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Climate models predict a possible increase in the frequency of strong climate events such as El Niño-Southern Oscillation (ENSO), which in parts of the tropics are the cause of exceptional droughts, these threaten global food production. Agroforestry systems are often suggested as promising diversification options to increase farmers' resilience to extreme climatic events. In the Northeastern state of Bahia, where most Brazilian cocoa is grown in wildlife-friendly agroforests, ENSOs cause severe droughts which negatively affect forest and agriculture. Cocoa (Theobroma cacao) is described as being sensitive to drought but there are no field-studies of the effect of ENSO-related drought on adult cocoa trees in the America's; there is one study of an experimentally-imposed drought in Indonesia which resulted in 10 to 46% yield loss. In our study, in randomly chosen farms in Bahia, Brazil, we measured the effect of the 2015–16 severe ENSO, which caused an unprecedented drought in cocoa agroforests. We show that drought caused high cocoa tree mortality (15%) and severely decreased cocoa yield (89%); the drought also increased infection rate of the chronic fungal disease witches' broom (Moniliophthora perniciosa). Ours findings showed that Brazilian cocoa agroforests are at risk and that increasing frequency of strong droughts are likely to cause decreased cocoa yields in the coming decades. Furthermore, because cocoa, like many crops, is grown somewhat beyond its climatic limits, it and other crops could be the 'canaries in the coalmine' warning of forthcoming major drought effects on semi-natural and natural vegetation.

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.

There is growing interest in understanding how declining soil fertility in the prolonged absence of major disturbance drives ecological processes, or ecosystem retrogression'. However, there are few well characterized study systems for exploring this phenomenon in the tropics, despite tropics occupying over 40% of the Earth's terrestrial surface. We studied two types of montane rain forest in the Blue Mountains of Jamaica that represent distinct stages in ecosystem development, i.e. an earlier stage with shallow organic matter and a late stage with deep organic matter (hereafter mull' and mor' stages). We characterized responses of soil fertility and plant, soil microbial and nematode communities to the transition from mull to mor and whether these responses were coupled. For soil abiotic properties, we found this transition led to lower amounts of both nitrogen (N) and phosphorus (P) and an enhanced N to P ratio. This led to shorter-statured and less diverse forest, and convergence of tree species composition among plots. At the whole community (but not individual species) level foliar and litter N and P diminished from mull to mor, while foliar N to P and resorption efficiency of P relative to N increased, indicating increasing P relative to N limitation. We also found impairment of soil microbes (but not nematodes) and an increasing role of fungi relative to bacteria during the transition. Our results show that retrogression phenomena involving increasing nutrient (notably P) limitation can be important drivers in tropical systems, and are likely to involve aboveground-belowground feedbacks whereby plants produce litter of diminishing quality, impairing soil microbial processes and thus reducing the supply of nutrients from the soil for plant growth. Such feedbacks between plants and the soil, mediated by plant litter and organic matter quality, may serve as major though often overlooked drivers of long term environmental change.

Summary 1. The cycling of nutrients in litterfall is considered a key mechanism in the maintenance of tropical forest fertility but its importance has rarely been quantified experimentally. 2. We carried out a long-term (5 years), large-scale litter manipulation experiment in lowland semi-evergreen tropical forest to determine how changes in litterfall affect forest nutrient cycling. We hypothesized that: (i) long-term litter removal would decrease the forest’s nutrient supply; (ii) litter addition would increase the forest’s nutrient supply; (iii) soil and foliar nutrient concentrations would change in response to litter manipulation and would eventually affect above-ground productivity. 3. To test our hypotheses, we measured trunk growth, litterfall, and nutrient concentrations in live leaves, litter and soil in plots where litter was removed once a month (L−), litter was added once a month (L+) and controls (CT). 4. After 5 years, the concentration of nitrate in the soil and soil stocks of inorganic nitrogen were higher in the L+ plots and lower in the L− plots compared to the controls. Ammonium concentrations in the soil were also lower in the L− plots. Nitrogen in leaves and litter and the annual nitrogen return by litter were higher in the L+ plots, while potassium return was lower in the L− plots. Surprisingly, our treatments had little effect on phosphorus in soil, leaves or litter, even though lowland tropical forests are generally thought to be largely phosphorus limited. 5. Trunk growth of large trees was not affected by litter manipulation but rainy season litterfall from 2003 to 2008 was 13% higher in the L+ plots compared to the controls. 6. Synthesis. Litter removal affected forest nutrient cycling and productivity less than expected, probably because the soil at our site is moderately fertile. However, litter addition increased litterfall indicating that some limitation of forest production was removed by litter addition. We expected strong effects o

Tropical forest floor characteristics such as depth and nutrient concentrations are highly heterogeneous even over small spatial scales and it is unclear how these differences contribute to patchiness in forest floor arthropod abundance and diversity. In a lowland tropical forest in Panama we experimentally increased litter standing crop by removing litter from five plots (L−) and adding it to five other plots (L+); we had five control plots. After 32 mo of treatments we investigated how arthropod abundance and diversity were related to differences in forest floor physical (mass, depth, water content) and chemical properties (pH, nutrient concentrations). Forest floor mass and total arthropod abundance were greater in L+ plots compared with controls. There were no treatment differences in nutrient concentrations, pH or water content of the organic horizons. Over all plots, the mass of the fermentation horizon (Oe) was greater than the litter horizon (Oi); arthropod diversity and biomass were also greater in the Oe horizon but nutrient concentrations tended to be higher in the Oi horizon. Arthropod abundance was best explained by forest floor mass, while arthropod diversity was best explained by phosphorus, calcium and sodium concentrations in the Oi horizon and by phosphorus concentrations in the Oe horizon. Differences in arthropod community composition between treatments and horizons correlated with phosphorus concentration and dry mass of the forest floor. We conclude that at a local scale, arthropod abundance is related to forest floor mass (habitat space), while arthropod diversity is related to forest floor nutrient concentrations (habitat quality).

The impact of elevated CO2 on leaf-litter and root exudate production may alter soil carbon storage capacities for the future. In particular when so-called ‘priming effects’, the counterintuitive loss of soil carbon following input of organic carbon substrates, are taken into consideration. Here we investigate the dynamics of priming effects and ask whether the source of primed carbon is microbial biomass or soil organic matter and whether specific microbial groups, as identified by phospholipid fatty acid (PLFA) biomarkers, may be important in causing them. We measured d13C within soil CO2 efflux and PLFA biomarkers following C3 soil priming effects caused by additions of C4 sugar-cane sucrose and maize (Zea mays L.) leaf-litter chopped and ground. All additions caused an initial pulse of priming effect CO2 and a later pulse of substrate-derived CO2, showing that priming effects can be induced rapidly following changes in substrate supply. Priming effects persisted over 32 days and led to a loss of soil carbon, with an increase in soil carbon decomposition of 169% following sucrose addition, 44% following chopped maize and 67% following ground maize additions. An increased concentration of soil-derived carbon within specific PLFA biomarkers provided evidence that a source of the primed carbon was soil organic matter. Certain Gram negative bacteria, identified by PLFA biomarkers (16:1v5, 16:1v7), showed increased uptake of soil carbon for both sucrose and maize treatments and may be directly linked to priming effects. Our study provides evidence that substrate carbon inputs to soil induce rapid changes in specific microbial groups, which in turn increase soil carbon metabolism.

Aboveground litter production in forests is likely to increase as a consequence of elevated atmospheric carbon dioxide (CO2) concentrations, rising temperatures, and shifting rainfall patterns. As litterfall represents a major flux of carbon from vegetation to soil, changes in litter inputs are likely to have wide-reaching consequences for soil carbon dynamics. Such disturbances to the carbon balance may be particularly important in the tropics because tropical forests store almost 30% of the global soil carbon, making them a critical component of the global carbon cycle; nevertheless, the effects of increasing aboveground litter production on belowground carbon dynamics are poorly understood. We used long-term, large-scale monthly litter removal and addition treatments in a lowland tropical forest to assess the consequences of increased litterfall on belowground CO2 production. Over the second to the fifth year of treatments, litter addition increased soil respiration more than litter removal decreased it; soil respiration was on average 20% lower in the litter removal and 43% higher in the litter addition treatment compared to the controls but litter addition did not change microbial biomass. We predicted a 9% increase in soil respiration in the litter addition plots, based on the 20% decrease in the litter removal plots and an 11% reduction due to lower fine root biomass in the litter addition plots. The 43% measured increase in soil respiration was therefore 34% higher than predicted and it is possible that this ‘extra’ CO2 was a result of priming effects, i.e. stimulation of the decomposition of older soil organic matter by the addition of fresh organic matter. Our results show that increases in aboveground litter production as a result of global change have the potential to cause considerable losses of soil carbon to the atmosphere in tropical forests.

The majority of above-ground carbon in tropical forests is stored in wood, which is returned to the atmosphere during decomposition of coarse woody debris. However, the factors controlling wood decomposition have not been experimentally manipulated over time scales comparable to the length of this process. We hypothesized that wood decomposition is limited by nutrient availability and tested this hypothesis in a long-term litter addition and removal experiment in a lowland tropical forest in Panama. Specifically, we quantified decomposition using a 15-year chronosequence of decaying boles, and measured respiration rates and nutrient limitation of wood decomposer communities. The long-term probability that a dead tree completely decomposed was decreased in plots where litter was removed, but did not differ between litter addition and control treatments. Similarly, respiration rates of wood decomposer communities were greater in control treatments relative to litter removal plots; litter addition treatments did not differ from either of the other treatments. Respiration rates increased in response to nutrient addition (nitrogen, phosphorus, and potassium) in the litter removal and addition treatments, but not in the controls. Established decreases in concentrations of soil nutrients in litter removal plots and increased respiration rates in response to nutrient addition suggest that reduced rates of wood decomposition after litter removal were caused by decreased nutrient availability. The effects of litter manipulations differed directionally from a previous short-term decomposition study in the same plots, and reduced rates of bole decomposition in litter removal plots did not emerge until after more than 6 years of decomposition. These differences suggest that litter-mediated effects on nutrient dynamics have complex interactions with decomposition over time.

Aboveground litter production in forests is likely to increase as a consequence of elevated atmospheric carbon dioxide (CO2) concentrations, rising temperatures, and shifting rainfall patterns. As litterfall represents a major flux of carbon from vegetation to soil, changes in litter inputs are likely to have wide-reaching consequences for soil carbon dynamics. Such disturbances to the carbon balance may be particularly important in the tropics because tropical forests store almost 30% of the global soil carbon, making them a critical component of the global carbon cycle; nevertheless, the effects of increasing aboveground litter production on belowground carbon dynamics are poorly understood. We used long-term, large-scale monthly litter removal and addition treatments in a lowland tropical forest to assess the consequences of increased litterfall on belowground CO2 production. Over the second to the fifth year of treatments, litter addition increased soil respiration more than litter removal decreased it; soil respiration was on average 20% lower in the litter removal and 43% higher in the litter addition treatment compared to the controls but litter addition did not change microbial biomass. We predicted a 9% increase in soil respiration in the litter addition plots, based on the 20% decrease in the litter removal plots and an 11% reduction due to lower fine root biomass in the litter addition plots. The 43% measured increase in soil respiration was therefore 34% higher than predicted and it is possible that this 'extra' CO2 was a result of priming effects, i.e. stimulation of the decomposition of older soil organic matter by the addition of fresh organic matter. Our results show that increases in aboveground litter production as a result of global change have the potential to cause considerable losses of soil carbon to the atmosphere in tropical forests.

We maintained a factorial nitrogen (N), phosphorus (P), and potassium (K) addition experiment for 11 years in a humid lowland forest growing on a relatively fertile soil in Panama to evaluate potential nutrient limitation of tree growth rates, fine-litter production, and fine-root biomass. We replicated the eight factorial treatments four times using 32 plots of 40x40 m each. The addition of K was associated with significant decreases in stand-level fine root biomass and, in a companion study of seedlings, decreases in allocation to roots and increases in height growth rates. The addition of K and N together was associated with significant increases in growth rates of saplings and poles (1–10 cm in diameter at breast height) and a further marginally significant decrease in stand-level fine-root biomass. The addition of P was associated with a marginally significant (P = 0.058) increase in fine-litter production that was consistent across all litter fractions. Our experiment provides evidence that N, P, and K all limit forest plants growing on a relatively fertile soil in the lowland tropics, with the strongest evidence for limitation by K among seedlings, saplings, and poles

The majority of above-ground carbon in tropical forests is stored in wood, which is returned to the atmosphere during decomposition of coarse woody debris. However, the factors controlling wood decomposition have not been experimentally manipulated over time scales comparable to the length of this process. We hypothesized that wood decomposition is limited by nutrient availability and tested this hypothesis in a long-term litter addition and removal experiment in a lowland tropical forest in Panama. Specifically, we quantified decomposition using a 15-year chronosequence of decaying boles, and measured respiration rates and nutrient limitation of wood decomposer communities. The long-term probability that a dead tree completely decomposed was decreased in plots where litter was removed, but did not differ between litter addition and control treatments. Similarly, respiration rates of wood decomposer communities were greater in control treatments relative to litter removal plots; litter addition treatments did not differ from either of the other treatments. Respiration rates increased in response to nutrient addition (nitrogen, phosphorus, and potassium) in the litter removal and addition treatments, but not in the controls. Established decreases in concentrations of soil nutrients in litter removal plots and increased respiration rates in response to nutrient addition suggest that reduced rates of wood decomposition after litter removal were caused by decreased nutrient availability. The effects of litter manipulations differed directionally from a previous short-term decomposition study in the same plots, and reduced rates of bole decomposition in litter removal plots did not emerge until after more than 6 years of decomposition. These differences suggest that litter-mediated effects on nutrient dynamics have complex interactions with decomposition over time.

Aboveground litter production in forests is likely to increase as a consequence of elevated atmospheric carbon dioxide (CO2) concentrations, rising temperatures, and shifting rainfall patterns. As litterfall represents a major flux of carbon from vegetation to soil, changes in litter inputs are likely to have wide-reaching consequences for soil carbon dynamics. Such disturbances to the carbon balance may be particularly important in the tropics because tropical forests store almost 30% of the global soil carbon, making them a critical component of the global carbon cycle; nevertheless, the effects of increasing aboveground litter production on belowground carbon dynamics are poorly understood. We used long-term, large-scale monthly litter removal and addition treatments in a lowland tropical forest to assess the consequences of increased litterfall on belowground CO2 production. Over the second to the fifth year of treatments, litter addition increased soil respiration more than litter removal decreased it; soil respiration was on average 20% lower in the litter removal and 43% higher in the litter addition treatment compared to the controls but litter addition did not change microbial biomass. We predicted a 9% increase in soil respiration in the litter addition plots, based on the 20% decrease in the litter removal plots and an 11% reduction due to lower fine root biomass in the litter addition plots. The 43% measured increase in soil respiration was therefore 34% higher than predicted and it is possible that this 'extra' CO2 was a result of priming effects, i.e. stimulation of the decomposition of older soil organic matter by the addition of fresh organic matter. Our results show that increases in aboveground litter production as a result of global change have the potential to cause considerable losses of soil carbon to the atmosphere in tropical forests.