Tropical Forest In Panama Research Articles

Tropical carbon sink accelerated by symbiotic dinitrogen fixation

A major uncertainty in the land carbon cycle is whether symbiotic nitrogen fixation acts to enhance the tropical forest carbon sink. Nitrogen-fixing trees can supply vital quantities of the growth-limiting nutrient nitrogen, but the extent to which the resulting carbon–nitrogen feedback safeguards ecosystem carbon sequestration remains unclear. We combine (i) field observations from 112 plots spanning 300 years of succession in Panamanian tropical forests, and (ii) a new model that resolves nitrogen and light competition at the scale of individual trees. Fixation doubled carbon accumulation in early succession and enhanced total carbon in mature forests by ~10% (~12MgC ha−1) through two mechanisms: (i) a direct fixation effect on tree growth, and (ii) an indirect effect on the successional sequence of non-fixing trees. We estimate that including nitrogen-fixing trees in Neotropical reforestation projects could safeguard the sequestration of 6.7 Gt CO2 over the next 20 years. Our results highlight the connection between functional diversity of plant communities and the critical ecosystem service of carbon sequestration for mitigating climate change.

Leaf reflectance spectroscopy captures variation in carboxylation capacity across species, canopy environment and leaf age in lowland moist tropical forests.

Understanding the pronounced seasonal and spatial variation in leaf carboxylation capacity (Vc,max ) is critical for determining terrestrial carbon cycling in tropical forests. However, an efficient and scalable approach for predicting Vc,max is still lacking. Here the ability of leaf spectroscopy for rapid estimation of Vc,max was tested. Vc,max was estimated using traditional gas exchange methods, and measured reflectance spectra and leaf age in leaves sampled from tropical forests in Panama and Brazil. These data were used to build a model to predict Vc,max from leaf spectra. The results demonstrated that leaf spectroscopy accurately predicts Vc,max of mature leaves in Panamanian tropical forests (R2 =0.90). However, this single-age model required recalibration when applied to broader leaf demographic classes (i.e. immature leaves). Combined use of spectroscopy models for Vc,max and leaf age enabled construction of the Vc,max -age relationship solely from leaf spectra, which agreed with field observations. This suggests that the spectroscopy technique can capture the seasonal variability in Vc,max , assuming sufficient sampling across diverse species, leaf ages and canopy environments. This finding will aid development of remote sensing approaches that can be used to characterize Vc,max in moist tropical forests and enable an efficient means to parameterize and evaluate terrestrial biosphere models.

Quantifying Leaf Phenology of Individual Trees and Species in a Tropical Forest Using Unmanned Aerial Vehicle (UAV) Images

Tropical forests exhibit complex but poorly understood patterns of leaf phenology. Understanding species- and individual-level phenological patterns in tropical forests requires datasets covering large numbers of trees, which can be provided by Unmanned Aerial Vehicles (UAVs). In this paper, we test a workflow combining high-resolution RGB images (7 cm/pixel) acquired from UAVs with a machine learning algorithm to monitor tree and species leaf phenology in a tropical forest in Panama. We acquired images for 34 flight dates over a 12-month period. Crown boundaries were digitized in images and linked with forest inventory data to identify species. We evaluated predictions of leaf cover from different models that included up to 14 image features extracted for each crown on each date. The models were trained and tested with visual estimates of leaf cover from 2422 images from 85 crowns belonging to eight species spanning a range of phenological patterns. The best-performing model included both standard color metrics, as well as texture metrics that quantify within-crown variation, with r2 of 0.84 and mean absolute error (MAE) of 7.8% in 10-fold cross-validation. In contrast, the model based only on the widely-used Green Chromatic Coordinate (GCC) index performed relatively poorly (r2 = 0.52, MAE = 13.6%). These results highlight the utility of texture features for image analysis of tropical forest canopies, where illumination changes may diminish the utility of color indices, such as GCC. The algorithm successfully predicted both individual-tree and species patterns, with mean r2 of 0.82 and 0.89 and mean MAE of 8.1% and 6.0% for individual- and species-level analyses, respectively. Our study is the first to develop and test methods for landscape-scale UAV monitoring of individual trees and species in diverse tropical forests. Our analyses revealed undescribed patterns of high intraspecific variation and complex leaf cover changes for some species.

The hydraulic efficiency-safety trade-off differs between lianas and trees.

Hydraulic traits are important for woody plant functioning and distribution. Associations among hydraulic traits, other leaf and stem traits, and species’ performance are relatively well understood for trees, but remain poorly studied for lianas. We evaluated the coordination among hydraulic efficiency (i.e., maximum hydraulic conductivity), hydraulic safety (i.e., cavitation resistance), a suite of eight morphological and physiological traits, and species’ abundances for saplings of 24 liana species and 27 tree species in wet tropical forests in Panama. Trees showed a strong trade‐off between hydraulic efficiency and hydraulic safety, whereas efficiency and safety were decoupled in lianas. Hydraulic efficiency was strongly and similarly correlated with acquisitive traits for lianas and trees (e.g., positively with gas exchange rates and negatively with wood density). Hydraulic safety, however, showed no correlations with other traits in lianas, but with several in trees (e.g., positively with leaf dry matter content and wood density and negatively with gas exchange rates), indicating that in lianas hydraulic efficiency is an anchor trait because it is correlated with many other traits, while in trees both efficiency and safety are anchor traits. Traits related to shade tolerance (e.g., low specific leaf area and high wood density) were associated with high local tree sapling abundance, but not with liana abundance. Our results suggest that different, yet unknown mechanisms determine hydraulic safety and local‐scale abundance for lianas compared to trees. For trees, the trade‐off between efficiency and safety will provide less possibilities for ecological strategies. For lianas, however, the uncoupling of efficiency and safety could allow them to have high hydraulic efficiency, and hence high growth rates, without compromising resistance to cavitation under drought, thus allowing them to thrive and outperform trees under drier conditions.

Connectivity explains local ant community structure in a Neotropical forest canopy: a large-scale experimental approach.

Understanding how habitat structure and resource availability affect local species distributions is a key goal of community ecology. Where habitats occur as a mosaic, variation in connectivity among patches influences both local species richness and composition, and connectivity is a key conservation concern in fragmented landscapes. Similarly, availability of limiting resources frequently determines species coexistence or exclusion. For primarily cursorial arthropods like ants, gaps between neighboring trees are a significant barrier to movement through the forest canopy. Competition for limited resources such as nest sites also promotes antagonistic interactions. Lianas (woody vines) connect normally isolated neighboring tree crowns and often have hollow stems inhabited by ants. We used two large-scale liana-removal experiments to determine how connectivity and nest site availability provided by lianas affect arboreal ant species richness, species composition, and β-diversity in a lowland tropical forest in Panama. Removing lianas from a tree crown reduced ant species richness up to 35%, and disproportionately affected species that require large foraging areas. Adding artificial connectivity to trees mitigated the effects of liana removal. Ant colonization of artificial nests was higher (73% occupied) in trees without lianas vs. trees with lianas (28% occupied). However, artificial nests typically were colonized by existing polydomous, resident ant species. As a result, nest addition did not affect ant community structure. Collectively, these results indicate that lianas are important to the maintenance of arboreal ant diversity specifically by providing connectivity among neighboring tree crowns. Anticipated increases in liana abundance in this forest could increase the local (tree-level) species richness of arboreal ants, with a compositional bias toward elevating the density of broad-ranging specialist predators.

Dead Wood Necromass in a Moist Tropical Forest: Stocks, Fluxes, and Spatiotemporal Variability

Woody debris (WD) stocks and fluxes are important components of forest carbon budgets and yet remain understudied, particularly in tropical forests. Here we present the most comprehensive assessment of WD stocks and fluxes yet conducted in a tropical forest, including one of the first tropical estimates of suspended WD. We rely on data collected over 8 years in an old-growth moist tropical forest in Panama to quantify spatiotemporal variability and estimate minimum sample sizes for different components. Downed WD constituted the majority of total WD mass (78%), standing WD contributed a substantial minority (21%), and suspended WD was the smallest component (1%). However, when considering sections of downed WD that are elevated above the soil, the majority of WD inputs and approximately 50% of WD stocks were disconnected from the forest floor. Branchfall and liana wood accounted for 17 and 2% of downed WD, respectively. Residence times averaged 1.9 years for standing coarse WD (CWD; > 20 cm diameter) and 3.6 years for downed CWD. WD stocks and inputs were highly spatially variable, such that the sampling efforts necessary to estimate true values within 10% with 95% confidence were > 130 km of transects for downed CWD and > 550 ha area for standing CWD. The vast majority of studies involve much lower sampling efforts, suggesting that considerably more data are required to precisely quantify tropical forest WD pools and fluxes. The demonstrated importance of elevated WD in our study indicates a need to understand how elevation above the ground alters decomposition rates and incorporate this understanding into models of forest carbon cycling.

Wood decomposition in aquatic and terrestrial ecosystems in the tropics: contrasting biotic and abiotic processes.

Wood decomposition, a critical process in carbon and nutrient cycles, is influenced by environmental conditions, decomposer communities and substrate composition. While these factors differ between land and stream habitats, across-habitat comparisons of wood decay processes are rare, limiting our ability to evaluate the context- dependency of the drivers of decay. Here we tracked wood decomposition of three tree species placed in stream and terrestrial habitats in a lowland tropical forest in Panama. At 3 and 11 months we measured mass loss, wood nitrogen and wood polymer concentrations, and sampled wood-associated fungal and bacterial communities. After 11 months of decay we found that mass loss occurred 9% faster in streams than on land, but loss of cellulose, hemicellulose and lignin did not differ between habitats. We also observed large differences in microbial decomposer communities between habitats. Overall, we found faster mass loss of wood in water, but no differences in biotic decay processes between habitats despite distinct microbial communities in streams and on land. Our research challenges the assumption that wood decays relatively slowly in water reflecting unfavorable environmental conditions and a limited capacity of aquatic microbial communities to effectively degrade wood polymers.

Reproductive traits as predictors of assembly chronosequence patterns in epiphyllous bryophyte metacommunities

Abstract Understanding the mechanisms underlying species assembly is a central focus of plant ecology and is crucial to revealing how plant communities are structured. However, the temporal limitations of most terrestrial plant communities preclude collection of species assembly data in a tractable time‐frame. The aim of this study is to investigate the importance of dispersal potential, as estimated by interspecific variation in sexual and asexual expression, as a predictor of patch chronosequence assembly for epiphyllous (leaf‐inhabiting) bryophytes. Secondarily, we investigate whether spatiotemporal patterns in patch turnover and compositional variation infer evidence of local (within‐patch) population processes. The frequency and distribution of 55 epiphyllous bryophytes across five leaf age classes were studied on an understorey shrub Piper grande (Piperaceae) in premontane Panamanian tropical forest. Logistical models were used to test whether interspecific variation in dispersal‐related life‐history traits predicts assembly order. Beta diversity and ordination analyses were employed to examine nestedness vs. turnover, and probe for repeatable patterns in patch chronosequence assembly, respectively. Our results emphasize the importance of dispersal potential on species assembly patterns as earlier arrivals exhibit greater probabilities of sexual and specialized asexual expression. High turnover coupled with temporal convergence in species composition also points to evidence of within‐patch biotic filtering. Synthesis. Epiphylls are locally diverse in humid tropical forests worldwide; however, the mechanisms which maintain their high local diversity are unknown. This study suggests that interspecific variation in dispersal capacity in combination with diffuse indirect effects is the principal contributor to the high alpha‐diversity of these ephemeral metacommunities.

Responses of arbuscular mycorrhizal fungi to long-term inorganic and organic nutrient addition in a lowland tropical forest.

Improved understanding of the nutritional ecology of arbuscular mycorrhizal (AM) fungi is important in understanding how tropical forests maintain high productivity on low-fertility soils. Relatively little is known about how AM fungi will respond to changes in nutrient inputs in tropical forests, which hampers our ability to assess how forest productivity will be influenced by anthropogenic change. Here we assessed the influence of long-term inorganic and organic nutrient additions and nutrient depletion on AM fungi, using two adjacent experiments in a lowland tropical forest in Panama. We characterised AM fungal communities in soil and roots using 454-pyrosequencing, and quantified AM fungal abundance using microscopy and a lipid biomarker. Phosphorus and nitrogen addition reduced the abundance of AM fungi to a similar extent, but affected community composition in different ways. Nutrient depletion(removal of leaf litter) had a pronounced effect on AM fungal community composition, affecting nearly as many OTUs as phosphorus addition. The addition of nutrients in organic form (leaf litter) had little effect on any AM fungal parameter. Soil AM fungal communities responded more strongly to changes in nutrient availability than communities in roots. This suggests that the 'dual niches' of AM fungi in soil versus roots are structured to different degrees by abiotic environmental filters, and biotic filters imposed by the plant host. Our findings indicate that AM fungal communities are fine-tuned to nutrient regimes, and support future studies aiming to link AM fungal community dynamics with ecosystem function.

Resource acquisition and reproductive strategies of tropical forest in response to the El Ni\xf1o\u2013Southern Oscillation

The El Niño–Southern Oscillation (ENSO) is the largest source of interannual climate variability in much of the tropics. We hypothesize that tropical plants exhibit interannual variation in reproduction and resource acquisition strategies driven by ENSO that mirrors their seasonal responses. We analyze the relationship of leaf and seed fall to climate variation over 30 years in a seasonally dry tropical forest in Panama where El Niño brings warm, dry, and sunny conditions. Elevated leaf fall precedes the onset of El Niño, and elevated seed production follows, paralleling associations with dry seasons. Our results provide evidence of a shift in allocation from leafing to fruiting in response to a warming phase of ENSO. This shift may enable plants to take advantage of higher light availability, while coping with higher atmospheric water demand and lower water supply. These findings might be an indicator of adaptive strategies to optimize reproduction and resource acquisition.

Decomposition of coarse woody debris in a long‐term litter manipulation experiment: A focus on nutrient availability

Abstract 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. A plain language summary is available for this article.

Ormyridae (Hymenoptera, Chalcidoidea) de Panamá asociados con agallas de cinípidos (Hymenoptera, Cynipidae) sobre especies de Quercus (Fagaceae)

Se estudiaron las especies de la familia Ormyridae emergidas de 15 de los 65 morfotipos de agallas inducidas por especies de la familia Cynipidae en árboles del género Quercus conocidas para Panamá. Las recolectas se realizaron entre enero de 2008 a diciembre de 2010 en los bosques nubosos de la Cordillera Central de Panamá. Se identificaron tres especies del género Ormyrus (Chalcidoidea: Ormyridae), O. hegeli, O. unifasciatipennis y O. venustus, parasitoides de las especies inductoras e inquilinas de las agallas hospedadoras. La especie más abundante fue O. venustus y, por primera vez, se reporta para el Neotrópico O. hegeli, que sólo había sido citada para Norteamérica. Los resultados amplían los registros de distribución de esta familia de avispas parasitoides y confirman la presunción de que la carencia de datos de algunos grupos de artrópodos en los bosques tropicales se debe más a la falta de muestreo y no a la ausencia de ellos en estas áreas.

Linking lidar and forest modeling to assess biomass estimation across scales and disturbance states

Light detection and ranging (lidar) is currently the state-of-the-art remote sensing technology for measuring the 3D structures of forests. Studies have shown that various lidar-derived metrics can be used to predict forest attributes, such as aboveground biomass. However, finding out which metric works best at which scale and under which conditions requires extensive field inventories as ground-truth data. The goal of our study was to overcome the limitations of inventory data by complementing field-derived data with virtual forest stands from a dynamic forest model. The simulated stands were used to compare 29 different lidar metrics for their utility as predictors of tropical forest biomass at different spatial scales. We used the process-based forest model FORMIND, developed a lidar simulation model, based on the Beer-Lambert law of light extinction, and applied it to a tropical forest in Panama. Simulation scenarios comprised undisturbed primary forests and stands exposed to logging and fire disturbance regimes, resulting in mosaics of different successional stages, totaling 3.7 million trees on 4200ha. The simulated forest was sampled with the lidar model. Several lidar metrics, in particular height metrics, showed good correlations with forest biomass, even for disturbed forest. Estimation errors (nRMSE) increased with decreasing spatial scale from <10% (200-m scale) to >30% (20-m scale) for the best metrics. At the often used 1-ha scale, the top-of-canopy height obtained from canopy height models with fine to relatively coarse pixel resolutions (1 to 10m) yielded the most accurate biomass predictions, with nRMSE<6% for undisturbed and nRMSE<9% for disturbed forests. This study represents the first time dynamic modeling of a tropical forest has been combined with lidar remote sensing to systematically investigate lidar-to-biomass relationships for varying lidar metrics, scales and disturbance states. In the future, this approach can be used to explore the potential of remote sensing of other forest attributes, e.g., carbon dynamics, and other remote sensing systems, e.g., spaceborne lidar and radar.

Soilborne fungi have host affinity and host-specific effects on seed germination and survival in a lowland tropical forest

The Janzen-Connell (JC) hypothesis provides a conceptual framework for explaining the maintenance of tree diversity in tropical forests. Its central tenet-that recruits experience high mortality near conspecifics and at high densities-assumes a degree of host specialization in interactions between plants and natural enemies. Studies confirming JC effects have focused primarily on spatial distributions of seedlings and saplings, leaving major knowledge gaps regarding the fate of seeds in soil and the specificity of the soilborne fungi that are their most important antagonists. Here we use a common garden experiment in a lowland tropical forest in Panama to show that communities of seed-infecting fungi are structured predominantly by plant species, with only minor influences of factors such as local soil type, forest characteristics, or time in soil (1-12 months). Inoculation experiments confirmed that fungi affected seed viability and germination in a host-specific manner and that effects on seed viability preceded seedling emergence. Seeds are critical components of reproduction for tropical trees, and the factors influencing their persistence, survival, and germination shape the populations of seedlings and saplings on which current perspectives regarding forest dynamics are based. Together these findings bring seed dynamics to light in the context of the JC hypothesis, implicating them directly in the processes that have emerged as critical for diversity maintenance in species-rich tropical forests.

Tree functional diversity affects litter decomposition and arthropod community composition in a tropical forest

AbstractDisturbance can alter tree species and functional diversity in tropical forests, which in turn could affect carbon and nutrient cycling via the decomposition of plant litter. However, the influence of tropical tree diversity on forest floor organisms and the processes they mediate are far from clear. We investigated the influence of different litter mixtures on arthropod communities and decomposition processes in a 60‐year‐old lowland tropical forest in Panama, Central America. We used litter mixtures representing pioneer and old growth tree species in experimental mesocosms to assess the links between litter types, decomposition rates, and litter arthropod communities. Overall, pioneer species litter decomposed most rapidly and old growth species litter decomposed the slowest but there were clear non‐additive effects of litter mixtures containing both functional groups. We observed distinct arthropod communities in different litter mixtures at 6 mo, with greater arthropod diversity and abundance in litter from old growth forest species. By comparing the decay of different litter mixtures in mesocosms and conventional litterbags, we demonstrated that our mesocosms represent an effective approach to link studies of litter decomposition and arthropod communities. Our results indicate that changes in the functional diversity of litter could have wider implications for arthropod communities and ecosystem functioning in tropical forests.

The role of soil chemistry and plant neighbourhoods in structuring fungal communities in three Panamanian rainforests

Summary Fungi play critical roles in ecosystem processes and interact with plant communities in mutualistic, pathogenic, and commensal ways. Fungal communities are thought to depend on both associated tree communities and soil properties. However, the relative importance of the biotic and abiotic drivers of soil fungal community structure and diversity in lowland tropical forests remains poorly understood. We examined the community structure of trees and fungi at different levels of phosphorus (0·17–16·3 mg kg−1) in moist tropical forests in Panama. We predicted that arbuscular mycorrhizal (AM) fungal composition would be more strongly associated with soil properties than with local tree communities while the composition of other fungal clades would be more strongly correlated with local tree communities than soil properties. We also predicted that fungal operational taxonomic unit (OTU) richness would be negatively correlated with soil fertility and positively correlated with tree species diversity within and among forests. We characterized soil chemistry, fine root biomass, and sequenced the ITS1 barcode region to describe fungal community composition from 70 soil cores across three 1‐ha tropical rainforest sites in Panama. The sites vary in soil chemistry, including P, and in tree species community composition, but experience similar annual rainfall. AM fungal community composition was partially correlated with soil chemistry (r = 0·32, P ≤ 0·001), but not with local tree communities, while non‐AM fungal communities were nearly equally correlated with soil chemistry (Partial Mantel test, r = 0·38, P ≤ 0·001) as with tree communities (r = 0·36, P ≤ 0·001). Linear models showed that AM OTU richness was not explained by any independent variable. For non‐AM fungi, phosphorus, pH, and soil moisture better predicted OTU richness across all cores than other biotic and abiotic factors. Synthesis. Our results show that AM fungal structure is driven primarily by soil chemistry. For non‐AM fungi, soil properties and the local tree community can play a joint role in structuring communities. Furthermore, we found that more diverse local tree communities did not harbour more fungal species. Our results suggest that soil properties act as an environmental filter for both trees and fungi, setting the stage for interactions between the two.

Liana effects on biomass dynamics strengthen during secondary forest succession.

Secondary forests are important carbon sinks, but their biomass dynamics vary markedly within and across landscapes. The biotic and abiotic drivers of this variation are still not well understood. We tested the effects of soil resource availability and competition by lianas on the biomass dynamics of young secondary tropical forests in Panama and assessed the extent to which liana effects were mediated by soil resource availability. Over a five-year period, growth, mortality, and recruitment of woody plants of ≥1cm diameter were monitored in 84 plots in 3-30-year-old secondary forests across the Agua Salud site in central Panama. Biomass dynamics and the effects of lianas and soil resources were examined using (generalized) linear mixed-effect models and a model averaging approach. There was strong spatial and temporal variation in liana biomass within and across the plots. The relative biomass of lianas had a strong negative effect on overall tree growth, growth of understory trees decreased with soil fertility and dry season soil water content, and the effect of lianas on tree mortality varied with soil fertility. Tree recruitment was not associated with any of the predictor variables. Our model indicates that tree biomass growth across our landscape was reduced with 22% due to competition with lianas, and that the effect of lianas increased during succession, from 19% after five years to 32% after 30years. The projected liana-induced growth reduction after 60years was 47%, which was consistent with data from a nearby site. Our study shows that the observed liana proliferation across tropical forests may reduce the sequestration and storage of carbon in young secondary forests, with important implications for the carbon balance of tropical forest landscapes and consequently for global climate change. Our study highlights the need to incorporate lianas and soil variables in research on the biomass dynamics of secondary forest across tropical landscapes, and the need for well-replicated longitudinal studies to cover landscape-level variability in the relevant abiotic and biotic components.

Insitu temperature response of photosynthesis of 42 tree and liana species in the canopy of two Panamanian lowland tropical forests with contrasting rainfall regimes.

Tropical forests contribute significantly to the global carbon cycle, but little is known about the temperature response of photosynthetic carbon uptake in tropical species, and how this varies within and across forests. We determined insitu photosynthetic temperature-response curves for upper canopy leaves of 42 tree and liana species from two tropical forests in Panama with contrasting rainfall regimes. On the basis of seedling studies, we hypothesized that species with high photosynthetic capacity - light-demanding, fast-growing species - would have a higher temperature optimum of photosynthesis (TOpt ) than species with low photosynthetic capacity - shade-tolerant, slow-growing species - and that, therefore, TOpt would scale with the position of a species on the slow-fast continuum of plant functional traits. TOpt was remarkably similar across species, regardless of their photosynthetic capacity and other plant functional traits. Community-average TOpt was almost identical to mean maximum daytime temperature, which was higher in the dry forest. Photosynthesis above TOpt appeared to be more strongly limited by stomatal conductance in the dry forest than in the wet forest. The observation that all species in a community shared similar TOpt values suggests that photosynthetic performance is optimized under current temperature regimes. These results should facilitate the scaling up of photosynthesis in relation to temperature from leaf to stand level in species-rich tropical forests.

A phosphorus threshold for mycoheterotrophic plants in tropical forests.

The majority of terrestrial plants associate with arbuscular mycorrhizal (AM) fungi, which typically facilitate the uptake of limiting mineral nutrients by plants in exchange for plant carbon. However, hundreds of non-photosynthetic plant species-mycoheterotrophs-depend entirely on AM fungi for carbon as well as mineral nutrition. Mycoheterotrophs can provide insight into the operation and regulation of AM fungal relationships, but little is known about the factors, fungal or otherwise, that affect mycoheterotroph abundance and distribution. In a lowland tropical forest in Panama, we conducted the first systematic investigation into the influence of abiotic factors on the abundance and distribution of mycoheterotrophs, to ask whether the availability of nitrogen and phosphorus altered the occurrence of mycoheterotrophs and their AM fungal partners. Across a natural fertility gradient spanning the isthmus of Panama, and also in a long-term nutrient-addition experiment, mycoheterotrophs were entirely absent when soil exchangeable phosphate concentrations exceeded 2 mg P kg-1 Experimental phosphorus addition reduced the abundance of AM fungi, and also reduced the abundance of the specific AM fungal taxa required by the mycoheterotrophs, suggesting that the phosphorus sensitivity of mycoheterotrophs is underpinned by the phosphorus sensitivity of their AM fungal hosts. The soil phosphorus concentration of 2 mg P kg-1 also corresponds to a marked shift in tree community composition and soil phosphatase activity across the fertility gradient, suggesting that our findings have broad ecological significance.

Arbuscular mycorrhizal fungal community composition is altered by long-term litter removal but not litter addition in a lowland tropical forest.

Tropical forest productivity is sustained by the cycling of nutrients through decomposing organic matter. Arbuscular mycorrhizal (AM) fungi play a key role in the nutrition of tropical trees, yet there has been little experimental investigation into the role of AM fungi in nutrient cycling via decomposing organic material in tropical forests. We evaluated the responses of AM fungi in a long-term leaf litter addition and removal experiment in a tropical forest in Panama. We described AM fungal communities using 454-pyrosequencing, quantified the proportion of root length colonised by AM fungi using microscopy, and estimated AM fungal biomass using a lipid biomarker. AM fungal community composition was altered by litter removal but not litter addition. Root colonisation was substantially greater in the superficial organic layer compared with the mineral soil. Overall colonisation was lower in the litter removal treatment, which lacked an organic layer. There was no effect of litter manipulation on the concentration of the AM fungal lipid biomarker in the mineral soil. We hypothesise that reductions in organic matter brought about by litter removal may lead to AM fungi obtaining nutrients from recalcitrant organic or mineral sources in the soil, besides increasing fungal competition for progressively limited resources.