Global patterns of tree density are contingent upon local determinants in the world's natural forests.

Previous attempts to quantify tree abundance at global scale have largely neglected the role of local competition in modulating the influence of climate and soils on tree density. Here, we evaluated whether mean tree size in the world's natural forests alters the effect of global productivity on tree density. In doing so, we gathered a vast set of forest inventories including >3000 sampling plots from 23 well-conserved areas worldwide to encompass (as much as possible) the main forest biomes on Earth. We evidence that latitudinal productivity patterns of tree density become evident as large trees become dominant. Global estimates of tree abundance should, therefore, consider dependencies of latitudinal sources of variability on local biotic influences to avoid underestimating the number of trees on Earth and to properly evaluate the functional and social consequences.

Las cabras como dispersores de semillas: aportes y limitaciones para la regeneración del bosque tropical estacionalmente seco de Ecuador / Goats as seed dispersers: contributions and limitations for the regeneration of the seasonally dry tropical forest in Ecuador

Resumen Introducción: Los bosques tropicales estacionalmente secos están sujetos a procesos de degradación crónica que ponen en riesgo su diversidad. La ganadería intensiva dentro de los remanentes de bosque se plantea como una de las principales causas de estos procesos de degradación. Sin embargo, el ganado también podría estar cumpliendo el rol de dispersor de semillas para algunas especies. Objetivo: Evaluar el rol de las cabras en la dispersión de semillas de especies leñosas y las posibles consecuencias de su comportamiento alimenticio sobre la estructura de la vegetación. Métodos: Entre diciembre 2016 y junio 2017 se recolectaron fecas de corrales (N = 38) y parcelas de vegetación (N = 42) de tres localidades de bosque seco en el Suroccidente de Ecuador. Todas las semillas encontradas en las fecas (N = 13 326) se registraron e identificaron taxonómicamente. Para evaluar el efecto de la ingestión de semillas sobre la germinación, se estableció un experimento de siembra de semillas extraídas de fecas procedentes de corrales y semillas recolectadas directamente de la planta. Resultados: Las cabras dispersaron 10 especies leñosas, de las cuales al menos el 50 % son leguminosas. Las semillas de Acacia macracantha representaron más del 70 % de semillas presentes en fecas de cabra. El paso de semillas por el tracto digestivo de las cabras mejoró significativamente el porcentaje y velocidad de germinación en Albizia multiflora, Piscidia carthagenensis y Ziziphus thyrsiflora, mientras que en Choroleucon mangense y Prosopis juliflora no se obtuvo germinación. No encontramos una correlación entre la riqueza de árboles establecidos y el número de especies encontradas en las fecas (χ2 = -0.23, P = 0.53). La composición de especies de semillas dispersadas en las fecas no mostró una dependencia de la localidad, a pesar de que la composición de la vegetación establecida cambia entre localidades. La abundancia de semillas en las fecas no mostró relación con la abundancia de árboles en la vegetación establecida. Conclusiones: Las cabras pueden suplir en cierta medida el rol de otros ungulados localmente extintos, mejorando la germinación de especies de leguminosas con testa dura. Sin embargo, su comportamiento alimenticio y la alta dominancia en la dispersión de ciertas especies puede tener importantes efectos en la estructura de la comunidad.

Abstract Introduction: Seasonally dry tropical forests are subject to chronic degradation processes, which has reduced the populations of some important animal dispersers. Intensive livestock farming within the forest remnants is considered one of the main causes of these degradation processes. However, domestic ungulates could also be fulfilling the role of seed dispersal for some wild species. Objective: To evaluate the role of goats as seed dispersers of woody species and the possible consequences of their feeding behavior on the vegetation structure. Methods: Between December 2016 and June 2017, we collected goat feces from pens (N = 38) and vegetation plots (N = 42) from three locations of dry forest in the Southwestern Ecuador. All the seeds found in the feces (N = 13 326) were recorded and taxonomically identified. To evaluate the effect of gut passage on seed germination, we sowed the seeds found in the goat feces from the pens and seeds collected directly from parent plants. Results: Goats dispersed seeds from ten species, of which at least 50 % are legumes. Acacia macracantha seeds represented ca. 70 % of seeds present in goat feces. The passage of seeds through the digestive tract of goats significantly improved the percentage and speed of germination in Albizia multiflora, Piscidia carthagenensis and Ziziphus thyrsiflora, while in Choroleucon mangense and Prosopis juliflora no germination was registered. We did not find a correlation between the richness of established trees and the number of species found in the goat feces (χ2 = -0.23, P = 0.53). The composition of dispersed seed species in the feces did not show a dependency on the locality, although the composition of the established vegetation changes between localities. The frequency of seeds in the feces did not show a relationship with the tree abundance in the established vegetation. Conclusions: Goats can play, at least partially, the role of locally extinct wild ungulates, improving the germination of legume species with hard coat. However, this positive effect can be blurred by their feeding behavior and high preference for particular species, which can modify the dominance of some species, and result in changes in the composition and structure of the vegetation.

A multifaceted approach to understanding bat community response to disturbance in a seasonally dry tropical forest.

Given widespread habitat degradation and loss, reliable indicators are needed that provide a comprehensive assessment of community response to anthropogenic disturbance. The family Phyllostomidae (Order Chiroptera) has frequently been the focus of research evaluating bats' response to habitat disturbance in seasonally dry tropical forests (SDTFs). However, few studies compare this family to the larger bat assemblage to assess its efficacy as a bioindicator. We compared community and species-specific attributes of understory phyllostomid and all understory bat species: (1) along a gradient of habitat disturbance within a human-modified SDTF landscape; and (2) between forest and riparian habitats within each disturbance level. We captured 290 individuals belonging to 13 species and 4 families. Phyllostomid species exhibited greater sensitivity to disturbance than the understory bat community as a whole based on richness and beta diversity. Both groups were more sensitive to disturbance in forest than riparian habitat, but phyllostomid species were more likely to be lost from highly disturbed forest habitat. The two dominant species declined in abundance with disturbance but variation in body condition was species-specific. These results suggest that Phyllostomidae are more effective indicators of human disturbance in SDTF than the understory bat community as a whole and evaluation of bats' response to disturbance is best accomplished with a multifaceted approach.

Biodiversity and ecosystem functions depend on environmental conditions and resources rather than the geodiversity of a tropical biodiversity hotspot.

Biodiversity and ecosystem functions are highly threatened by global change. It has been proposed that geodiversity can be used as an easy-to-measure surrogate of biodiversity to guide conservation management. However, so far, there is mixed evidence to what extent geodiversity can predict biodiversity and ecosystem functions at the regional scale relevant for conservation planning. Here, we analyse how geodiversity computed as a compound index is suited to predict the diversity of four taxa and associated ecosystem functions in a tropical mountain hotspot of biodiversity and compare the results with the predictive power of environmental conditions and resources (climate, habitat, soil). We show that combinations of these environmental variables better explain species diversity and ecosystem functions than a geodiversity index and identified climate variables as more important predictors than habitat and soil variables, although the best predictors differ between taxa and functions. We conclude that a compound geodiversity index cannot be used as a single surrogate predictor for species diversity and ecosystem functions in tropical mountain rain forest ecosystems and is thus little suited to facilitate conservation management at the regional scale. Instead, both the selection and the combination of environmental variables are essential to guide conservation efforts to safeguard biodiversity and ecosystem functions.

Climate reverses directionality in the richness-abundance relationship across the World's main forest biomes.

More tree species can increase the carbon storage capacity of forests (here referred to as the more species hypothesis) through increased tree productivity and tree abundance resulting from complementarity, but they can also be the consequence of increased tree abundance through increased available energy (more individuals hypothesis). To test these two contrasting hypotheses, we analyse the most plausible pathways in the richness-abundance relationship and its stability along global climatic gradients. We show that positive effect of species richness on tree abundance only prevails in eight of the twenty-three forest regions considered in this study. In the other forest regions, any benefit from having more species is just as likely (9 regions) or even less likely (6 regions) than the effects of having more individuals. We demonstrate that diversity effects prevail in the most productive environments, and abundance effects become dominant towards the most limiting conditions. These findings can contribute to refining cost-effective mitigation strategies based on fostering carbon storage through increased tree diversity. Specifically, in less productive environments, mitigation measures should promote abundance of locally adapted and stress tolerant tree species instead of increasing species richness.

Linking patterns and processes of tree community assembly across spatial scales in tropical montane forests.

Many studies have tried to assess the role of both deterministic and stochastic processes in community assembly, yet a lack of consensus exists on which processes are more prevalent and at which spatial scales they operate. To shed light on this issue, we tested two nonmutually exclusive, scale-dependent hypotheses: (1) that competitive exclusion dominates at small spatial scales; and (2) that environmental filtering does so at larger ones. To accomplish this, we studied the functional patterns of tropical montane forest communities along two altitudinal gradients, in Ecuador and Peru, using floristic and functional data from 60 plots of 0.1 ha. We found no evidence of either functional overdispersion or clustering at small spatial scales, but we did find functional clustering at larger ones. The observed pattern of clustering, consistent with an environmental filtering process, was more evident when maximizing the environmental differences among any pair of plots. To strengthen the link between the observed community functional pattern and the underlying process of environmental filtering, we explored differences in the climatic preferences of the most abundant species found at lower and higher elevations and examined whether their abundances shifted along the elevation gradient. We found (1) that greater community functional differences (observed between lower and upper tropical montane forest assemblies) were mostly the result of strong climatic preferences, maintained across the Neotropics; and (2) that the abundances of such species shifted along the elevational gradient. Our findings support the conclusion that, at large spatial scales, environmental filtering is the overriding mechanism for community assembly, because the pattern of functional clustering was linked to species' similarities in their climatic preferences, which ultimately resulted in shifts in species abundances along the gradient. However, there was no evidence of competitive exclusion at more homogeneous, smaller spatial scales, where plant species effectively compete for resources.

Trade-Offs Among Aboveground, Belowground, and Soil Organic Carbon Stocks Along Altitudinal Gradients in Andean Tropical Montane Forests.

Tropical montane forests (TMFs) play an important role as a carbon reservoir at a global scale. However, there is a lack of a comprehensive understanding on the variation in carbon storage across TMF compartments [namely aboveground biomass (AGB), belowground biomass (BGB), and soil organic matter] along altitudinal and environmental gradients and their potential trade-offs. This study aims to: 1) understand how carbon stocks vary along altitudinal gradients in Andean TMFs, and; 2) determine the influence of climate, particularly precipitation seasonality, on the distribution of carbon stocks across different forest compartments. The study was conducted in sixty 0.1 ha plots along two altitudinal gradients at the Podocarpus National Park (Ecuador) and Río Abiseo National Park (Peru). At each plot, we calculated the amount of carbon in AGB (i.e. aboveground carbon stock, AGC), BGB (i.e. belowground carbon stock, BGC), and soil organic matter (i.e. soil organic carbon stock, SOC). The mean total carbon stock was 244.76 ± 80.38 Mg ha-1 and 211.51 ± 46.95 Mg ha-1 in the Ecuadorian and Peruvian plots, respectively. Although AGC, BGC, and SOC showed different partitioning patterns along the altitudinal gradient both in Ecuador and Peru, total carbon stock did not change with altitude in either site. The combination of annual mean temperature and precipitation seasonality explained differences in the observed patterns of carbon stocks across forest compartments between the two sites. This study suggests that the greater precipitation seasonality of colder, higher altitudes may promote faster turnover rates of organic matter and nutrients and, consequently, less accumulation of SOC but greater AGC and BGC, compared to those sites with lesser precipitation seasonality. Our results demonstrate the capacity of TMFs to store substantial amounts of carbon and suggest the existence of a trade-off in carbon stocks among forest compartments, which could be partly driven by differences in precipitation seasonality, especially under the colder temperatures of high altitudes.

Mapping tree species vulnerability to multiple threats as a guide to restoration and conservation of tropical dry forests.

Understanding the vulnerability of tree species to anthropogenic threats is important for the efficient planning of restoration and conservation efforts. We quantified and compared the effects of future climate change and four current threats (fire, habitat conversion, overgrazing and overexploitation) on the 50 most common tree species of the tropical dry forests of northwestern Peru and southern Ecuador. We used an ensemble modelling approach to predict species distribution ranges, employed freely accessible spatial datasets to map threat exposures, and developed a trait-based scoring approach to estimate species-specific sensitivities, using differentiated trait weights in accordance with their expected importance in determining species sensitivities to specific threats. Species-specific vulnerability maps were constructed from the product of the exposure maps and the sensitivity estimates. We found that all 50 species face considerable threats, with an average of 46% of species' distribution ranges displaying high or very high vulnerability to at least one of the five threats. Our results suggest that current levels of habitat conversion, overexploitation and overgrazing pose larger threats to most of the studied species than climate change. We present a spatially explicit planning strategy for species-specific restoration and conservation actions, proposing management interventions to focus on (a) in situ conservation of tree populations and seed collection for tree planting activities in areas with low vulnerability to climate change and current threats; (b) ex situ conservation or translocation of populations in areas with high climate change vulnerability; and (c) active planting or assisted regeneration in areas under high current threat vulnerability but low climate change vulnerability, provided that interventions are in place to lower threat pressure. We provide an online, user-friendly tool to visualize both the vulnerability maps and the maps indicating priority restoration and conservation actions.

Plant height and hydraulic vulnerability to drought and cold.

Understanding how plants survive drought and cold is increasingly important as plants worldwide experience dieback with drought in moist places and grow taller with warming in cold ones. Crucial in plant climate adaptation are the diameters of water-transporting conduits. Sampling 537 species across climate zones dominated by angiosperms, we find that plant size is unambiguously the main driver of conduit diameter variation. And because taller plants have wider conduits, and wider conduits within species are more vulnerable to conduction-blocking embolisms, taller conspecifics should be more vulnerable than shorter ones, a prediction we confirm with a plantation experiment. As a result, maximum plant size should be short under drought and cold, which cause embolism, or increase if these pressures relax. That conduit diameter and embolism vulnerability are inseparably related to plant size helps explain why factors that interact with conduit diameter, such as drought or warming, are altering plant heights worldwide.

Comment on "The extent of forest in dryland biomes".

The study by Bastin et al (Reports, 12 May 2017, p. 635) is based on an incomplete delimitation of dry forest distribution and on an old and incorrect definition of drylands. Its sampling design includes many plots located in humid ecosystems and ignores critical areas for the conservation of dry forests. Therefore, its results and conclusions may be unreliable.

Climate and soil attributes determine plant species turnover in global drylands.

AIM: Geographic, climatic, and soil factors are major drivers of plant beta diversity, but their importance for dryland plant communities is poorly known. This study aims to: i) characterize patterns of beta diversity in global drylands, ii) detect common environmental drivers of beta diversity, and iii) test for thresholds in environmental conditions driving potential shifts in plant species composition. LOCATION: 224 sites in diverse dryland plant communities from 22 geographical regions in six continents. METHODS: Beta diversity was quantified with four complementary measures: the percentage of singletons (species occurring at only one site), Whittake's beta diversity (ß(W)), a directional beta diversity metric based on the correlation in species occurrences among spatially contiguous sites (ß(R2)), and a multivariate abundance-based metric (ß(MV)). We used linear modelling to quantify the relationships between these metrics of beta diversity and geographic, climatic, and soil variables. RESULTS: Soil fertility and variability in temperature and rainfall, and to a lesser extent latitude, were the most important environmental predictors of beta diversity. Metrics related to species identity (percentage of singletons and ß(W)) were most sensitive to soil fertility, whereas those metrics related to environmental gradients and abundance ((ß(R2)) and ß(MV)) were more associated with climate variability. Interactions among soil variables, climatic factors, and plant cover were not important determinants of beta diversity. Sites receiving less than 178 mm of annual rainfall differed sharply in species composition from more mesic sites (> 200 mm). MAIN CONCLUSIONS: Soil fertility and variability in temperature and rainfall are the most important environmental predictors of variation in plant beta diversity in global drylands. Our results suggest that those sites annually receiving ~ 178 mm of rainfall will be especially sensitive to future climate changes. These findings may help to define appropriate conservation strategies for mitigating effects of climate change on dryland vegetation.

Functional traits determine plant co-occurrence more than environment or evolutionary relatedness in global drylands.

Plant-plant interactions are driven by environmental conditions, evolutionary relationships (ER) and the functional traits of the plants involved. However, studies addressing the relative importance of these drivers are rare, but crucial to improve our predictions of the effects of plant-plant interactions on plant communities and of how they respond to differing environmental conditions. To analyze the relative importance of -and interrelationships among- these factors as drivers of plant-plant interactions, we analyzed perennial plant co-occurrence at 106 dryland plant communities established across rainfall gradients in nine countries. We used structural equation modeling to disentangle the relationships between environmental conditions (aridity and soil fertility), functional traits extracted from the literature, and ER, and to assess their relative importance as drivers of the 929 pairwise plant-plant co-occurrence levels measured. Functional traits, specifically facilitated plants' height and nurse growth form, were of primary importance, and modulated the effect of the environment and ER on plant-plant interactions. Environmental conditions and ER were important mainly for those interactions involving woody and graminoid nurses, respectively. The relative importance of different plant-plant interaction drivers (ER, functional traits, and the environment) varied depending on the region considered, illustrating the difficulty of predicting the outcome of plant-plant interactions at broader spatial scales. In our global-scale study on drylands, plant-plant interactions were more strongly related to functional traits of the species involved than to the environmental variables considered. Thus, moving to a trait-based facilitation/competition approach help to predict that: 1) positive plant-plant interactions are more likely to occur for taller facilitated species in drylands, and 2) plant-plant interactions within woody-dominated ecosystems might be more sensitive to changing environmental conditions than those within grasslands. By providing insights on which species are likely to better perform beneath a given neighbour, our results will also help to succeed in restoration practices involving the use of nurse plants.

Decoupling of soil nutrient cycles as a function of aridity in global drylands.

The biogeochemical cycles of carbon (C), nitrogen (N) and phosphorus (P) are interlinked by primary production, respiration and decomposition in terrestrial ecosystems. It has been suggested that the C, N and P cycles could become uncoupled under rapid climate change because of the different degrees of control exerted on the supply of these elements by biological and geochemical processes. Climatic controls on biogeochemical cycles are particularly relevant in arid, semi-arid and dry sub-humid ecosystems (drylands) because their biological activity is mainly driven by water availability. The increase in aridity predicted for the twenty-first century in many drylands worldwide may therefore threaten the balance between these cycles, differentially affecting the availability of essential nutrients. Here we evaluate how aridity affects the balance between C, N and P in soils collected from 224 dryland sites from all continents except Antarctica. We find a negative effect of aridity on the concentration of soil organic C and total N, but a positive effect on the concentration of inorganic P. Aridity is negatively related to plant cover, which may favour the dominance of physical processes such as rock weathering, a major source of P to ecosystems, over biological processes that provide more C and N, such as litter decomposition. Our findings suggest that any predicted increase in aridity with climate change will probably reduce the concentrations of N and C in global drylands, but increase that of P. These changes would uncouple the C, N and P cycles in drylands and could negatively affect the provision of key services provided by these ecosystems.

Plant species richness and ecosystem multifunctionality in global drylands.

Experiments suggest that biodiversity enhances the ability of ecosystems to maintain multiple functions, such as carbon storage, productivity, and the buildup of nutrient pools (multifunctionality). However, the relationship between biodiversity and multifunctionality has never been assessed globally in natural ecosystems. We report here on a global empirical study relating plant species richness and abiotic factors to multifunctionality in drylands, which collectively cover 41% of Earth's land surface and support over 38% of the human population. Multifunctionality was positively and significantly related to species richness. The best-fitting models accounted for over 55% of the variation in multifunctionality and always included species richness as a predictor variable. Our results suggest that the preservation of plant biodiversity is crucial to buffer negative effects of climate change and desertification in drylands.