Tropical cyclones are expected to intensify under a warming climate, with uncertain effects on tropical forests. One key challenge to predicting how more intense storms will influence these ...ecosystems is to attribute impacts specifically to storm meteorology rather than differences in forest characteristics. Here we compare tree damage data collected in the same forest in Puerto Rico after Hurricanes Hugo (1989, category 3), Georges (1998, category 3), and María (2017, category 4). María killed twice as many trees as Hugo, and for all but two species, broke 2- to 12-fold more stems than the other two storms. Species with high density wood were resistant to uprooting, hurricane-induced mortality, and were protected from breakage during Hugo but not María. Tree inventories and a wind exposure model allow us to attribute these differences in impacts to storm meteorology. A better understanding of risk factors associated with tree species susceptibility to severe storms is key to predicting the future of forest ecosystems under climate warming.
Topography affects abiotic conditions which can influence the structure, function, and dynamics of ecological communities. An increasing number of studies have demonstrated biological consequences of ...fine-scale topographic heterogeneity but we have a limited understanding of how We merged high-resolution (1 sq. meter) data on topography and canopy height derived from airborne lidar with ground-based data from 15 forest plots in Puerto Rico distributed along a precipitation gradient spanning ca. 800 to 3,500 mm yr(exp -1). Ground-based data included species composition, estimated above-ground biomass (AGB), and two key functional traits (wood density and leaf mass per area, LMA) that reflect resource-use strategies and a trade-off between hydraulic safety and hydraulic efficiency. We used hierarchical Bayesian models to evaluate how the interaction between topography climate is related to metrics of forest structure (i.e., canopy height and AGB), as well as taxonomic and functional alpha- and beta-diversity. Fine-scale topography (characterized with the topographic wetness index, TWI) significantly affected forest structure and the strength (and in some cases direction) of these effects varied across the precipitation gradient. In all plots, canopy height increased with topographic wetness but the effect was much stronger in dry compared to wet forest plots. In dry forest plots, topographically wetter microsites also had higher levels of AGB but in wet forest plots, topographically drier microsites had higher AGB. Fine-scale topography influenced functional composition but had only weak or non-significant effects on taxonomic and functional alpha- and beta-diversity. For instance, community-weighted wood density followed a similar pattern to AGB across plots. We also found a marginally significant association between variation of wood density and topographic heterogeneity that depended on climate context. Synthesis: The effects of fine-scale topographic heterogeneity on tropical forest structure and composition depend on the climate context. Our study demonstrates how a stronger integration of topographic heterogeneity across precipitation gradients could improve estimates of forest structure and biomass, and may provide insight to the ways that topography might mediate species responses to drought and climate change.
Predicting the fate of tropical forests under a changing climate requires understanding species responses to climatic variability and extremes. Seedlings may be particularly vulnerable to climatic ...stress given low stored resources and undeveloped roots; they also portend the potential effects of climate change on future forest composition. Here we use data for ca. 50,000 tropical seedlings representing 25 woody species to assess (i) the effects of interannual variation in rainfall and solar radiation between 2007 and 2016 on seedling survival over 9 years in a subtropical forest; and (ii) how spatial heterogeneity in three environmental factors—soil moisture, understory light, and conspecific neighborhood density—modulate these responses. Community‐wide seedling survival was not sensitive to interannual rainfall variability but interspecific variation in these responses was large, overwhelming the average community response. In contrast, community‐wide responses to solar radiation were predominantly positive. Spatial heterogeneity in soil moisture and conspecific density were the predominant and most consistent drivers of seedling survival, with the majority of species exhibiting greater survival at low conspecific densities and positive or nonlinear responses to soil moisture. This environmental heterogeneity modulated impacts of rainfall and solar radiation. Negative conspecific effects were amplified during rainy years and at dry sites, whereas the positive effects of radiation on survival were more pronounced for seedlings existing at high understory light levels. These results demonstrate that environmental heterogeneity is not only the main driver of seedling survival in this forest but also plays a central role in buffering or exacerbating impacts of climate fluctuations on forest regeneration. Since seedlings represent a key bottleneck in the demographic cycle of trees, efforts to predict the long‐term effects of a changing climate on tropical forests must take into account this environmental heterogeneity and how its effects on regeneration dynamics play out in long‐term stand dynamics.
Seedlings of Prestoea acuminata var. montana on the forest floor.
Species number, functional traits, and phylogenetic history all contribute to characterizing the biological diversity in plant communities. The phylogenetic component of diversity has been ...particularly difficult to quantify in species-rich tropical tree assemblages. The compilation of previously published (and often incomplete) data on evolutionary relationships of species into a composite phylogeny of the taxa in a forest, through such programs as Phylomatic, has proven useful in building community phylogenies although often of limited resolution. Recently, DNA barcodes have been used to construct a robust community phylogeny for nearly 300 tree species in a forest dynamics plot in Panama using a supermatrix method. In that study sequence data from three barcode loci were used to generate a well-resolved species-level phylogeny.
Here we expand upon this earlier investigation and present results on the use of a phylogenetic constraint tree to generate a community phylogeny for a diverse, tropical forest dynamics plot in Puerto Rico. This enhanced method of phylogenetic reconstruction insures the congruence of the barcode phylogeny with broadly accepted hypotheses on the phylogeny of flowering plants (i.e., APG III) regardless of the number and taxonomic breadth of the taxa sampled. We also compare maximum parsimony versus maximum likelihood estimates of community phylogenetic relationships as well as evaluate the effectiveness of one- versus two- versus three-gene barcodes in resolving community evolutionary history.
As first demonstrated in the Panamanian forest dynamics plot, the results for the Puerto Rican plot illustrate that highly resolved phylogenies derived from DNA barcode sequence data combined with a constraint tree based on APG III are particularly useful in comparative analysis of phylogenetic diversity and will enhance research on the interface between community ecology and evolution.
Our understanding of the long-lasting effects of human land use on soil fungal communities in tropical forests is limited. Yet, over 70% of all remaining tropical forests are growing in former ...agricultural or logged areas. We investigated the relationship among land use history, biotic and abiotic factors, and soil fungal community composition and diversity in a second-growth tropical forest in Puerto Rico. We coupled high-throughput DNA sequencing with tree community and environmental data to determine whether land use history had an effect on soil fungal community descriptors. We also investigated the biotic and abiotic factors that underlie such differences and asked whether the relative importance of biotic (tree diversity, basal tree area, and litterfall biomass) and abiotic (soil type, pH, iron, and total carbon, water flow, and canopy openness) factors in structuring soil fungal communities differed according to land use history. We demonstrated longlasting effects of land use history on soil fungal communities. At our research site, most of the explained variation in soil fungal composition (R2 = 18.6%), richness (R2 = 11.4%), and evenness (R2 = 10%) was associated with edaphic factors. Areas previously subject to both logging and farming had a soil fungal community with lower beta diversity and greater evenness of fungal operational taxonomic units (OTUs) than areas subject to light logging. Yet, fungal richness was similar between the two areas of historical land use. Together, these results suggest that fungal communities in disturbed areas are more homogeneous and diverse than in areas subject to light logging. Edaphic factors were the most strongly correlated with soil fungal composition, especially in areas subject to light logging, where soils are more heterogenous. High functional tree diversity in areas subject to both logging and farming led to stronger correlations between biotic factors and fungal composition than in areas subject to light logging. In contrast, fungal richness and evenness were more strongly correlated with biotic factors in areas of light logging, suggesting that these metrics might reflect long-term associations in old-growth forests. The large amount of unexplained variance in fungal composition suggests that these communities are structured by both stochastic and niche assemblage processes.
Leaf litter represents an important link between tree community composition, forest productivity and biomass, and ecosystem processes. In forests, the spatial distribution of trees and ...species-specific differences in leaf litter production and quality are likely to cause spatial heterogeneity in nutrient returns to the forest floor and, therefore, in the redistribution of soil nutrients. Using mapped trees and leaf litter data for 12 tree species in a subtropical forest with a well-documented history of land use, we: (1) parameterized spatially explicit models of leaf litter biomass and nutrient deposition; (2) assessed variation in leaf litter inputs across forest areas with different land use legacies; and (3) determined the degree to which the quantity and quality of leaf litter inputs and soil physical characteristics are associated with spatial heterogeneity in soil nutrient ratios (C:N and N:P). The models captured the effects of tree size and location on spatial variation in leaf litterfall (
R
2
= 0.31-0.79). For all 12 focal species, most of the leaf litter fell less than 5 m away from the source trees, generating fine-scale spatial heterogeneity in leaf litter inputs. Secondary forest species, which dominate areas in earlier successional stages, had lower leaf litter C:N ratios and produced less litter biomass than old-growth specialists. In contrast, P content and N:P ratios did not vary consistently among successional groups. Interspecific variation in leaf litter quality translated into differences in the quantity and quality (C:N) of total leaf litter biomass inputs and among areas with different land use histories. Spatial variation in leaf litter C:N inputs was the major factor associated with heterogeneity in soil C:N ratios relative to soil physical characteristics. In contrast, spatial variation soil N:P was more strongly associated with spatial variation in topography than heterogeneity in leaf litter inputs. The modeling approach presented here can be used to generate prediction surfaces for leaf litter deposition and quality onto the forest floor, a useful tool for understanding soil-vegetation feedbacks. A better understanding of the role of leaf litter inputs from secondary vegetation in restoring soil nutrient stocks will also assist in managing expanding secondary forests in tropical regions.
The degree to which turnover in biological communities is structured by deterministic or stochastic factors and the identities of influential deterministic factors are fundamental, yet unresolved, ...questions in ecology. Answers to these questions are particularly important for projecting the fate of forests with diverse disturbance histories worldwide. To uncover the processes governing turnover we use species-level molecular phylogenies and functional trait data sets for two long-term tropical forest plots with contrasting disturbance histories: one forest is older-growth, and one was recently disturbed. Having both phylogenetic and functional information further allows us to parse out the deterministic influences of different ecological filters. With the use of null models we find that compositional turnover was random with respect to phylogeny on average, but highly nonrandom with respect to measured functional traits. Furthermore, as predicted by a deterministic assembly process, the older-growth and disturbed forests were characterized by less than and greater than expected functional turnover, respectively. These results suggest that the abiotic environment, which changes due to succession in the disturbed forest, strongly governs the temporal dynamics of disturbed and undisturbed tropical forests. Predicting future changes in the composition of disturbed and undisturbed forests may therefore be tractable when using a functional-trait-based approach.
A number of recent studies have documented long-term declines in abundances of important arthropod groups, primarily in Europe and North America. These declines are generally attributed to habitat ...loss, but a recent study B.C. Lister, A. Garcia,
115, E10397-E10406 (2018) from the Luquillo Experimental Forest (LEF) in Puerto Rico attributed declines to global warming. We analyze arthropod data from the LEF to evaluate long-term trends within the context of hurricane-induced disturbance, secondary succession, and temporal variation in temperature. Our analyses demonstrate that responses to hurricane-induced disturbance and ensuing succession were the primary factors that affected total canopy arthropod abundances on host trees, as well as walkingstick abundance on understory shrubs. Ambient and understory temperatures played secondary roles for particular arthropod species, but populations were just as likely to increase as they were to decrease in abundance with increasing temperature. The LEF is a hurricane-mediated system, with major hurricanes effecting changes in temperature that are larger than those induced thus far by global climate change. To persist, arthropods in the LEF must contend with the considerable variation in abiotic conditions associated with repeated, large-scale, and increasingly frequent pulse disturbances. Consequently, they are likely to be well-adapted to the effects of climate change, at least over the short term. Total abundance of canopy arthropods after Hurricane Maria has risen to levels comparable to the peak after Hurricane Hugo. Although the abundances of some taxa have declined over the 29-y period, others have increased, reflecting species turnover in response to disturbance and secondary succession.
Projected increases in cyclonic storm intensity under a warming climate will have profound effects on forests, potentially changing these ecosystems from carbon sinks to sources. Forecasting storm ...impacts on these ecosystems requires consideration of risk factors associated with storm meteorology, landscape structure, and forest attributes. Here we evaluate risk factors associated with damage severity caused by Hurricanes María and Irma across Puerto Rican forests. Using field and remote sensing data, total forest aboveground biomass (AGB) lost to the storms was estimated at 10.44 (±2.33) Tg, ca. 23% of island-wide pre-hurricane forest AGB. Storm-related rainfall was a stronger predictor of forest damage than maximum wind speeds. Soil water storage capacity was also an important risk factor, corroborating the influence of rainfall on forest damage. Expected increases of 20% in hurricane-associated rainfall in the North Atlantic highlight the need to consider how such shifts, together with high speed winds, will affect terrestrial ecosystems.
Collaboration between ecologists and learning scientists can give rise to powerful models for scientific outreach within ecology. This paper presents a process by which learning scientists and ...ecologists codesigned a science curriculum that invites students to join an ecological community of practice. In the Journey to El Yunque middle school science curriculum, students engage with simulation models generated from data gathered by Luquillo Long Term Ecological Research (LUQ LTER) scientists. Journey to El Yunque students can explore post‐hurricane population changes in yagrumo (Cecropia schreberiana), tabonuco (Dacryodes excelsa), coquís (Eleutherodactylus coquí), snails (Caracolus caracola), anoles (Anolis stratulus and A. gundlachi), veiled stinkhorn mushrooms (Dictyophora indusiata), and caterpillars (Historis odius). Ecology‐based revisions to Journey to El Yunque have included adding models of the effects of repeated hurricanes on limiting factors, based in part on findings from a canopy trimming experiment. Revisions based on classroom testing include simplifying student‐facing model controls to allow students to focus on the essential model components. The ongoing collaboration that keeps the Journey to El Yunque curriculum on the cutting edge of ecological and educational advances has been sustained for over two decades. We attribute the longevity of this work to (1) the long‐term nature of LUQ LTER, (2) a sustained interdisciplinary collaboration, and (3) our long‐term relationships with schools.
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