Defaunation impacts on seed survival and its effect on the biomass of future tropical forests

Large animal species, which provide important ecological functions such as dispersal of seeds or top–down control of seed predators, are very vulnerable in fragmented forests, being unable to survive in small fragments, and facing increasing hunting pressure. The loss of large animals affects two main ecological processes crucial for the tree reproductive cycle: seed dispersal of large seeds (e.g. provided by tapirs) and control of seed predator population (e.g. provided by large cats). The changes in both processes are expected to increase seed mortality since seeds are not dispersed away from conspecifics (causing increased pre‐dispersal mortality due to negative density dependent effects) and/or face increased predation after a dispersal event (post‐dispersal mortality). Although an extensive body of empirical knowledge exists on seed predation, the link between seed loss and adult tree community composition and structure is not well established, as well as the temporal scale seed changes affect adults. Using an individual‐based forest model (FORMIND), we evaluate the long‐term consequences of increased pre and post‐dispersal seed mortality on the future forest biomass retention of a Brazilian northeastern Atlantic forest. Our results show that forest biomass is significantly affected after 80–93% pre‐dispersal loss of large seeds, or post‐dispersal predation densities of 20–25 predators per parent tree. Large‐seeded tree species are at increased risk of local extinction causing up to 26.2% loss of forest biomass when both pre and post‐dispersal processes are combined. However, these changes can last up to 100 years after the occurrence of defaunation. In summary we conclude that large animal loss has the potential to reduce future forest biomass and tree species‐richness by impacting seed survival, and should be considered in the planning of biodiversity friendly landscapes as well as in calculations of the global carbon budget.     

Hunting pressure modulates the composition and size structure of terrestrial and arboreal vertebrates in Amazonian forests

Overhunting is a leading contemporary driver of tropical forest wildlife loss. The absence or extremely low densities of large-bodied vertebrates disrupts plant-animal mutualisms and consequently degrades key ecosystem services. Understanding patterns of defaunation is therefore crucial given that most tropical forests worldwide are now “half-empty”. Here we investigate changes in vertebrate community composition and size structure along a gradient of marked anthropogenic hunting pressure in the Médio Juruá region of western Brazilian Amazonia. Using a novel camera trapping grid design deployed both in the understorey and the forest canopy, we estimated the aggregate biomass of several functional groups of terrestrial and arboreal species at 28 sites along the hunting gradient. Generalized linear models (GLMs) identified hunting pressure as the most important driver of aggregate biomass for game, terrestrial, and arboreal species, as well as nocturnal rodents, frugivores, and granivores. Local hunting pressure affected vertebrate community structure as shown by both GLM and ordination analyses. The size structure of vertebrate fauna changed in heavily hunted areas due to population declines in large-bodied species and apparent compensatory increases in nocturnal rodents. Our study shows markedly altered vertebrate community structure even in remote but heavily settled areas of continuous primary forest. Depletion of frugivore and granivore populations, and concomitant density-compensation by seed predators, likely affect forest regeneration in persistently overhunted tropical forests. These findings contribute to a better understanding of how cascading effects induced by historical defaunation operate, informing wildlife management policy in tropical peri-urban, rural and wilderness areas.

     

Changes in tree community structure in defaunated forests are not driven only by dispersal limitation

1. Bushmeat hunting has reduced population sizes of large frugivorous vertebrates throughout the tropics, thereby reducing the dispersal of seeds. This is believed to affect tree population dynamics, and therefore community composition, because the seed dispersal of large‐seeded trees depends upon large‐bodied vertebrates.
2. We report on a long‐running study of the effect of defaunation on a tropical tree community. In three censuses over 11 years, we compared sapling recruitment between a hunted and a nonhunted site, which are nearby and comparable to one another, to determine the extent to which species composition has changed through time following defaunation. We expected to find a reduced abundance of tree species that rely on large frugivores for dispersal at the hunted site and altered community structure as a consequence.
3. Although community composition at the hunted site diverged from that at the nonhunted site, the changes were independent of dispersal syndrome, with no trend toward a decline in species that are dispersed by large, hunted vertebrates. Moreover, the loss of large‐bodied dispersers did not generate the changes in tree community composition that we hypothesized. Some species presumed to rely on large‐bodied frugivores for dispersal are effectively recruiting despite the absence of their dispersers.
4. Synthesis: The presumption that forests depleted of large‐bodied dispersers will experience rapid, directional compositional change is not fully supported by our results. Altered species composition in the sapling layer at the hunted site, however, indicates that defaunation may be connected with changes to the tree community, but that the nature of these changes is not unidirectional as previously assumed. It remains difficult to predict how defaunation will affect tree community composition without a deeper understanding of the driving mechanisms at play.

     

Improving predictions of tropical forest response to climate change through integration of field studies and ecosystem modeling

Tropical forests play a critical role in carbon and water cycles at a global scale. Rapid climate change is anticipated in tropical regions over the coming decades and, under a warmer and drier climate, tropical forests are likely to be net sources of carbon rather than sinks. However, our understanding of tropical forest response and feedback to climate change is very limited. Efforts to model climate change impacts on carbon fluxes in tropical forests have not reached a consensus. Here, we use the Ecosystem Demography model (ED2) to predict carbon fluxes of a Puerto Rican tropical forest under realistic climate change scenarios. We parameterized ED2 with species‐specific tree physiological data using the Predictive Ecosystem Analyzer workflow and projected the fate of this ecosystem under five future climate scenarios. The model successfully captured interannual variability in the dynamics of this tropical forest. Model predictions closely followed observed values across a wide range of metrics including aboveground biomass, tree diameter growth, tree size class distributions, and leaf area index. Under a future warming and drying climate scenario, the model predicted reductions in carbon storage and tree growth, together with large shifts in forest community composition and structure. Such rapid changes in climate led the forest to transition from a sink to a source of carbon. Growth respiration and root allocation parameters were responsible for the highest fraction of predictive uncertainty in modeled biomass, highlighting the need to target these processes in future data collection. Our study is the first effort to rely on Bayesian model calibration and synthesis to elucidate the key physiological parameters that drive uncertainty in tropical forests responses to climatic change. We propose a new path forward for model‐data synthesis that can substantially reduce uncertainty in our ability to model tropical forest responses to future climate.     

Fingerprinting the impacts of global change on tropical forests

Recent observations of widespread changes in mature tropical forests such as increasing tree growth, recruitment and mortality rates and increasing above-ground biomass suggest that ''global change'' agents may be causing predictable changes in tropical forests. However, consensus over both the robustness of these changes and the environmental drivers that may be causing them is yet to emerge. This paper focuses on the second part of this debate. We review (i) the evidence that the physical, chemical and biological environment that tropical trees grow in has been altered over recent decades across large areas of the tropics, and (ii) the theoretical, experimental and observational evidence regarding the most likely effects of each of these changes on tropical forests. Ten potential widespread drivers of environmental change were identified: temperature, precipitation, solar radiation, climatic extremes (including El Niño-Southern Oscillation events), atmospheric CO2 concentrations, nutrient deposition, O3/acid depositions, hunting, land-use change and increasing liana numbers. We note that each of these environmental changes is expected to leave a unique ''fingerprint'' in tropical forests, as drivers directly force different processes, have different distributions in space and time and may affect some forests more than others (e.g. depending on soil fertility). Thus, in the third part of the paper we present testable a priori predictions of forest responses to assist ecologists in attributing particular changes in forests to particular causes across multiple datasets. Finally, we discuss how these drivers may change in the future and the possible consequences for tropical forests.     

Large herbivores regulate the spatial recruitment of a hyperdominant Neotropical palm

Large mammalian herbivores play an important role in shaping the diversity of tropical forests by affecting the survival of seedlings and saplings beneath parent plants. The white‐lipped peccary (Tayassu pecari) accounts for the largest herbivore biomass that controls seed and seedling survival in Neotropical ecosystems. However, hunting and habitat loss has driven peccaries to local extinction for most of their original distribution, so it is likely that their absence will affect plant recruitment dynamics. We tested the effects of peccary local extinction on the density and spatial distribution of the hyperdominant palm Euterpe edulis by performing a fine‐scale characterization of its spatial recruitment in six forest sites in the Brazilian Atlantic forest. We compared the age structure and the spatial patterns of seedlings, saplings, and adults as well as the relationship between them. We found that while under the presence of peccaries there was a decrease in recruitment rates under adults, the local extinction of these large mammals led to a more clumped process of spatial recruitment. Despite such contrasting spatial patterns of recruitment dynamics, neither age structure nor the random spatial distribution of adults was affected by the presence or absence of peccaries, indicating that their early effects on these palm populations are mitigated as recruitment advances. Our findings highlight the role of large‐bodied forest‐dwelling herbivores in regulating the fine‐scale spatial recruitment of plants and advance our understanding on the effects of defaunation in tropical forests. Abstract in Portuguese is available with online material.     

Logging and indigenous hunting impacts on persistence of large Neotropical animals

Areas allocated for industrial logging and community‐owned forests account for over 50% of all remaining tropical forests. Landscape‐scale conservation strategies that include these forests are expected to have substantial benefits for biodiversity, especially for large mammals and birds that require extensive habitat but that are susceptible to extirpation due to synergies between logging and hunting. In addition, their responses to logging alone are poorly understood due to their cryptic behavior and low densities. In this study, we assessed the effects of logging and hunting on detection and occupancy rates of large vertebrates in a multiple‐use forest on the Guiana Shield. Our study site was certified as being responsibly managed for timber production and indigenous communities are legally guaranteed use‐rights to the forest. We coupled camera‐trap data for wildlife detection with a spatially explicit dataset on indigenous hunting. A multi‐species occupancy model found a weak positive effect of logging on occupancy and detection rates, while hunting had a weak negative effect. Model predictions of species richness were also higher in logged forest sites compared to unlogged forest sites. Density estimates for jaguars and ocelots in our multiple‐use area were similar to estimates reported for fully protected areas. Involvement of local communities in forest management, control of forest access, and nesting production forests in a landscape that includes protected areas seemed important for these positive biodiversity outcomes. The maintenance of vertebrate species bodes well for both biodiversity and the humans that depend on multiple‐use forests.     

Allometric constraints and competition enable the simulation of size structure and carbon fluxes in a dynamic vegetation model of tropical forests (LM3PPA‐TV)

Tropical forests are a key determinant of the functioning of the Earth system, but remain a major source of uncertainty in carbon cycle models and climate change projections. In this study, we present an updated land model (LM3PPA‐TV) to improve the representation of tropical forest structure and dynamics in Earth system models (ESMs). The development and parameterization of LM3PPA‐TV drew on extensive datasets on tropical tree traits and long‐term field censuses from Barro Colorado Island (BCI), Panama. The model defines a new plant functional type (PFT) based on the characteristics of shade‐tolerant, tropical tree species, implements a new growth allocation scheme based on realistic tree allometries, incorporates hydraulic constraints on biomass accumulation, and features a new compartment for tree branches and branch fall dynamics. Simulation experiments reproduced observed diurnal and seasonal patterns in stand‐level carbon and water fluxes, as well as mean canopy and understory tree growth rates, tree size distributions, and stand‐level biomass on BCI. Simulations at multiple sites captured considerable variation in biomass and size structure across the tropical forest biome, including observed responses to precipitation and temperature. Model experiments suggested a major role of water limitation in controlling geographic variation forest biomass and structure. However, the failure to simulate tropical forests under extreme conditions and the systematic underestimation of forest biomass in Paleotropical locations highlighted the need to incorporate variation in hydraulic traits and multiple PFTs that capture the distinct floristic composition across tropical domains. The continued pressure on tropical forests from global change demands models which are able to simulate alternative successional pathways and their pace to recovery. LM3PPA‐TV provides a tool to investigate geographic variation in tropical forests and a benchmark to continue improving the representation of tropical forests dynamics and their carbon storage potential in ESMs.     

No evidence for consistent long‐term growth stimulation of 13 tropical tree species: results from tree‐ring analysis

The important role of tropical forests in the global carbon cycle makes it imperative to assess changes in their carbon dynamics for accurate projections of future climate–vegetation feedbacks. Forest monitoring studies conducted over the past decades have found evidence for both increasing and decreasing growth rates of tropical forest trees. The limited duration of these studies restrained analyses to decadal scales, and it is still unclear whether growth changes occurred over longer time scales, as would be expected if CO₂‐fertilization stimulated tree growth. Furthermore, studies have so far dealt with changes in biomass gain at forest‐stand level, but insights into species‐specific growth changes – that ultimately determine community‐level responses – are lacking. Here, we analyse species‐specific growth changes on a centennial scale, using growth data from tree‐ring analysis for 13 tree species (~1300 trees), from three sites distributed across the tropics. We used an established (regional curve standardization) and a new (size‐class isolation) growth‐trend detection method and explicitly assessed the influence of biases on the trend detection. In addition, we assessed whether aggregated trends were present within and across study sites. We found evidence for decreasing growth rates over time for 8–10 species, whereas increases were noted for two species and one showed no trend. Additionally, we found evidence for weak aggregated growth decreases at the site in Thailand and when analysing all sites simultaneously. The observed growth reductions suggest deteriorating growth conditions, perhaps due to warming. However, other causes cannot be excluded, such as recovery from large‐scale disturbances or changing forest dynamics. Our findings contrast growth patterns that would be expected if elevated CO₂ would stimulate tree growth. These results suggest that commonly assumed growth increases of tropical forests may not occur, which could lead to erroneous predictions of carbon dynamics of tropical forest under climate change.     

Large tree mortality and the decline of forest biomass following Amazonian wildfires

Surface fires in Amazonian forests could contribute as much as 5% of annual carbon emissions from all anthropogenic sources during severe El Niño years. However, these estimates are based on short‐term figures of post‐burn tree mortality, when large thicker barked trees (representing a disproportionate amount of the forest biomass) appear to resist the fires. On the basis of a longer term study, we report that the mortality of large trees increased markedly between 1 and 3 years, more than doubling current estimates of biomass loss and committed carbon emissions from low‐intensity fires in tropical forests.     

Ecological responses to el Niño-induced surface fires in central Brazilian Amazonia: management implications for flammable tropical forests

Over the past 20 years the combined effects of El Niño-induced droughts and land-use change have dramatically increased the frequency of fire in humid tropical forests. Despite the potential for rapid ecosystem alteration and the current prevalence of wildfire disturbance, the consequences of such fires for tropical forest biodiversity remain poorly understood. We provide a pan-tropical review of the current state of knowledge of these fires, and include data from a study in a seasonally dry terra firme forest of central Brazilian Amazonia. Overall, this study supports predictions that rates of tree mortality and changes in forest structure are strongly linked to burn severity. The potential consequences for biomass loss and carbon emissions are explored. Despite the paucity of data on faunal responses to tropical forest fires, some trends are becoming apparent; for example, large canopy frugivores and understorey insectivorous birds appear to be highly sensitive to changes in forest structure and composition during the first 3 years after fires. Finally, we appraise the management implications of fires and evaluate the viability of techniques and legislation that can be used to reduce forest flammability, prevent anthropogenic ignition sources from coming into contact with flammable forests and aid the post-fire recovery process.     

Tree Species Richness Strengthens Relationships between Ants and the Functional Composition of Spider Assemblages in a Highly Diverse Forest

In species‐rich ecosystems, such as subtropical and tropical forests, higher trophic level interactions are key mediators of ecosystem functioning. Plant species loss may alter these interactions, but the effects of plant diversity might be modified by intraguild interactions, particularly among predators. We analyzed the relationships between spiders and ants, two dominant predatory arthropod taxa, on tree saplings across a gradient from medium to high woody plant species richness in a subtropical forest in Southeast China. Neither ant nor spider total biomass was significantly related to plant species richness. By contrast, the biomass distribution of web‐building and hunting spiders changed and spider family richness increased in the presence of ants, resulting in more web builder‐dominated assemblages. However, these relationships depended on the plant communities, and were stronger in plots with higher plant species richness. Our results indicate that in addition to potential effects of ants on hunting spiders in particular, ants could indirectly influence intraguild interactions within spider assemblages. The observed shifts in the spider assemblages with increasing ant presence and plant species richness may have functional consequences, as web‐building and hunting spiders have distinct prey spectra. The relationships among ants, spiders, and plant species richness might contribute to explaining the non‐significant relationship between the overall effects of predators and plant diversity previously observed in the same forest plots. Our findings thus give insight into the complexity of biotic interactions in such species‐rich ecosystems.     

Defaunation and fragmentation erode small mammal diversity dimensions in tropical forests

Forest fragmentation and defaunation are considered the main drivers of biodiversity loss, yet the synergistic effects of landscape changes and biotic interactions on assemblage structure have been poorly investigated. Here, we use an extensive dataset of 283 assemblages and 105 species of small mammals to understand how defaunation of medium and large mammals and forest fragmentation change the community composition and diversity of rodents and marsupials in tropical forests of South America. We used structured equation models to investigate the relationship between small mammal species, functional and phylogenetic diversity with forest size, forest cover and the occurrence of medium and large mammals. The best‐fit model showed that defaunation reduced functional diversity, and that species diversity of small mammals increased with forest patch size. Forest cover did not affect functional and phylogenetic diversity. Our results indicate that occurrence of medium and large sized mammals (probably acting as predators, or competitors of small mammals) and forest patch size help to retain species and functional diversity in small mammal communities. Further, the number of species in a small mammal community was critical to the maintenance of phylogenetic diversity, and may have a pronounced influence on the ecological functions played by small mammals. Identifying how phylogenetic and functional diversity change in function of human pressures allows us to better understand the contribution of extant lineages to ecosystem functioning in tropical forests.     

Pantropical geography of lightning‐caused disturbance and its implications for tropical forests

Lightning is a major agent of disturbance, but its ecological effects in the tropics are unquantified. Here we used ground and satellite sensors to quantify the geography of lightning strikes in terrestrial tropical ecosystems, and to evaluate whether spatial variation in lightning frequency is associated with variation in tropical forest structure and dynamics. Between 2013 and 2018, tropical terrestrial ecosystems received an average of 100.4 million lightning strikes per year, and the frequency of strikes was spatially autocorrelated at local‐to‐continental scales. Lightning strikes were more frequent in forests, savannas, and urban areas than in grasslands, shrublands, and croplands. Higher lightning frequency was positively associated with woody biomass turnover and negatively associated with aboveground biomass and the density of large trees (trees/ha) in forests across Africa, Asia, and the Americas. Extrapolating from the only tropical forest study that comprehensively assessed tree damage and mortality from lightning strikes, we estimate that lightning directly damages c. 832 million trees in tropical forests annually, of which c. 194 million die. The similarly high lightning frequency in tropical savannas suggests that lightning also influences savanna tree mortality rates and ecosystem processes. These patterns indicate that lightning‐caused disturbance plays a major and largely unappreciated role in pantropical ecosystem dynamics and global carbon cycling.     

Variations in dung beetle assemblages across a gradient of hunting in a tropical forest

Populations of large mammals are severely depleted by hunting in tropical forests, with direct effects on plant regeneration. But indirect consequences on commensal taxa depending on them for food resources, like coprophagous beetles, are less documented. Cascading effects of species loss across Scarabaeinae are expected, with likely significant negative implications for ecosystem functions. We examined dung beetle assemblages using pitfall traps at three rain forest sites in French Guiana ranging from intact mammalian fauna (Nouragues) to moderate (Kaw) and heavy (Matoury) defaunation. The site with the most depauperate mammalian fauna showed significantly lower dung beetle species richness than the two other two sites, which were not different from each other. Mean abundance and biomass per trap were not different across sites whereas community composition strongly differed among sites. A positive correlation was observed between body size and the individual contribution to dissimilarity between Nouragues and Kaw. The species contributing the most to dissimilarity were large. By contrast, one medium-sized species, dominant in Matoury, contributed the most to dissimilarity between Matoury and other sites. Diurnal genera of large tunnellers showed a higher diversity and abundance in Nouragues compared to other sites, whereas a nocturnal genus showed no differences. Large rollers were more abundant in Kaw compared to other sites. None of the groups of small beetles but one were affected by defaunation. Our results suggest that loss of large mammal populations affects dung beetle assemblage structure and causes decreasing abundance or disappearance of large tunnellers species that have a major impact on several dung beetle-mediated ecological processes.     

A climate change‐induced threat to the ecological resilience of a subtropical monsoon evergreen broad‐leaved forest in Southern China

Recent studies have suggested that tropical forests may not be resilient against climate change in the long term, primarily owing to predicted reductions in rainfall and forest productivity, increased tree mortality, and declining forest biomass carbon sinks. These changes will be caused by drought‐induced water stress and ecosystem disturbances. Several recent studies have reported that climate change has increased tree mortality in temperate and boreal forests, or both mortality and recruitment rates in tropical forests. However, no study has yet examined these changes in the subtropical forests that account for the majority of China's forested land. In this study, we describe how the monsoon evergreen broad‐leaved forest has responded to global warming and drought stress using 32 years of data from forest observation plots. Due to an imbalance in mortality and recruitment, and changes in diameter growth rates between larger and smaller trees and among different functional groups, the average DBH of trees and forest biomass have decreased. Sap flow measurements also showed that larger trees were more stressed than smaller trees by the warming and drying environment. As a result, the monsoon evergreen broad‐leaved forest community is undergoing a transition from a forest dominated by a cohort of fewer and larger individuals to a forest dominated by a cohort of more and smaller individuals, with a different species composition, suggesting that subtropical forests are threatened by their lack of resilience against long‐term climate change.     

Global‐scale impacts of nitrogen deposition on tree carbon sequestration in tropical,temperate, and boreal forests: A meta‐analysis

Elevated nitrogen (N) deposition may increase net primary productivity in N‐limited terrestrial ecosystems and thus enhance the terrestrial carbon (C) sink. To assess the magnitude of this N‐induced C sink, we performed a meta‐analysis on data from forest fertilization experiments to estimate N‐induced C sequestration in aboveground tree woody biomass, a stable C pool with long turnover times. Our results show that boreal and temperate forests responded strongly to N addition and sequestered on average an additional 14 and 13 kg C per kg N in aboveground woody biomass, respectively. Tropical forests, however, did not respond significantly to N addition. The common hypothesis that tropical forests do not respond to N because they are phosphorus‐limited could not be confirmed, as we found no significant response to phosphorus addition in tropical forests. Across climate zones, we found that young forests responded more strongly to N addition, which is important as many previous meta‐analyses of N addition experiments rely heavily on data from experiments on seedlings and young trees. Furthermore, the C–N response (defined as additional mass unit of C sequestered per additional mass unit of N addition) was affected by forest productivity, experimental N addition rate, and rate of ambient N deposition. The estimated C–N responses from our meta‐analysis were generally lower that those derived with stoichiometric scaling, dynamic global vegetation models, and forest growth inventories along N deposition gradients. We estimated N‐induced global C sequestration in tree aboveground woody biomass by multiplying the C–N responses obtained from the meta‐analysis with N deposition estimates per biome. We thus derived an N‐induced global C sink of about 177 (112–243) Tg C/year in aboveground and belowground woody biomass, which would account for about 12% of the forest biomass C sink (1,400 Tg C/year).     

Tree allometry and improved estimation of carbon stocks and balance in tropical forests

Tropical forests hold large stores of carbon, yet uncertainty remains regarding their quantitative contribution to the global carbon cycle. One approach to quantifying carbon biomass stores consists in inferring changes from long-term forest inventory plots. Regression models are used to convert inventory data into an estimate of aboveground biomass (AGB). We provide a critical reassessment of the quality and the robustness of these models across tropical forest types, using a large dataset of 2,410 trees ≥ 5 cm diameter, directly harvested in 27 study sites across the tropics. Proportional relationships between aboveground biomass and the product of wood density, trunk cross-sectional area, and total height are constructed. We also develop a regression model involving wood density and stem diameter only. Our models were tested for secondary and old-growth forests, for dry, moist and wet forests, for lowland and montane forests, and for mangrove forests. The most important predictors of AGB of a tree were, in decreasing order of importance, its trunk diameter, wood specific gravity, total height, and forest type (dry, moist, or wet). Overestimates prevailed, giving a bias of 0.5–6.5% when errors were averaged across all stands. Our regression models can be used reliably to predict aboveground tree biomass across a broad range of tropical forests. Because they are based on an unprecedented dataset, these models should improve the quality of tropical biomass estimates, and bring consensus about the contribution of the tropical forest biome and tropical deforestation to the global carbon cycle. Electronic Supplementary Material Supplementary material is available for this article at     

Tree diversity,composition, forest structure and aboveground biomass dynamics after single and repeated fire in a Bornean rain forest

Forest fires remain a devastating phenomenon in the tropics that not only affect forest structure and biodiversity, but also contribute significantly to atmospheric CO2. Fire used to be extremely rare in tropical forests, leaving ample time for forests to regenerate to pre-fire conditions. In recent decades, however, tropical forest fires occur more frequently and at larger spatial scales than they used to. We studied forest structure, tree species diversity, tree species composition, and aboveground biomass during the first 7 years since fire in unburned, once burned and twice burned forest of eastern Borneo to determine the rate of recovery of these forests. We paid special attention to changes in the tree species composition during burned forest regeneration because we expect the long-term recovery of aboveground biomass and ecosystem functions in burned forests to largely depend on the successful regeneration of the pre-fire, heavy-wood, species composition. We found that forest structure (canopy openness, leaf area index, herb cover, and stem density) is strongly affected by fire but shows quick recovery. However, species composition shows no or limited recovery and aboveground biomass, which is greatly reduced by fire, continues to be low or decline up to 7 years after fire. Consequently, large amounts of the C released to the atmosphere by fire will not be recaptured by the burned forest ecosystem in the near future. We also observed that repeated fire, with an inter-fire interval of 15 years, does not necessarily lead to a huge deterioration in the regeneration potential of tropical forest. We conclude that burned forests are valuable and should be conserved and that long-term monitoring programs in secondary forests are necessary to determine their recovery rates, especially in relation to aboveground biomass accumulation.     

Changes of community composition at multiple trophic levels due to hunting in Nigerian tropical forests

Hunting in tropical forests decimates large mammals, and this may have direct and indirect effects on other trophic levels and lead to trophic cascades. We compared replicated sites of hunted and protected forests in southeastern Nigeria, with respect to community composition of primates, other mammals, birds, plant seedlings, and mature trees. We make predictions regarding the community composition at the different trophic levels. In forests where large primates are rare, we hypothesize that their ecological role will not be fully compensated for by small frugivores. We apply multivariate methods to assess changes in community composition of mammals, birds, and seedlings, controlling for any differences between sites in the other groups, including mature trees. Medium and large (4–180 kg) primates were much rarer in hunted sites, while porcupine and rock hyrax increased in abundance with hunting. In contrast, the community composition of birds was similar in both types of forests. Seedling communities were significantly related to the community composition of mammals, and thus strongly affected by hunting. In protected forests primate dispersed plant seedling species dominated, whereas in hunted forests the seedling community was shifted towards one dominated by abiotically dispersed species. This was probably both a consequence of reduced seed dispersal by primates, and increased seed predation by rodents and hyrax. Hence we found no evidence for buffering effects on tree regeneration through functional compensation by non‐hunted animals (such as birds). Our results highlight how seedling communities are changed by the complex plant–animal intera ctions, triggered by the loss of seed dispersers. The results predict a rarity of primate‐dispersed trees in future tropical forest canopies; a forest less diverse in timber and non‐timber resources.