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  1. Slik JWF, Franklin J, Arroyo-Rodríguez V, Field R, Aguilar S, Aguirre N, et al.
    Proc Natl Acad Sci U S A, 2018 02 20;115(8):1837-1842.
    PMID: 29432167 DOI: 10.1073/pnas.1714977115
    Knowledge about the biogeographic affinities of the world's tropical forests helps to better understand regional differences in forest structure, diversity, composition, and dynamics. Such understanding will enable anticipation of region-specific responses to global environmental change. Modern phylogenies, in combination with broad coverage of species inventory data, now allow for global biogeographic analyses that take species evolutionary distance into account. Here we present a classification of the world's tropical forests based on their phylogenetic similarity. We identify five principal floristic regions and their floristic relationships: (i) Indo-Pacific, (ii) Subtropical, (iii) African, (iv) American, and (v) Dry forests. Our results do not support the traditional neo- versus paleotropical forest division but instead separate the combined American and African forests from their Indo-Pacific counterparts. We also find indications for the existence of a global dry forest region, with representatives in America, Africa, Madagascar, and India. Additionally, a northern-hemisphere Subtropical forest region was identified with representatives in Asia and America, providing support for a link between Asian and American northern-hemisphere forests.
  2. Slik JW, Arroyo-Rodríguez V, Aiba S, Alvarez-Loayza P, Alves LF, Ashton P, et al.
    Proc Natl Acad Sci U S A, 2015 Jun 16;112(24):7472-7.
    PMID: 26034279 DOI: 10.1073/pnas.1423147112
    The high species richness of tropical forests has long been recognized, yet there remains substantial uncertainty regarding the actual number of tropical tree species. Using a pantropical tree inventory database from closed canopy forests, consisting of 657,630 trees belonging to 11,371 species, we use a fitted value of Fisher's alpha and an approximate pantropical stem total to estimate the minimum number of tropical forest tree species to fall between ∼ 40,000 and ∼ 53,000, i.e., at the high end of previous estimates. Contrary to common assumption, the Indo-Pacific region was found to be as species-rich as the Neotropics, with both regions having a minimum of ∼ 19,000-25,000 tree species. Continental Africa is relatively depauperate with a minimum of ∼ 4,500-6,000 tree species. Very few species are shared among the African, American, and the Indo-Pacific regions. We provide a methodological framework for estimating species richness in trees that may help refine species richness estimates of tree-dependent taxa.
  3. Mo L, Zohner CM, Reich PB, Liang J, de Miguel S, Nabuurs GJ, et al.
    Nature, 2023 Dec;624(7990):92-101.
    PMID: 37957399 DOI: 10.1038/s41586-023-06723-z
    Forests are a substantial terrestrial carbon sink, but anthropogenic changes in land use and climate have considerably reduced the scale of this system1. Remote-sensing estimates to quantify carbon losses from global forests2-5 are characterized by considerable uncertainty and we lack a comprehensive ground-sourced evaluation to benchmark these estimates. Here we combine several ground-sourced6 and satellite-derived approaches2,7,8 to evaluate the scale of the global forest carbon potential outside agricultural and urban lands. Despite regional variation, the predictions demonstrated remarkable consistency at a global scale, with only a 12% difference between the ground-sourced and satellite-derived estimates. At present, global forest carbon storage is markedly under the natural potential, with a total deficit of 226 Gt (model range = 151-363 Gt) in areas with low human footprint. Most (61%, 139 Gt C) of this potential is in areas with existing forests, in which ecosystem protection can allow forests to recover to maturity. The remaining 39% (87 Gt C) of potential lies in regions in which forests have been removed or fragmented. Although forests cannot be a substitute for emissions reductions, our results support the idea2,3,9 that the conservation, restoration and sustainable management of diverse forests offer valuable contributions to meeting global climate and biodiversity targets.
  4. Ma H, Crowther TW, Mo L, Maynard DS, Renner SS, van den Hoogen J, et al.
    Nat Plants, 2023 Nov;9(11):1795-1809.
    PMID: 37872262 DOI: 10.1038/s41477-023-01543-5
    Understanding what controls global leaf type variation in trees is crucial for comprehending their role in terrestrial ecosystems, including carbon, water and nutrient dynamics. Yet our understanding of the factors influencing forest leaf types remains incomplete, leaving us uncertain about the global proportions of needle-leaved, broadleaved, evergreen and deciduous trees. To address these gaps, we conducted a global, ground-sourced assessment of forest leaf-type variation by integrating forest inventory data with comprehensive leaf form (broadleaf vs needle-leaf) and habit (evergreen vs deciduous) records. We found that global variation in leaf habit is primarily driven by isothermality and soil characteristics, while leaf form is predominantly driven by temperature. Given these relationships, we estimate that 38% of global tree individuals are needle-leaved evergreen, 29% are broadleaved evergreen, 27% are broadleaved deciduous and 5% are needle-leaved deciduous. The aboveground biomass distribution among these tree types is approximately 21% (126.4 Gt), 54% (335.7 Gt), 22% (136.2 Gt) and 3% (18.7 Gt), respectively. We further project that, depending on future emissions pathways, 17-34% of forested areas will experience climate conditions by the end of the century that currently support a different forest type, highlighting the intensification of climatic stress on existing forests. By quantifying the distribution of tree leaf types and their corresponding biomass, and identifying regions where climate change will exert greatest pressure on current leaf types, our results can help improve predictions of future terrestrial ecosystem functioning and carbon cycling.
  5. Luo XS, Imai N, Dorigatti I
    Travel Med Infect Dis, 2020 01 26;33:101562.
    PMID: 31996323 DOI: 10.1016/j.tmaid.2020.101562
    BACKGROUND: No large-scale Zika epidemic has been observed to date in Southeast Asia following the 2015-16 Latin American and the Caribbean epidemic. One hypothesis is Southeast Asian populations' partial immunity to Zika.

    METHOD: We estimated the two conditions for a Zika outbreak emergence in Southeast Asia: (i) the risk of Zika introduction from Latin America and the Caribbean and, (ii) the risk of autochthonous transmission under varying assumptions on population immunity. We also validated the model used to estimate the risk of introduction by comparing the estimated number of Zika seeds introduced into the United States with case counts reported by the Centers for Disease Control and Prevention (CDC).

    RESULTS: There was good agreement between our estimates and case counts reported by the CDC. We thus applied the model to Southeast Asia and estimated that, on average, 1-10 seeds were introduced into Indonesia, Malaysia, the Philippines, Singapore, Thailand and Vietnam. We also found increasing population immunity levels from 0 to 90% reduced probability of autochthonous transmission by 40% and increasing individual variation in transmission further reduced the outbreak probability.

    CONCLUSIONS: Population immunity, combined with heterogeneity in transmission, can explain why no large-scale outbreak was observed in Southeast Asia during the 2015-16 epidemic.

  6. Imai N, White MT, Ghani AC, Drakeley CJ
    PLoS Negl Trop Dis, 2014 Jul;8(7):e2978.
    PMID: 25058400 DOI: 10.1371/journal.pntd.0002978
    INTRODUCTION: Plasmodium knowlesi is now recognised as a leading cause of malaria in Malaysia. As humans come into increasing contact with the reservoir host (long-tailed macaques) as a consequence of deforestation, assessing the potential for a shift from zoonotic to sustained P. knowlesi transmission between humans is critical.

    METHODS: A multi-host, multi-site transmission model was developed, taking into account the three areas (forest, farm, and village) where transmission is thought to occur. Latin hypercube sampling of model parameters was used to identify parameter sets consistent with possible prevalence in macaques and humans inferred from observed data. We then explore the consequences of increasing human-macaque contact in the farm, the likely impact of rapid treatment, and the use of long-lasting insecticide-treated nets (LLINs) in preventing wider spread of this emerging infection.

    RESULTS: Identified model parameters were consistent with transmission being sustained by the macaques with spill over infections into the human population and with high overall basic reproduction numbers (up to 2267). The extent to which macaques forage in the farms had a non-linear relationship with human infection prevalence, the highest prevalence occurring when macaques forage in the farms but return frequently to the forest where they experience higher contact with vectors and hence sustain transmission. Only one of 1,046 parameter sets was consistent with sustained human-to-human transmission in the absence of macaques, although with a low human reproduction number (R(0H) = 1.04). Simulations showed LLINs and rapid treatment provide personal protection to humans with maximal estimated reductions in human prevalence of 42% and 95%, respectively.

    CONCLUSION: This model simulates conditions where P. knowlesi transmission may occur and the potential impact of control measures. Predictions suggest that conventional control measures are sufficient at reducing the risk of infection in humans, but they must be actively implemented if P. knowlesi is to be controlled.

  7. Imai N, Samejima H, Langner A, Ong RC, Kita S, Titin J, et al.
    PLoS One, 2009;4(12):e8267.
    PMID: 20011516 DOI: 10.1371/journal.pone.0008267
    Sustainable forest management (SFM), which has been recently introduced to tropical natural production forests, is beneficial in maintaining timber resources, but information about the co-benefits for biodiversity conservation and carbon sequestration is currently lacking.
  8. Imai N, Furukawa T, Tsujino R, Kitamura S, Yumoto T
    PLoS One, 2018;13(5):e0197391.
    PMID: 29763452 DOI: 10.1371/journal.pone.0197391
    While many tropical countries are experiencing rapid deforestation, some have experienced forest transition (FT) from net deforestation to net reforestation. Numerous studies have identified causative factors of FT, among which forest scarcity has been considered as a prerequisite for FT. In fact, in SE Asia, the Philippines, Thailand and Viet Nam, which experienced FT since 1990, exhibited a lower remaining forest area (30±8%) than the other five countries (68±6%, Cambodia, Indonesia, Laos, Malaysia, and Myanmar) where forest loss continues. In this study, we examined 1) the factors associated with forest scarcity, 2) the proximate and/or underlying factors that have driven forest area change, and 3) whether causative factors changed across FT phases (from deforestation to net forest gain) during 1980-2010 in the eight SE Asian countries. We used production of wood, food, and export-oriented food commodities as proximate causes and demographic, social, economic and environmental factors, as well as land-use efficiency, and wood and food trade as underlying causes that affect forest area change. Remaining forest area in 1990 was negatively correlated with population density and potential land area of lowland forests, while positively correlated with per capita wood production. This implies that countries rich in accessible and productive forests, and higher population pressures are the ones that have experienced forest scarcity, and eventually FT. Food production and agricultural input were negatively and positively correlated, respectively, with forest area change during 1980-2009. This indicates that more food production drives deforestation, but higher efficiency of agriculture is correlated with forest gain. We also found a U-shaped response of forest area change to social openness, suggesting that forest gain can be achieved in both open and closed countries, but deforestation might be accelerated in countries undergoing societal transition. These results indicate the importance of environmental, agricultural and social variables on forest area dynamics, and have important implications for predicting future tropical forest change.
  9. Hirano Y, Kitayama K, Imai N
    Ecol Evol, 2022 Feb;12(3):e8669.
    PMID: 35309744 DOI: 10.1002/ece3.8669
    Soil organic phosphorus (P) compounds can be the main P source for plants in P-limited tropical rainforests. Phosphorus occurs in diverse chemical forms, including monoester P, diester P, and phytate, which require enzymatic hydrolysis by phosphatase into inorganic P before assimilation by plants. The interactions between plant interspecific differences in organic P acquisition strategies via phosphatase activities with root morphological traits would lead to P resource partitioning, but they have not been rigorously evaluated. We measured the activities of three classes of phosphatases (phosphomonoesterase, PME; phosphodiesterase, PDE; and phytase, PhT), specific root length (SRL), root diameter, and root tissue density in mature tree species with different mycorrhizal associations (ectomycorrhizal [ECM] or arbuscular mycorrhizal [AM]) and different successional status (climax or pioneer species) in Sabah, Malaysia. We studied nitrogen (N)- and P-fertilized plots to evaluate the acquisition strategies for organic P under P-limited conditions 7 years after fertilization was initiated. P fertilization reduced the PME activity in all studied species and reduced PhT and PDE activities more in climax species than in the two pioneer species, irrespective of the mycorrhizal type. PDE activity increased in some climax species after N fertilization, suggesting that these species allocate excess N to the synthesis of PDE. Moreover, PME and PhT activities, but not PDE activity, correlated positively with SRL. We suggest that climax species tend to be more strongly dependent on recalcitrant organic P (i.e., phytate and/or diester P) than pioneer species, regardless of the mycorrhizal type. We also suggest that trees in which root PME or PhT activity is enhanced can increase their SRL to acquire P efficiently. Resource partitioning of soil organic P would occur among species through differences in their phosphatase activities, which plays potentially ecologically important role in reducing the competition among coexisting tree species in lowland tropical rainforests.
  10. Delavaux CS, Crowther TW, Zohner CM, Robmann NM, Lauber T, van den Hoogen J, et al.
    Nature, 2023 Sep;621(7980):773-781.
    PMID: 37612513 DOI: 10.1038/s41586-023-06440-7
    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.
  11. Delavaux CS, Crowther TW, Zohner CM, Robmann NM, Lauber T, van den Hoogen J, et al.
    Nature, 2023 Oct;622(7982):E2.
    PMID: 37752352 DOI: 10.1038/s41586-023-06654-9
  12. Cooper DLM, Lewis SL, Sullivan MJP, Prado PI, Ter Steege H, Barbier N, et al.
    Nature, 2024 Jan;625(7996):728-734.
    PMID: 38200314 DOI: 10.1038/s41586-023-06820-z
    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.
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