Plant-Mammal Interactions in Tropical Bolivian Forests with Different Hunting Pressures

Abstract: We studied plant-animal interactions and vegetation structure in two geographically close tropical Bolivian forests subjected to different hunting intensities. We hypothesized that reduction of mammals openface> 1 kg in an "intensively hunted forest," compared with an "occasionally hunted forest," should correlate with decreased seed predation and seedling trampling, increased seedling survival and density, and decreased tree-species diversity at the seedling stage in relation to the adult stage. The occasionally hunted forest held 1.7 times as many mammalian species as the intensively hunted forest. As predicted, predation of Astrocaryum murumuru seeds was 34.2% lower in the intensively hunted forest. Similarly, trampling of model seedlings was 5.4 times lower and seedling survival was 1.15 times greater in the intensively hunted forest than that in the occasionally hunted forest. But the intensively hunted forest displayed lower seedling densities and a higher ratio of seedling diversity to tree diversity than did the occasionally hunted forest. Reduction of peccaries from the intensively hunted forest may explain much of the between-site differences in seed predation, trampling, and seedling survival. Lack of consistent differences in seedling density and diversity could mean that reduced granivory and trampling may be counteracted by reduced seed dispersal. The reduction of mammalian populations may produce a complex mosaic of forest patches with distinct degrees of structural change, depending on the intensity of defaunation.     

Meta‐Analysis of the Effects of Human Disturbance on Seed Dispersal by Animals

Animal‐mediated seed dispersal is important for sustaining biological diversity in forest ecosystems, particularly in the tropics. Forest fragmentation, hunting, and selective logging modify forests in myriad ways and their effects on animal‐mediated seed dispersal have been examined in many case studies. However, the overall effects of different types of human disturbance on animal‐mediated seed dispersal are still unknown. We identified 35 articles that provided 83 comparisons of animal‐mediated seed dispersal between disturbed and undisturbed forests; all comparisons except one were conducted in tropical or subtropical ecosystems. We assessed the effects of forest fragmentation, hunting, and selective logging on seed dispersal of fleshy‐fruited tree species. We carried out a meta‐analysis to test whether forest fragmentation, hunting, and selective logging affected 3 components of animal‐mediated seed dispersal: frugivore visitation rate, number of seeds removed, and distance of seed dispersal. Forest fragmentation, hunting, and selective logging did not affect visitation rate and were marginally associated with a reduction in seed‐dispersal distance. Hunting and selective logging, but not fragmentation, were associated with a large reduction in the number of seeds removed. Fewer seeds of large‐seeded than of small‐seeded tree species were removed in hunted or selectively logged forests. A plausible explanation for the consistently negative effects of hunting and selective logging on large‐seeded plant species is that large frugivores, as the predominant seed dispersers for large‐seeded plant species, are the first animals to be extirpated from hunted or logged forests. The reduction in forest area after fragmentation appeared to have weaker effects on frugivore communities and animal‐mediated seed dispersal than hunting and selective logging. The differential effects of hunting and selective logging on large‐ and small‐seeded tree species underpinned case studies that showed disrupted plant‐frugivore interactions could trigger a homogenization of seed traits in tree communities in hunted or logged tropical forests. Meta Análisis de los Efectos de la Perturbación Humana sobre la Dispersión de Semillas por Animales     

Ecological consequences of forest elephant declines for Afrotropical forests

Poaching is rapidly extirpating African forest elephants (Loxodonta cyclotis) from most of their historical range, leaving vast areas of elephant‐free tropical forest. Elephants are ecological engineers that create and maintain forest habitat; thus, their loss will have large consequences for the composition and structure of Afrotropical forests. Through a comprehensive literature review, we evaluated the roles of forest elephants in seed dispersal, nutrient recycling, and herbivory and physical damage to predict the cascading ecological effects of their population declines. Loss of seed dispersal by elephants will favor tree species dispersed abiotically and by smaller dispersal agents, and tree species composition will depend on the downstream effects of changes in elephant nutrient cycling and browsing. Loss of trampling and herbivory of seedlings and saplings will result in high tree density with release from browsing pressures. Diminished seed dispersal by elephants and high stem density are likely to reduce the recruitment of large trees and thus increase homogeneity of forest structure and decrease carbon stocks. The loss of ecological services by forest elephants likely means Central African forests will be more like Neotropical forests, from which megafauna were extirpated thousands of years ago. Without intervention, as much as 96% of Central African forests will have modified species composition and structure as elephants are compressed into remaining protected areas. Stopping elephant poaching is an urgent first step to mitigating these effects, but long‐term conservation will require land‐use planning that incorporates elephant habitat into forested landscapes that are being rapidly transformed by industrial agriculture and logging.     

Seed predation by Neotropical rain forest mammals increases diversity in seedling recruitment

Seed dispersal and seedling recruitment (the transition of seeds to seedlings) set the spatiotemporal distribution of new individuals in plant communities. Many terrestrial rain forest mammals consume post-dispersal seeds and seedlings, often inflicting density-dependent mortality. In part because of density-dependent mortality, diversity often increases during seedling recruitment, making it a critical stage for species coexistence. We determined how mammalian predators, adult tree abundance, and seed mass interact to affect seedling recruitment in a western Amazonian rain forest. We used exclosures that were selectively permeable to three size classes of mammals: mice and spiny rats (weighing <1 kg), medium-sized rodents (1-12 kg), and large mammals (20-200 kg). Into each exclosure, we placed seeds of 13 tree species and one canopy liana, which varied by an order of magnitude in adult abundance and seed mass. We followed the fates of the seeds and resulting seedlings for at least 17 months. We assessed the effect of each mammalian size class on seed survival, seedling survival and growth, and the density and diversity of the seedlings that survived to the end of the experiment. Surprisingly, large mammals had no detectable effect at any stage of seedling recruitment. In contrast, small- and medium-sized mammals significantly reduced seed survival, seedling survival, and seedling density. Furthermore, predation by small mammals increased species richness on a per-stem basis. This increase in diversity resulted from their disproportionately intense predation on common species and large-seeded species. Small mammals thereby generated a rare-species advantage in seedling recruitment, the critical ingredient for frequency dependence. Predation by small (and to a lesser extent, medium-sized) mammals on seeds and seedlings significantly increases tree species diversity in tropical forests. This is the first long-term study to dissect the effects of various mammalian predators on the recruitment of a diverse set of tree species.     

Effects of Forest Fragmentation on Seed Dispersal and Seedling Establishment in Ornithochorous Trees

Abstract: Habitat fragmentation increases seed dispersal limitation across the landscape and may also affect subsequent demographic stages such as seedling establishment. Thus, the development of adequate plans for forest restoration requires an understanding of mechanisms by which fragmentation hampers seed delivery to deforested areas and knowledge of how fragmentation affects the relationship between seed‐deposition patterns and seedling establishment. We evaluated the dispersal and recruitment of two bird‐dispersed, fleshy‐fruited tree species (Crataegus monogyna and Ilex aquifolium) in fragmented secondary forests of northern Spain. Forest fragmentation reduced the probability of seed deposition for both trees because of decreased availability of woody perches and fruit‐rich neighborhoods for seed dispersers, rather than because of reductions in tree cover by itself. The effects of fragmentation went beyond effects on the dispersal stage in Crataegus because seedling establishment was proportional to the quantities of bird‐dispersed seeds arriving at microsites. In contrast, postdispersal mortality in Ilex was so high that it obscured the seed‐to‐seedling transition. These results suggest that the effects of fragmentation are not necessarily consistent across stages of recruitment across species. Habitat management seeking to overcome barriers to forest recovery must include the preservation, and even the planting, of fleshy‐fruited trees in the unforested matrix as a measure to encourage frugivorous birds to enter into open and degraded areas. An integrative management strategy should also explicitly consider seed‐survival expectancies at microhabitats to preserve plant‐population dynamics and community structure in fragmented landscapes.     

Effects of Local Disturbance of Tropical Forests on Frugivores and Seed Removal of a Small-Seeded Afrotropical Tree

Abstract:  Small-scale, local disturbance of tropical forests, for example from selective logging, is widespread, but its effects on biodiversity and ecosystem function have rarely been studied. In 3 East African tropical rainforests, we investigated the effect of different levels of local forest disturbance on the frugivore community and on tree visitation and fruit removal rates of the small-seeded tree Celtis durandii. We quantified birds and primates in little and heavily disturbed sites, distinguishing between forest specialists, forest generalists, and forest visitors. We quantified frugivorous tree visitors and seed removal rates of C. durandii trees in the same sites. Forest disturbance reduced the species richness and density of the frugivore community and of forest specialists. Frugivorous species and individuals visiting the study trees were reduced significantly, which led to a marginally significant decline in fruit removal by all frugivores and a significant reduction in removal by forest specialists. Reduction in fruit removal by forest specialists was not compensated for by increases in removal by forest generalists or visitors. Results did not differ among the 3 rainforests, which suggests they were consistent at a regional scale. So local forest disturbance led to a loss of frugivores and their seed removal services. This suggests that large-seeded tree species and trees with small fruits are losing seed dispersers. Thus, local forest disturbance appears to have a more general negative impact on frugivores and their seed dispersal services than anticipated previously.     

Experimental manipulation of seed shadows of an Afrotropical tree determines drivers of recruitment

The loss of animals in tropical forests may alter seed dispersal patterns and reduce seedling recruitment of tree species, but direct experimental evidence is scarce. We manipulated dispersal patterns of Manilkara mabokeensis, a monkey-dispersed tree, to assess the extent to which spatial distributions of seeds drive seedling recruitment. Based on the natural seed shadow, we created seed distributions with seeds deposited under the canopy ("no dispersal"), with declining density from the tree ("natural dispersal"), and at uniform densities ("good dispersal"). These distributions mimicked dispersal patterns that could occur with the extirpation of monkeys, low levels of hunting, and high rates of seed dispersal. We monitored seedling emergence and survival for 18 months and recorded the number of leaves and damage to leaves. "Good dispersal" increased seedling survival by 26%, and "no dispersal" decreased survival by 78%, relative to "natural dispersal." Using a mixed-effects survival model, we decoupled the distance and density components of the seed shadow: seedling survival depended on the seed density, but not on the distance from the tree. Although community seedling diversity tended to decrease with longer dispersal distances, we found no conclusive evidence that patterns of seed dispersal influence the diversity of the seedling community. Local seed dispersal does affect seedling recruitment and survival, with better dispersal resulting in higher seedling recruitment; hence the loss of dispersal services that comes with the reduction or extirpation of seed dispersers will decrease regeneration of some tree species.     

Vertebrate herbivory impacts seedling recruitment more than niche partitioning or density-dependent mortality

In tropical forests, resource-based niches and density-dependent mortality are mutually compatible mechanisms that can act simultaneously to limit seedling populations. Differences in the strengths of these mechanisms will determine their roles in maintaining species coexistence. In the first assessment of these mechanisms in a Congo Basin forest, we quantified their relative strengths and tested the extent to which density-dependent mortality is driven by the distance-dependent behavior of seed and seedling predators predicted by the Janzen-Connell hypothesis. We conducted a large-scale seed addition experiment for five randomly selected tropical tree species, caging a subset of seed addition quadrats against vertebrate predators. We then developed models to assess the mechanisms that determine seedling emergence (three months after seed addition) and survival (two years after seed addition). As predicted, both niche differentiation and density-dependent mortality limited seedling recruitment, but predation had the strongest effects on seedling emergence and survival. Seedling species responded differently to naturally occurring environmental variation among sites, including variation in light levels and soil characteristics, supporting predictions of niche-based theories of tropical tree species coexistence. The addition of higher densities of seeds into quadrats initially led to greater seedling emergence, but survival to two years decreased with seed density. Seed and seedling predation reduced recruitment below levels maintained by density-dependent mortality, an indication that predators largely determine the population size of tree seedlings. Seedling recruitment was unrelated to the distance to or density of conspecific adult trees, suggesting that recruitment patterns are generated by generalist vertebrate herbivores rather than the specialized predators predicted by the Janzen-Connell hypothesis. If the role of seed and seedling predation in limiting seedling recruitment is a general phenomenon, then the relative abundances of tree species might largely depend on species-specific adaptations to avoid, survive, and recover from damage induced by vertebrate herbivores. Likewise, population declines of herbivorous vertebrate species (many of which are large and hunted) may trigger shifts in species composition of tropical forests.     

Fragmentation drives tropical forest fragments to early successional states: A modelling study for Brazilian Atlantic forests

Land use leads to massive habitat destruction and fragmentation in tropical forests. Despite its global dimensions the effects of fragmentation on ecosystem dynamics are not well understood due to the complexity of the problem. We present a simulation analysis performed by the individual-based model FORMIND. The model was applied to the Brazilian Atlantic Forest, one of the world's biodiversity hot spots, at the Plateau of São Paulo. This study investigates the long-term effects of fragmentation processes on structure and dynamics of different sized remnant tropical forest fragments (1-100 ha) at community and plant functional type (PFT) level. We disentangle the interplay of single effects of different key fragmentation processes (edge mortality, increased mortality of large trees, local seed loss and external seed rain) using simulation experiments in a full factorial design.Our analysis reveals that particularly small forest fragments below 25 ha suffer substantial structural changes, biomass and biodiversity loss in the long term. At community level biomass is reduced up to 60%. Two thirds of the mid- and late-successional species groups, especially shade-tolerant (late successional climax) species groups are prone of extinction in small fragments. The shade-tolerant species groups were most strongly affected; its tree number was reduced more than 60% mainly by increased edge mortality. This process proved to be the most powerful of those investigated, explaining alone more than 80% of the changes observed for this group. External seed rain was able to compensate approximately 30% of the observed fragmentation effects for shade-tolerant species.Our results suggest that tropical forest fragments will suffer strong structural changes in the long term, leading to tree species impoverishment. They may reach a new equilibrium with a substantially reduced subset of the initial species pool, and are driven towards an earlier successional state. The natural regeneration potential of a landscape scattered with forest fragments appears to be limited, as external seed rain is not able to fully compensate for the observed fragmentation-induced changes. Our findings suggest basic recommendations for the management of fragmented tropical forest landscapes.     

Functional nonredundancy of elephants in a disturbed tropical forest

Conservation efforts are often motivated by the threat of global extinction. Yet if conservationists had more information suggesting that extirpation of individual species could lead to undesirable ecological effects, they might more frequently attempt to protect or restore such species across their ranges even if they were not globally endangered. Scientists have seldom measured or quantitatively predicted the functional consequences of species loss, even for large, extinction‐prone species that theory suggests should be functionally unique. We measured the contribution of Asian elephants (Elephas maximus) to the dispersal of 3 large‐fruited species in a disturbed tropical moist forest and predicted the extent to which alternative dispersers could compensate for elephants in their absence. We created an empirical probability model with data on frugivory and seed dispersal from Buxa Tiger Reserve, India. These data were used to estimate the proportion of seeds consumed by elephants and other frugivores that survive handling and density‐dependent processes (Janzen‐Connell effects and conspecific intradung competition) and germinate. Without compensation, the number of seeds dispersed and surviving density‐dependent effects decreased 26% (Artocarpus chaplasha), 42% (Careya arborea), and 72% (Dillenia indica) when elephants were absent from the ecosystem. Compensatory fruit removal by other animals substantially ameliorated these losses. For instance, reductions in successful dispersal of D. indica were as low as 23% when gaur (Bos gaurus) persisted, but median dispersal distance still declined from 30% (C. arborea) to 90% (A. chaplasha) without elephants. Our results support the theory that the largest animal species in an ecosystem have nonredundant ecological functionality and that their extirpation is likely to lead to the deterioration of ecosystem processes such as seed dispersal. This effect is likely accentuated by the overall defaunation of many tropical systems.     

Effects of howler monkey reintroduction on ecological interactions and processes

Rewilding has been an increasingly popular tool to restore plant–animal interactions and ecological processes impaired by defaunation. However, the reestablishment of such processes has seldom been assessed. We investigated the restoration of ecological interactions following the reintroduction of the brown howler monkey (Alouatta guariba) to a defaunated Atlantic forest site. We expected the reintroduction to restore plant–animal interactions and interactions between howlers and dung beetles, which promote secondary seed dispersal. We estimated the number of interactions expected to be restored by the reintroduction to provide the baseline interaction richness that could be restored. We followed the reintroduced howler monkeys twice a week for 24 months (337 hours total) to assess their diet. We used howler monkey dung in secondary seed dispersal experiments with 2484 seed mimics to estimate the removal rates by dung beetles and collected the beetles to assess community attributes. We compared the potential future contribution of howler monkeys and other frugivores to seed dispersal based on the seed sizes they disperse in other areas where they occur. In 2 years, howler monkeys consumed 60 animal-dispersed plant species out of the 330 estimated. Twenty-one dung beetle species were attracted to experimentally provided dung; most of them were tunnelers, nocturnal, and large-sized (>10 mm). On average 30% (range 0–100%) of the large seed mimics (14 mm) were moved by dung beetles. About 91% of the species consumed by howlers (size range 0.3–34.3 mm) overlapped in seed size with those removed by dung beetles. In our study area, howler monkeys may consume more large-seeded fruit species than most other frugivores, highlighting their potential to affect forest regeneration. Our results show reintroductions may effectively restore ecological links and enhance ecological processes.     

Neutral theory and the evolution of ecological equivalence

Since the publication of the unified neutral theory in 2001, there has been much discussion of the theory, pro and con. The hypothesis of ecological equivalence is the fundamental yet controversial idea behind neutral theory. Assuming trophically similar species are demographically alike (symmetric) on a per capita basis is only an approximation, but it is equivalent to asking: How many of the patterns of ecological communities are the result of species similarities, rather than of species differences? The strategy behind neutral theory is to see how far one can get with the simplification of assuming ecological equivalence before introducing more complexity. In another paper, I review the empirical evidence that led me to hypothesize ecological equivalence among many of the tree species in the species-rich tropical forest on Barro Colorado Island (BCI). In this paper, I develop a simple model for the evolution of ecological equivalence or niche convergence, using as an example evolution of the suite of life history traits characteristic of shade tolerant tropical tree species. Although the model is simple, the conclusions from it seem likely to be robust. I conclude that ecological equivalence for resource use are likely to evolve easily and often, especially in species-rich communities that are dispersal and recruitment limited. In the case of the BCI forest, tree species are strongly dispersal- and recruitment-limited, not only because of restricted seed dispersal, but also because of low recruitment success due to heavy losses of the seedling stages to predators and pathogens and other abiotic stresses such as drought. These factors and the high species richness of the community strongly reduce the potential for competitive exclusion of functionally equivalent or nearly equivalent species.     

The memory of spatial patterns: changes in local abundance and aggregation in a tropical forest

The current spatial pattern of a population is the result of previous individual birth, death, and dispersal events. We present a simple model followed by a comparative analysis for a species-rich plant community to show how the current spatial aggregation of a population may hold information about recent population dynamics. Previous research has shown how locally restricted seed dispersal often leads to stronger aggregation in less abundant populations than it does in more abundant populations. In contrast, little is known about how changes in the local abundance of a species may affect the spatial distribution of individuals. If the level of aggregation within a species depends to some extent on the abundance of the species, then changes in abundance should lead to subsequent changes in aggregation. However, an overall change of spatial pattern relies on many individual birth and death events, and a surplus of deaths or births may have short-term effects on aggregation that are opposite to the long-term change predicted by the change in abundance. The change in aggregation may therefore lag behind the change in abundance, and consequently, the current aggregation may hold information about recent population dynamics. Using an individual-based simulation model with local dispersal and density-dependent competition, we show that, on average, recently growing populations should be more aggregated than shrinking populations of the same current local abundance. We tested this hypothesis using spatial data on individuals from a long-term tropical rain forest plot, and find support for this relationship in canopy trees, but not in understory and shrub species. On this basis we argue that current spatial aggregation is an important characteristic that contains information on recent changes in local abundance, and may be applied to taxonomic groups where dispersal is limited and within-species aggregation is observed.     

Long-term CO2 enrichment of a forest ecosystem: implications for forest regeneration and succession.

The composition and successional status of a forest affect carbon storage and net ecosystem productivity, yet it remains unclear whether elevated atmospheric carbon dioxide (CO2) will impact rates and trajectories of forest succession. We examined how CO2 enrichment (+200 microL CO2/L air differential) affects forest succession through growth and survivorship of tree seedlings, as part of the Duke Forest free-air CO2 enrichment (FACE) experiment in North Carolina, USA. We planted 2352 seedlings of 14 species in the low light forest understory and determined effects of elevated CO2 on individual plant growth, survival, and total sample biomass accumulation, an integrator of plant growth and survivorship over time, for six years. We used a hierarchical Bayes framework to accommodate the uncertainty associated with the availability of light and the variability in growth among individual plants. We found that most species did not exhibit strong responses to CO2. Ulmus alata (+21%), Quercus alba (+9.5%), and nitrogen-fixing Robinia pseudoacacia (+230%) exhibited greater mean annual relative growth rates under elevated CO2 than under ambient conditions. The effects of CO2 were small relative to variability within populations; however, some species grew better under low light conditions when exposed to elevated CO2 than they did under ambient conditions. These species include shade-intolerant Liriodendron tulipifera and Liquidambar styraciflua, intermediate-tolerant Quercus velutina, and shade-tolerant Acer barbatum, A. rubrum, Prunus serotina, Ulmus alata, and Cercis canadensis. Contrary to our expectation, shade-intolerant trees did not survive better with CO2 enrichment, and population-scale responses to CO2 were influenced by survival probabilities in low light. CO2 enrichment did not increase rates of sample biomass accumulation for most species, but it did stimulate biomass growth of shade-tolerant taxa, particularly Acer barbatum and Ulmus alata. Our data suggest a small CO2 fertilization effect on tree productivity, and the possibility of reduced carbon accumulation rates relative to today's forests due to changes in species composition.     

Meta‐Analysis of the Effects of Forest Fragmentation on Interspecific Interactions

Forest fragmentation dramatically alters species persistence and distribution and affects many ecological interactions among species. Recent studies suggest that mutualisms, such as pollination and seed dispersal, are more sensitive to the negative effects of forest fragmentation than antagonisms, such as predation or herbivory. We applied meta‐analytical techniques to evaluate this hypothesis and quantified the relative contributions of different components of the fragmentation process (decreases in fragment size, edge effects, increased isolation, and habitat degradation) to the overall effect. The effects of fragmentation on mutualisms were primarily driven by habitat degradation, edge effects, and fragment isolation, and, as predicted, they were consistently more negative on mutualisms than on antagonisms. For the most studied interaction type, seed dispersal, only certain components of fragmentation had significant (edge effects) or marginally significant (fragment size) effects. Seed size modulated the effect of fragmentation: species with large seeds showed stronger negative impacts of fragmentation via reduced dispersal rates. Our results reveal that different components of the habitat fragmentation process have varying impacts on key mutualisms. We also conclude that antagonistic interactions have been understudied in fragmented landscapes, most of the research has concentrated on particular types of mutualistic interactions such as seed dispersal, and that available studies of interspecific interactions have a strong geographical bias (arising mostly from studies carried out in Brazil, Chile, and the United States). Meta‐Análisis de los Efectos de la Fragmentación del Bosque sobre las Interacciones Interespecíficas     

Nutrient regulation of organic matter decomposition in a tropical rain forest

Terrestrial biosphere-atmosphere CO2 exchange is dominated by tropical forests, so understanding how nutrient availability affects carbon (C) decomposition in these ecosystems is central to predicting the global C cycle's response to environmental change. In tropical rain forests, phosphorus (P) limitation of primary production and decomposition is believed to be widespread, but direct evidence is rare. We assessed the effects of nitrogen (N) and P fertilization on litter-layer organic matter decomposition in two neighboring tropical rain forests in southwest Costa Rica that are similar in most ways, but that differ in soil P availability. The sites contain 100-200 tree species per hectare and between species foliar nutrient content is variable. To control for this heterogeneity, we decomposed leaves collected from a widespread neotropical species, Brosimum utile. Mass loss during decomposition was rapid in both forests, with B. utile leaves losing >80% of their initial mass in <300 days. High organic matter solubility throughout decomposition combined with high rainfall support a model of litter-layer decomposition in these rain forests in which rapid mass loss in the litter layer is dominated by leaching of dissolved organic matter (DOM) rather than direct CO2 mineralization. While P fertilization did not significantly affect mass loss in the litter layer, it did stimulate P immobilization in decomposing material, leading to increased P content and a lower C:P ratio in soluble DOM. In turn, increased P content of leached DOM stimulated significant increases in microbial mineralization of DOM in P-fertilized soil. These results show that, while nutrients may not affect mass loss during decomposition in nutrient-poor, wet ecosystems, they may ultimately regulate CO2 losses (and hence C storage) by limiting microbial mineralization of DOM leached from the litter layer to soil.     

The impact of fragmentation and density regulation on forest succession in the Atlantic rain forest

The Atlantic Rain Forest, an important biodiversity hot spot, has faced severe habitat loss since the last century which has resulted in a highly fragmented landscape with a large number of small forest patches (<100 ha). For conservation planning it is essential to understand how current and future forest regeneration depends on ecological processes, fragment size and the connection to the regional seed pool. We have investigated the following questions by applying the forest growth simulation model FORMIND to the situation of the Atlantic Forest in the state of São Paulo, SE Brazil: (1) which set of parameters describing the local regeneration and level of density regulation can reproduce the biomass distribution and stem density of an old growth forest in a reserve? (2) Which additional processes apart from those describing the dynamics of an old growth forest, drive forest succession of small isolated fragments? (3) Which role does external seed input play during succession? Therefore, more than 300 tree species have been classified into nine plant functional types (PFTs), which are characterized by maximum potential height and shade tolerance. We differentiate between two seed dispersal modes: (i) local dispersal, i.e. all seedlings originated from fertile trees within the simulated area and (ii) external seed rain. Local seed dispersal has been parameterized following the pattern oriented approach, using biomass estimates of old growth forest. We have found that moderate density regulation is essential to achieve coexistence for a broad range of regeneration parameters. Considering the expected uncertainty and variability in the regeneration processes it is important that the forest dynamics are robust to variations in the regeneration parameters. Furthermore, edge effects such as increased mortality at the border and external seed rain have been necessary to reproduce the patterns for small isolated fragments. Overall, simulated biomass is much lower in the fragments compared to the continuous forest, whereas shade tolerant species are affected most strongly by fragmentation. Our simulations can supplement empirical studies by extrapolating local knowledge on edge effects of fragments to larger temporal and spatial scales. In particular our results show the importance of external seed rain and therefore highlight the importance of structural connectivity between regenerating fragments and mature forest stands.     

Responses of Seed‐Dispersing Birds to Amount of Rainforest in the Landscape around Fragments

Habitat loss and fragmentation alter the composition of bird assemblages in rainforest. Because birds are major seed dispersers in rainforests, fragmentation‐induced changes to frugivorous bird assemblages are also likely to alter the ecological processes of seed dispersal and forest regeneration, but the specific nature of these changes is poorly understood. We assessed the influence of fragment size and landscape forest cover on the abundance, species composition, and functional properties of the avian seed disperser community in an extensively cleared, former rainforest landscape of subtropical Australia. Bird surveys of fixed time and area in 25 rainforest fragments (1–139 ha in size across a 1800 km² region) provided bird assemblage data which were coupled with prior knowledge of bird species’ particular roles in seed dispersal to give measurements of seven different attributes of the seed disperser assemblage. We used multimodel regression to assess how patch size and surrounding forest cover (within 200 m, 1000 m, and 5000 m radii) influenced variation in the abundance of individual bird species and of functional groups based on bird species’ responses to fragmentation and their roles in seed dispersal. Surrounding forest cover, specifically rainforest cover, generally had a greater effect on frugivorous bird assemblages than fragment size. Amount of rainforest cover within 200 m of fragments was the main factor positively associated with abundances of frugivorous birds that are both fragmentation sensitive and important seed dispersers. Our results suggest a high proportion of local rainforest cover is required for the persistence of seed‐dispersing birds and the maintenance of seed dispersal processes. Thus, even small rainforest fragments can function as important parts of habitat networks for seed‐dispersing birds, whether or not they are physically connected by vegetation. Respuestas de Aves Dispersoras de Semillas al Incremento de Selvas en el Paisaje Alrededor de Fragmentos     

Directional changes in the species composition of a tropical forest

Long-term studies have revealed that the structure and dynamics of many tropical forests are changing, but the causes and consequences of these changes remain debated. To learn more about the forces driving changes within tropical forests, we investigated shifts in tree species composition over the past 25 years within the 50-ha Forest Dynamics Plot on Barro Colorado Island (BCI), Panama, and examined how observed patterns relate to predictions of (1) random population fluctuations, (2) carbon fertilization, (3) succession from past disturbance, (4) recovery from an extreme El Ni?o drought at the start of the study period, and (5) long-term climate change. We found that there have been consistent and directional changes in the tree species composition. These shifts have led to increased relative representations of drought-tolerant species as determined by the species' occurrence both across a gradient of soil moisture within BCI and across a wider precipitation gradient from a dry forest near the Pacific coast of Panama to a wet forest near its Caribbean coast. These nonrandom changes cannot be explained by stochastic fluctuations or carbon fertilization. They may be the legacy of the El Ni?o drought, or alternatively, potentially reflect increased aridity due to long-term climate change. By investigating compositional changes, we increased not only our understanding of the ecology of tropical forests and their responses to large-scale disturbances, but also our ability to predict how future global change will impact some of the critical services provided by these important ecosystems.     

Changes in microbial community characteristics and soil organic matter with nitrogen additions in two tropical forests

Microbial communities and their associated enzyme activities affect the amount and chemical quality of carbon (C) in soils. Increasing nitrogen (N) deposition, particularly in N-rich tropical forests, is likely to change the composition and behavior of microbial communities and feed back on ecosystem structure and function. This study presents a novel assessment of mechanistic links between microbial responses to N deposition and shifts in soil organic matter (SOM) quality and quantity. We used phospholipid fatty acid (PLFA) analysis and microbial enzyme assays in soils to assess microbial community responses to long-term N additions in two distinct tropical rain forests. We used soil density fractionation and 13C nuclear magnetic resonance (NMR) spectroscopy to measure related changes in SOM pool sizes and chemical quality. Microbial biomass increased in response to N fertilization in both tropical forests and corresponded to declines in pools of low-density SOM. The chemical quality of this soil C pool reflected ecosystem-specific changes in microbial community composition. In the lower-elevation forest, there was an increase in gram-negative bacteria PLFA biomass, and there were significant losses of labile C chemical groups (O-alkyls). In contrast, the upper-elevation tropical forest had an increase in fungal PLFAs with N additions and declines in C groups associated with increased soil C storage (alkyls). The dynamics of microbial enzymatic activities with N addition provided a functional link between changes in microbial community structure and SOM chemistry. Ecosystem-specific changes in microbial community composition are likely to have far-reaching effects on soil carbon storage and cycling. This study indicates that microbial communities in N-rich tropical forests can be sensitive to added N, but we can expect significant variability in how ecosystem structure and function respond to N deposition among tropical forest types.