Annual variation in predation and dispersal of Arolla pine (Pinus cembra L.) seeds by Eurasian red squirrels and other seed-eaters

Seed predation and dispersal are key processes in the survival and distribution of plant species. Many animals cache seeds for later consumption, and, failing to recover some of these seeds, act as seed dispersers, influencing post-dispersal seed and seedling survival. Both animal and plant benefit from scatterhoarding and natural selection of seed characteristics and adaptations of seed predators (and dispersers) is one of the most important examples of co-evolution and mutualism. We studied the producer–consumer Arolla pine (Pinus cembra)–red squirrel (Sciurus vulgaris) system in a subalpine forest in the Italian Alps. Arolla pine produced large seed-crops (masting) at irregular intervals, followed by years with poor or moderate seed production. Squirrel density fluctuated in synchrony with the food resource, eliminating the time-lag normally present when resources are produced in pulses. In all years except 2009 (a mast-crop year), all Arolla pine cones were harvested (their seeds consumed and/or cached) by September to late October by different species. Both squirrels and nutcrackers (Nucifraga caryocatactes) fed on seeds, and their relative pre-dispersal predation rates (on cones in the canopy) differed between years. Overall, nutcrackers consumed more seeds between July and October than squirrels, but in 1 year squirrels took the largest number of seeds. Pre-dispersal seed predation by squirrels tended to be lower in years with large seed-crop size and there was a positive correlation, over the entire study period, between density of recovered hoards and Arolla pine seed density of the previous year. We conclude that (i) squirrels and nutcrackers are important pre-dispersal seed predators and seeds dispersers; (ii) squirrels are also post-dispersal seed predators, and (iii) the proportion of cached seeds consumed by squirrels increased with the size of the Arolla pine seed-crop, suggesting that red squirrel is a conditional mutualistic scatterhoarder of Arolla pine seeds.     

Flying vs. climbing: Factors controlling arboreal seed removal in oak-beech forests

Nuts are heavy and nutritious seeds that need animals to be successfully dispersed. Most studies address nut removal by a single animal species once seeds fall onto the ground. However, nuts are also accessible before the seed drop and usually to a wide guild of seed foragers. This study examines the factors controlling arboreal seed removal in oak-beech forests within the whole guild of nut foragers. We found that seed-dispersing rodents (Apodemus sylvaticus) were the main acorn removers in the oaks (up to 3.75 m height), with a rapid seed encounter and a high removal rate. However, rodents did not climb the beech trees, probably due to their smoother bark in comparison to oak bark and/or the lower nutritional value of beechnuts with regard to acorns. Jays (Garrulus glandarius) were more abundant in oak stands (both dense and scattered) and clearly preferred acorns to beechnuts whereas nuthatches (Sitta europaea) were more abundant in beech stands and preferred beechnuts to acorns. Non-storing birds such as great tits (Parus major) also removed acorns and beechnuts, especially in the stands where oaks are dominant. Jays and rodents preferred sound seeds over insect-infested seeds but such a preference was not found for nuthatches. This study highlights that pure beech stands showed a reduced guild of arboreal nut foragers in comparison to oak stands. This different guild could probably affect the spatial patterns of seed dispersal, with a proportionally higher number of long dispersal events for acorns (mostly jay-dispersed) than for beechnuts (mostly nuthatch-dispersed). Long-distance dispersal of beechnuts (by jays) is determined by the presence of other preferred species (oaks) and their frequency of non-mast years. Seed location in different habitats strongly determines the contribution of different arboreal removers (including climbing rodents) and their removal speed, leading to a differential seed fate that will eventually affect tree regeneration. As nuthatches are sedentary birds, it is important to maintain old and dead trees where they can breed (crevices), forage (arthropods) and store seeds in order to favor beechnut dispersal and gene flow. By maintaining or favoring oak trees within beech stands we will ensure a wider guild of arboreal nut dispersers.     

Intrapopulation variation in flowering phenology and fecundity of silver birch, implications for adaptability to changing climate

Earlier modeling has suggested that long distance gene flow is of importance in increasing the adaptability of tree populations in a changing climate. In times of warming temperatures, early flowering phenotypes may be favored, because early flowering may be connected to early onset of growth. Long term direct measurements of flowering, pollen cloud and intrapopulation fecundity variations are needed to test this hypothesis. Having one of the furthest transported types of pollen grains, birch may have good potential for long distance gene flow. Our daily observations over eight years, of 30 silver birches (Betula pendula Roth), showed that the onset and duration of flowering phenology was determined by the accumulation of spring temperature sums, but the window for potential long distance gene flow was narrow. The interannual phenology variations were large, the order of timing of female and male catkins in various trees of this monoecious species tended to differ, and the timing of generative and vegetative spring phenologies were not correlated at tree level. Early flowering trees tended to have a higher variation in germinability than later flowering trees. No other connection between timing of phenology of flowering and seed quantity or quality was found. Although stochastic climatic conditions produce interannually variable phenological windows, probability for long distance gene flow from areas differing in timing of spring temperature accumulations may be low in silver birch. Intertree variations in fecundity were high, and the majority of seeds and pollen were produced by only a few trees. Moreover, the amount of seeds produced was positively related to seed germinability, thus large phenotypic fecundity variations may decrease interannual genetic variations in seed sets.     

Liana habitat and host preferences in northern temperate forests

Lianas and other climbers are important ecological and structural components of forest communities. Like other plants, their abundance in a given habitat depends on a variety of factors, such as light, soil moisture and nutrients. However, since lianas require external support, host tree characteristics also influence their distribution. Lianas are conspicuous life forms in tropical regions, but in temperate areas, where they are less prominent, little is known about factors that control their distributions in these forests. We surveyed the climbing plant species in 20 mature (100 years and greater) forested habitats in the Midwest USA at a variety of levels from simple presence/absence, to ground layer abundances, to those species that had ascended trees. We also examined attributes of the tree species with climbers attached to them. Using cluster analysis, we distinguished five different tree communities in our survey locations. We determined that 25% of the trees we surveyed had one or more lianas attached to it, with Parthenocissus quinquefolia (Virginia creeper) the most common climbing species encountered. Canopy cover and soil attributes both influenced climber species presence/absence and ground layer climber abundance. The proportion of liana species of a given climbing type (roots, stem twiner, tendril climber) was significantly related to the DBH of the host tree, with more root climbers and fewer stem and tendril climbers on large trees. In general, the DBH of climbing lianas had a significant positive relationship to the DBH of the host tree; however this varied by the identity of the liana and the tree species. The greater the DBH of the host tree, the higher the probability that it was colonized by one or more lianas, with tree species such as Pinus banksiana (jack pine) and Quercus alba (white oak) being more susceptible to liana colonization than others. Finally, some liana species such as Celastrus scandens (American bittersweet) showed a preference for certain tree species (i.e., P. banksiana) as hosts. The information obtained about the relationship between the tree and climber community in this study provides insight into some of the factors that influence liana distributions in understudied temperate forest habitats and how lianas contribute to the structure of these mature forests. In addition, these data can provide a point of comparison to other liana communities in both temperate and tropical regions.     

Density-dependent selfing and its effects on seed performance in a tropical canopy tree species, Shorea acuminata (Dipterocarpaceae)

In the conservation and management practices of natural forests, sound reproduction and regeneration form the basis of the maintenance and viability of the tree populations. To obtain and serve biological information for sustainable forest management, we investigated reproductive biology and inbreeding depression in seeds of an important dipterocarp tree species, Shorea acuminata (Dipterocarpaceae), by both field and laboratory experiments. Results of parental analysis of immature and mature seeds showed that selfing rates varied greatly, from 7.6 to 88.4% among eight mother trees, and the mean overall selfing rate was 38.3%. Observed outcrossing events within a 40-ha study plot were predominantly (76.5%) short-distance events with a mating distance (md) ≤ 100 m. Since the selfing rate sharply decreased with increase in the number of flowering conspecifics (i.e., individuals of the same species) within a 100-m radius from the mother trees, the local density of flowering conspecifics appears to be the key factor determining the outcrossing rate in S. acuminata. However, the extremely high selfing rate (88.4%) observed for one tree could not be simply explained by the low local density of flowering conspecifics. Instead, differences in its flowering phenology (its flowering peaked ca. a week earlier than most of the other examined individuals) may have severely limited its receipt of pollen from other conspecifics, and thus promoted selfing. Since there were no significant differences in the proportion of selfed progeny between immature and mature seed stages, there was no evidence of selective abortion of selfed seeds during seed development. However, the seed mass of outcrossed progeny was heavier than that of selfed progeny, and heavier seeds showed higher success rates at germination and seedling establishment. These results suggest that inbreeding depression resulted in reductions in seed mass and may reduce the fitness of selfed seeds in S. acuminata. In addition, the outcrossing rate of S. acuminata was more sensitive to low local conspecific flowering-tree densities than that of a sympatric bee-pollinated dipterocarp species with greater pollination distances. These results suggest that the management of local adult-tree densities is important for avoiding selfing and inbreeding depression in future generations, especially in a species like S. acuminata with predominantly short-distance pollination.     

Effects of bamboo stands on seed rain and seed limitation in a rainforest

Bamboos often negatively affect tree recruitment, survival, and growth, leading to arrested tree regeneration in forested habitats. Studies so far have focused on the effects of bamboos on the performance of seedlings and saplings, but the influence of bamboos on forest dynamics may start very early in the forest regeneration process by altering seed rain patterns. We tested the prediction that the density and composition of the seed rain are altered and seed limitation is higher in stands of Guadua tagoara (B or bamboo stands), a large-sized woody bamboo native from the Brazilian Atlantic Forest, compared to forest patches without bamboos (NB or non-bamboo stands). Forty 1 m² seed traps were set in B and NB stands, and the seed rain was monitored monthly for 1 year. The seed rain was not greatly altered by the presence of bamboos: rarefied seed species richness was higher for B stands, patterns of dominance and density of seeds were similar between stands, and differences in overall composition were slight. Seed limitation, however, was greater at B stands, likely as a resulted of reduced tree density. Despite such reduced density, the presence of trees growing amidst and over the bamboos seems to play a key role in keeping the seeds falling in B stands because they serve as food sources for frugivores or simply as perches for them. The loss of such trees may lead to enhanced seed limitation, contributing ultimately to the self-perpetuating bamboo disturbance cycle.     

Enrichment planting as a silvicultural option in the eastern Amazon: Case study of Fazenda Cauaxi

Liana-dominated forest patches constitute 15–20% of old-growth forests in the Eastern Amazon but are generally excluded from management for timber production. Here we ask if liana-dominated patches may be brought into production by clearing lianas and conducting enrichment planting (EP) of native timber species. We present growth results from 8 years of such EP trials. Rapid growth and low mortality of all species in this study suggest that EP in cleared liana patches can contribute to timber stocks in second and third harvests of managed forests. The most vigorous individuals of Parkiagigantocarpa and Schizolobium amazonicum in each enrichment site grew more than 1 cm diameter per year (rates were initially >2 cm yr⁻¹), and attained dominant canopy positions and diameters equal to those of small canopy trees in the surrounding forest within 8 years of planting (mean dbh ∼18 cm and ∼20 cm, respectively, at year 8). Limited data on Ceiba pentandra plantings indicate a similar trajectory for this species (dbh ∼40 cm in 8 years). The most vigorous Swietenia macrophylla grew at least 1 cm per year in enrichment plots (mean dbh ∼10 cm in 8 years), but take longer to attain dominant positions. Tabebuia serratifolia may take much longer to reach the canopy than other species tested (rates <1 m yr⁻¹). We attribute the excellent performance to light availability; planting in intact soil with minimal compaction and abundant organic material; and low competition rates maintained by periodic thinning of competing vegetation.     

Recent infestation of forest stands by spruce beetles does not predict habitat use by little brown bats (Myotis lucifugus) in southwestern Yukon, Canada

Insect outbreaks affect forest structure which may have significant effects on the habitat of other animals. Forest-dwelling insectivorous bats are likely affected by associated changes in the abundance of roost trees and insect prey, altered foraging and flying efficiency, and predation risk. We examined the short-term effects (3-13 years post-infestation) of an outbreak of spruce beetles (Dendroctonus rufipennis) on the habitat use of little brown bats (Myotis lucifugus) in the boreal forest of the southwestern Yukon, Canada. We measured bat activity, using Anabat II bat detectors, in 90 forested stands that had experienced from 0 to 90% tree mortality due to spruce beetles. We used generalized linear models to assess whether bat activity varied with tree mortality, season, tree density, canopy closure, or distance to the nearest lake or town. Bat activity did not vary significantly with tree mortality, season, or canopy closure, but decreased with increasing tree density. Bat activity was significantly greater in areas close to both the nearest lake and nearest town, and was low in areas that were far from either. Our results indicate that in the short-term, habitat use by little brown bats was not related to the severity of spruce beetle infestation, but suggest that in the long-term, bats may be positively affected by decreased tree density as beetle-killed trees fall down.     

Restoration of dry tropical forests in Central America: A review of pattern and process

Much information on restoration and management exists for wet tropical forests of Central America but comparatively little work has been done in the dry forests of this region. Such information is critical for reforestation efforts that are now occurring throughout Central America. This paper describes processes of degradation due to land use and provides a conceptual framework for the restoration of dry tropical forest. Most of this forest type was initially harvested for timber and then cleared for cattle in the last century (1930-1970). Only 1.7% remains largely restricted to infertile soils and remote areas on the Pacific coastal side of Panama, Costa Rica, Nicaragua and Mexico. These cleared areas are again in a state of transition due to a combination of decreasing land productivity, and land speculation for tourism development. Some farms have been sold to new landowners who are interested in reforesting to increase biodiversity and forest cover. Attempts have therefore been made to reforest by protecting the land from fire and cattle, by supplementing natural regrowth with enrichment planting, or through use of tree plantations. Experimental studies have demonstrated the ability of these lands to grow back to forests because of native species ability to sprout after cutting, and the capacity of remnant trees in field and riparian zones to provide seeds and to moderate edge environment for seed germination and seedling establishment. However, research also shows that on sites with long histories of land clearance, species diversity will remain low with functional groups missing unless some active management occurs. Under-planting with late-successional native tree species can add structure and diversity; enrichment planting with large-fruited shade-intolerant species can initiate new islands of more diverse regeneration beneath their canopies; and plantings of fast-growing, nitrogen-fixing trees that provide light canopy shade can moderate the environment below, promoting regeneration establishment of late-successional species. Plantations are the only option for lands that have lost almost all remnants of native forest, and where soils and vegetation have changed to new states of structure and function. Conversion of pastures to tree plantations that can facilitate natural regeneration beneath them is appropriate when pastures are prone to fire and/or lack immediate seed sources nearby. After the grasses have been shaded out, natural recruitment can slowly occur over a 10-15 years period. Under-planting of shade-tolerant late-successional species can supplement species composition and structure.     

Effect of patch size and isolation on mating patterns and seed production in an urban population of Chinese pine (Pinus tabulaeformis Carr.)

Fragmentation is a critical issue for tree populations because the creation of small patches can reduce local population size and increase isolation, both of which can promote inbreeding and its negative consequences, as well as loss of genetic diversity. To test the hypothesis that patch size and isolation influence mating patterns or seed production in forest trees, we utilize the spatial array of trees of an urban population of Chinese Pine (Pinus tabulaeformis Carr.) planted in patches around Beijing. Our design includes 28 urban patches, with patch size ranging from 1 to 2000 adult trees and isolation (edge distance index) ranging from 37.5 m to 245.8 m. We examined the average number of seeds per cone and percentage of viable seed per cone for each patch as measures of seed production. By utilizing seven paternally inherited chloroplast microsatellite loci, we estimated the mating pattern parameters for each patch, including the level of selfing, the amount of immigrant pollen and the effective number of pollen sources (N_ep). Using a general linear model selection procedure based on AIC value, we found patch size was the best predictor of the selfing and immigration rate; smaller patches had a higher selfing and immigration rate. Small patches with one adult had relatively high N_ep which indicates connectivity among urban patches. However, due to the reduced amount of immigrant pollen and limited diversity of local pollen, intermediate sized patches (with 5–10 adults) had the lowest N_ep among the study patches. For patches with more than 10 adults, N_ep was increased with patch size. The percentage of viable seeds per cone significantly decreased with patch size, indicating a possible negative consequence of inbreeding. The effect of patch size on mating patterns and seed production suggests that the patches of trees experience less connectivity than trees within continuous forest. These findings indicate that forest management practices should emphasize the maintenance of an optimal patch size because, despite the fact that tree species show the potential for long distance pollen movement, the number of local trees strongly influences the mating patterns.     

Phenology and pollination of Manilkara zapota in forest and homegardens

Knowledge on reproductive phenology and pollination biology are basic elements that should be considered in the management and exploitation of plant species that offer non-timber products. The tropical tree Manilkara zapota is a species from which non-timber products have been obtained for centuries by Mayan communities in Mexico. Nevertheless, there are no quantitative reports on its reproductive biology and the factors that limit fruit production. The present study describes the reproductive phenology, breeding system and pollination of this species in two contrasting environments: medium-height, subdeciduous forest, and homegardens (“solares”) in a Mayan community in the state of Yucatan. Significant differences were found between environments both in the temporal distribution of flower and mature fruit production, as well as in the proportion of mature fruits. Homegarden trees showed the greatest fruit production, although flower production did not differ between environments. Mature fruits were of better quality (i.e., greater fresh weight) in homegardens. Hand pollination experiments showed that M. zapota is self-compatible, and that there is pollinator limitation for fruit production in trees that grow in homegardens. We propose that water and soil nutrients are the main factors limiting M. zapota fruit production in forests, while in homegardens the main factor appears to be pollinator availability.     

Seed dispersal and its implications for silviculture of African mahogany (Entandrophragma spp.) in undisturbed forest in the Central African Republic

The study of seed dispersal has received considerable attention in tropical ecology with emphasis placed on its role in the maintenance of species diversity. Since seeds of numerous high-value tropical timber trees are nominally or actively wind dispersed, the subject is also of interest to tropical foresters. In central Africa, up to 70% of harvested timber trees are wind dispersed, including African mahoganies of the genus Entandrophragma. The dual goals of managing many remote areas in central Africa for both biodiversity and timber production values has led to the consideration of low impact logging techniques. In the Central African Republic, increasing light downwind of Entandrophragma has been proposed as a way of increasing regeneration and recruitment of these trees. Mean wind direction at the time of dehiscence of Entandrophragma angolense and Entandrophragma cylindricum was compared with compass orientation of seed shadows for five individuals in mature, undisturbed forest. Mean wind direction was found to be toward the northeast for morning, afternoon, and daily winds and was significantly different from the orientation of seed shadows for all trees. Mean branch direction did not vary significantly from a random pattern, but matched the direction of the seed shadow better than wind direction did. It is concluded that harvesting interventions aimed at increasing light downwind of Entandrophragma will not necessarily benefit seedlings of the species at the study site and further silvicultural research is needed to determine appropriate management interventions, particularly in remote areas of Africa where conservation values are important.     

Intra and inter-annual variation in seed rain in a secondary dry tropical forest excluded from chronic disturbance

Domestic livestock grazing and firewood extraction are chronic and pervasive disturbances in dry secondary forest that reduce diversity and modify composition. Exclosure of such disturbances may allow the reestablishment of original diversity. To investigate potential vegetation recovery following 30 years of perturbation, seed rain was evaluated monthly during 3 years in 96 0.78 m² seed traps located in eight exclosures established in 2006 in a species-poor secondary dry forest in central Mexico. Twelve additional seed traps were located outside the exclosures in 2010 for comparison purposes. Baseline diversity of seed sources (trees, ?5 cm DBH) was recorded to compare standing tree composition with seed rain. Also, baseline diversity of potential primary (birds) and secondary (ants) dispersal vectors were registered to infer current plant-animal interactions. Total seed rain over 3 years within the exclosures comprised 180,375 seeds from 61 woody species, an average of 1.12 seeds/m²/month. Half of these species (31) and 95% of the seeds falling within the exclosures belonged to trees typical of perturbed sites. Sixty percent of all recorded seeds belonged to an early-successional, bird-dispersed shrub (Hamelia patens Jacq., Rubiaceae). Seed rain at one non-excluded plot was dominated by the wind-dispersed early-successional tree Heliocarpus sp. (Tiliaceae). Richness and seed density increased significantly in the second year of exclusion; however, by the third year, richness remained constant while seed density decreased. Seed rain was richer and more abundant during the dry season and was dominated by wind-dispersed species. Phenology, successional status and the dispersal modes of woody species explained most variance in richness and seed density of seed rain in early-successional dry forest. Seeds of early-successional species were dispersed in equal measure by abiotic and biotic vectors, whereas biotic vectors dispersed a higher density of late-successional seeds. The amount and timing of fall of abiotically dispersed seeds of the three most abundant species in the standing tree community may prolong their current domination of the landscape. However, even when richness in the seed rain within exclosures remained constant through time, composition was gradually shifting towards animal-dispersed species which may start a positive feedback producing an increase in animal visitation and a further increase of richness.     

Establishment of native tropical timber trees in monoculture and mixed-species plantations: Small-scale effects on tree performance and insect herbivory

Plantations of native timber species established on former pasture are a potential strategy to reduce the logging pressure on remnant natural forests in the tropics. Such plantations may help to mitigate or reverse the negative impacts of land degradation, and they may contribute to the long-term livelihood of livestock farmers. Planting native trees is, however, perceived as a risky activity due to limited knowledge of their performance and due to marked losses of newly established seedlings attributed to insect pests. Our study focuses on the small-scale effects of environmental heterogeneity, tree diversity and insecticide treatment on the performance of three native Central American timber species two years after establishment, and on damage inflicted by insect herbivores. Growth, survival and herbivore leaf damage were quantified for Anacardium excelsum (Anacardiaceae), Cedrela odorata (Meliaceae) and Tabebuia rosea (Bignoniaceae) planted in an experiment in Panama as (1) monocultures, (2) mixed stands, and (3) mixed stands protected by insecticides. Our study revealed that small-scale effects can have a substantial impact on the success of timber trees planted on former pasture. Growth performance and survival of the three species was strongly affected by small-scale environmental heterogeneity, which was expressed as significant differences in growth and survival among different plantation plots at the same study site. Establishment of trees in mixed stands did not have significant effects on tree survival and growth compared to pure stands, although it reduced herbivore pressure in one of the studied tree species. All tree species grew best and suffered lowest leaf damage when protected by insecticides, indicating a general influence of insect herbivory on growth of newly established trees. In contrast to growth performance, survival was not significantly affected by different management practices or herbivory. The large variability among plots in tree survival and growth, and also in the effects of management practices such as planting design and insecticide treatment, emphasizes the importance of small-scale environmental heterogeneity on tree survival and growth.     

Silvicultural treatments enhance growth rates of future crop trees in a tropical dry forest

Silvicultural treatments are often needed in selectively logged tropical forest to enhance the growth rates of many commercial tree species and, consequently, for recovering a larger proportion of the initial volume harvested over the next cutting cycle. The available data in the literature suggest, however, that the effect of silvicultural treatments on tree growth is smaller in dry forests than in humid forest tree species. In this study, we analyze the effect of logging and application of additional silvicultural treatments (liana cutting and girdling of competing trees) on the growth rates of future crop trees (FCTs; i.e., trees of current and potentially commercial timber species with adequate form and apparent growth potential). The study was carried out in a tropical dry forest in Bolivia where a set of 21.25-ha plots were monitored for 4 years post-logging. Plots received one of four treatments that varied in intensity of both logging and silvicultural treatments as follows: normal (reduced-impact) logging; normal logging and low-intensity silviculture; increased logging intensity and high-intensity silviculture; and, unlogged controls. The silvicultural treatments applied to FCTs involved liberation from lianas and overtopping trees. Results showed that rates of FCT stem diameter growth increased with light availability, logging intensity, and intensity of silvicultural treatments, and decrease with liana infestation degree. Growth rate increment was larger in the light and intensive silvicultural treatment (22–27%). Long-lived pioneer species showed the strongest response to intensive silviculture (50% increase) followed by total shade-tolerant species (24%) and partial shade-tolerant species (10%). While reduced-impact logging is often not sufficient to guarantee the sustainability of timber yields, application of silvicultural treatments that substantially enhanced the growth rates of FCTs will help move the management of these forests closer to the goal of sustained yield.     

Seed mass effects on performance of Pinus halepensis Mill. seedlings sown after fire

Pinus halepensis is a widely distributed species in the Mediterranean basin. It is generally well adapted to regenerate after wildfire, except when fire intervals are too short (≤15 years). In these latter situations, direct seeding could be a good alternative for restoring pine woodland after fire. Under dry climate, low seedling growth and survival can greatly limit the use of seeding. Early studies have shown that seedlings grown from large seeds have higher seedling establishment, growth and survival. Seed size grading may however reduce the genetic diversity of a seed lot by eliminating part or all of the families with relatively small seeds. An alternative to improve seed lot quality without losing genetic variability could be collecting and sieving seeds from each family separately. In order to explore the influence of seed mass on P. halepensis seedling performance, seeds from five half-sib families differing in mean seed mass were sown under greenhouse and field conditions. Final seedling emergence was unrelated to seed mass and half-sib family. The time of emergence was also unrelated to seed mass but it varied among families. Seed mass showed a positive effect on seedling height and diameter, both at population level and within family throughout the study period (9 months under greenhouse conditions and 20 months under field conditions). We also observed a negative relationship between seed mass and relative growth rate for seedling diameter, but it was not high enough to fully compensate the initial differences due to seed mass at the end of the 20-month study period. Seedling predation had a considerable impact on seedling survival, and it was not related to seed mass. When seedlings killed by predation were excluded from the survival analysis, larger seedlings, coming from larger seeds, showed slightly better survival, but only during the first growing period. The small advantages obtained from large seed mass in seedling development do not seem to justify the increased operational costs derived from seed mass selection for each family.     

Consequences of dry-season seed dispersal on seedling establishment of dry forest trees: Should we store seeds until the rains?

We examined the following hypotheses: (i) seeds of dry forest trees have high pre- and post-germination mortality by desiccation due to the time between seed dispersal and germination and to irregular rains at the onset of the rainy season; (ii) seedlings from seeds dispersed in the dry season which survive the dry spells are larger at the end of the first rainy season than those dispersed in the rainy season because the former have more time to grow. We evaluated the possible trade-off between few large seedlings (resulting from natural dispersal) × many small seedlings (resulting from delayed dispersal) on seedling survival during the dry season. We sowed eight tree species in a greenhouse in September, simulating the natural dispersal timing (before the rains), and in November, when rains are more constant. Because shading can counteract the effects of desiccation, we applied three levels of shade (10%, 40% and 72% of PPFD). From September 2005 to December 2006, we provided the daily precipitation of a median year from a major patch of dry forest in Central Brazil. At the end of the rainy season, a subset of seedlings was collected for growth measurements (dry mass) and the remainder was left to follow the dry season survivorship. The lower germination expected for seeds dispersed in the dry season and in full sun was not confirmed for species that had some dormancy. The delayed dispersal was advantageous for the initial establishment of fast germinating species, but it was irrelevant or even disadvantageous for others. Also, the greenhouse weather was certainly milder than the natural environment, reducing the potential for mortality by desiccation. The growth of the four species of higher dormancy were not affected by timing of seed dispersal, while three out of four fast germinating species had higher root biomass when dispersed in the dry season. The growth during the rainy season did not affect seedling survival during the dry season. Keeping seeds to sow when rain is constant might be a good strategy to increase the establishment of fast-germinating tree species.     

Seed biology and germination ecophysiology of Camellia nitidissima

Camellia nitidissima Chi (Theaceae) is a famous ornamental species with golden-yellow flowers. Due to deforestation and destructive collection of seedlings, its natural population size has been reduced greatly in recent decades. To provide basic information for its conservation, we studied the morphological and physiological characteristics of seeds and the environmental factors influencing germination. The seed production of C. nitidissima was of low yield, which caused lack of seed resource. The seeds were big and heavy, so they did not spread far from the mother trees and did not enter soil after falling down. Optimal temperatures for seed germination were 25–30 °C. The fresh-harvested seeds had high moisture content and weak desiccation-tolerance, so they tended to be recalcitrant. Soaking for 24 h in 10–30% PEG solution could significantly reduce the seed germination percentage. The imbibed seeds placed on the soil surface mostly could germinate, but some of them could not develop into seedlings because the embryo roots did not grow into soil. When the seeds were sowed at the depth of 1 cm, the seedling emergence reached over 70%, but then decreased with increased sowing depth, and reached about 50% when at the depth of 5 cm. The seeds sowed in sandy soil had higher emergence than those in clayed soil. It is feasible for seedling nursing to harvest seeds in time, harvesting prematurely or postmaturely may reduce seed quality.     

Effects of logging, liana tangles and pasture on seed fate of dry forest tree species in Central Brazil

We assessed seed fate of six tree species in a seasonally deciduous forest of Central Brazil. Seed germination, predation, removal and death caused by pathogens or dessication were followed in an undisturbed forest site, a logged forest site, and an active pasture. In both forest types we sowed seeds under patches of liana tangles (Low Forest habitat) and under patches of mature forest (High Forest habitat). Seeds were monitored weekly for up to 8 months. Overall, small-seeded species (Astronium fraxinifolium and Tabebuia impetiginosa) had high germination, although germination was lower in the pasture due to insect predation. The hard-coated seeds of Erythrina sp. had higher germination rates in the pasture. The large-seeded Cavanillesia arborea and Swartzia multijuga had high seed removal (ca. 90%) in the pasture and under High Forest, but only 23–34% under Low Forest in the logged site. Lower removal of large seeds in Low Forest suggests that this habitat conceals the seeds and constrains the movement of large mammals. Both species were favored in Low Forest, suggesting that caution should be used when managing lianas. When protected from predation, seeds with a soft coat and high water content, such as the seeds of Eugenia dysenterica and S. multijuga, died from desiccation in the pasture. Restoration efforts must take into consideration high mortality caused by desiccation or insect predation in abandoned pastures of dry forests.     

Intensive tree planting facilitates tropical forest biodiversity and biomass accumulation in Kibale National Park, Uganda

The extensive area of degraded tropical land and the calls to conserve forest biodiversity and sequester carbon to offset climate change demonstrate the need to restore forest in the tropics. Deforested land is sometimes replanted with fast-growing trees; however, the consequences of intensive replanting on biomass accumulation or plant and animal diversity are poorly understood. The purpose of this study was to determine how intensive replanting affected tropical forest regeneration and biomass accumulation over ten years. We studied reforested sites in Kibale National Park, Uganda, that were degraded in the 1970s and replanted with five native tree species in 1995. We identified and measured the size of planted versus naturally regenerating trees, and felled and weighed matched trees outside the park to calculate region-specific allometric equations for above-ground tree biomass. The role of shrubs and grasses in facilitating or hindering the establishment of trees was evaluated by correlating observed estimates of percent cover to tree biomass. We found 39 tree species naturally regenerating in the restored area in addition to the five originally planted species. Biomass was much higher for planted (15,675 kg/ha) than naturally regenerated trees (4560 kg/ha), but naturally regenerating tree regrowth was an important element of the landscape. The establishment of tree seedlings initially appeared to be facilitated by shrubs, primarily Acanthus pubescens and the invasive Lantana camara; however, both are expected to hinder tree recruitment in the long-term. Large and small-seeded tree species were found in the replanted area, indicating that bird and mammal dispersers contributed to natural forest restoration. These results demonstrate that intensive replanting can accelerate the natural accumulation of biomass and biodiversity and facilitate the restoration of tropical forest communities. However, the long-term financial costs and ecological benefits of planting and maintaining reforested areas need to be weighed against other potential restoration strategies.