Uncertainties about controls on tree mortality make forest responses to land-use and climate change difficult to predict. We tracked biomass of tree functional groups in tropical forest inventories ...across Puerto Rico and the U.S. Virgin Islands, and with random forests we ranked 86 potential predictors of small tree survival (young or mature stems 2.5-12.6 cm diameter at breast height). Forests span dry to cloud forests, range in age, geology and past land use and experienced severe drought and storms. When excluding species as a predictor, top predictors are tree crown ratio and height, two to three species traits and stand to regional factors reflecting local disturbance and the system state (widespread recovery, drought, hurricanes). Native species, and species with denser wood, taller maximum height, or medium typical height survive longer, but short trees and species survive hurricanes better. Trees survive longer in older stands and with less disturbed canopies, harsher geoclimates (dry, edaphically dry, e.g., serpentine substrates, and highest-elevation cloud forest), or in intervals removed from hurricanes. Satellite image phenology and bands, even from past decades, are top predictors, being sensitive to vegetation type and disturbance. Covariation between stand-level species traits and geoclimate, disturbance and neighboring species types may explain why most neighbor variables, including introduced vs. native species, had low or no importance, despite univariate correlations with survival. As forests recovered from a hurricane in 1998 and earlier deforestation, small trees of introduced species, which on average have lighter wood, died at twice the rate of natives. After hurricanes in 2017, the total biomass of trees ≥12.7 cm dbh of the introduced species Spathodea campanulata spiked, suggesting that more frequent hurricanes might perpetuate this light-wooded species commonness. If hurricane recovery favors light-wooded species while drought favors others, climate change influences on forest composition and ecosystem services may depend on the frequency and severity of extreme climate events.
Celotno besedilo
Dostopno za:
DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
Understanding the heterogeneity of biomass accumulation in second-growth tropical forests following land use abandonment is important for informing ecosystem carbon models and forest restoration ...efforts. There is an urgent need for a broad sample of second-growth forests to enhance our knowledge of carbon accumulation in human-dominated landscapes, especially for older forests. Puerto Rico has predominantly second-growth forests, ranging in age from approximately 25 to more than 80 years. We used an island-wide sample of airborne lidar from the NASA Goddard Lidar, Hyperspectral, and Thermal (G-LiHT) Airborne Imager collected on March 2017, forest inventory data, and data on forest age, precipitation, soils, and land use to estimate aboveground biomass stocks in moist and wet, second-growth tropical forests. Biomass accumulation rates in Puerto Rico were lower, on average, than in other Neotropical forests. Median biomass across >16,700 ha of older second-growth forests was 105 Mg ha−1, and sampled biomass rarely surpassed 250 Mg ha−1. Differences in biomass by age were large and persistent across different substrates and land uses, with a plateau in the pattern of island-wide biomass accumulation after about 33 years. A spatial regression model showed that multiple factors were related to biomass accumulation, including time since abandonment, geologic substrate, past land use as coffee or pasture, precipitation, topographic wetness index, and slope. Our findings have important consequences for the total carbon storage and expected climate mitigation benefits of large-scale reforestation efforts, and highlight the value of airborne lidar for quantifying biomass variability in complex tropical landscapes.
Understanding the role of alien species in forest communities, and how native and alien species interact to shape the composition and structure of contemporary forests, is of critical importance to ...invasion ecology and natural resource management. We used vegetation data collected over a 20‐year period in 341 permanent plots representing remnants of closed‐canopy forests and post‐agricultural secondary forests across Puerto Rico to compare changes in the composition and abundance of native and alien woody species in plots with and without aliens across different forest types and to assess whether aliens and natives show divergence or convergence regarding functional roles and ecological strategies. We also tested the applicability of Grime's CSR (competitive, stress‐tolerant, and ruderal strategies) theory to explain naturalization success. Species richness and abundance of natives are consistently lower in plots in which aliens are present compared with those without them. This negative association between aliens and natives has been consistent over the 20 years and across all forest types. Both native and total richness slightly increased over the 20 years, but the increase in native species richness was three times lower in plots with aliens relative to those without aliens. The CSR classification provided insight into the naturalization success of aliens. Corroborating the “join the locals” hypothesis, aliens use the same functional spaces as natives. The exception is in dry forests, where aliens and natives differ in the use of functional spaces, a result that corroborates the “try harder” hypothesis. Generally, aliens were better competitors compared with natives, and natives were more stress‐tolerant than aliens. Our combined results suggest that alien species may inhibit population growth or even drive local changes in native plant communities by transforming the assembly and dynamics of tropical forests. Ultimately, modifications linked to invasive species may have significant implications for local forests, affecting their regeneration and productivity. More definitive conclusions require additional plot censuses, and analyses of disturbance regimes and stand‐age structure to reveal the long‐term implications of alien species on regenerating tropical forests, including their vulnerability, resilience, and adaptive capacity to cope with various aspects of climate change.
Aim: Seasonally dry tropical forest (SDTF) of the Caribbean Islands (primarily West Indies) is floristically distinct from Neotropical SDTF in Central and South America. We evaluate whether tree ...species composition was associated with climatic gradients or geographical distance. Turnover (dissimilarity) in species composition of different islands or among more distant sites would suggest communities structured by speciation and dispersal limitations. A nested pattern would be consistent with a steep resource gradient. Correlation of species composition with climatic variation would suggest communities structured by broad-scale environmental filtering. Location: The West Indies (The Bahamas, Cuba, Hispaniola, Jamaica, Puerto Rico, US Virgin Islands, Guadeloupe, Martinique, St. Lucia), Providencia (Colombia), south Florida (USA) and Florida Keys (USA). Taxon: Seed plants—woody taxa (primarily trees). Methods: We compiled 572 plots from 23 surveys conducted between 1969 and 2016. Hierarchical clustering of species in plots, and indicator species analysis for the resulting groups of sites, identified geographical patterns of turnover in species composition. Nonparametric analysis of variance, applied to principal components of bioclimatic variables, determined the degree of covariation in climate with location. Nestedness versus turnover in species composition was evaluated using beta diversity partitioning. Generalized dissimilarity modelling partitioned the effect of climate versus geographical distance on species composition. Results: Despite a set of commonly occurring species, SDTF tree community composition was distinct among islands and was characterized by spatial turnover on climatic gradients that covaried with geographical gradients. Greater Antillean islands were characterized by endemic indicator species. Northern subtropical areas supported distinct, rather than nested, SDTF communities in spite of low levels of endemism. Main conclusions: The SDTF species composition was correlated with climatic variation. SDTF on large Greater Antillean islands (Hispaniola, Jamaica and Cuba) was characterized by endemic species, consistent with their geological history and the biogeography of plant lineages. These results suggest that both environmental filtering and speciation shape Caribbean SDTF tree communities.
During the mid‐1900s, most of the island of Puerto Rico was deforested, but a shift in the economy from agriculture to small industry beginning in the 1950s resulted in the abandonment of ...agricultural lands and recovery of secondary forest. This unique history provides an excellent opportunity to study secondary forest succession and suggest strategies for tropical forest restoration. To determine the pattern of secondary succession, we describe the woody vegetation in 71 abandoned pastures and forest sites in four regions of Puerto Rico. The density, basal area, aboveground biomass, and species richness of the secondary forest sites were similar to those of the old growth forest sites (>80 yr) after approximately 40 years. The dominant species that colonized recently abandoned pastures occurred over a broad elevational range and are widespread in the neotropics. The species richness of Puerto Rican secondary forests recovered rapidly, but the species composition was quite different in comparison with old growth forest sites, suggesting that enrichment planting will be necessary to restore the original composition. Exotic species were some of the most abundant species in the secondary forest, but their long‐term impact depended on life history characteristics of each species. These data demonstrate that one restoration strategy for tropical forest in abandoned pastures is simply to protect the areas from fire, and allow natural regeneration to produce secondary forest. This strategy will be most effective if remnant forest (i.e., seed sources) still exist in the landscape and soils have not been highly degraded. Patterns of forest recovery also suggest strategies for accelerating natural recovery by planting a suite of generalist species that are common in recently abandoned pastures in Puerto Rico and throughout much of the neotropics.
Fine-resolution satellite imagery is needed for characterizing dry-season phenology in tropical forests since many tropical forests are very spatially heterogeneous due to their diverse species and ...environmental background. However, fine-resolution satellite imagery, such as Landsat, has a 16-day revisit cycle that makes it hard to obtain a high-quality vegetation index time series due to persistent clouds in tropical regions. To solve this challenge, this study explored the feasibility of employing a series of advanced technologies for reconstructing a high-quality Landsat time series from 2005 to 2009 for detecting dry-season phenology in tropical forests; Puerto Rico was selected as a testbed. We combined bidirectional reflectance distribution function (BRDF) correction, cloud and shadow screening, and contaminated pixel interpolation to process the raw Landsat time series and developed a thresholding method to extract 15 phenology metrics. The cloud-masked and gap-filled reconstructed images were tested with simulated clouds. In addition, the derived phenology metrics for grassland and forest in the tropical dry forest zone of Puerto Rico were evaluated with ground observations from PhenoCam data and field plots. Results show that clouds and cloud shadows are more accurately detected than the Landsat cloud quality assessment (QA) band, and that data gaps resulting from those clouds and shadows can be accurately reconstructed (R2 = 0.89). In the tropical dry forest zone, the detected phenology dates (such as greenup, browndown, and dry-season length) generally agree with the PhenoCam observations (R2 = 0.69), and Landsat-based phenology is better than MODIS-based phenology for modeling aboveground biomass and leaf area index collected in field plots (plot size is roughly equivalent to a 3 × 3 Landsat pixels). This study suggests that the Landsat time series can be used to characterize the dry-season phenology of tropical forests after careful processing, which will help to improve our understanding of vegetation–climate interactions at fine scales in tropical forests.
Height-diameter relationship models, denoted as H-D models, have important applications in sustainable forest management which include studying the vertical structure of a forest stand, understanding ...the habitat heterogeneity for wildlife niches, analyzing the growth rate pattern for making decisions regarding silvicultural treatments. Compared to monocultures, characterizing allometric relationships for uneven-aged, mixed-species forests, especially tropical forests, is more challenging and has historically received less attention. Modeling how the competitive interactions between trees of varying sizes and multiple species affects these relationships adds a high degree of complexity. In this study, five regression methods and five distance-independent competition indices were evaluated for temperate and pantropical tree species in different physiographic regions. A total of 163,922 individual tree measurements from the US Department of Agriculture, Forest Inventory and Analysis (FIA) database were used in analyses, which cover Appalachian plateau (AP) and Ridge and Valley (VR) in the southeastern US, as well as Caribbean (CAR) and Pacific (PAC) islands. Results indicated that the generalized additive model (GAM) and the Pearl and Reed model provided more accurate predictions than other regression methods examined. Models with competition indices had a varying level of predictability, while diameter ratio, cumulative distribution function and partitioned stand density index (PSDI) were found to improve the prediction accuracy for AP, VR and CAR. The results of this work provide additional insights on modeling H-D relationships for a variety of species in temperate and pantropical forests.
We mapped native, endemic, and introduced (i.e., exotic) tree species counts, relative basal areas of functional groups, species basal areas, and forest biomass from forest inventory data, satellite ...imagery, and environmental data for Puerto Rico and the Virgin Islands. Imagery included time series of Landsat composites and Moderate Resolution Imaging Spectroradiometer (MODIS)-based phenology. Environmental data included climate, land-cover, geology, topography, and road distances. Large-scale deforestation and subsequent forest regrowth are clear in the resulting maps decades after large-scale transition back to forest. Stand age, climate, geology, topography, road/urban locations, and protection are clearly influential. Unprotected forests on more accessible or arable lands are younger and have more introduced species and deciduous and nitrogen-fixing basal areas, fewer endemic species, and less biomass. Exotic species are widespread—except in the oldest, most remote forests on the least arable lands, where shade-tolerant exotics may persist. Although the maps have large uncertainty, their patterns of biomass, tree species diversity, and functional traits suggest that for a given geoclimate, forest age is a core proxy for forest biomass, species counts, nitrogen-fixing status, and leaf longevity. Geoclimate indicates hard-leaved species commonness. Until global wall-to-wall remote sensing data from specialized sensors are available, maps from multispectral image time series and other predictor data should help with running ecosystem models and as sustainable development indicators. Forest attribute models trained with a tree species ordination and mapped with nearest neighbor substitution (Phenological Gradient Nearest Neighbor method, PGNN) yielded larger correlation coefficients for observed vs. mapped tree species basal areas than Cubist regression tree models trained separately on each species. In contrast, Cubist regression tree models of forest structural and functional attributes yielded larger such correlation coefficients than the ordination-trained PGNN models.
Seasonally dry tropical forests are distributed across Latin America and the Caribbean and are highly threatened, with less than 10% of their original extent remaining in many countries. Using 835 ...inventories covering 4660 species of woody plants, we show marked floristic turnover among inventories and regions, which may be higher than in other neotropical biomes, such as savanna. Such high floristic turnover indicates that numerous conservation areas across many countries will be needed to protect the full diversity of tropical dry forests. Our results provide a scientific framework within which national decision-makers can contextuaiize the floristic significance of their dry forest at a regional and continental scale.
Remotely-sensed estimates of forest biomass are usually based on various measurements of canopy height, area, volume or texture, as derived from LiDAR, radar or fine spatial resolution imagery. These ...measurements are then calibrated to estimates of stand biomass that are primarily based on tree stem diameters. Although humid tropical forest seasonality can have low amplitudes compared with temperate regions, seasonal variations in growth-related factors like temperature, humidity, rainfall, wind speed and day length affect both tropical forest deciduousness and tree height-diameter relationships. Consequently, seasonal patterns in spectral measures of canopy greenness derived from satellite imagery should be related to tree height-diameter relationships and hence to estimates of forest biomass or biomass growth that are based on stand height or canopy area. In this study, we tested whether satellite image-based measures of tropical forest phenology, as estimated by indices of seasonal patterns in canopy greenness constructed from Landsat satellite images, can explain the variability in forest deciduousness, forest biomass and net biomass growth after already accounting for stand height or canopy area. Models of forest biomass that added phenology variables to structural variables similar to those that can be estimated by LiDAR or very high-spatial resolution imagery, like canopy height and crown area, explained up to 12% more variation in biomass. Adding phenology to structural variables explained up to 25% more variation in Net Biomass Growth (NBG). In all of the models, phenology contributed more as interaction terms than as single-effect terms. In addition, models run on only fully-forested plots performed better than models that included partially-forested plots. For forest NBG, the models produced better results when only those plots with a positive growth, from Inventory Cycle 1 to Inventory Cycle 2, were analyzed, as compared to models that included all plots