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Global change exposes forest ecosystems to many risks including novel climatic conditions, increased frequency of climatic extremes and sudden emergence and spread of pests and pathogens. At the same time, forest landscape restoration has regained global attention as an integral strategy for climate change mitigation. Owing to unpredictable future risks and the need for new forests that provide multiple ecosystem services, mixed-species forests have been advocated for this purpose. However, the successful establishment of mixed forests requires intrinsic knowledge of biodiversity's role for forest ecosystem functioning. In this respect, a better understanding of tree-tree interactions and how they contribute to observed positive tree species richness effects on key ecosystem functions is critical. Here, we review the current knowledge of the underlying mechanisms of tree-tree interactions and argue that positive net biodiversity effects at the community scale may emerge from the dominance of positive over negative interactions at the local neighbourhood scale. In a second step, we demonstrate how tree-tree interactions and the immediate tree neighbourhood's role can be systematically assessed in a tree diversity experiment. The expected results will improve predictions about the effects of tree interactions on ecosystem functioning based on general principles. We argue that this knowledge is urgently required to guide the design of tree species mixtures for the successful establishment of newly planted forests.

Plant-soil feedbacks have been recognised as playing a key role in a range of ecological processes, including succession, invasion, species coexistence and population dynamics. However, there is substantial variation between species in the strength of plant-soil feedbacks and predicting this variation remains challenging. Here, we propose an original concept to predict the outcome of plant-soil feedbacks. We hypothesize that plants with different root traits culture different proportions of pathogens and mutualists and this contributes to differences in performance between home soils (cultured by conspecifics) versus away soils (cultured by heterospecifics). We use the recently described root economics spectrum, which identifies two gradients in root traits. A conservation gradient distinguishes fast vs. slow species, and from the growth defence theory we predict that these species culture different amounts of pathogens in their soils. A collaboration gradient distinguishes species that associate with mycorrhizae to outsource soil nutrient acquisition vs. those which use a “do it yourself” strategy and capture nutrients without mycorrhizae. We provide a framework, which predicts that the strength and direction of feedback between a pair of species is determined by the functional distance between them along each axis. We then use data from two case studies to show how to apply the framework, by analysing the response of plant-soil feedbacks to measures of distance and position along each axis and find support for our predictions. Finally, we highlight further areas where our framework could be developed and propose study designs that would help to fill current research gaps.

Plant‐soil feedbacks (PSFs) have been shown to strongly affect plant performance under controlled conditions, and PSFs are thought to have far reaching consequences for plant population dynamics and the structuring of plant communities. However, thus far the relationship between PSF and plant species abundance in the field is not consistent. Here, we synthesize PSF experiments from tropical forests to semiarid grasslands, and test for a positive relationship between plant abundance in the field and PSFs estimated from controlled bioassays. We meta‐analyzed results from 22 PSF experiments and found an overall positive correlation (0.12 ≤ r¯ ≤ 0.32) between plant abundance in the field and PSFs across plant functional types (herbaceous and woody plants) but also variation by plant functional type. Thus, our analysis provides quantitative support that plant abundance has a general albeit weak positive relationship with PSFs across ecosystems. Overall, our results suggest that harmful soil biota tend to accumulate around and disproportionately impact species that are rare. However, data for the herbaceous species, which are most common in the literature, had no significant abundance‐PSFs relationship. Therefore, we conclude that further work is needed within and across biomes, succession stages and plant types, both under controlled and field conditions, while separating PSF effects from other drivers (e.g., herbivory, competition, disturbance) of plant abundance to tease apart the role of soil biota in causing patterns of plant rarity versus commonness., Across studies, we detected a small but significant positive correlation between plant‐soil feedbacks measured during controlled experiments and plant field abundance. Plants that are rare are seemingly burdened more by the negative effects of harmful soil biota than common plants.

Aim Palms are an iconic, diverse and often abundant component of tropical ecosystems that provide many ecosystem services. Being monocots, tree palms are evolutionarily, morphologically and physiologically distinct from other trees, and these differences have important consequences for ecosystem services (e.g., carbon sequestration and storage) and in terms of responses to climate change. We quantified global patterns of tree palm relative abundance to help improve understanding of tropical forests and reduce uncertainty about these ecosystems under climate change. Location Tropical and subtropical moist forests. Time period Current. Major taxa studied Palms (Arecaceae). Methods We assembled a pantropical dataset of 2,548 forest plots (covering 1,191ha) and quantified tree palm (i.e., ≥10cm diameter at breast height) abundance relative to co‐occurring non‐palm trees. We compared the relative abundance of tree palms across biogeographical realms and tested for associations with palaeoclimate stability, current climate, edaphic conditions and metrics of forest structure. Results On average, the relative abundance of tree palms was more than five times larger between Neotropical locations and other biogeographical realms. Tree palms were absent in most locations outside the Neotropics but present in >80% of Neotropical locations. The relative abundance of tree palms was more strongly associated with local conditions (e.g., higher mean annual precipitation, lower soil fertility, shallower water table and lower plot mean wood density) than metrics of long‐term climate stability. Life‐form diversity also influenced the patterns; palm assemblages outside the Neotropics comprise many non‐tree (e.g., climbing) palms. Finally, we show that tree palms can influence estimates of above‐ground biomass, but the magnitude and direction of the effect require additional work. Conclusions Tree palms are not only quintessentially tropical, but they are also overwhelmingly Neotropical. Future work to understand the contributions of tree palms to biomass estimates and carbon cycling will be particularly crucial in Neotropical forests.

Agriculture and the exploitation of natural resources have transformed tropical mountain ecosystems across the world, and the consequences of these transformations for biodiversity and ecosystem functioning are largely unknown1-3. Conclusions that are derived from studies in non-mountainous areas are not suitable for predicting the effects of land-use changes on tropical mountains because the climatic environment rapidly changes with elevation, which may mitigate or amplify the effects of land use4,5. It is of key importance to understand how the interplay of climate and land use constrains biodiversity and ecosystem functions to determine the consequences of global change for mountain ecosystems. Here we show that the interacting effects of climate and land use reshape elevational trends in biodiversity and ecosystem functions on Africa's largest mountain, Mount Kilimanjaro (Tanzania). We find that increasing land-use intensity causes larger losses of plant and animal species richness in the arid lowlands than in humid submontane and montane zones. Increases in land-use intensity are associated with significant changes in the composition of plant, animal and microorganism communities; stronger modifications of plant and animal communities occur in arid and humid ecosystems, respectively. Temperature, precipitation and land use jointly modulate soil properties, nutrient turnover, greenhouse gas emissions, plant biomass and productivity, as well as animal interactions. Our data suggest that the response of ecosystem functions to land-use intensity depends strongly on climate; more-severe changes in ecosystem functioning occur in the arid lowlands and the cold montane zone. Interactions between climate and land use explained-on average-54% of the variation in species richness, species composition and ecosystem functions, whereas only 30% of variation was related to single drivers. Our study reveals that climate can modulate the effects of land use on biodiversity and ecosystem functioning, and points to a lowered resistance of ecosystems in climatically challenging environments to ongoing land-use changes in tropical mountainous regions.

Many experiments have shown that biodiversity enhances ecosystem functioning. However, we have little understanding of how environmental heterogeneity shapes the effect of diversity on ecosystem functioning and to what extent this diversity effect is mediated by variation in species richness or species turnover. This knowledge is crucial to scaling up the results of experiments from local to regional scales. Here we quantify the diversity effect and its components-that is, the contributions of variation in species richness and species turnover-for 22 ecosystem functions of microorganisms, plants and animals across 13 major ecosystem types on Mt Kilimanjaro, Tanzania. Environmental heterogeneity across ecosystem types on average increased the diversity effect from explaining 49% to 72% of the variation in ecosystem functions. In contrast to our expectation, the diversity effect was more strongly mediated by variation in species richness than by species turnover. Our findings reveal that environmental heterogeneity strengthens the relationship between biodiversity and ecosystem functioning and that species richness is a stronger driver of ecosystem functioning than species turnover. Based on a broad range of taxa and ecosystem functions in a non-experimental system, these results are in line with predictions from biodiversity experiments and emphasize that conserving biodiversity is essential for maintaining ecosystem functioning.

BackgroundRecent evidence suggests that soil microbial communities can regulate plant community dynamics. In addition, the drought tolerance of plants can be enhanced by soil microbes. So far, few studies have assessed the variation in the microbiome of specific plant species along environmental gradients. Yet understanding these dynamics is essential to improve predictions of plant-soil feedbacks and the consequences of ongoing climate changes. Here we characterized the soil microbiome of two co-occurring Mediterranean oaks along a precipitation gradient, using amplicon sequencing of phylogenetic marker genes for prokaryotes and fungi. Additionally, we identified tree-specific and locally-specific microbes potentially responsible for tree community dynamics.ResultsWe show that two co-occurring, evergreen Mediterranean oak species harbor distinct microbiomes along a precipitation gradient. The soil microbial diversity increased along the precipitation gradient, for prokaryotic α and β diversity and for fungal β diversity. Quercus ilex harbored richer fungal communities than Quercus suber, and host-specific taxa more often belonged to fungi than to prokaryotes. Notably, the microbial communities at the dry end of the precipitation gradient harbored more locally-specific prokaryotic and fungal taxa than the microbial communities with a higher diversity, at the wet end of the gradient, suggesting higher specialization in drier areas.ConclusionsEven congeneric tree species, belonging to the same functional group, can harbor distinct and specific soil microbiomes. These microbiomes become more similar and consist of more specialized taxa under drier compared with wetter conditions. With this, our study offers a step towards a better understanding of the context-dependency of plant-soil feedbacks by going beyond α and β diversities and focusing on specialized taxa potentially driving community changes along environmental gradients. We hope that our study will stimulate future research assessing the importance of context-dependency of interactions between plants and soil communities in a changing world.

11 páginas.-- 3 figuras.-- 2 tablas.-- 47 referencias.-- Electronic supplementary material The online version of this article ( https://doi.org/10.1007/s11104-018-3567-z) contains supplementary material, which is available to authorized users, Aims: Plant-soil feedbacks (PSFs) have been shown to be relevant drivers of forest community dynamics. However, few studies have explored variation of PSFs along environmental gradients. In a framework of climate change, there is a great need to understand how interactions between plants and soil microbes respond along climatic gradients. Therefore, we compared PSFs along a precipitation gradient in Mediterranean oak forests and included trait responses. Following the Stress Gradient Hypothesis (SGH), we expected less negative or even positive PSFs in the physically harsh dry end of our gradient and more negative PSFs in the wettest end. Methods: We grew Quercus ilex and Quercus suber acorns on soil inoculated with microbes sampled under adults of both species in six sites ranging in annual precipitation. After 4 months, we measured shoot biomass and allocation and morphological traits above and belowground. Results: We found negative PSFs for Q. ilex independent of precipitation, whereas for Q. suber PSFs ranged from positive in dry sites to negative in wet sites, in agreement with the SGH. The leaf allocation showed patterns similar to shoot biomass, but belowground allocation and morphological traits revealed responses which could not be detected aboveground. Conclusions: We provide first evidence for context-dependent PSFs along a precipitation gradient. Moreover, we show that measuring root traits can help improve our understanding of climate-dependent PSFs. Such understanding helps to predict plant soil microbe interactions, and their role as drivers of plant community dynamics under ongoing climate change., This study was supported by the Swiss National Science Foundation (SNSF) in the context of a mobility fellowship granted to G.R. (P2BEP3_162092). L.G.A. acknowledges support from the MICINN project INTERCAPA (CGL-2014-56739-R).

Significance Identifying and explaining regional differences in tropical forest dynamics, structure, diversity, and composition are critical for anticipating region-specific responses to global environmental change. Floristic classifications are of fundamental importance for these efforts. Here we provide a global tropical forest classification that is explicitly based on community evolutionary similarity, resulting in identification of five major tropical forest regions and their relationships: (i) Indo-Pacific, (ii) Subtropical, (iii) African, (iv) American, and (v) Dry forests. African and American forests are grouped, reflecting their former western Gondwanan connection, while Indo-Pacific forests range from eastern Africa and Madagascar to Australia and the Pacific. The connection between northern-hemisphere Asian and American forests is confirmed, while Dry forests are identified as a single tropical biome., Knowledge about the biogeographic affinities of the world’s tropical forests helps to better understand regional differences in forest structure, diversity, composition, and dynamics. Such understanding will enable anticipation of region-specific responses to global environmental change. Modern phylogenies, in combination with broad coverage of species inventory data, now allow for global biogeographic analyses that take species evolutionary distance into account. Here we present a classification of the world’s tropical forests based on their phylogenetic similarity. We identify five principal floristic regions and their floristic relationships: (i) Indo-Pacific, (ii) Subtropical, (iii) African, (iv) American, and (v) Dry forests. Our results do not support the traditional neo- versus paleotropical forest division but instead separate the combined American and African forests from their Indo-Pacific counterparts. We also find indications for the existence of a global dry forest region, with representatives in America, Africa, Madagascar, and India. Additionally, a northern-hemisphere Subtropical forest region was identified with representatives in Asia and America, providing support for a link between Asian and American northern-hemisphere forests.

Research in savannas has focused on tree-grass interactions, whereas tree species coexistence received little attention. A leading hypothesis to explain tree coexistence is the Janzen-Connell model, which proposes an accumulation of host-specific enemies, e.g., soil organisms. While it has been shown in several non-savanna case studies that seedlings dispersed away from the mother perform better than seedlings that stay close (home-away effect), few studies tested whether foreign seedling species can replace own seedlings under conspecific adults (replacement effect). Some studies additionally tested for negative effects of conspecific biota (conspecific effect) to demonstrate the accumulation of enemies. We tested these effects by reciprocally growing seedlings of four tree species on soil collected beneath adults of all species, with and without applying a soil sterilization treatment. We found negative home-away effects suggesting that dispersal is advantageous and negative replacement effects suggesting species replacement under adults. While negative conspecific effects indicate accumulated enemies, positive heterospecific effects indicate an accumulation of mutualists rather than enemies for some species. We suggest that plant-soil feedbacks may well contribute to tree coexistence in savannas due to both negative conspecific and positive heterospecific feedbacks.

Agriculture and the exploitation of natural resources have transformed tropical mountain ecosystems across the world, and the consequences of these transformations for biodiversity and ecosystem functioning are largely unknown

The montane forests of Mount Kilimanjaro in Tanzania have been subjected to a long history of selective logging. However, since 1984 logging of indigenous trees is prohibited. Today, these forests allow us to evaluate the long-term effects of selective logging. We mapped the height and diameter at breast height (DBH) of all trees >10 cm DBH on 10 sites of 0.25 ha. Five sites represent non-logged forests, another five selectively logged forests. We tested whether forests were still visibly affected 30–40 years after selective logging in terms of their forest structure and tree diversity. Additionally we compared tree densities of different species guilds, including disturbance-indicator species, late-successional species and main timber species. Furthermore, we specifically compared the community size distributions of selectively logged and non-logged forests, first across all species and then for the most important timber species, Ocotea usambarensis, alone. 30–40 years after selective logging forests still showed a higher overall stem density, mainly due to higher relative abundances of small trees (

The factors determining gradients of biodiversity are a fundamental yet unresolved topic in ecology. While diversity gradients have been analysed for numerous single taxa, progress towards general explanatory models has been hampered by limitations in the phylogenetic coverage of past studies. By parallel sampling of 25 major plant and animal taxa along a 3.7 km elevational gradient on Mt. Kilimanjaro, we quantify cross-taxon consensus in diversity gradients and evaluate predictors of diversity from single taxa to a multi-taxa community level. While single taxa show complex distribution patterns and respond to different environmental factors, scaling up diversity to the community level leads to an unambiguous support for temperature as the main predictor of species richness in both plants and animals. Our findings illuminate the influence of taxonomic coverage for models of diversity gradients and point to the importance of temperature for diversification and species coexistence in plant and animal communities., Explaining species richness patterns is a key question in ecology. Peters et al. sample diverse plant and animal groups across elevation on Mt. Kilimanjaro to show that, while disparate factors drive distributions of individual taxa, diversity overall decreases with elevation, mostly driven by effects of temperature.

Natural enemies, especially host-specific enemies, are hypothesised to facilitate the coexistence of plant species by disproportionately inflicting more damage at increasing host abundance. However, few studies have assessed such Janzen‐Connell mechanisms on a scale relevant for coexistence and no study has evaluated potential topdown influences on the specialized pests. We quantified seed predation by specialist invertebrates and generalist vertebrates, as well as larval predation on these invertebrates, for the Neotropical palm Attalea butyracea across ten 4-ha plots spanning 20-fold variation in palm density. As palm density increased, seed attack by bruchid beetles increased, whereas seed predation by rodents held constant. But because rodent predation on bruchid larvae increased disproportionately with increasing palm density, bruchid emergence rates and total seed predation by rodents and bruchids combined were both density-independent. Our results demonstrate that topdown effects can limit the potential of host-specific insects to induce negative-density dependence in plant populations.

The protection and sustainable management of forest carbon stocks, particularly in the tropics, is a key factor in the mitigation of global change effects. However, our knowledge of how land use and elevation affect carbon stocks in tropical ecosystems is very limited. We compared aboveground biomass of trees, shrubs and herbs for eleven natural and human-influenced habitat types occurring over a wide elevation gradient (866–4550 m) at the world's highest solitary mountain, Mount Kilimanjaro. Thanks to the enormous elevation gradient, we covered important natural habitat types, e.g., savanna woodlands, montane rainforest and afro-alpine vegetation, as well as important land-use types such as maize fields, grasslands, traditional home gardens, coffee plantations and selectively logged forest. To assess tree and shrub biomass with pantropical allometric equations, we measured tree height, diameter at breast height and wood density and to assess herbaceous biomass, we sampled destructively. Among natural habitats, tree biomass was highest at intermediate elevation in the montane zone (340 Mg ha−1), shrub biomass declined linearly from 7 Mg ha−1 at 900 m to zero above 4000 m, and, inverse to tree biomass, herbaceous biomass was lower at mid-elevations (1 Mg ha−1) than in savannas (900 m, 3 Mg ha−1) or alpine vegetation (above 4000 m, 6 Mg ha−1). While the various land-use types dramatically decreased woody biomass at all elevations, though to various degrees, herbaceous biomass was typically increased. Our study highlights tropical montane forest biomass as important aboveground carbon stock and quantifies the extent of the strong aboveground biomass reductions by the major land-use types, common to East Africa. Further, it shows that elevation and land use differently affect different vegetation strata, and thus the matrix for other organisms.

Negative density dependence (NDD) of recruitment is pervasive in tropical tree species. We tested the hypotheses that seed dispersal is NDD, due to intraspecific competition for dispersers, and that this contributes to NDD of recruitment. We compared dispersal in the palm Attalea butyracea across a wide range of population density on Barro Colorado Island in Panama and assessed its consequences for seed distributions. We found that frugivore visitation, seed removal and dispersal distance all declined with population density of A. butyracea, demonstrating NDD of seed dispersal due to competition for dispersers. Furthermore, as population density increased, the distances of seeds from the nearest adult decreased, conspecific seed crowding increased and seedling recruitment success decreased, all patterns expected under poorer dispersal. Unexpectedly, however, our analyses showed that NDD of dispersal did not contribute substantially to these changes in the quality of the seed distribution; patterns with population density were dominated by effects due solely to increasing adult and seed density.

Natural enemies, especially host-specific enemies, are hypothesised to facilitate the coexistence of plant species by disproportionately inflicting more damage at increasing host abundance. However, few studies have assessed such Janzen-Connell mechanisms on a scale relevant for coexistence and no study has evaluated potential top-down influences on the specialized pests. We quantified seed predation by specialist invertebrates and generalist vertebrates, as well as larval predation on these invertebrates, for the Neotropical palm Attalea butyracea across ten 4-ha plots spanning 20-fold variation in palm density. As palm density increased, seed attack by bruchid beetles increased, whereas seed predation by rodents held constant. But because rodent predation on bruchid larvae increased disproportionately with increasing palm density, bruchid emergence rates and total seed predation by rodents and bruchids combined were both density-independent. Our results demonstrate that top-down effects can limit the potential of host-specific insects to induce negative-density dependence in plant populations.

Tropical forests are carbon-dense and highly productive ecosystems. Consequently, they play an important role in the global carbon cycle. In the present study we used an individual-based forest model (FORMIND) to analyze the carbon balances of a tropical forest. The main processes of this model are tree growth, mortality, regeneration, and competition. Model parameters were calibrated using forest inventory data from a tropical forest at Mt. Kilimanjaro. The simulation results showed that the model successfully reproduces important characteristics of tropical forests (aboveground biomass, stem size distribution and leaf area index). The estimated aboveground biomass (385 t/ha) is comparable to biomass values in the Amazon and other tropical forests in Africa. The simulated forest reveals a gross primary production of 24 tcha(-1) yr(-1). Modeling above- and belowground carbon stocks, we analyzed the carbon balance of the investigated tropical forest. The simulated carbon balance of this old-growth forest is zero on average. This study provides an example of how forest models can be used in combination with forest inventory data to investigate forest structure and local carbon balances.