Soil seed burial and competition with surrounding plants determine the emergence and development of seedling of an endangered species Horsfieldia hainanensis Merr. in China.

Three well-conserved Horsfieldia hainanensis Merr. populations were used to investigate their soil seed bank and seedling regeneration characteristics and their relationship to environmental factors. The results showed that the seed reserves were low in the H. hainanensis soil seed bank (16.93~24.74 seed/m2). The distribution pattern for the seeds and seedlings in the H. hainanensis populations was aggregated, and they were mainly found around 2-3 m from the mother plant. The seeds in the litter layer and the 5-10 cm soil layer showed no vigor, and only 25.7%~33.3% of the total seeds in the 0-5 cm soil layer were viable affected by the high temperature and humidity, the animals' eating and poisoning. Affected by the height and coverage of the surrounding herbaceous layer and shrub layer, the seedlings of H. hainanensis could not obtain enough light and nutrients in the competition, resulting in the survival competitiveness of 1- to 3-year-old (1-3a) seedlings in the habitat had been in a weak position and a large number of seedlings died. It would take at least four years for seedlings to develop under the current environmental constraints. It can be concluded that the low seed reserve in the soil seed bank and high mortality of seedlings of H. hainanensis lead to slow or even stagnation of population regeneration, which was an important reason for the endangered of H. hainanensis. Therefore, the next research focus is to explore the influence mechanism of environmental factors on seed germination and seedling growth of H. hainanensis.

Evidence of within-species specialization by soil microbes and the implications for plant community diversity.

Microbes are thought to maintain diversity in plant communities by specializing on particular species, but it is not known whether microbes that specialize within species (i.e., on genotypes) affect diversity or dynamics in plant communities. Here we show that soil microbes can specialize at the within-population level in a wild plant species, and that such specialization could promote species diversity and seed dispersal in plant communities. In a shadehouse experiment in Panama, we found that seedlings of the native tree species, Virola surinamensis (Myristicaceae), had reduced performance in the soil microbial community of their maternal tree compared with in the soil microbial community of a nonmaternal tree from the same population. Performance differences were unrelated to soil nutrients or to colonization by mycorrhizal fungi, suggesting that highly specialized pathogens were the mechanism reducing seedling performance in maternal soils. We then constructed a simulation model to explore the ecological and evolutionary consequences of genotype-specific pathogens in multispecies plant communities. Model results indicated that genotype-specific pathogens promote plant species coexistence-albeit less strongly than species-specific pathogens-and are most effective at maintaining species richness when genetic diversity is relatively low. Simulations also revealed that genotype-specific pathogens select for increased seed dispersal relative to species-specific pathogens, potentially helping to create seed dispersal landscapes that allow pathogens to more effectively promote diversity. Combined, our results reveal that soil microbes can specialize within wild plant populations, affecting seedling performance near conspecific adults and influencing plant community dynamics on ecological and evolutionary time scales.

Female fruit production depends on female flower production and crown size rather than male density in a continuous population of a tropical dioecious tree (Virola surinamensis).

PREMISE OF THE STUDY: Factors related to pollen and resource limitation were evaluated to predict female fruit production in a tropical dioecious tree. Pollen limitation via variation in the male density at local scales is expected to limit female reproduction success in dioecious plants. METHODOLOGY: We modeled the roles of local male density, female crown size, crown illumination, and female flower production on female fruit initiation and mature fruit production in a continuous population (62 ha plot) of a tropical dioecious tree (Virola surinamensis). In addition, we used microsatellites to describe the scale of effective pollen flow, the male effective population size, and the spatial genetic structure within/between progenies and males. KEY RESULTS: The local male density was not related to female fruit initiation or mature fruit production. Female floral production had a positive effect on fruit initiation. The female crown size was positively related to fruit maturation. Seeds from the same female and seeds from different but spatially proximal females were generally half-siblings; however, proximal females showed greater variation. Proximal male-female adult pairs were not significantly more genetically related than distant pairs. The probability of paternity was negatively affected by the distance between seeds and males; most effective pollen dispersal events (∼85%) occurred from males located less than 150 m from females. The number of males siring progenies was greater than the number of males found at local scales. CONCLUSIONS: Female fecundity in this continuous population of Virola surinamensis is not limited by the availability of pollen from proximal males. Rather, resource allocation to floral production may ultimately determine female reproductive success.

Photosynthesis of seedlings of Otoba novogranatensis (Myristicaceae) and Ruagea glabra (Meliaceae) in abandoned pasture, secondary forest and plantation habitats in Costa Rica.

Enrichment planting in naturally recovering secondary forests or in tree plantations is increasingly being used as strategy to restore later-successional, large-seeded tropical forest trees. We seeded two tree species (Otoba novogranatensis and Ruagea glabra) in three agricultural sites in Southern Costa Rica: abandoned pastures, eight to ten year old secondary forests and three year old tree plantations (containing two N-fixing of four total tree species). We measured micrometeorological conditions, soil water content, plant water potential, leaf area, foliar C and N, and photosynthesis to better understand mechanistic responses of seedlings to conditions in the different successional habitats. Micrometeorological conditions, soil water content, and plant water potential were generally similar across habitats. Certain aspects of leaves (such as Specific Leaf Area and foliar N content), and photosynthesis (e.g. quantum yield and electron transport rate) were highest in the plantations, intermediate in the secondary forests, and lowest in abandoned pastures. Enhanced rates of photosynthetic biochemistry (such as Vxmax and Jmax) and Photosystem II efficiency (e.g. thermal energy dissipation) occurred in leaves from the plantations compared to the abandoned pastures, which may be related to higher leaf %N content. Results suggest that foliar N may be of greater importance than soil water content and micrometeorological factors in driving differences in photosynthetic processes across planting habitats. Planting seeds of these two species in plantations containing three year old trees (including two N-fixing species) enhances certain aspects of their photosynthesis and growth, compared to seedlings in abandoned pastures with non-native grasses, and thus can help increase forest recovery on abandoned agricultural lands.

Photosynthesis of seedlings of Otoba novogranatensis (Myristicaceae) and Ruagea glabra (Meliaceae) in abandoned pasture, secondary forest and plantation habitats in Costa Rica / Fotosíntesis de plántulas de Otoba novogranatensis (Myristicaceae) y Ruagea glabra (Meliaceae) en pastizales abandonados, bosques secundarios y hábitats de plantaciones en Costa Rica

Enrichment planting in naturally recovering secondary forests or in tree plantations is increasingly being used as strategy to restore later-successional, large-seeded tropical forest trees. We seeded two tree species (Otoba novogranatensis and Ruagea glabra) in three agricultural sites in Southern Costa Rica: abandoned pastures, eight to ten year old secondary forests and three year old tree plantations (containing two N-fixing of four total tree species). We measured micrometeorological conditions, soil water content, plant water potential, leaf area, foliar C and N, and photosynthesis to better understand mechanistic responses of seedlings to conditions in the different successional habitats. Micrometeorological conditions, soil water content, and plant water potential were generally similar across habitats. Certain aspects of leaves (such as Specific Leaf Area and foliar N content), and photosynthesis (e.g. quantum yield and electron transport rate) were highest in the plantations, intermediate in the secondary forests, and lowest in abandoned pastures. Enhanced rates of photosynthetic biochemistry (such as Vcmax and Jmax) and Photosystem II efficiency (e.g. thermal energy dissipation) occurred in leaves from the plantations compared to the abandoned pastures, which may be related to higher leaf %N content. Results suggest that foliar N may be of greater importance than soil water content and micrometeorological factors in driving differences in photosynthetic processes across planting habitats. Planting seeds of these two species in plantations containing three year old trees (including two N-fixing species) enhances certain aspects of their photosynthesis and growth, compared to seedlings in abandoned pastures with non-native grasses, and thus can help increase forest recovery on abandoned agricultural lands.

El enriquecimiento de bosques secundarios o plantaciones forestales en proceso de regeneración natural por medio de la siembra de plántulas es una práctica cada vez más utilizada para restaurar bosques tropicales en estado de sucesión tardía. Sembramos dos especies de árboles (Otoba novogranatensis y Ruagea glabra) en pastizales abandonados, bosques secundarios de ocho a diez años de edad y plantaciones forestales de tres años de edad (con dos especies fijadoras de Nitrógeno de un total de cuatro especies) en tres sitios agrícolas en el Sur de Costa Rica. Medimos condiciones micrometeorológicas, contenido de agua del suelo, potencial hídrico de las plantas, área foliar, C y N foliar, y fotosíntesis para entender de una mejor manera las respuestas funcionales de las plántulas ante condiciones de distintos estadíos sucesionales. Las condiciones micrometeorológicas, contenido hídrico del suelo y el potencial hídrico de las plantas fueron mayoritariamente similares entre hábitats. Algunos aspectos de las hojas (como Área Foliar Específica y contenido de N foliar) y fotosíntesis (ej.: rendimiento cuántico y tasa de transporte de electrones) presentaron valores mayores en las plantaciones, intermedios en los bosques secundarios y menores en los pastizales abandonados. Se obtuvo un aumento en las tasas fotosintéticas bioquímicas (como Vcmax, Jmax) y la eficiencia del Fotosistema II (ej.: disipación de energía térmica) en hojas provenientes de las plantaciones comparado a las de los pastizales, posiblemente relacionado a un mayor %N foliar. Los resultados sugieren que el N foliar puede ser más importante que el contenido de agua del suelo y que los factores micrometeorológicos para marcar diferencias en los procesos fotosintéticos entre hábitats. Las plántulas de estas dos especies en las plantaciones con árboles de tres años de edad (incluyendo dos fijadoras de N) incrementaron ciertos aspectos de su fotosíntesis y crecimiento comparado a las plántulas en los pastizales abandonados de especies exóticas, por lo tanto, esta práctica puede ayudar a incrementar la recuperación de los bosques en áreas agrícolas abandonadas.

Palms, peccaries and perturbations: widespread effects of small-scale disturbance in tropical forests.

BACKGROUND: Disturbance is an important process structuring ecosystems worldwide and has long been thought to be a significant driver of diversity and dynamics. In forests, most studies of disturbance have focused on large-scale disturbance such as hurricanes or tree-falls. However, smaller sub-canopy disturbances could also have significant impacts on community structure. One such sub-canopy disturbance in tropical forests is abscising leaves of large arborescent palm (Arececeae) trees. These leaves can weigh up to 15 kg and cause physical damage and mortality to juvenile plants. Previous studies examining this question suffered from the use of static data at small spatial scales. Here we use data from a large permanent forest plot combined with dynamic data on the survival and growth of > 66,000 individuals over a seven-year period to address whether falling palm fronds do impact neighboring seedling and sapling communities, or whether there is an interaction between the palms and peccaries rooting for fallen palm fruit in the same area as falling leaves. We tested the wider generalisation of these hypotheses by comparing seedling and sapling survival under fruiting and non-fruiting trees in another family, the Myristicaceae. RESULTS: We found a spatially-restricted but significant effect of large arborescent fruiting palms on the spatial structure, population dynamics and species diversity of neighbouring sapling and seedling communities. However, these effects were not found around slightly smaller non-fruiting palm trees, suggesting it is seed predators such as peccaries rather than falling leaves that impact on the communities around palm trees. Conversely, this hypothesis was not supported in data from other edible species, such as those in the family Myristicaceae. CONCLUSIONS: Given the abundance of arborescent palm trees in Amazonian forests, it is reasonable to conclude that their presence does have a significant, if spatially-restricted, impact on juvenile plants, most likely on the survival and growth of seedlings and saplings damaged by foraging peccaries. Given the abundance of fruit produced by each palm, the widespread effects of these small-scale disturbances appear, over long time-scales, to cause directional changes in community structure at larger scales.

Taxonomic scale-dependence of habitat niche partitioning and biotic neighbourhood on survival of tropical tree seedlings.

In order to differentiate between mechanisms of species coexistence, we examined the relative importance of local biotic neighbourhood, abiotic habitat factors and species differences as factors influencing the survival of 2330 spatially mapped tropical tree seedlings of 15 species of Myristicaceae in two separate analyses in which individuals were identified first to species and then to genus. Using likelihood methods, we selected the most parsimonious candidate models as predictors of 3 year seedling survival in both sets of analyses. We found evidence for differential effects of abiotic niche and neighbourhood processes on individual survival between analyses at the genus and species levels. Niche partitioning (defined as an interaction of taxonomic identity and abiotic neighbourhood) was significant in analyses at the genus level, but did not differentiate among species in models of individual seedling survival. By contrast, conspecific and congeneric seedling and adult density were retained in the minimum adequate models of seedling survival at species and genus levels, respectively. We conclude that abiotic niche effects express differences in seedling survival among genera but not among species, and that, within genera, community and/or local variation in adult and seedling abundance drives variation in seedling survival. These data suggest that different mechanisms of coexistence among tropical tree taxa may function at different taxonomic or phylogenetic scales. This perspective helps to reconcile perceived differences of importance in the various non-mutually exclusive mechanisms of species coexistence in hyper-diverse tropical forests.

Neighborhood and community interactions determine the spatial pattern of tropical tree seedling survival.

Factors affecting survival and recruitment of 3531 individually mapped seedlings of Myristicaceae were examined over three years in a highly diverse neotropical rain forest, at spatial scales of 1-9 m and 25 ha. We found convincing evidence of a community compensatory trend (CCT) in seedling survival (i.e., more abundant species had higher seedling mortality at the 25-ha scale), which suggests that density-dependent mortality may contribute to the spatial dynamics of seedling recruitment. Unlike previous studies, we demonstrate that the CCT was not caused by differences in microhabitat preferences or life history strategy among the study species. In local neighborhood analyses, the spatial autocorrelation of seedling survival was important at small spatial scales (1-5 m) but decayed rapidly with increasing distance. Relative seedling height had the greatest effect on seedling survival. Conspecific seedling density had a more negative effect on survival than heterospecific seedling density and was stronger and extended farther in rare species than in common species. Taken together, the CCT and neighborhood analyses suggest that seedling mortality is coupled more strongly to the landscape-scale abundance of conspecific large trees in common species and the local density of conspecific seedlings in rare species. We conclude that negative density dependence could promote species coexistence in this rain forest community but that the scale dependence of interactions differs between rare and common species.

Relative importance of photosynthetic physiology and biomass allocation for tree seedling growth across a broad light gradient.

Studies of tree seedling physiology and growth under field conditions provide information on the mechanisms underlying inter- and intraspecific differences in growth and survival at a critical period during forest regeneration. I compared photosynthetic physiology, growth and biomass allocation in seedlings of three shade-tolerant tree species, Virola koschynii Warb., Dipteryx panamensis (Pittier) Record & Mell and Brosimum alicastrum Swartz., growing across a light gradient created by a forest-pasture edge (0.5 to 67% diffuse transmittance (%T)). Most growth and physiological traits showed nonlinear responses to light availability, with the greatest changes occurring between 0.5 and 20 %T. Specific leaf area (SLA) and nitrogen per unit leaf mass (N mass) decreased, maximum assimilation per unit leaf area (A area) and area-based leaf N concentration (N area) increased, and maximum assimilation per unit leaf mass (A mass) did not change with increasing irradiance. Plastic responses in SLA were important determinants of leaf N and A area across the gradient. Species differed in magnitude and plasticity of growth; B. alicastrum had the lowest relative growth rates (RGR) and low plasticity. Its final biomass varied only 10-fold across the light gradient. In contrast, the final biomass of D. panamensis and V. koschynii varied by 100- and 50-fold, respectively, and both had higher RGR than B. alicastrum. As light availability increased, all species decreased biomass allocation to leaf tissue (mass and area) and showed a trade-off between allocation to leaf area at a given plant mass (LAR) and net gain in mass per unit leaf area (net assimilation rate, NAR). This trade-off largely reflected declines in SLA with increasing light. Finally, A area was correlated with NAR and both were major determinants of intraspecific variation in RGR. These data indicate the importance of plasticity in photosynthetic physiology and allocation for variation in tree seedling growth among habitats that vary in light availability.