Indirect interactions among tropical tree species through shared rodent seed predators: a novel mechanism of tree species coexistence

The coexistence of numerous tree species in tropical forests is commonly explained by negative dependence of recruitment on the conspecific seed and tree density due to specialist natural enemies that attack seeds and seedlings (‘Janzen–Connell’ effects). Less known is whether guilds of shared seed predators can induce a negative dependence of recruitment on the density of different species of the same plant functional group. We studied 54 plots in tropical forest on Barro Colorado Island, Panama, with contrasting mature tree densities of three coexisting large seeded tree species with shared seed predators. Levels of seed predation were far better explained by incorporating seed densities of all three focal species than by conspecific seed density alone. Both positive and negative density dependencies were observed for different species combinations. Thus, indirect interactions via shared seed predators can either promote or reduce the coexistence of different plant functional groups in tropical forest.     

The relative importance of above- versus belowground competition for tree growth during early succession of a tropical moist forest

Competition between neighboring plants plays a major role in the population dynamics of tree species in the early phases of humid tropical forest succession. We evaluated the relative importance of above- versus below-ground competition during the first years of old-field succession on soil with low fertility in Southern Mexico, using the premise that competition for light is size-asymmetric, unlike competition for nutrients. Plant growth is thus expected to be disproportionally impeded by larger neighbors. We studied how growth and survival of 3.5–5.5 m tall saplings of Cecropia peltata and Trichospermum mexicanum, two pioneer species that dominate the secondary forests in the study region, varied with the abundance and size of neighboring trees in 1–2 year old secondary vegetation. We found that local neighborhood basal area varied 10-fold (3 to 30 cm² m^-2) and explained most of the variation in diameter and height growth of the target saplings. Most growth variables were strongly affected by the neighbors bigger than the focal trees with no significant additive effect of the smaller neighbors, indicating asymmetric competition. Smaller neighbors did have a small but significant additive effect on the diameter growth of Cecropia saplings and stem slenderness of Trichospermum saplings. We conclude that competition for light was more important than belowground competition in this initial phase of moist tropical forest successional, despite the low soil fertility.     

Spatial patterns of herbivory by gall-forming insects: a test of the soil fertility hypothesis in a Mexican tropical dry forest

It has been proposed that fertile soils reduce the incidence of gall-forming insect (GFI) species in plant communities. This is known as the soil fertility hypothesis. The main objective of this study was to analyze the spatial distribution of GFI species under different habitats in a tropical dry forest at the Chamela-Cuixmala Biosphere Reserve, Mexico. Eight habitats which differ in soil type, topography, nutrient availability and vegetation were chosen. We found that 38 GFI species specialize on their host plant species. GFI species richness was negatively correlated with phosphorous and nitrogen availability. Using phosphorous as an indicator of soil fertility, we found low frequency and density of GFI on fertile soils. Our study indicates that soil fertility is one of the factors that negatively affects the patterns of spatial distribution of species richness, incidence and abundance of GFI at the community level in two different ways: i) indirectly affecting GFI species richness in plants adapted to infertile soils and ii) directly affecting GFI responses to plant traits of hosts found in a fertility gradient.     

山地土壤肥力与植物群落次生演替关系研究

从土壤物理、化学和生物性质出发,通过土壤肥力综合指标体系,利用多元统计方法,对重庆缙云山亚热带常绿阔叶林不同演替阶段山地土壤肥力的变化特征进行综合评价。并分析了缙云山亚热带常绿阔叶林群落演替与山地土壤肥力变化的关系。     

Plant functional assembly is mediated by rainfall and soil conditions in a seasonally dry tropical forest

Understanding how species assembly is influenced by the interplay of climate, local environmental conditions and human-caused disturbances remains a central question in ecology and conservation. Here, we assess how plant species abundance is determined by combinations of functional traits (ecological strategies) and interacting gradients of rainfall, soil conditions (fertility and field capacity) and chronic anthropogenic disturbance in a Caatinga dry tropical forest, Brazil. We tested for trait–environment relationships using multivariate methods (RLQ) accounting for groups of species sharing similar responses to gradients and similar expression of multiple traits (i.e. response groups). Overall, species’ abundances changed predictably in response to rainfall and soil fertility, and were mediated by functional traits, i.e. species with particular trait combinations tended to respond similarly to multifactorial conditions. Briefly, three ecological strategies emerged: species with low wood density and soft (i.e. lower dry matter content), thick leaves converged into a trait syndrome characterizing a drought-avoidance strategy through water storage. They were particularly abundant under extremely low precipitation and relatively high soil field capacity. Under conditions of increasing rainfall and decreasing soil field capacity, species with high wood density were favored, consistent with a drought-tolerance strategy. However, these species fell into two groups relative to leaf-investment: more conservative leaves (low SLA) on relatively fertile soils vs. thinner and softer (i.e. high SLA) leaves on unfertile soils. In seasonally dry tropical forests, low SLA on relatively fertile soils may represent a water conservation strategy. Unexpectedly, no ecological strategy emerged in response to disturbance. The patterns we uncovered help to understand the interplay between precipitation, soil fertility and anthropogenic disturbance in plant species filtering in seasonally dry tropical forests. Moreover, our results underline that impacts of future climate change will depend on how rainfall patterns covary with finer-scale environmental factors such as soil fertility and field capacity.     

热带森林植物多样性及其维持机制

热带森林具有地球上最丰富的植物多样性。关于热带森林植物多样性的维持机制,虽有众多假说,但均未形成完善的理论体系。不同的学者从不同的角度出发得出了许多结论,但也引起了不少争议。本文主要简述了4种经常被用来解释热带森林高植物多样性的机制：生态位分化、取食压力、生活史负相关和随机竞争,每一种机制都有大量的证据支持。热带森林植物沿微环境梯度的非随机空间分布表明其生态位分化很明显,并对其多样性起重要作用。动物的取食降低母树周围同种幼苗的生长率和存活率,为其他物种存活提供了机会,这就是取食压力假说,这是一个极有前景但仍需大量实验验证的假说。生活史负相关使得热带森林的许多植物能够共存。冠层植物的抑制使得随机性在林下植物的建立过程中起决定作用。     

Responses to Fire in Selected Tropical Dry Forest Trees1

Fire is a frequent disturbance in the tropical dry forests of Central America, yet very little is known about how native species respond to such events. We conducted an experimental burn in a tropical dry forest of western Nicaragua to evaluate plant responses to fire with respect to survivorship and recruitment. Measurements of woody vegetation of all size classes were carried out prior to the prescribed burn and three successive years post fire. We selected the 15 most abundant species <10 cm DBH to assess percent survivorship and sprouting responses post fire. Changes in seedling densities for these 15 most abundant species and the 15 least abundant species were analyzed using a repeated measure ANOVA. We also assessed changes in seedling densities for three species of international conservation concern. We found three major fire‐coping strategies among common dry forests plants: resisters (low fire‐induced mortality), resprouters (vigorous sprouting), and recruiters (increased seeding post‐fire). While survivorship was generally high relative to tropical moist forest species, those species with lower survivorship used either seeding or sprouting as an alternative strategy for persisting in the forest community. Seed dispersal mechanisms, particularly wind dispersal, appear to be an important factor in recruitment success post‐fire. Burn treatment led to a significant increase in the density of seedlings for two species of conservation concern: Guaiacum sanctum and Swietenia humilis. Results of this study suggest that common dry forest species in western Nicaragua are fire tolerant. Further study of individual species and their fire responses is merited.     

Effects of Competition and Facilitation on Species Assemblage in Two Types of Tropical Cloud Forest

Competition and facilitation between tree individuals are two kinds of non-random processes influencing the structure and functioning of forest communities, but how these two plant-plant interactions change along gradient of resources or environments remains very much a matter of debate. We developed a null model to test the size-distance regression, and assessed the effects of competition and facilitation (including interspecific interactions, intraspecific interactions and overall species interactions) on each adult tree species assemblage [diameter at breast height (dbh) ≥5 cm] across two types of tropical cloud forest with different environmental and resource regimes. The null model test revealed that 17% to 27% tree species had positive dbh-distance correlations while 11% to 19% tree species showed negative dbh-distance correlations within these two forest types, indicating that both competition and facilitation processes existed during the community assembly. The importance of competition for heterospecific species, and the intensity of competition for both heterospecific and overall species increased from high to low resources for all the shared species spanning the two forests. The importance of facilitation for conspecific and overall species, as well as that the intensity of facilitation for both heterospecific and conspecific species increased with increasing low air temperature stress for all the shared species spanning the two forests. Our results show that both competition and facilitation processes simultaneously affect parts of species assemblage in the tropical cloud forests. Moreover, the fact that nearly 50% species assemblage is not detected with our approaches suggest that tree species in these tropical forest systems are assembled with multiple ecological processes, and that there is a need to explore the processes other than the two biotic interactions in further researches.     

Woody life-form composition and association on rainfall and soil fertility gradients in Ghana

The hypothesis that treelets/shrubs are more sensitive than trees to drought and low soil fertility was tested by analysing species densities (species per unit area) in 155 tropical forest plots (0.0625 ha) in Ghana, West Africa. Species densities of both life forms decreased with increasing dryness and with increasing soil fertility, which were correlated. Goodness of fit tests detected species preferences for four combinations of rainfall (dry: <1500 mm yr^-1, wet: 1500 mm yr^-1) and soil fertility (two classes based on a composite variable with high weightings for total exchangeable bases and pH), and a meta-analysis of these results compared life forms. The results contradicted the hypothesised response of life forms in two ways: (i) species densities of treelets/shrubs were comparably or relatively less reduced by dry conditions than were tree species densities, and (ii) treelet/shrub species did not show a more frequent preference for wet or fertile sites than did tree species. Light availability and rooting depth are discussed as explanations for the sensitivity of understorey plants to water and nutrient stress, and for the data's failure to support the hypothesis.     

Termite (Insecta: Isoptera) Species Composition in a Primary Rain Forest and Agroforests in Central Amazonia

Termites play important roles in organic matter decomposition, nutrient cycling, and soil structure in tropical rain forests. When forests are replaced by agriculture, termite species richness, abundance, and function often decline. We compared the termite assemblage of a primary forest site with that of a low plant diversity, palm-based agroforest (five plant species) and a high plant diversity, home-garden agroforest (10 plant species) using a rapid biodiversity assessment protocol. In comparing the primary forest termite species composition to previously published studies, we found soil feeders and the Apicotermitinae to be more dominant than previously reported in Amazonia. Thirty percent of the species belonged to the Apicotermitinae, and an unusually high percentage (57%) of species were soil feeders. Unexpectedly, the palm-based agroforest, despite its lower plant diversity, was closer to primary forest in termite species composition, rate of species accumulation, and proportions of species in taxonomic and functional classes than was the home-garden agroforest. This suggests that particular plant attributes may better determine the termite assemblage than plant diversity alone in these agroecosystems. Unlike other agroecosystems reported in the literature, Apicotermitinae and soil feeders were proportionally more abundant in these agroforests than in primary forest. The ability of agroforests to support populations of soil feeders has a potentially positive effect on soil fertility in these agroecosystems; insomuch as feeding guild is a proxy for function, these closed-canopy agroforests may be able to sustain the same termite-mediated functions as primary forest.     

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.     

Spatial Scales of Pollen and Seed-Mediated Gene Flow in Tropical Rain Forest Trees

Gene flow via seed and pollen is a primary determinant of genetic and species diversity in plant communities at different spatial scales. This paper reviews studies of gene flow and population genetic structure in tropical rain forest trees and places them in ecological and biogeographic context. Although much pollination is among nearest neighbors, an increasing number of genetic studies report pollination ranging from 0.5–14 km for canopy tree species, resulting in extensive breeding areas in disturbed and undisturbed rain forest. Direct genetic measures of seed dispersal are still rare; however, studies of fine scale spatial genetic structure (SGS) indicate that the bulk of effective seed dispersal occurs at local scales, and we found no difference in SGS (Sp statistic) between temperate (N?=?24 species) and tropical forest trees (N?=?15). Our analysis did find significantly higher genetic differentiation in tropical trees (F _ST?=?0.177; N?=?42) than in temperate forest trees (F _ST?=?0.116; N?=?82). This may be due to the fact that tropical trees experience low but significant rates of self-fertilization and bi-parental inbreeding, whereas half of the temperate tree species in our survey are wind pollinated and are more strictly allogamous. Genetic drift may also be more pronounced in tropical trees due to the low population densities of most species.     

Weak Competition Among Tropical Tree Seedlings: Implications for Species Coexistence

The intensity of competition among forest tree seedlings is poorly understood, but has important ramifications for their recruitment and for the maintenance of species diversity. Intense competition among seedlings could allow competitively dominant species to exclude subordinate species. Alternatively, the low density and small stature of forest tree seedlings could preclude intense interseedling competition. In this case, other processes, such as size-asymmetric competition with adults, interactions with consumers, or neutral dynamics would prevail as those structuring the forest understory. We tested the intensity of, and potential for, intraspecific competition among tree seedlings of three species ( Brosimum alicastrum , Matisia cordata , and Pouteria reticulata ) in two Neotropical rain forests. We reduced stem densities by up to 90 percent and monitored individual growth and survival rates for up to 24 mo. Individual growth and survival rates were generally unrelated to stem density. Contrary to the predicted behavior of intensely competing plant populations, the distribution of individual heights did not become more left-skewed with time for any species, regardless of plot density; i.e. , excesses of short, suppressed individuals did not accumulate in high-density plots. We further measured the overlap of zones of influence (ZOIs) to assess the potential for resource competition. Seedling ZOIs overlapped only slightly in extremely dense monodominant plots, and even less in ambient-density plots of mixed composition. Our results thus suggest that interseedling competition was weak. Given the low density of tree seedlings in Neotropical forests, we infer that resource competition among seedlings may be irrelevant to their recruitment.     

Population structure and adult plant performance of forest herbs in three contrasting habitats

In Flanders, forests have been subjected to subsequent periods of reclamation and reforestation, leading to a mosaic of forest patches of various ages and with different abiotic conditions, depending on the duration and intensity of the past agricultural activities. While extensive research has been done on the relative importance of dispersal limitation and establishment limitation in explaining colonization patterns of so-called ancient forest plant species, little is known about the viability of populations of these species once they have colonized recent forests. Therefore, a comparative study on population structure and plant performance was carried out for three forest plant species with different colonization capacity ( Primula elatior , Geum urbanum and Arum maculatum ) in three different habitat types: ancient forests, recently established forests and small landscape elements (hedgerows). Contrary to the expectation, it was shown that adult plant performance – a combination of vegetative as well as reproductive characteristics – of the three species was generally higher in recent forests and small landscape elements. This also led to higher seedling densities and proportions in these habitat types. Adult densities, however, were extremely low in newly established forests, suggesting reduced seedling and juvenile survival under these circumstances, probably due to higher light availability and hence increased competition from species like Urtica dioica . Although there might be no immediate establishment limitation, installing a shrub layer in recent forests might enhance survival and result in a stable population structure. Hedgerows on the other hand seemed to contain viable populations of certain forest plant species and can therefore function as an important colonization source when incorporated in recently established forests.     

The negative effect of lianas on tree growth varies with tree species and season

Lianas reduce tree growth, reproduction, and survival in tropical forests. Liana competition can be particularly intense in isolated forest fragments, where liana densities are high, and thus, host tree infestation is common. Furthermore, lianas appear to grow particularly well during seasonal drought, when they may compete particularly intensely with trees. Few studies, however, have experimentally quantified the seasonal effects of liana competition on multiple tree species in tropical forests. We used a liana removal experiment in a forest fragment in southeastern Brazil to test whether the effects of lianas on tree growth vary with season and tree species identity. We conducted monthly diameter measurements using dendrometer bands on 88 individuals of five tree species for 24 months. We found that lianas had a stronger negative effect on some tree species during the wet season compared to the dry season. Furthermore, lianas significantly reduced the diameter growth of two tree species but had no effect on the other three tree species. The strong negative effect of lianas on some trees, particularly during the wet season, indicates that the effect of lianas on trees varies both seasonally and with tree species identity. Abstract in Portuguese is available with online material.     

High density of white-faced capuchins (Cebus capucinus) and habitat quality in the Taboga Forest of Costa Rica

Across the globe, primates are threatened by human activities. This is especially true for species found in tropical dry forests, which remain largely unprotected. Our ability to predict primate abundance in the face of human activity depends on different species' sensitivities as well as on the characteristics of the forest itself. We studied plant and primate distribution and abundance in the Taboga Forest, a 516-ha tropical dry forest surrounded by agricultural fields in northwestern Costa Rica. We found that the density of white-faced capuchins (Cebus capucinus) at Taboga is 2–6 times higher than reported for other long-term white-faced capuchin sites. Using plant transects, we also found relatively high species richness, diversity, and equitability compared with other tropical dry forests. Edge transects (i.e., within 100 m from the forest boundary) differed from interior transects in two ways: (a) tree species associated with dry forest succession were well-established in the edge and (b) canopy cover in the edge was maintained year-round, while the interior forest was deciduous. Sighting rates for capuchins were higher near water sources but did not vary between the edge and interior forest. For comparison, we also found the same to be true for the only other primate in the Taboga Forest, mantled howler monkeys (Alouatta palliata). Year-round access to water might explain why some primate species can flourish even alongside anthropogenic disturbance. Forest fragments like Taboga may support high densities of some species because they provide a mosaic of habitats and key resources that buffer adverse ecological conditions.     

Post‐Establishment Seedling Success in the Brazilian Cerrado: A Comparison of Savanna and Forest Species1

Due to frequent fire, low nutrient availability, and prolonged drought, tropical savanna is a stressful environment for the survival and growth of woody plant seedlings. To understand why forest species do not succeed in this environment while savanna species are able to persist, the effects of fire and woody cover on seedlings of these two functional groups were investigated in the Brazilian Cerrado. Seedlings were established in experimental plots under three densities of woody cover, in sites protected from fire and sites to be subjected to fire. There was a clear difference in the ability of savanna and forest species to survive fire. None of the three forest species were able to survive fire during the first two years of life, whereas eight of the nine savanna species were able to resprout following fire. The small seed size of the ninth savanna species, Miconia albicans, predisposed its seedlings to be sensitive to fire, because there was a strong positive correlation between seed size and survivorship. Savanna species were less dependent on woody cover than were forest species, which exhibited higher growth and survival under tree canopies than in open grassland. The low rates of establishment and survival of forest trees in savanna, combined with high sensitivity to fire, appear sufficient to prevent the expansion of forest into savanna under current fire regimes in the Cerrado.     

Plant diversity in tropical forests: a review of mechanisms of species coexistence

Evidence concerning mechanisms hypothesized to explain species coexistence in hyper-diverse communities is reviewed for tropical forest plants. Three hypotheses receive strong support. Niche differences are evident from non-random spatial distributions along micro-topographic gradients and from a survivorship-growth tradeoff during regeneration. Host-specific pests reduce recruitment near reproductive adults (the Janzen-Connell effect), and, negative density dependence occurs over larger spatial scales among the more abundant species and may regulate their populations. A fourth hypothesis, that suppressed understory plants rarely come into competition with one another, has not been considered before and has profound implications for species coexistence. These hypotheses are mutually compatible. Infrequent competition among suppressed understory plants, niche differences, and Janzen-Connell effects may facilitate the coexistence of the many rare plant species found in tropical forests while negative density dependence regulates the few most successful and abundant species.     

Land‐use Change Dynamics,Soil Type and Species Forming Mono‐dominant Patches: the Case of Pteridium aquilinum in a Neotropical Rain Forest Region

Deforestation and agricultural land degradation in tropical regions can create conditions for growth of perennial plant species forming mono‐dominated patches (MDP). Such species might limit forest regeneration, and their proliferation forces the abandonment of fields and subsequent deforestation to establish new fields. Therefore, identifying factors fostering MDP species is critical for biodiversity conservation in human‐modified landscapes. Here, we propose a conceptual framework to identify such factors and apply it to the case of Pteridium aquilinum (bracken fern), a light‐demanding species, tolerant of low soil fertility and fire. We hypothesize that bracken proliferation is promoted by land‐use changes that increase light availability, especially in sites with low soil fertility and land uses involving fire. We assessed this idea using agricultural fields in southeastern Mexico with different land‐use change histories and quantifying prevalence and cover of bracken. Five different land‐use change histories resulted from transitions among forest, crop, pasture, and fallow field stages. Of the 133 fields sampled, 71 percent had P. aquilinum; regression tree analysis indicated that 65 percent of inter‐field variation in prevalence and 90 percent in cover was explained by land‐use change history and soil type. Maximum prevalence, cover, and rates of increase in bracken were found on fields with low fertility sandy/clay soils, which had been used for crops and pasture, were frequently burned, and had high levels of light. Fields on fertile alluvial soil never used for pasture were bracken‐free. Agriculture promoting high light environments on less fertile soils is a major cause of bracken proliferation and likely that of other MDP species.     

Comparing forest structure and biodiversity on private and public land: secondary tropical dry forests in Costa Rica

Secondary forests constitute a substantial proportion of tropical forestlands. These forests occur on both public and private lands and different underlying environmental variables and management regimes may affect post‐abandonment successional processes and resultant forest structure and biodiversity. We examined whether differences in ownership led to differences in forest structure, tree diversity, and tree species composition across a gradient of soil fertility and forest age. We collected soil samples and surveyed all trees in 82 public and 66 private 0.1‐ha forest plots arrayed across forest age and soil gradients in Guanacaste, Costa Rica. We found that soil fertility appeared to drive the spatial structure of public vs. private ownership; public conservation lands appeared to be non‐randomly located on areas of lower soil fertility. On private lands, areas of crops/pasture appeared to be non‐randomly located on higher soil fertility areas while forests occupied areas of lower soil fertility. We found that forest structure and tree species diversity did not differ significantly between public and private ownership. However, public and private forests differed in tree species composition: 11 percent were more prevalent in public forest and 7 percent were more prevalent in private forest. Swietenia macrophylla, Cedrela odorata, and Astronium graveolens were more prevalent in public forests likely because public forests provide stronger protection for these highly prized timber species. Guazuma ulmifolia was the most abundant tree in private forests likely because this species is widely consumed and dispersed by cattle. Furthermore, some compositional differences appear to result from soil fertility differences due to non‐random placement of public and private land holdings with respect to soil fertility. Land ownership creates a distinctive species composition signature that is likely the result of differences in soil fertility and management between the ownership types. Both biophysical and social variables should be considered to advance understanding of tropical secondary forest structure and biodiversity.