a l...

We examined the influence of small-scale gap disturbances on stand development and tree species coexistence in an old-growth Fagus sylvatica–Abies alba forest in the Dinaric Mountains of Bosnia–Herzegovina. The structure and composition of tree regeneration in gaps were compared to the forest as a whole, and the influence of gap size on the density and composition of regeneration was assessed. Transition probabilities were also calculated from gapfillers in different life stages to examine canopy replacement patterns. The structure and composition of tree regeneration were similar between gaps and the forest as a whole, and there was no relationship between overall regeneration density and gap size, indicating most individuals established prior to gap formation. Likewise, there was no strong evidence of gap-size partitioning for shade tolerant F. sylvatica and A. alba, although less tolerant Acer pseudoplatanus only recruited to taller life stages in larger gaps. Transition probabilities calculated from the seedling and sapling data suggest that most gaps will be captured by F. sylvatica, while probabilities based on pole-sized gapmakers indicate both A. alba and F. sylvatica will be maintained in the canopy. We suggest that gaps primarily play a role in reorganizing advance regeneration, and that coexistence of shade tolerant F. sylvatica and A. alba is more likely related to their differential ability to tolerate shaded understory conditions, particularly during larger life stages, rather than gap-size partitioning.

Maximum Rubisco activities (V(cmax)), rates of photosynthetic electron transport (J(max)), and leaf nitrogen and chlorophyll concentrations were studied along a light gradient in the canopies of four temperate deciduous species differing in shade tolerance according to the ranking: Populus tremula L. < Fraxinus excelsior L. < Tilia cordata Mill. = Corylus avellana L. Long-term light environment at the canopy sampling locations was characterized by the fractional penetration of irradiance in the photosynthetically active spectral region (I(sum)). We used a process-based model to distinguish among photosynthesis limitations resulting from variability in fractional nitrogen investments in Rubisco (P(R)), bioenergetics (P(B), N in rate-limiting proteins of photosynthetic electron transport) and light harvesting machinery (P(L), N in chlorophyll and thylakoid chlorophyll-protein complexes). On an area basis, V(cmax) and J(max) (V(a) (cmax) and J(a) (max)) increased with increasing ...

Species dynamics in an abandoned urban forest of Central Japan is described in this paper. The dominant species in the urban plantation were Cryptomeria japonica and Chamaecyparis obtusa. A variety of eight patches of the canopy was produced by previous forest management practices. Progressive and retrogressive species dynamics within these eight patches are investigated in this paper. The study elucidates the deterministic role of patchiness in the nature of species colonization and the maintenance of species diversity in an urban forest. Altogether 139 native and/or naturalized species, including 23 shrub and 35 tree species, were recorded in the study area of 3.2 ha. The performance of species varied according to their successional attributes indicating a selective canopy influence. Twenty percent of the tree species were shade-intolerant pioneers (e.g. Cornus spp., Rhus javanica var. roxburghii) re-established under selective tree-felling. Thirty percent were shade-tolerant climax species (e.g. Neolitsea sericea, Persea thunbergii) dominant in remnant closed patches. The remaining 50% belonged to various seral types with aggressive deciduous species (e.g. Aphananthe aspera, Celtis sinensis) in most of the patches. Some ruderal herbaceous species dominated heavily disturbed clear-felled patches. This study suggests that canopy modification influences the subsequent colonization pattern. Furthermore, heterogeneous patches contribute to greater species diversity and dynamics in isolated woodlands.

Enrichment planting is commonly used for increasing the density of desired tree species in secondary forests often characterized by a prevalence of low commercial species. The objectives of the study were to identify the optimal enrichment planting method vis-à-vis gap and line planting, and to evaluate the performance of two dipterocarps (Vatica cinerea and Dipterocarpus alatus) and three legumes (Afzelia xylocarpa, Pterocarpus macrocarpus, Dalbergia cochinchinensis) planted in logged-over mixed deciduous forest of Laos. The enrichment planting trial was arranged in a randomized complete block design with seven replications. Survival, height and diameter were measured seven years after planting and subjected to analysis of variance. Survival rate of planted seedlings did not vary between enrichment planting (p > 0.05). However, diameter and height growth was favored more in gaps than in planting lines. This was related to rapid canopy closure in both gaps and lines, albeit more remarkable in planting lines. Significant inter-species variation was detected for survival rate, height and diameter. The shade-tolerant dipterocarps had better survival and growth than the light-demanding leguminous species (p < 0.0001). The size class distribution of individuals was irregular, accentuating uneven light condition in the understory. Given the difficulty to maintain constant line width and even light condition, the cost of annual clean operation and the rigid geometric patterns of planting lines, the use of logged-over gaps for enrichment planting is recommended. Dipterocarpus alatus and V. cinerea are recommended for enrichment planting in mixed deciduous forests. A. xylocarpa, D. cochichinensis and P. macrocarpus can be planted in wider gaps, lines and open sites as plantation of mixed species or under canopy of young swidden forests.

Large-scale patterns of Amazonian biodiversity have until now been obscured by a sparse and scattered inventory record. Here we present the first comprehensive spatial model of tree α-diversity and tree density in Amazonian rainforests, based on the largest-yet compilation of forest inventories and bolstered by a spatial interpolation technique that allows us to estimate diversity and density in areas that have never been inventoried. These data were then compared to continent-wide patterns of rainfall seasonality. We find that dry season length, while only weakly correlated with average tree α-diversity, is a strong predictor of tree density and of maximum tree α-diversity. The most diverse forests for any given DSL are concentrated in a narrow latitudinal band just south of the equator, while the least diverse forests for any given DSL are found in the Guayana Shield and Amazonian Bolivia. Denser forests are more diverse than sparser forests, even when we used a measure of diversity that corrects for sample size. We propose that rainfall seasonality regulates tree α-diversity and tree density by affecting shade tolerance and subsequently the number of different functional types of trees that can persist in an area.

Tree regeneration outcomes are challenging to generalize and difficult to predict. Many tree species can establish new propagules in a variety of post-disturbance environments and many different reproductive mechanisms may be used. In order to develop conceptual models that accurately reflect reproductive potential, we need a better understanding of the similarities in regeneration ecology among species. We used information from the forest ecology literature to evaluate the reproductive attributes of sixty-two tree species in the central hardwood region of the eastern United States. Each species was classified categorically for features such as flowering, seed production and dispersal, seed dormancy, germination requirements, seedling characteristics, and vegetative reproduction. Cluster analysis (Jaccard's similarity coefficient, complete linkage method) and ordination (homogeneity analysis) were used to separate nine groups (guilds) of species that had similar reproductive attributes. Individual attributes that had high variance in the first and second dimensions included: seed banking, seed dispersal, seedling shade tolerance, and seedbed requirements. Members of each guild had similar levels of reproductive specialization and guilds were either pioneer-like, opportunistic, or persistent. Pioneer guilds included: short-lived or fugitive species that colonize sites rapidly and are too shade intolerant to replace themselves; shade-tolerant species that colonize frequently disturbed sites; and stress-tolerant pioneers that survive on dry or nutrient-poor sites. Opportunistic guilds contained species that are remarkably versatile in their reproductive effort. The most flexible opportunists can colonize new sites, maintain seed in a seed bank, sprout from existing stems and persist as a seedling or sapling bank. Persistent guilds contain species that develop and maintain advance regeneration. These include: species with moderate understory tolerance that regenerate via cycles of dieback and resprouting; and more tolerant species that maintain seedling or sapling banks. Our regeneration guilds may provide a useful approach for more realistically representing large and diverse sets of tree species in forest ecosystem models.

We contrasted the regeneration success of angiosperm canopy species with that of Podocarpus latifolius to test whether this conifer was sufficiently shade tolerant to be favoured on infrequently disturbed and well-shaded sites. Seedling and sapling population structures were measured in several habitats representing a light gradient in warm temperate forest in the Drakenberg mountains of South Africa. The angiosperm-dominated forest