Biogeographic history and environmental gradients modulate non-stationary patterns of tropical tree diversity.

Here we studied the entire Atlantic Forest hotspot to investigate whether the effect of different environmental predictors depends on the phylogenetic extension and the biogeographical history of different Atlantic Forest sectors. We used occurrence data of 3,183 plant species with arboreal or arborescent habits. We reconstructed climatic stability across 120,000 years using the Random Forest method. Then, we compared the effect of biogeographical history, topographic, and climatic variables on species richness and phylogenetic diversity using Geographically Weighted Regression (GWR) models. Niche conservatism drives the strength and direction of environmental correlates with tree diversity, interacting with the biogeographical and phylogenetic extension considered. Low current climate seasonalities were the main drivers of species richness and phylogenetic diversity variation across the Atlantic Forest. Whereas in higher phylogenetic extension, topographic heterogeneity increased the number of tree species independent of the sector, deep-past climate stability favored phylogenetic diversity by increasing relict lineages of distant clades in all forests, but with anomalies in the southern sector. This investigation yields substantial evidence that the response of the northern and southern sectors of the Atlantic Forest to identical environmental conditions diverges significantly, providing compelling support for the imprint of phylogenetic heritage in generating non-linear diversity patterns.

Latitudinal patterns in stabilizing density dependence of forest communities.

Numerous studies have shown reduced performance in plants that are surrounded by neighbours of the same species1,2, a phenomenon known as conspecific negative density dependence (CNDD)3. A long-held ecological hypothesis posits that CNDD is more pronounced in tropical than in temperate forests4,5, which increases community stabilization, species coexistence and the diversity of local tree species6,7. Previous analyses supporting such a latitudinal gradient in CNDD8,9 have suffered from methodological limitations related to the use of static data10-12. Here we present a comprehensive assessment of latitudinal CNDD patterns using dynamic mortality data to estimate species-site-specific CNDD across 23 sites. Averaged across species, we found that stabilizing CNDD was present at all except one site, but that average stabilizing CNDD was not stronger toward the tropics. However, in tropical tree communities, rare and intermediate abundant species experienced stronger stabilizing CNDD than did common species. This pattern was absent in temperate forests, which suggests that CNDD influences species abundances more strongly in tropical forests than it does in temperate ones13. We also found that interspecific variation in CNDD, which might attenuate its stabilizing effect on species diversity14,15, was high but not significantly different across latitudes. Although the consequences of these patterns for latitudinal diversity gradients are difficult to evaluate, we speculate that a more effective regulation of population abundances could translate into greater stabilization of tropical tree communities and thus contribute to the high local diversity of tropical forests.

Consistent patterns of common species across tropical tree communities.

Trees structure the Earth's most biodiverse ecosystem, tropical forests. The vast number of tree species presents a formidable challenge to understanding these forests, including their response to environmental change, as very little is known about most tropical tree species. A focus on the common species may circumvent this challenge. Here we investigate abundance patterns of common tree species using inventory data on 1,003,805 trees with trunk diameters of at least 10 cm across 1,568 locations1-6 in closed-canopy, structurally intact old-growth tropical forests in Africa, Amazonia and Southeast Asia. We estimate that 2.2%, 2.2% and 2.3% of species comprise 50% of the tropical trees in these regions, respectively. Extrapolating across all closed-canopy tropical forests, we estimate that just 1,053 species comprise half of Earth's 800 billion tropical trees with trunk diameters of at least 10 cm. Despite differing biogeographic, climatic and anthropogenic histories7, we find notably consistent patterns of common species and species abundance distributions across the continents. This suggests that fundamental mechanisms of tree community assembly may apply to all tropical forests. Resampling analyses show that the most common species are likely to belong to a manageable list of known species, enabling targeted efforts to understand their ecology. Although they do not detract from the importance of rare species, our results open new opportunities to understand the world's most diverse forests, including modelling their response to environmental change, by focusing on the common species that constitute the majority of their trees.

Native diversity buffers against severity of non-native tree invasions.

Determining the drivers of non-native plant invasions is critical for managing native ecosystems and limiting the spread of invasive species1,2. Tree invasions in particular have been relatively overlooked, even though they have the potential to transform ecosystems and economies3,4. Here, leveraging global tree databases5-7, we explore how the phylogenetic and functional diversity of native tree communities, human pressure and the environment influence the establishment of non-native tree species and the subsequent invasion severity. We find that anthropogenic factors are key to predicting whether a location is invaded, but that invasion severity is underpinned by native diversity, with higher diversity predicting lower invasion severity. Temperature and precipitation emerge as strong predictors of invasion strategy, with non-native species invading successfully when they are similar to the native community in cold or dry extremes. Yet, despite the influence of these ecological forces in determining invasion strategy, we find evidence that these patterns can be obscured by human activity, with lower ecological signal in areas with higher proximity to shipping ports. Our global perspective of non-native tree invasion highlights that human drivers influence non-native tree presence, and that native phylogenetic and functional diversity have a critical role in the establishment and spread of subsequent invasions.

Tree diversity increases decadal forest soil carbon and nitrogen accrual.

Increasing soil carbon and nitrogen storage can help mitigate climate change and sustain soil fertility1,2. A large number of biodiversity-manipulation experiments collectively suggest that high plant diversity increases soil carbon and nitrogen stocks3,4. It remains debated, however, whether such conclusions hold in natural ecosystems5-12. Here we analyse Canada's National Forest Inventory (NFI) database with the help of structural equation modelling (SEM) to explore the relationship between tree diversity and soil carbon and nitrogen accumulation in natural forests. We find that greater tree diversity is associated with higher soil carbon and nitrogen accumulation, validating inferences from biodiversity-manipulation experiments. Specifically, on a decadal scale, increasing species evenness from its minimum to maximum value increases soil carbon and nitrogen in the organic horizon by 30% and 42%, whereas increasing functional diversity enhances soil carbon and nitrogen in the mineral horizon by 32% and 50%, respectively. Our results highlight that conserving and promoting functionally diverse forests could promote soil carbon and nitrogen storage, enhancing both carbon sink capacity and soil nitrogen fertility.

Effects of moisture and density-dependent interactions on tropical tree diversity.

Tropical tree diversity increases with rainfall1,2. Direct physiological effects of moisture availability and indirect effects mediated by biotic interactions are hypothesized to contribute to this pantropical increase in diversity with rainfall2-6. Previous studies have demonstrated direct physiological effects of variation in moisture availability on tree survival and diversity5,7-10, but the indirect effects of variation in moisture availability on diversity mediated by biotic interactions have not been shown11. Here we evaluate the relationships between interannual variation in moisture availability, the strength of density-dependent interactions, and seedling diversity in central Panama. Diversity increased with soil moisture over the first year of life across 20 annual cohorts. These first-year changes in diversity persisted for at least 15 years. Differential survival of moisture-sensitive species did not contribute to the observed changes in diversity. Rather, negative density-dependent interactions among conspecifics were stronger and increased diversity in wetter years. This suggests that moisture availability enhances diversity indirectly through moisture-sensitive, density-dependent conspecific interactions. Pathogens and phytophagous insects mediate interactions among seedlings in tropical forests12-18, and many of these plant enemies are themselves moisture-sensitive19-27. Changes in moisture availability caused by climate change and habitat degradation may alter these interactions and tropical tree diversity.

Widespread herbivory cost in tropical nitrogen-fixing tree species.

Recent observations suggest that the large carbon sink in mature and recovering forests may be strongly limited by nitrogen1-3. Nitrogen-fixing trees (fixers) in symbiosis with bacteria provide the main natural source of new nitrogen to tropical forests3,4. However, abundances of fixers are tightly constrained5-7, highlighting the fundamental unanswered question of what limits new nitrogen entering tropical ecosystems. Here we examine whether herbivory by animals is responsible for limiting symbiotic nitrogen fixation in tropical forests. We evaluate whether nitrogen-fixing trees experience more herbivory than other trees, whether herbivory carries a substantial carbon cost, and whether high herbivory is a result of herbivores targeting the nitrogen-rich leaves of fixers8,9. We analysed 1,626 leaves from 350 seedlings of 43 tropical tree species in Panama and found that: (1) although herbivory reduces the growth and survival of all seedlings, nitrogen-fixing trees undergo 26% more herbivory than non-fixers; (2) fixers have 34% higher carbon opportunity costs owing to herbivory than non-fixers, exceeding the metabolic cost of fixing nitrogen; and (3) the high herbivory of fixers is not driven by high leaf nitrogen. Our findings reveal that herbivory may be sufficient to limit tropical symbiotic nitrogen fixation and could constrain its role in alleviating nitrogen limitation on the tropical carbon sink.

When and where to protect forests.

Ongoing deforestation poses a major threat to biodiversity1,2. With limited resources and imminent threats, deciding when as well as where to conserve is a fundamental question. Here we use a dynamic optimization approach to identify an optimal sequence for the conservation of plant species in 458 forested ecoregions globally over the next 50 years. The optimization approach includes species richness in each forested ecoregion, complementarity of species across ecoregions, costs of conservation that rise with cumulative protection in an ecoregion, the existing degree of protection, the rate of deforestation and the potential for reforestation in each ecoregion. The optimal conservation strategy for this formulation initially targets a small number of ecoregions where further deforestation leads to large reductions in species and where the costs of conservation are low. In later years, conservation efforts spread to more ecoregions, and invest in both expanded protection of primary forest and reforestation. The largest gains in species conservation come in Melanesia, South and Southeast Asia, the Anatolian peninsula, northern South America and Central America. The results highlight the potentially large gains in conservation that can be made with carefully targeted investments.

Warm springs alter timing but not total growth of temperate deciduous trees.

As the climate changes, warmer spring temperatures are causing earlier leaf-out1-3 and commencement of CO2 uptake1,3 in temperate deciduous forests, resulting in a tendency towards increased growing season length3 and annual CO2 uptake1,3-7. However, less is known about how spring temperatures affect tree stem growth8,9, which sequesters carbon in wood that has a long residence time in the ecosystem10,11. Here we show that warmer spring temperatures shifted stem diameter growth of deciduous trees earlier but had no consistent effect on peak growing season length, maximum growth rates, or annual growth, using dendrometer band measurements from 440 trees across two forests. The latter finding was confirmed on the centennial scale by 207 tree-ring chronologies from 108 forests across eastern North America, where annual ring width was far more sensitive to temperatures during the peak growing season than in the spring. These findings imply that any extra CO2 uptake in years with warmer spring temperatures4,5 does not significantly contribute to increased sequestration in long-lived woody stem biomass. Rather, contradicting projections from global carbon cycle models1,12, our empirical results imply that warming spring temperatures are unlikely to increase woody productivity enough to strengthen the long-term CO2 sink of temperate deciduous forests.

Even modest climate change may lead to major transitions in boreal forests.

The sensitivity of forests to near-term warming and associated precipitation shifts remains uncertain1-9. Herein, using a 5-year open-air experiment in southern boreal forest, we show divergent responses to modest climate alteration among juveniles of nine co-occurring North American tree species. Warming alone (+1.6 °C or +3.1 °C above ambient temperature) or combined with reduced rainfall increased the juvenile mortality of all species, especially boreal conifers. Species differed in growth responses to warming, ranging from enhanced growth in Acer rubrum and Acer saccharum to severe growth reductions in Abies balsamea, Picea glauca and Pinus strobus. Moreover, treatment-induced changes in both photosynthesis and growth help explain treatment-driven changes in survival. Treatments in which species experienced conditions warmer or drier than at their range margins resulted in the most adverse impacts on growth and survival. Species abundant in southern boreal forests had the largest reductions in growth and survival due to climate manipulations. By contrast, temperate species that experienced little mortality and substantial growth enhancement in response to warming are rare throughout southern boreal forest and unlikely to rapidly expand their density and distribution. Therefore, projected climate change will probably cause regeneration failure of currently dominant southern boreal species and, coupled with their slow replacement by temperate species, lead to tree regeneration shortfalls with potential adverse impacts on the health, diversity and ecosystem services of regional forests.

High exposure of global tree diversity to human pressure.

Safeguarding Earth's tree diversity is a conservation priority due to the importance of trees for biodiversity and ecosystem functions and services such as carbon sequestration. Here, we improve the foundation for effective conservation of global tree diversity by analyzing a recently developed database of tree species covering 46,752 species. We quantify range protection and anthropogenic pressures for each species and develop conservation priorities across taxonomic, phylogenetic, and functional diversity dimensions. We also assess the effectiveness of several influential proposed conservation prioritization frameworks to protect the top 17% and top 50% of tree priority areas. We find that an average of 50.2% of a tree species' range occurs in 110-km grid cells without any protected areas (PAs), with 6,377 small-range tree species fully unprotected, and that 83% of tree species experience nonnegligible human pressure across their range on average. Protecting high-priority areas for the top 17% and 50% priority thresholds would increase the average protected proportion of each tree species' range to 65.5% and 82.6%, respectively, leaving many fewer species (2,151 and 2,010) completely unprotected. The priority areas identified for trees match well to the Global 200 Ecoregions framework, revealing that priority areas for trees would in large part also optimize protection for terrestrial biodiversity overall. Based on range estimates for >46,000 tree species, our findings show that a large proportion of tree species receive limited protection by current PAs and are under substantial human pressure. Improved protection of biodiversity overall would also strongly benefit global tree diversity.

Tropical tree mortality has increased with rising atmospheric water stress.

Evidence exists that tree mortality is accelerating in some regions of the tropics1,2, with profound consequences for the future of the tropical carbon sink and the global anthropogenic carbon budget left to limit peak global warming below 2 °C. However, the mechanisms that may be driving such mortality changes and whether particular species are especially vulnerable remain unclear3-8. Here we analyse a 49-year record of tree dynamics from 24 old-growth forest plots encompassing a broad climatic gradient across the Australian moist tropics and find that annual tree mortality risk has, on average, doubled across all plots and species over the last 35 years, indicating a potential halving in life expectancy and carbon residence time. Associated losses in biomass were not offset by gains from growth and recruitment. Plots in less moist local climates presented higher average mortality risk, but local mean climate did not predict the pace of temporal increase in mortality risk. Species varied in the trajectories of their mortality risk, with the highest average risk found nearer to the upper end of the atmospheric vapour pressure deficit niches of species. A long-term increase in vapour pressure deficit was evident across the region, suggesting that thresholds involving atmospheric water stress, driven by global warming, may be a primary cause of increasing tree mortality in moist tropical forests.

The number of tree species on Earth.

One of the most fundamental questions in ecology is how many species inhabit the Earth. However, due to massive logistical and financial challenges and taxonomic difficulties connected to the species concept definition, the global numbers of species, including those of important and well-studied life forms such as trees, still remain largely unknown. Here, based on global ground-sourced data, we estimate the total tree species richness at global, continental, and biome levels. Our results indicate that there are ∼73,000 tree species globally, among which ∼9,000 tree species are yet to be discovered. Roughly 40% of undiscovered tree species are in South America. Moreover, almost one-third of all tree species to be discovered may be rare, with very low populations and limited spatial distribution (likely in remote tropical lowlands and mountains). These findings highlight the vulnerability of global forest biodiversity to anthropogenic changes in land use and climate, which disproportionately threaten rare species and thus, global tree richness.

Demographic and life history traits explain patterns in species vulnerability to extinction.

As ecosystems face disruption of community dynamics and habitat loss, the idea of determining ahead of time which species can become extinct is an important subject in conservation biology. A species' vulnerability to extinction is dependent upon both intrinsic (life-history strategies, genetics) and extrinsic factors (environment, anthropogenic threats). Studies linking intrinsic traits to extinction risk have shown variable results, and to our knowledge, there has not been a systematic analysis looking at how demographic patterns in stage-specific survival and reproductive rates correlate to extinction risk. We used matrix projection models from the COMPADRE and COMADRE matrix databases and IUCN Red List status as our proxy of extinction risk to investigate if some demographic patterns are more vulnerable to extinction than others. We obtained data on demographic rates, phylogeny, and IUCN status for 159 species of herbaceous plants, trees, mammals, and birds. We calculated 14 demographic metrics related to different aspects of life history and elasticity values and analyzed whether they differ based on IUCN categories using conditional random forest analysis and phylogenetic generalized least square regressions. We mapped all species within the database, both with IUCN assessment and without, and overlaid them with biodiversity hotspots to investigate if there is bias within the assessed species and how many of the non-assessed species could use the demographic information recorded in COMPADRE and COMADRE for future IUCN assessments. We found that herbaceous perennials are more vulnerable when they mature early and have high juvenile survival rates; birds are more vulnerable with high progressive growth and reproduction; mammals are more vulnerable when they have longer generation times. These patterns may be used to assess relative vulnerability across species when lacking abundance or trend data.

Tree structure and diversity of a Humid Mountain Forest in the protected natural area La Martinica, Veracruz, Mexico / Estructura y diversidad arbórea de un bosque húmedo de montaña en el Área Natural Protegida La Martinica, Veracruz, México

Abstract Introduction: The Humid Mountain Forest (HMF) has the highest number of plants per unit of surface, whose vegetation grows under heterogeneous environmental conditions and possess a high floristic variation. HMF conservation is important due to the biodiversity it harbors and the environmental services it provides. Objective: This work evaluated the effect of the terrain aspect and density of the forest canopy on the structure and tree diversity in La Martinica Protected Natural Area, Mexico. Methods: Stratified sampling was performed in four terrain aspects and two canopy density conditions. Twenty five sampling units of 20 x 25 m were considered, in which the normal diameter (ND), total height and the largest and smallest diameters of the crown of the individuals with a ND ≥ 10 cm were registered. The diversity was estimated by rarefaction curves and the structure was analyzed through the importance value index (IVI) and the forest value index (FVI). Results: 37 species belonging to 30 genera and 24 families were recorded. Greater diversity was observed in the north terrain aspect and in the closed canopy. Tree species with the highest structural values were different between terrain aspect and canopy types; Carpinus tropicalis presented the highest values in the zenithal terrain aspect, Lippia myriocephala in the east and south terrain aspect, and Liquidambar styraciflua in the north. In both canopy types Lippia myriocephala obtained the highest IVI values and FVI in the open canopy; Carpinus tropicalis reached a higher FVI in the closed canopy. Conclusions: Tree structure was different in the four terrain aspects and two canopy conditions studied. The greatest difference in species composition and diversity was observed between the north and east terrain aspects; the north presented the highest richness values, frequent and dominant species.

Resumen Introducción: El Bosque Húmedo de Montaña (BHM) posee el mayor número de especies vegetales por unidad de superficie, cuya vegetación se desarrolla en condiciones ambientales muy heterogéneas y presenta una alta variación florística. La conservación del BHM es importante debido a la biodiversidad que alberga y los servicios ambientales que proporciona. Objetivo: Este trabajo evaluó el efecto de la orientación del terreno y la densidad del dosel del bosque sobre la estructura y diversidad arbórea en el Área Natural Protegida La Martinica, México. Métodos: Se efectuó un muestreo estratificado en cuatro orientaciones del terreno y dos condiciones de densidad del dosel; se consideraron 25 unidades de muestreo de 20 x 25 m, en las que se registró: el diámetro normal (DN), la altura total y los diámetros mayor y menor de la copa de los individuos con un DN ≥10 cm. La diversidad se estimó por medio de curvas de rarefacción y la estructura se analizó con el índice de valor de importancia (IVI) y el índice de valor forestal (IVF). Resultados: Se registraron 37 especies pertenecientes a 30 géneros y 24 familias. Se observó una mayor diversidad en la orientación norte y en el dosel cerrado. Las especies arbóreas con valores estructurales más altos fueron diferentes entre orientaciones y tipos de dosel; Carpinus tropicalis presentó los valores más elevados en la orientación cenital, Lippia myriocephala en las orientaciones este y sur, y Liquidambar styraciflua en la norte. En ambos tipos de dosel Lippia myriocephala obtuvo los valores más altos del IVI e IVF en el dosel abierto; Carpinus tropicalis alcanzó un IVF más elevado en el dosel cerrado. Conclusiones: La estructura arbórea fue diferente en las cuatro orientaciones estudiadas y en las dos condiciones del dosel. La mayor diferencia en composición de especies y diversidad se observó entre las orientaciones norte y este, de ellas, la norte presentó los valores más altos de riqueza, especies frecuentes y dominantes.

Applied aerial spectroscopy: A case study on remote sensing of an ancient and semi-natural woodland.

An area of ancient and semi-natural woodland (ASNW) has been investigated by applied aerial spectroscopy using an unmanned aerial vehicle (UAV) with multispectral image (MSI) camera. A novel normalised difference spectral index (NDSI) algorithm was developed using principal component analysis (PCA). This novel NDSI was then combined with a simple segmentation method of thresholding and applied for the identification of native tree species as well as the overall health of the woodland. Using this new approach allowed the identification of trees at canopy level, across 7.4 hectares (73,934 m2) of ASNW, as oak (53%), silver birch (37%), empty space (9%) and dead trees (1%). This UAV derived data was corroborated, for its accuracy, by a statistically valid ground-level field study that identified oak (47%), silver birch (46%) and dead trees (7.4%). This simple innovative approach, using a low-cost multirotor UAV with MSI camera, is both rapid to deploy, was flown around 100 m above ground level, provides useable high resolution (5.3 cm / pixel) data within 22 mins that can be interrogated using readily available PC-based software to identify tree species. In addition, it provides an overall oversight of woodland health and has the potential to inform a future woodland regeneration strategy.

1.36 million years of Mediterranean forest refugium dynamics in response to glacial-interglacial cycle strength.

The sediment record from Lake Ohrid (Southwestern Balkans) represents the longest continuous lake archive in Europe, extending back to 1.36 Ma. We reconstruct the vegetation history based on pollen analysis of the DEEP core to reveal changes in vegetation cover and forest diversity during glacial-interglacial (G-IG) cycles and early basin development. The earliest lake phase saw a significantly different composition rich in relict tree taxa and few herbs. Subsequent establishment of a permanent steppic herb association around 1.2 Ma implies a threshold response to changes in moisture availability and temperature and gradual adjustment of the basin morphology. A change in the character of G-IG cycles during the Early-Middle Pleistocene Transition is reflected in the record by reorganization of the vegetation from obliquity- to eccentricity-paced cycles. Based on a quantitative analysis of tree taxa richness, the first large-scale decline in tree diversity occurred around 0.94 Ma. Subsequent variations in tree richness were largely driven by the amplitude and duration of G-IG cycles. Significant tree richness declines occurred in periods with abundant dry herb associations, pointing to aridity affecting tree population survival. Assessment of long-term legacy effects between global climate and regional vegetation change reveals a significant influence of cool interglacial conditions on subsequent glacial vegetation composition and diversity. This effect is contrary to observations at high latitudes, where glacial intensity is known to control subsequent interglacial vegetation, and the evidence demonstrates that the Lake Ohrid catchment functioned as a refugium for both thermophilous and temperate tree species.

Climate change may induce connectivity loss and mountaintop extinction in Central American forests.

The tropical forests of Central America serve a pivotal role as biodiversity hotspots and provide ecosystem services securing human livelihood. However, climate change is expected to affect the species composition of forest ecosystems, lead to forest type transitions and trigger irrecoverable losses of habitat and biodiversity. Here, we investigate potential impacts of climate change on the environmental suitability of main plant functional types (PFTs) across Central America. Using a large database of occurrence records and physiological data, we classify tree species into trait-based groups and project their suitability under three representative concentration pathways (RCPs 2.6, 4.5 and 8.5) with an ensemble of state-of-the-art correlative modelling methods. Our results forecast transitions from wet towards generalist or dry forest PFTs for large parts of the study region. Moreover, suitable area for wet-adapted PFTs is projected to latitudinally diverge and lose connectivity, while expected upslope shifts of montane species point to high risks of mountaintop extinction. These findings underline the urgent need to safeguard the connectivity of habitats through biological corridors and extend protected areas in the identified transition hotspots.