Local-scale drivers of tree survival in a temperate forest

Tree survival plays a central role in forest ecosystems. Although many factors such as tree size, abiotic and biotic neighborhoods have been proposed as being important in explaining patterns of tree survival, their contributions are still subject to debate. We used generalized linear mixed models to examine the relative importance of tree size, local abiotic conditions and the density and identity of neighbors on tree survival in an old-growth temperate forest in northeastern China at three levels (community, guild and species). Tree size and both abiotic and biotic neighborhood variables influenced tree survival under current forest conditions, but their relative importance varied dramatically within and among the community, guild and species levels. Of the variables tested, tree size was typically the most important predictor of tree survival, followed by biotic and then abiotic variables. The effect of tree size on survival varied from strongly positive for small trees (1-20 cm dbh) and medium trees (20-40 cm dbh), to slightly negative for large trees (>40 cm dbh). Among the biotic factors, we found strong evidence for negative density and frequency dependence in this temperate forest, as indicated by negative effects of both total basal area of neighbors and the frequency of conspecific neighbors. Among the abiotic factors tested, soil nutrients tended to be more important in affecting tree survival than topographic variables. Abiotic factors generally influenced survival for species with relatively high abundance, for individuals in smaller size classes and for shade-tolerant species. Our study demonstrates that the relative importance of variables driving patterns of tree survival differs greatly among size classes, species guilds and abundance classes in temperate forest, which can further understanding of forest dynamics and offer important insights into forest management.     

长白山次生针阔混交林乔木幼苗存活的影响因素分析

基于长白山次生针阔混交林样地, 以520个1 m × 1 m幼苗样方中胸径小于1 cm的乔木幼苗为研究对象, 选取2013年和2014年的幼苗调查数据, 运用广义线性混合模型(GLMM)分析了生物邻体和生境异质性对幼苗存活的影响, 探讨了次生针阔混交林幼苗存活影响因素及物种共存机制。结果表明: (1)适宜大树生长的局域生境同样也适宜幼苗的存活, 幼苗存活率与土壤含水量和有效氮等土壤养分显著正相关。(2)幼苗个体之间存在明显的竞争, 较多的幼苗邻体显著降低幼苗的存活率。同种大树邻体和同种幼苗邻体与幼苗存活显著负相关, 表明存在负密度制约效应。(3)随着幼苗年龄的增加, 生境异质性对幼苗存活的影响逐渐增大。该研究证实了密度制约效应和生境异质性对幼苗存活有着重要影响, 其相对重要性随着幼苗年龄级、功能群以及物种种类而变化。     

Effects of biotic neighbors and habitat heterogeneity on tree seedling survival in a secondary mixed conifer and broad-leaved forest in Changbai Mountain

Aims Our study aimed to understand the effects and the relative importance of biotic neighborhood and habitat heterogeneity for tree seedling survival in a secondary mixed conifer and broad-leaved forest in Changbai Mountain, north-eastern China.
Methods The generalized mixed linear model was used to examine the relative effects of biotic neighborhood and habitat heterogeneity on seedling survival over two years.
Important findings Our results showed that both biotic neighborhood and habitat heterogeneity had significant effects on the seedling survival at community level. The local environment suitable for the adult growth was also suitable for seedling survival. The soil moisture and soil available nitrogen exhibited significant positive effects on seedling survival. On the other hand, seedling density had significant negative effects on seedling survival due to the individual competition. Particularly, we found significant negative density-dependent effects on seedling survival which was caused by conspecific adult and seedling neighbors. As expected, with the increasing of seedling survival age, the habitat heterogeneity became more important on seedling survival. These results suggest that both local biotic neighborhood and habitat heterogeneity drive seedling survival in this temperate forest, and their relative importance varies with different seedling age classes and species traits.     

Effects of local biotic neighbors and habitat heterogeneity on tree and shrub seedling survival in an old-growth temperate forest

Seedling dynamics play a crucial role in determining species distributions and coexistence. Exploring causes of variation in seedling dynamics can therefore provide key insights into the factors affecting these phenomena. We examined the relative importance of biotic neighborhood processes and habitat heterogeneity using survival data for 5,827 seedlings in 39 tree and shrub species over 2?years from an old-growth temperate forest in northeastern China. We found significant negative density-dependence effects on survival of tree seedlings, and limited effects of habitat heterogeneity (edaphic and topographic variables) on survival of shrub seedlings. The importance of negative density dependence on young tree seedling survival was replaced by habitat in tree seedlings ??4?years old. As expected, negative density dependence was more apparent in gravity-dispersed species compared to wind-dispersed and animal-dispersed species. Moreover, we found that a community compensatory trend existed for trees. Therefore, although negative density dependence was not as pervasive as in other forest communities, it is an important mechanism for the maintenance of community diversity in this temperate forest. We conclude that both negative density dependence and habitat heterogeneity drive seedling survival, but their relative importance varies with seedling age classes and species traits.     

Seedling interactions in a tropical forest in Panama

Competition is believed to be a central force limiting local diversity and controlling the structure of plant communities. However, it has been proposed that the stressed understory environment limits total understory plant density to such low levels that competitive exclusion cannot be an important factor limiting the local diversity of understory plants. To evaluate the importance of inter-seedling competition, we performed a seedling competition experiment with five shade-tolerant species in a tropical moist forest in Panama. Three-month-old seedlings were transplanted into the forest singly or with their roots intertwined with a single conspecific or heterospecific seedling in all pairwise species combinations. If competition is important, performance (survival, stem height, and number of leaves after one and six years) would be expected to be lowest with a conspecific neighbor and greatest without a neighbor. The experiment was replicated in five 0.24-m² plots at each of 20 sites in tall secondary forest. To test whether seedling performance differed among treatments we fitted linear mixed models (LMM) and generalized linear mixed models (GLMM), treating species identity and microsite (site and plot) as random effects. The five shade-tolerant study species all experienced good establishment with relatively high survival and growth rates. The neighbor treatment consistently affected seedling performance, but the effect was always very small, both in absolute terms and relative to the much stronger species and microsite effects. Seedlings with a conspecific neighbor consistently performed worse than seedlings with a heterospecific neighbor, but having no neighbor generally did not cause superior performance relative to the other treatments. We conclude that direct competitive interactions are relatively unimportant among understory plants in humid tropical forests.     

Determinants of tree survival at local scale in a sub-tropical forest

Tree survival is a critical driver of stand dynamics, influencing forest structure and composition. Many local-scale drivers (tree size, abiotic and biotic factors) have been proposed as being important in explaining patterns of tree survival, but their contributions are still unknown. We examined the relative importance of these local drivers on tree survival using generalized linear mixed models in an old-growth sub-tropical forest in south China at three levels (community, guild, and species). Among the variables tested, tree size was typically the most important driver of tree survival, followed by abiotic and then biotic variables. Tree size has a strongly positive effect on tree survival for small trees (10–30 cm dbh) and shade-tolerant tree species. Of the abiotic factors tested, elevation tended to be more important in affecting tree survival than other topographic variables. Abiotic factors generally influenced survival of species with relatively high abundances, for individuals in smaller size classes and for mid-tolerant species. Among biotic factors, we found that the mortality of tree species was not driven by density- and frequency-dependent effects in this sub-tropical forest, as indicated by the results of both total basal area of neighbors and the proportion of conspecific neighbors in our study. We conclude that the relative importance of variables driving patterns of tree survival varied greatly among tree size classes, species guilds, shade tolerance, density, and abundance classes in this sub-tropical forest. These results also provide critical information for future studies of forest dynamics and offer insight into forest management in this region.     

Drivers of seedling survival in a temperate forest and their relative importance at three stages of succession

Negative density dependence (NDD) and niche partitioning have been perceived as important mechanisms for the maintenance of species diversity. However, little is known about their relative contributions to seedling survival. We examined the effects of biotic and abiotic neighborhoods and the variations of biotic neighborhoods among species using survival data for 7503 seedlings belonging to 22 woody species over a period of 2 years in three different forest types, a half‐mature forest (HF), a mature forest (MF), and an old‐growth forest (OGF), each of these representing a specific successional stage in a temperate forest ecosystem in northeastern China. We found a convincing evidence for the existence of NDD in temperate forest ecosystems. The biotic and abiotic variables affecting seedlings survival change with successional stage, seedling size, and age. The strength of NDD for the smaller (<20 cm in height) and younger seedlings (1–2 years) as well as all seedlings combined varies significantly among species. We found no evidence that a community compensatory trend (CCT) existed in our study area. The results of this study demonstrate that the relative importance of NDD and habitat niche partitioning in driving seedling survival varies with seedling size and age and that the biotic and abiotic factors affecting seedlings survival change with successional stage.     

Density dependence and habitat preference shape seedling survival in a subtropical forest in central China

Aims Seedlings are vulnerable to many kinds of fatal abiotic and biotic agents, and examining the causes of seedling dynamics can help understand mechanisms of species coexistence. To disentangle the relative importance of neighborhood densities, habitat factors and phylogenetic relatedness on focal seedling survival, we monitored the survival of 5306 seedlings of 104 species>15 months. We address the following questions: (i) How do neighborhood densities, habitat variables and phylogenetic relatedness affect seedling survival? What is the relative importance of conspecific densities, habitat variables and phylogenetic relatedness to seedling survival? (ii) Does the importance of the neighborhood densities, habitat variables and phylogenetic relatedness vary among growth forms, leaf habits or dispersal modes? Specially, does the conspecific negative density dependence inhibit tree and deciduous seedlings more compared with shrub and evergreen species? Does density dependence affect the wind and animal-dispersed species equally?Methods We established 135 census stations to monitor seedling dynamics in a 25-ha subtropical forest plot in central China. Conspecific and heterospecific seedling density in the 1-m 2 seedling plot and adult basal area within a 20-m radius provided neighborhood density variables. Mean elevation, convexity and aspect of every 5- × 5-m grid with seedling plots were used to quantify habitat characteristics. We calculated the relative average phylodiversity between focal seedling and heterospecific neighbors to quantify the species relatedness in the neighborhood. Eight candidate generalized linear mixed models with binominal error distribution were used to compare the relative importance of these variables to seedling survival. Akaike's information criteria were used to identify the most parsimonious models.Important findings At the community level, both the neighborhood densities and phylogenetic relatedness were important to seedling survival. We found negative effects of increasing conspecific seedlings, which suggested the existence of species-specific density-dependent mortality. Phylodiversity of heterospecific neighbors was negatively related to survival of focal seedlings, indicating similar habitat preference shared among phylogenetically closely related species may drive seedling survival. The relative importance of neighborhood densities, habitat variables and phylogenetic relatedness varied among ecological guilds. Conspecific densities had significant negative effect for deciduous and wind-dispersed species, and marginally significant for tree seedlings>10cm tall and animal-dispersed species. Habitat variables had limited effects on seedling survival, and only elevation was related to the survival of evergreen species in the best-fit model. We conclude that both negative density-dependent mortality and habitat preference reflected by the phylogenetic relatedness shape the species coexistence at seedling stage in this forest.     

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.     

Density‐dependent Survival in Seedlings Differs among Woody Life‐forms in Tropical Wet Forests of a Caribbean Island

Negative density dependence contributes to seedling dynamics in forested ecosystems, but the relative importance of this factor for different woody plant life‐forms is not well‐understood. We used 1 yr of seedling survivorship data for woody seedlings in 17 different plots of lower to mid‐montane rain forests on the island of Dominica to examine how seedling height, abiotic factors, and biotic factors such as negative density dependence are related to seedling survival of five different life‐forms (canopy, midstory, and understory trees; shrubs; and lianas). Across 64 species, taller seedlings in seedling plots with higher canopy openness, greater seedling density, lower relative abundance of conspecific seedlings, and lower relative abundance of conspecific adults generally had a greater probability of surviving. Height was the strongest predictor of seedling survival for all life‐forms except lianas. Greater seedling density was positively related to survival for canopy and midstory trees but negatively related to survival for the other life‐forms. For trees, the relative abundance of conspecific seedling and adult neighbors had weak and strong negative effects on survival respectively. Neither shrub nor liana seedling survival was affected by the relative abundance of conspecific neighbors. Thus, negative density dependence is confirmed as an important structuring mechanism for tree seedling communities but does not seem to be important for lianas and shrubs in Dominican rain forests. These results represent the first direct assessment of controls on seedling survival of all woody life‐forms – an important step in understanding the dynamics and structure of the entire woody plant community.     

Environmental heterogeneity and biotic interactions mediate climate impacts on tropical forest regeneration

Predicting the fate of tropical forests under a changing climate requires understanding species responses to climatic variability and extremes. Seedlings may be particularly vulnerable to climatic stress given low stored resources and undeveloped roots; they also portend the potential effects of climate change on future forest composition. Here we use data for ca. 50,000 tropical seedlings representing 25 woody species to assess (i) the effects of interannual variation in rainfall and solar radiation between 2007 and 2016 on seedling survival over 9 years in a subtropical forest; and (ii) how spatial heterogeneity in three environmental factors—soil moisture, understory light, and conspecific neighborhood density—modulate these responses. Community‐wide seedling survival was not sensitive to interannual rainfall variability but interspecific variation in these responses was large, overwhelming the average community response. In contrast, community‐wide responses to solar radiation were predominantly positive. Spatial heterogeneity in soil moisture and conspecific density were the predominant and most consistent drivers of seedling survival, with the majority of species exhibiting greater survival at low conspecific densities and positive or nonlinear responses to soil moisture. This environmental heterogeneity modulated impacts of rainfall and solar radiation. Negative conspecific effects were amplified during rainy years and at dry sites, whereas the positive effects of radiation on survival were more pronounced for seedlings existing at high understory light levels. These results demonstrate that environmental heterogeneity is not only the main driver of seedling survival in this forest but also plays a central role in buffering or exacerbating impacts of climate fluctuations on forest regeneration. Since seedlings represent a key bottleneck in the demographic cycle of trees, efforts to predict the long‐term effects of a changing climate on tropical forests must take into account this environmental heterogeneity and how its effects on regeneration dynamics play out in long‐term stand dynamics.     

Do differences in understory light contribute to species distributions along a tropical rainfall gradient?

In tropical forests, regional differences in annual rainfall correlate with differences in plant species composition. Although water availability is clearly one factor determining species distribution, other environmental variables that covary with rainfall may contribute to distributions. One such variable is light availability in the understory, which decreases towards wetter forests due to differences in canopy density and phenology. We established common garden experiments in three sites along a rainfall gradient across the Isthmus of Panama in order to measure the differences in understory light availability, and to evaluate their influence on the performance of 24 shade-tolerant species with contrasting distributions. Within sites, the effect of understory light availability on species performance depended strongly on water availability. When water was not limiting, either naturally in the wetter site or through water supplementation in drier sites, seedling performance improved at higher light. In contrast, when water was limiting at the drier sites, seedling performance was reduced at higher light, presumably due to an increase in water stress that affected mostly wet-distribution species. Although wetter forest understories were on average darker, wet-distribution species were not more shade-tolerant than dry-distribution species. Instead, wet-distribution species had higher absolute growth rates and, when water was not limiting, were better able to take advantage of small increases in light than dry-distribution species. Our results suggest that in wet forests the ability to grow fast during temporary increases in light may be a key trait for successful recruitment. The slower growth rates of the dry-distribution species, possibly due to trade-offs associated with greater drought tolerance, may exclude these species from wetter forests.     

海南岛热带低地雨林刀耕火种弃耕地自然恢复过程中的群落构建

研究群落构建机制是群落生态学的一个重要目标, 群落动态过程中的构建规律对于了解群落演替机理有重要的作用。该文以海南岛刀耕火种干扰后自然恢复的10 hm ²热带低地雨林为研究对象, 通过比较不同恢复阶段的次生林(15年、30年和60年)和老龄林在幼苗、幼树和成年树群落的物种组成, 揭示次生演替过程中的群落构建规律。研究结果表明, 老龄林中不同径级群落的物种多样性及不同径级间的物种相似度显著高于各恢复阶段的次生林, 但优势种在群落中的比例低于各恢复阶段的次生林。随着自然恢复过程的进行, 次生林群落物种组成与老龄林的相似性也逐渐增大, 支持演替平衡理论。所有恢复阶段样地中幼苗的个体、物种丰富度和基于多度涵盖估计量(ACE)都低于幼树和成年树群落, 幼苗层物种组成与幼树、成年树也有较大差异, 说明新增到幼苗群落可能是一个难于预测的过程。研究结果说明了确定过程和随机过程共同决定了次生演替的群落构建。     

长白山阔叶红松林乔木幼苗组成及多度格局的影响因素

以长白山原始阔叶红松林样地为平台,以样方中胸径小于1 cm的乔木幼苗为研究对象,基于2018年两次的幼苗调查数据,利用广义线性混合模型分析影响群落乔木幼苗多度的生物邻体和生境因素。结果表明:该样地所有幼苗样方共调查到10064株乔木幼苗,累计17个物种,分属9科9属,水曲柳幼苗多度极高,在乔木树种幼苗中占优势地位;在群落水平,幼苗多度与异种大树效应、草本密度和林冠开阔度呈显著正相关,与草本盖度和土壤含水量呈显著负相关;在物种水平,水曲柳和紫椴幼苗多度的影响因素与群落水平筛选后的结果一致,红松幼苗多度与异种大树效应呈正相关,与同种大树效应及林冠开阔度呈负相关。该研究证实了生物邻体和生境因素共同影响幼苗多度格局,并且生物邻体和生境因素的相对重要性随幼苗物种种类不同而变化。     

On the adaptive value of large seeds for tropical moist forest trees: A review and synthesis

In moist temperate and tropical environments species that typically become established in closed, shaded habitats tend to have larger seeds than those that regenerate in open, secondary habitats. Despite this common pattern and the frequency with which benefits of small seed size for early successional species (large number, enhanced dispersal potential) have been discussed, little attention has focused on the advantages of large seeds for species that regenerate in closed, late successional associations. It is generally considered that large seeds enhance seedling survivorship at low light intensities. However, light intensity is only one of several factors that differ between shaded and sunlit habitats. This review examines microclimatic and biotic differences between shaded subcanopy habitats in mature tropical forests and those in sunlit, light gap habitats in which the early stages of tropical forest succession occur. Each factor is examined as a possible selective agent responsible for maintaining seed size differences between two guilds of tropical rainforest trees; the pioneer species that have small seeds and typically become established in large, sunlit gaps in the forest canopy and the persistent, relatively shade-tolerant species that have larger seeds and produce seedlings that survive for variable periods of time in the shade beneath the forest canopy. Three microclimatic factors that differ in subcanopy and gap habitats are examined; temperature, moisture, and light intensity. It is unlikely that temperature has been an important selective agent in maintaining the differences in seed sizes observed between the pioneer and persistent tree guilds. However, greater desiccation stress in light gaps might prevent successful regeneration of larger seeds in this habitat and thus might impose the smaller mean seed sizes of pioneer species. Reduced light intensities in subcanopy habitats also could favor larger seeds in the persistent guild because large seed reserves might 1) enhance the abilities of seeds to persist until suitable light (or moisture) conditions arise by providing for metabolic requirements of seeds during quiescent periods, 2) provide secondary compounds for defense of persistent seedlings against pathogens and predators during periods of low energy availability, 3) provide energy for construction of large amounts of photosynthetic tissue needed to maintain a positive net energy balance when light conditions are just above the leaf light compensation point of the plant, 4) provide energy for growth into higher light intensity strata, and 5) provide nutrients for replacement of lost or damaged tissues in persistent seedlings. Differences between soils in light gaps and subcanopy habitats are considered briefly. It is concluded that too little is known for predictions to be made regarding the probable effects of soil differences on the sizes of seeds able to survive in each habitat. Finally, differences between the two habitats in four biotic factors (competition, predation, pathogens, and mycorrhizal availability) are considered. Of these, greater competition for nutrients in the subcanopy habitat, and competition among co-germinating seedlings for light could have been important in favoring large seeds in the guild of persistent species. Pathogens are known to be more effective in shaded habitats, but data on seedling resistance to pathogens do not provide support for a role of seed size in enhancing resistance. Although differences in predation intensity and in mycorrhizal abundance in the two habitats have not been evaluated in the field, potential roles of these two factors in maintaining the seed size differential between these two guilds of forest trees are discussed. Despite the existence of numerous potential benefits of large seed reserves, seed sizes often must reflect compromises between conflicting selective pressures. Environmental conditions (e.g., moisture availability) can impose upper limits on seed size. Enhanced dispersal potential and greater total propagule numbers from maternal energy reserves are benefits of small seed size that can counterbalance selection for large seed reserves. The interactions between selective forces in molding seed sizes are discussed in a final section.     

Light gradient partitioning by tropical tree seedlings in the absence of canopy gaps

To explore the importance of light availability for seedling growth in low light environments, we examined light-dependent growth, biomass allocation and mortality of tree seedlings growing in sites with 0.2-6.5% full sun, the range of light commonly encountered in the understory of closed canopy, lowland tropical forests. We transplanted seedlings of the canopy tree species, Dipteryx panamensis, Virola koschnyii, and Brosimum alicastrum into second-growth forest and native tree plantations at La Selva Biological Station, Costa Rica. We assessed seedling survival, growth, and seedling light environments bimonthly for 14 months. Plants were harvested at the end of the study to assess leaf area, total biomass, biomass partitioning and root growth. Survivorship of all species exceeded 60% across all microsites, although both D. panamensis and B. alicastrum had lower probabilities of survival in the darkest microsites. All species showed a strong positive relationship between light availability and growth, increasing in total biomass as light increased. However, the strength of the growth response differed among species causing a change in the rank order of species growth rates as light availability increased. Although D. panamensis showed the lowest growth rates in the darkest microsites, a strong response to increasing light led to a cross-over in performance, such that D. panamensis had the highest growth rate at the highest light levels studied. These data suggest that resource gradient partitioning could occur even in low light environments (0.2-6.5%). Given the limited range of light regimes sampled (i.e., non-gap microsites), our data demonstrate that growth of tropical tree seedlings beneath closed canopies is highly sensitive to light availability and that shade-tolerant species vary in these responses. Our results show that understory light heterogeneity, in the absence of canopy gaps, can significantly affect recruitment processes for shade-tolerant tree species.     

The confounding effects of logging on tree seedling growth and herbivory in Central Amazon

Light availability is an important modulator of seedling growth and plant–herbivore dynamics. Logging increases light levels in forests, potentially altering herbivore–plant interactions that drive seedling establishment. We conducted a transplant experiment to evaluate how logging and herbivory affect seedling growth and survival in three shade‐tolerant tree species, at paired canopy gap and understory sites in logged forest and an adjacent unlogged area in central Amazonia (Brazil). Seedlings were either left exposed to naturally occurring insect herbivores or protected from insects by a fine netting structure. We measured the herbivore damage and growth rate of seedlings after 18 mo. In general, logged areas received more light than unlogged sites. Growth and herbivory rates were positively influenced by light, and herbivory was also influenced positively by logging. In gaps, increased growth mitigated foliar damage. Logging resulted in a loss of foliar tissue due to increased herbivory. Herbivory rates were higher in the understory of logged sites than in that of unlogged understory sites, but growth was similar in these areas. Thus, the understory of logged areas provided the least favorable sites for shade‐tolerant tree regeneration, due to higher herbivory rates. The effect of logging on biotic interactions can extend beyond the gaps it creates into untouched understory sites. To our knowledge, this is the first time such a pattern has been observed, highlighting the importance of evaluating the impact of logging on biotic interactions.     

Dispersal mode,shade tolerance,and phytogeographical affinity of tree species during secondary succession in tropical montane cloud forest

Secondary succession following land abandonment, represented by a chronosequence of 15 old fields (0–80 years old) and two old-growth forests, was studied in the tropical montane cloud forest region of Veracruz, Mexico. The objective was to determine successional trajectories in forest structure and species richness of trees ≥5 cm DBH, in terms of differences in seed dispersal mode, shade tolerance, and phytogeographical affinity. Data were analyzed using AIC model selection and logistic regressions. Mean and maximum canopy height reached values similar to old-growth forest at 35 and 80 years, respectively. Species richness and diversity values were reached earlier (15 and 25 years, respectively) while basal area and stem density tended to reach old-growth forest values within 80 years. Along the chronosequence, the proportion of species and individuals of wind-dispersed trees declined, that of bird dispersed small seeded trees remained constant, while that of gravity and animal dispersed large seeded trees increased; shade-intolerant species and individuals declined, while intermediate and shade-tolerant trees increased. Shade-tolerant canopy trees were rare during succession, even in the old-growth forest. Tropical tree species were more frequent than temperate ones throughout the chronosequence, but temperate tree individuals became canopy dominants at intermediate and old-growth forest stages.     

Seasonal and habitat differences affect the impact of food and predation on herbivores: a comparison between gaps and understory of a tropical forest

Herbivore populations are influenced by a combination of food availability and predator pressure, the relative contribution of which is hypothesized to vary across a productivity gradient. In tropical forests, treefall gaps are pockets of high productivity in the otherwise less productive forest understory. Thus, we hypothesize that higher light availability in gaps will increase plant resources, thereby decreasing resource limitation of herbivores relative to the understory. As a result, predators should regulate herbivore populations in gaps, whereas food should limit herbivores in the understory. We quantified potential food availability and compared arthropod herbivore and predator densities in large forest light gaps and in the intact understory in Panama. Plants, young leaves, herbivores and predators were significantly more abundant per ground area in gaps than in the understory. This pattern was similar when we focused on seven gap specialist plant species and 15 shade-tolerant species growing in gaps and understory. Consistent with the hypothesis, herbivory rates were higher in gaps than the understory. Per capita predation rates on artificial caterpillars indicated higher predation pressure in gaps in both the dry and late wet seasons. These diverse lines of evidence all suggest that herbivores experience higher predator pressure in gaps and more food limitation in the understory.     

Complementary Models of Tree Species–Soil Relationships in Old-Growth Temperate Forests

Ecosystem-level studies identify plant–soil feedbacks as important controls on soil nutrient availability, particularly for nitrogen and phosphorus. Although site- and species-specific studies of tree species–soil relationships are relatively common, comparatively fewer studies consider multiple co-existing species in old-growth forests across a range of sites that vary in underlying soil fertility. We characterized patterns in forest floor and mineral soil nutrients associated with four common tree species across eight undisturbed old-growth forests in Oregon, USA, and used two complementary conceptual models to assess tree species–soil relationships. Plant–soil feedbacks that could reinforce site-level differences in nutrient availability were assessed using the context-dependent relationships model, whereby relative species-based differences in each soil nutrient diverged or converged as nutrient status changed across sites. Tree species–soil relationships that did not reflect strong feedbacks were evaluated using a site-independent relationships model, whereby forest floor and surface mineral soil nutrient pools differed consistently by tree species across sites, without variation in deeper mineral soils. We found that the organically cycled elements carbon, nitrogen, and phosphorus exhibited context-dependent differences among species in both forest floor and mineral soil, and most often followed a divergence model, whereby species differences were greatest at high-nutrient sites. These patterns are consistent with theory emphasizing biotic control of these elements through plant–soil feedback mechanisms. Site-independent species differences were strongest for pools of the weatherable cations calcium, magnesium, potassium, as well as phosphorus, in mineral soils. Site-independent species differences in forest floor nutrients were attributable to one species that displayed significantly greater forest floor mass accumulation. Our findings confirm that site-independent and context-dependent tree species-soil relationships occur simultaneously in old-growth temperate forests, with context-dependent relationships strongest for organically cycled elements, and site-independent relationships strongest for weatherable elements with inorganic cycling phases. These models provide complementary explanations for patterns of nutrient accumulation and cycling in mixed-species old-growth temperate forests.