Tropical tree ectomycorrhiza are distributed independently of soil nutrients.

Mycorrhizae, a form of plant-fungal symbioses, mediate vegetation impacts on ecosystem functioning. Climatic effects on decomposition and soil quality are suggested to drive mycorrhizal distributions, with arbuscular mycorrhizal plants prevailing in low-latitude/high-soil-quality areas and ectomycorrhizal (EcM) plants in high-latitude/low-soil-quality areas. However, these generalizations, based on coarse-resolution data, obscure finer-scale variations and result in high uncertainties in the predicted distributions of mycorrhizal types and their drivers. Using data from 31 lowland tropical forests, both at a coarse scale (mean-plot-level data) and fine scale (20 × 20 metres from a subset of 16 sites), we demonstrate that the distribution and abundance of EcM-associated trees are independent of soil quality. Resource exchange differences among mycorrhizal partners, stemming from diverse evolutionary origins of mycorrhizal fungi, may decouple soil fertility from the advantage provided by mycorrhizal associations. Additionally, distinct historical biogeographies and diversification patterns have led to differences in forest composition and nutrient-acquisition strategies across three major tropical regions. Notably, Africa and Asia's lowland tropical forests have abundant EcM trees, whereas they are relatively scarce in lowland neotropical forests. A greater understanding of the functional biology of mycorrhizal symbiosis is required, especially in the lowland tropics, to overcome biases from assuming similarity to temperate and boreal regions.

Flower production decreases with warmer and more humid atmospheric conditions in a western Amazonian forest.

Climate models predict that everwet western Amazonian forests will face warmer and wetter atmospheric conditions, and increased cloud cover. It remains unclear how these changes will impact plant reproductive performance, such as flowering, which plays a central role in sustaining food webs and forest regeneration. Warmer and wetter nights may cause reduced flower production, via increased dark respiration rates or alteration in the reliability of flowering cue-based processes. Additionally, more persistent cloud cover should reduce the amounts of solar irradiance, which could limit flower production. We tested whether interannual variation in flower production has changed in response to fluctuations in irradiance, rainfall, temperature, and relative humidity over 18 yrs in an everwet forest in Ecuador. Analyses of 184 plant species showed that flower production declined as nighttime temperature and relative humidity increased, suggesting that warmer nights and greater atmospheric water saturation negatively impacted reproduction. Species varied in their flowering responses to climatic variables but this variation was not explained by life form or phylogeny. Our results shed light on how plant communities will respond to climatic changes in this everwet region, in which the impacts of these changes have been poorly studied compared with more seasonal Neotropical areas.

Seasonality of reproduction in an ever-wet lowland tropical forest in Amazonian Ecuador.

Flowering and fruiting phenology have been infrequently studied in the ever-wet hyperdiverse lowland forests of northwestern equatorial Amazonía. These Neotropical forests are typically called aseasonal with reference to climate because they are ever-wet, and it is often assumed they are also aseasonal with respect to phenology. The physiological limits to plant reproduction imposed by water and light availability are difficult to disentangle in seasonal forests because these variables are often temporally correlated, and both are rarely studied together, challenging our understanding of their relative importance as drivers of reproduction. Here we report on the first long-term study (18 years) of flowering and fruiting phenology in a diverse equatorial forest, Yasuní in eastern Ecuador, and the first to include a full suite of on-site monthly climate data. Using twice monthly censuses of 200 traps and >1000 species, we determined whether reproduction at Yasuní is seasonal at the community and species levels and analyzed the relationships between environmental variables and phenology. We also tested the hypothesis that seasonality in phenology, if present, is driven primarily by irradiance. Both the community- and species-level measures demonstrated strong reproductive seasonality at Yasuní. Flowering peaked in September-November and fruiting peaked in March-April, with a strong annual signal for both phenophases. Irradiance and rainfall were also highly seasonal, even though no month on average experienced drought (a month with <100 mm rainfall). Flowering was positively correlated with current or near-current irradiance, supporting our hypothesis that the extra energy available during the period of peak irradiance drives the seasonality of flowering at Yasuní. As Yasuní is representative of lowland ever-wet equatorial forests of northwestern Amazonía, we expect that reproductive phenology will be strongly seasonal throughout this region.

La fenología de floración y fructificación ha sido poco estudiada en los bosques bajos, lluviosos e hiperdiversos de la Amazonía noroccidental. Estos bosques neotropicales son típicamente llamados no estacionales debido a su clima siempre lluvioso y se asume que son no estacionales con respecto a la fenología. Los límites fisiológicos a la reproducción de las plantas impuestos por la disponibilidad de agua y luz en estos bosques son difíciles de desentrañar debido a que estas variables están a menudo correlacionadas temporalmente y las dos se estudian usualmente por separado, lo que desafía nuestra comprensión de su importancia relativa como desencadenantes de la reproducción. Este es el primer estudio de largo plazo (18 años) de la fenología de floración y fructificación en un bosque hiperdiverso de la Amazonía noroccidental ecuatorial, Yasuní, ubicado al este de Ecuador, y el primero en incluir un completo set de datos climáticos mensuales. Usando censos quincenales de 200 trampas y > 1000 especies, examinamos si la reproducción en Yasuní es estacional a nivel de comunidad y de especies y analizamos las relaciones de las variables ambientales con la fenología. También nos interesaba probar si la estacionalidad en la fenología, en caso de que esté presente está causada por la irradiancia. Tanto a nivel de comunidad como de especies, los datos demuestran una fuerte estacionalidad reproductiva en Yasuní. La floración alcanzó un máximo en septiembre-noviembre y la fructificación alcanzó un máximo en marzo-abril, con una fuerte y consistente señal anual en las dos fenofases. A su vez, la irradiancia y la lluvia fueron también marcadamente estacionales, aunque ningún mes en promedio experimentó sequía (i.e. <100 mm de lluvia). La floración fue positivamente correlacionada con la irradiación, apoyando nuestra hipótesis de que la energía extra disponible durante los periodos de mayor irradiación causa la estacionalidad de la floración en Yasuní. Debido a que Yasuní representa a los bosques ecuatoriales lluviosos de tierras bajas de la Amazonía noroccidental, esperamos que la fenología reproductiva sea fuertemente estacional a lo largo de esta región.

Long-term fire and vegetation change in northwestern Amazonia.

Amazonian forest plots are used to quantify biodiversity and carbon sequestration, and provide the foundation for much of what is known about tropical ecology. Many plots are assumed to be undisturbed, but recent work suggests that past fire, forest openings, and cultivation created vegetation changes that have persisted for decades to centuries (ecological legacies). The Yasuní Forest Dynamics plot is one of the most biodiverse places on earth, yet its human history remains unknown. Here, we use charcoal and phytolith analysis to investigate the fire and vegetation history of the Yasuní forest plot, and compare results with nearby forest plots in Colombia (Amacayacu) and Peru (Medio Putumayo-Algodón [MPA]) to explore the spatial variability of past disturbances and ecological legacies in northwestern Amazonia. Three 14C dated charcoal fragments provided evidence for a modern (1956 CE) and a past fire event ca. 750 years ago at Yasuní, compared with fire ages of 1000-1600 years ago documented at Amacayacu and MPA. Small-scale disturbances and localized canopy openings also occurred in the Yasuní plot. Phytolith assemblages from Yasuní and Amacayacu showed more variability in past vegetation change than MPA. Low-intensity, non-continuous disturbances occurred at all three plots in the past, and our results highlight the variability of past human activities both in space and time in northwestern Amazonia. Our data also suggest that post-Columbian human disturbances from the Rubber Boom (AD 1850-1920) and subsequent oil exploration have likely left stronger ecological legacies than those left by pre-Columbian peoples in our studied regions.

Multiscale phenological niches of seed fall in diverse Amazonian plant communities.

Phenology has long been hypothesized as an avenue for niche partitioning or interspecific facilitation, both promoting species coexistence. Tropical plant communities exhibit striking diversity in reproductive phenology, but many are also noted for large synchronous reproductive events. Here we study whether the phenology of seed fall in such communities is nonrandom, the temporal scales of phenological patterns, and ecological factors that drive reproductive phenology. We applied multivariate wavelet analysis to test for phenological synchrony versus compensatory dynamics (i.e., antisynchronous patterns where one species' decline is compensated by the rise of another) among species and across temporal scales. We used data from long-term seed rain monitoring of hyperdiverse plant communities in the western Amazon. We found significant synchronous whole-community phenology at multiple timescales, consistent with shared environmental responses or positive interactions among species. We also observed both compensatory and synchronous phenology within groups of species (confamilials) likely to share traits and seed dispersal mechanisms. Wind-dispersed species exhibited significant synchrony at ~6-month scales, suggesting these species might share phenological niches to match the seasonality of wind. Our results suggest that community phenology is shaped by shared environmental responses but that the diversity of tropical plant phenology may partly result from temporal niche partitioning. The scale-specificity and time-localized nature of community phenology patterns highlights the importance of multiple and shifting drivers of phenology.

Damage to living trees contributes to almost half of the biomass losses in tropical forests.

Accurate estimates of forest biomass stocks and fluxes are needed to quantify global carbon budgets and assess the response of forests to climate change. However, most forest inventories consider tree mortality as the only aboveground biomass (AGB) loss without accounting for losses via damage to living trees: branchfall, trunk breakage, and wood decay. Here, we use ~151,000 annual records of tree survival and structural completeness to compare AGB loss via damage to living trees to total AGB loss (mortality + damage) in seven tropical forests widely distributed across environmental conditions. We find that 42% (3.62 Mg ha-1 year-1 ; 95% confidence interval [CI] 2.36-5.25) of total AGB loss (8.72 Mg ha-1 year-1 ; CI 5.57-12.86) is due to damage to living trees. Total AGB loss was highly variable among forests, but these differences were mainly caused by site variability in damage-related AGB losses rather than by mortality-related AGB losses. We show that conventional forest inventories overestimate stand-level AGB stocks by 4% (1%-17% range across forests) because assume structurally complete trees, underestimate total AGB loss by 29% (6%-57% range across forests) due to overlooked damage-related AGB losses, and overestimate AGB loss via mortality by 22% (7%-80% range across forests) because of the assumption that trees are undamaged before dying. Our results indicate that forest carbon fluxes are higher than previously thought. Damage on living trees is an underappreciated component of the forest carbon cycle that is likely to become even more important as the frequency and severity of forest disturbances increase.

Accelerating the discovery of rare tree species in Amazonian forests: integrating long monitoring tree plot data with metabolomics and phylogenetics for the description of a new species in the hyperdiverse genus Inga Mill.

In species-rich regions and highly speciose genera, the need for species identification and taxonomic recognition has led to the development of emergent technologies. Here, we combine long-term plot data with untargated metabolomics, and morphological and phylogenetic data to describe a new rare species in the hyperdiverse genus of trees Inga Mill. Our combined data show that Inga coleyana is a new lineage splitting from their closest relatives I. coruscans and I. cylindrica. Moreover, analyses of the chemical defensive profile demonstrate that I. coleyana has a very distinctive chemistry from their closest relatives, with I. coleyana having a chemistry based on saponins and I. cylindrica and I. coruscans producing a series of dihydroflavonols in addition to saponins. Finally, data from our network of plots suggest that I. coleyana is a rare and probably endemic taxon in the hyper-diverse genus Inga. Thus, the synergy produced by different approaches, such as long-term plot data and metabolomics, could accelerate taxonomic recognition in challenging tropical biomes.

Demographic composition, not demographic diversity, predicts biomass and turnover across temperate and tropical forests.

The growth and survival of individual trees determine the physical structure of a forest with important consequences for forest function. However, given the diversity of tree species and forest biomes, quantifying the multitude of demographic strategies within and across forests and the way that they translate into forest structure and function remains a significant challenge. Here, we quantify the demographic rates of 1961 tree species from temperate and tropical forests and evaluate how demographic diversity (DD) and demographic composition (DC) differ across forests, and how these differences in demography relate to species richness, aboveground biomass (AGB), and carbon residence time. We find wide variation in DD and DC across forest plots, patterns that are not explained by species richness or climate variables alone. There is no evidence that DD has an effect on either AGB or carbon residence time. Rather, the DC of forests, specifically the relative abundance of large statured species, predicted both biomass and carbon residence time. Our results demonstrate the distinct DCs of globally distributed forests, reflecting biogeography, recent history, and current plot conditions. Linking the DC of forests to resilience or vulnerability to climate change, will improve the precision and accuracy of predictions of future forest composition, structure, and function.

Soil nitrogen concentration mediates the relationship between leguminous trees and neighbor diversity in tropical forests.

Legumes provide an essential service to ecosystems by capturing nitrogen from the atmosphere and delivering it to the soil, where it may then be available to other plants. However, this facilitation by legumes has not been widely studied in global tropical forests. Demographic data from 11 large forest plots (16-60 ha) ranging from 5.25° S to 29.25° N latitude show that within forests, leguminous trees have a larger effect on neighbor diversity than non-legumes. Where soil nitrogen is high, most legume species have higher neighbor diversity than non-legumes. Where soil nitrogen is low, most legumes have lower neighbor diversity than non-legumes. No facilitation effect on neighbor basal area was observed in either high or low soil N conditions. The legume-soil nitrogen positive feedback that promotes tree diversity has both theoretical implications for understanding species coexistence in diverse forests, and practical implications for the utilization of legumes in forest restoration.

Temporal population variability in local forest communities has mixed effects on tree species richness across a latitudinal gradient.

Among the local processes that determine species diversity in ecological communities, fluctuation-dependent mechanisms that are mediated by temporal variability in the abundances of species populations have received significant attention. Higher temporal variability in the abundances of species populations can increase the strength of temporal niche partitioning but can also increase the risk of species extinctions, such that the net effect on species coexistence is not clear. We quantified this temporal population variability for tree species in 21 large forest plots and found much greater variability for higher latitude plots with fewer tree species. A fitted mechanistic model showed that among the forest plots, the net effect of temporal population variability on tree species coexistence was usually negative, but sometimes positive or negligible. Therefore, our results suggest that temporal variability in the abundances of species populations has no clear negative or positive contribution to the latitudinal gradient in tree species richness.

Distribution and Community Assembly of Trees Along an Andean Elevational Gradient.

Highlighting patterns of distribution and assembly of plants involves the use of community phylogenetic analyses and complementary traditional taxonomic metrics. However, these patterns are often unknown or in dispute, particularly along elevational gradients, with studies finding different patterns based on elevation. We investigated how patterns of tree diversity and structure change along an elevation gradient using taxonomic and phylogenetic diversity metrics. We sampled 595 individuals (36 families; 53 genera; 88 species) across 15 plots along an elevational gradient (2440-3330 m) in Ecuador. Seventy species were sequenced for the rbcL and matK gene regions to generate a phylogeny. Species richness, Shannon-Weaver diversity, Simpson's Dominance, Simpson's Evenness, phylogenetic diversity (PD), mean pairwise distance (MPD), and mean nearest taxon distance (MNTD) were evaluated for each plot. Values were correlated with elevation and standardized effect sizes (SES) of MPD and MNTD were generated, including and excluding tree fern species, for comparisons across elevation. Taxonomic and phylogenetic metrics found that species diversity decreases with elevation. We also found that overall the community has a non-random phylogenetic structure, dependent on the presence of tree ferns, with stronger phylogenetic clustering at high elevations. Combined, this evidence supports the ideas that tree ferns have converged with angiosperms to occupy the same habitat and that an increased filtering of clades has led to more closely related angiosperm species at higher elevations.

Physical, but not chemical, antiherbivore defense expression is related to the clustered spatial distribution of tropical trees in an Amazonian forest.

The conspecific negative density dependence hypothesis states that mortality of young trees (seedlings and saplings) is higher near conspecific adults due to mechanisms such as allelopathy, intraspecific competition, and pest facilitation, explaining why in the tropics, most of plant species tend to be rare and live dispersed. However, there are some tree species that defy this expectation and grow in large clusters of conspecific juveniles and adults. We hypothesize that conspecifics living in clusters show higher and/or more variable defensive profiles than conspecifics with dispersed distributions.We evaluated our hypothesis by assessing the expression of physical leaf traits (thickness, and the resistance to punch, tear and shear) and leaf chemical defenses for six clustered and six non-clustered tree species in Yasuní National Park, Ecuadorian Amazon. We ask ourselves whether (a) clustered species have leaves with higher physical resistance to damage and more chemical defenses variability than non-clustered species; (b) saplings of clustered species may show higher physical resistance to damage and higher variation on chemical leaf defenses than their conspecific adults, and (c) saplings of non-clustered species show lower resistance to physical damage and lower variation in chemical defenses compared to conspecific adults.Overall, our study did not support any of our hypotheses. Remarkably, we found that soluble metabolites were significantly species-specific.Our study suggests that plants physical but not chemical leaf antiherbivore defenses may be a crucial strategy for explaining survivorship of clustered species. Trees in Yasuní may also fall along a suite of tolerance/escape/defense strategies based on limitations of each species physiological constraints for survival and establishment. We conclude that other mechanisms, such as those related to indirect defenses, soil nutrient exploitation efficiency, volatile organic compounds, delayed leaf-greening, and seed dispersal mechanisms, shall be evaluated to understand conspecific coexistence in this forest.

Disentangling the functional trait correlates of spatial aggregation in tropical forest trees.

Environmental filtering and dispersal limitation can both maintain diversity in plant communities by aggregating conspecifics, but parsing the contribution of each process has proven difficult empirically. Here, we assess the contribution of filtering and dispersal limitation to the spatial aggregation patterns of 456 tree species in a hyperdiverse Amazonian forest and find distinct functional trait correlates of interspecific variation in these processes. Spatial point process model analysis revealed that both mechanisms are important drivers of intraspecific aggregation for the majority of species. Leaf drought tolerance was correlated with species topographic distributions in this aseasonal rainforest, showing that future increases in drought severity could significantly impact community structure. In addition, seed mass was associated with the spatial scale and density of dispersal-related aggregation. Taken together, these results suggest environmental filtering and dispersal limitation act in concert to influence the spatial and functional structure of diverse forest communities.

Climate sensitive size-dependent survival in tropical trees.

Survival rates of large trees determine forest biomass dynamics. Survival rates of small trees have been linked to mechanisms that maintain biodiversity across tropical forests. How species survival rates change with size offers insight into the links between biodiversity and ecosystem function across tropical forests. We tested patterns of size-dependent tree survival across the tropics using data from 1,781 species and over 2 million individuals to assess whether tropical forests can be characterized by size-dependent life-history survival strategies. We found that species were classifiable into four 'survival modes' that explain life-history variation that shapes carbon cycling and the relative abundance within forests. Frequently collected functional traits, such as wood density, leaf mass per area and seed mass, were not generally predictive of the survival modes of species. Mean annual temperature and cumulative water deficit predicted the proportion of biomass of survival modes, indicating important links between evolutionary strategies, climate and carbon cycling. The application of survival modes in demographic simulations predicted biomass change across forest sites. Our results reveal globally identifiable size-dependent survival strategies that differ across diverse systems in a consistent way. The abundance of survival modes and interaction with climate ultimately determine forest structure, carbon storage in biomass and future forest trajectories.

Topography and neighborhood crowding can interact to shape species growth and distribution in a diverse Amazonian forest.

Abiotic constraints and biotic interactions act simultaneously to shape communities. However, these community assembly mechanisms are often studied independently, which can limit understanding of how they interact to affect species dynamics and distributions. We develop a hierarchical Bayesian neighborhood modeling approach to quantify the simultaneous effects of topography and crowding by neighbors on the growth of 124,704 individual stems ≥1 cm DBH for 1,047 tropical tree species in a 25-ha mapped rainforest plot in Amazonian Ecuador. We build multi-level regression models to evaluate how four key functional traits (specific leaf area, maximum tree size, wood specific gravity and seed mass) mediate tree growth response to topography and neighborhood crowding. Tree growth is faster in valleys than on ridges and is reduced by neighborhood crowding. Topography and crowding interact to influence tree growth in ~10% of the species. Specific leaf area, maximum tree size and seed mass are associated with growth responses to topography, but not with responses to neighborhood crowding or with the interaction between topography and crowding. In sum, our study reveals that topography and neighborhood crowding each influence tree growth in tropical forests, but act largely independently in shaping species distributions. While traits were associated with species response to topography, their role in species response to neighborhood crowding was less clear, which suggests that trait effects on neighborhood dynamics may depend on the direction (negative/positive) and degree of symmetry of biotic interactions. Our study emphasizes the importance of simultaneously assessing the individual and interactive role of multiple mechanisms in shaping species dynamics in high diversity tropical systems.

Temporal coexistence mechanisms contribute to the latitudinal gradient in forest diversity.

The tropical forests of Borneo and Amazonia may each contain more tree species diversity in half a square kilometre than do all the temperate forests of Europe, North America, and Asia combined. Biologists have long been fascinated by this disparity, using it to investigate potential drivers of biodiversity. Latitudinal variation in many of these drivers is expected to create geographic differences in ecological and evolutionary processes, and evidence increasingly shows that tropical ecosystems have higher rates of diversification, clade origination, and clade dispersal. However, there is currently no evidence to link gradients in ecological processes within communities at a local scale directly to the geographic gradient in biodiversity. Here, we show geographic variation in the storage effect, an ecological mechanism that reduces the potential for competitive exclusion more strongly in the tropics than it does in temperate and boreal zones, decreasing the ratio of interspecific-to-intraspecific competition by 0.25% for each degree of latitude that an ecosystem is located closer to the Equator. Additionally, we find evidence that latitudinal variation in climate underpins these differences; longer growing seasons in the tropics reduce constraints on the seasonal timing of reproduction, permitting lower recruitment synchrony between species and thereby enhancing niche partitioning through the storage effect. Our results demonstrate that the strength of the storage effect, and therefore its impact on diversity within communities, varies latitudinally in association with climate. This finding highlights the importance of biotic interactions in shaping geographic diversity patterns, and emphasizes the need to understand the mechanisms underpinning ecological processes in greater detail than has previously been appreciated.

No strong evidence for increasing liana abundance in the Myristicaceae of a Neotropical aseasonal rain forest.

The "liana dominance hypothesis" posits that lianas are increasing in abundance in tropical forests, thereby potentially reducing tree biomass due to competitive interactions between trees and lianas. This scenario has implications not only for forest ecosystem function and species composition, but also climate change given the mass of carbon stored in tropical trees. In 2003 and 2013, all Myristicaceae trees in the 50-ha Yasuní Forest Dynamics Plot, Ecuador, were surveyed for liana presence and load in their crowns. We tested the hypothesis that the proportion of trees with lianas increased between 2003 and 2013 in line with the liana dominance hypothesis. Contrary to expectations, the total proportion of trees with lianas decreased from 35% to 32%, and when only trees ≥10 cm diameter at breast height were considered liana incidence increased 44-48%. Liana load was dynamic with a large proportion of trees losing or gaining lianas over the 10-yr period; large trees with intermediate liana loads increased in proportion at the expense of those with low and high loads. Lianas also impacted performance: trees with 26-75% crown cover by lianas in 2003 had reduced growth rates of 80% compared to of liana-free trees, and trees with >75% crown cover had 33% the growth rate and a log odds of mortality eight times that of liana-free trees. We suggest that the lack of strong support found for the liana dominance hypothesis is likely due to the aseasonal climate of Yasuní, which limits the competitive advantage lianas maintain over trees during dry seasons due to their efficient capture and use of water. We propose further research of long-term liana dynamics from aseasonal forests is required to determine the generality of the increasing liana dominance hypothesis in Neotropical forests.

Stability in a changing world - palm community dynamics in the hyperdiverse western Amazon over 17 years.

Are the hyperdiverse local forests of the western Amazon undergoing changes linked to global and local drivers such as climate change, or successional dynamics? We analyzed local climatic records to assess potential climatic changes in Yasuní National Park, Ecuador, and compared two censuses (1995, 2012) of a palm community to assess changes in community structure and composition. Over 17 years, the structure and composition of this palm community remained remarkably stable. Soil humidity was significantly lower and canopy conditions were significantly more open in 2012 compared to 1995, but local climatic records showed that no significant changes in precipitation, temperature or river level have occurred during the last decade. Thus, we found no evidence of recent directional shifts in climate or the palm community in Yasuní. The absence of changes in local climate and plant community dynamics in Yasuní contrasts with recent findings from eastern Amazon, where environmental change is driving significant changes in ecosystem dynamics. Our findings suggest that until now, local forests in the northwest Amazon may have escaped pressure from climate change. The stability of this rich palm community embedded in the hyperdiverse Yasuní National Park underlines its uniqueness as a sanctuary for the protection of Amazonian diversity from global change impacts.

Ant Mutualism Increases Long-Term Growth and Survival of a Common Amazonian Tree.

How ecological context shapes mutualistic relationships remains poorly understood. We combined long-term tree census data with ant censuses in a permanent 25-ha Amazonian forest dynamics plot to evaluate the effect of the mutualistic ant Myrmelachista schumanni (Formicinae) on the growth and survival of the common Amazonian tree Duroia hirsuta (Rubiaceae), considering its interactions with tree growth, population structure, and habitat. We found that the mutualist ant more than doubled tree relative growth rates and increased odds of survival. However, host tree size and density of conspecific neighbors modified the effect of the ant. Smaller trees hosting the mutualist ant consistently grew faster when surrounded by higher densities of conspecifics, suggesting that the benefit to the tree outweighs any negative effects of high conspecific densities. Moreover, our findings suggest that the benefit afforded by the ant diminishes with plant age and also depends on the density of conspecific neighbors. We provide the first long-term large-scale evidence of how mutualism affects the population biology of an Amazonian tree species.

Limited carbon and biodiversity co-benefits for tropical forest mammals and birds.

The conservation of tropical forest carbon stocks offers the opportunity to curb climate change by reducing greenhouse gas emissions from deforestation and simultaneously conserve biodiversity. However, there has been considerable debate about the extent to which carbon stock conservation will provide benefits to biodiversity in part because whether forests that contain high carbon density in their aboveground biomass also contain high animal diversity is unknown. Here, we empirically examined medium to large bodied ground-dwelling mammal and bird (hereafter "wildlife") diversity and carbon stock levels within the tropics using camera trap and vegetation data from a pantropical network of sites. Specifically, we tested whether tropical forests that stored more carbon contained higher wildlife species richness, taxonomic diversity, and trait diversity. We found that carbon stocks were not a significant predictor for any of these three measures of diversity, which suggests that benefits for wildlife diversity will not be maximized unless wildlife diversity is explicitly taken into account; prioritizing carbon stocks alone will not necessarily meet biodiversity conservation goals. We recommend conservation planning that considers both objectives because there is the potential for more wildlife diversity and carbon stock conservation to be achieved for the same total budget if both objectives are pursued in tandem rather than independently. Tropical forests with low elevation variability and low tree density supported significantly higher wildlife diversity. These tropical forest characteristics may provide more affordable proxies of wildlife diversity for future multi-objective conservation planning when fine scale data on wildlife are lacking.