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Microbial communities and their associated enzyme activities affect the amount and chemical quality of carbon (C) in soils. Increasing nitrogen (N) deposition, particularly in N-rich tropical forests, is likely to change the composition and behavior of microbial communities and feed back on ecosystem structure and function. This study presents a novel assessment of mechanistic links between microbial responses to N deposition and shifts in soil organic matter (SOM) quality and quantity. We used phospholipid fatty acid (PLFA) analysis and microbial enzyme assays in soils to assess microbial community responses to long-term N additions in two distinct tropical rain forests. We used soil density fractionation and 13 C nuclear magnetic resonance (NMR) spectroscopy to measure related changes in SOM pool sizes and chemical quality. Microbial biomass increased in response to N fertilization in both tropical forests and corresponded to declines in pools of low-density SOM. The chemical quality of this soil C pool reflected ecosystem-specific changes in microbial community composition. In the lower-elevation forest, there was an increase in gram-negative bacteria PLFA biomass, and there were significant losses of labile C chemical groups (O-alkyls). In contrast, the upper-elevation tropical forest had an increase in fungal PLFAs with N additions and declines in C groups associated with increased soil C storage (alkyls). The dynamics of microbial enzymatic activities with N addition provided a functional link between changes in microbial community structure and SOM chemistry. Ecosystem-specific changes in microbial community composition are likely to have far-reaching effects on soil carbon storage and cycling. This study indicates that microbial communities in N-rich tropical forests can be sensitive to added N, but we can expect significant variability in how ecosystem structure and function respond to N deposition among tropical forest types.

Soil redox conditions exert substantial influence on biogeochemical processes in terrestrial ecosystems. Humid tropical forest soils are often characterized by fluctuating redox, yet how these dynamics affect patterns of organic matter decomposition and associated CO₂ fluxes remains poorly understood. We used a ¹³C-label incubation experiment in a humid tropical forest soil to follow the decomposition of plant litter and soil organic matter (SOM) in response to four redox regimes—static oxic or anoxic, and two oscillating treatments. We used high-resolution mass spectrometry to characterize the relative composition of organic compound classes in the water extractable OM. CO₂ production from litter and SOM showed different responses to redox treatments. While cumulative production of SOM-derived CO₂ was positively correlated with the length of oxic exposure (r = 0.89, n = 20), cumulative ¹³C-litterderived CO₂ production was not linked to oxygen availability. Litter-derived CO₂ production was highest under static anoxic conditions in the first half of the experiment, and later dropped to the lowest rate amongst the treatments. In anoxic soils, we observed depletion of more oxidized water-extractable OM (especially amino sugar-, carbohydrate-, and protein-like compounds) over the second half of the experiment, which likely served as substrates for anaerobic CO₂ production. Results from two-pool kinetic modeling showed that more frequent anoxic exposure limited decomposition of a slow-cycling C pool, but not a fast-cycling pool. These results suggest that aerobic and anaerobic heterotrophs were equally effective at degrading labile substrates released from fresh plant litter in this humid tropical forest soil, while aerobic decomposers were more effective in breaking down the potentially refractory compounds found in SOM.

Fine roots play an important role in plant nutrition, as well as in carbon, water, and nutrient cycling. Fine roots account for a third of terrestrial net primary production (NPP), and inclusion of their structure and function in global carbon models should improve predictions of ecosystem responses to climate change. However, studies focusing on underground plant components are much less frequent than those on aboveground structure. This is more marked in the tropics, where one-third of the planet's terrestrial NPP is produced. Some tropical forests have been more represented in the literature than others, as demonstrated in the collective studies in Puerto Rico. This Caribbean island's biodiversity, frequency of natural disturbances, ease of access to forests, and long-term plots have created an ideal place for the study of tropical ecological processes. This review of the literature emphasizes 50 years of root research and patterns revealed around Puerto Rico. The data in this review were compiled from scientific publications, conference reports, symposiums, and raw data shared by some researches. Emergent patterns include the shallow distribution of fine roots, the great variation in root biomass among different forest types, little variation in root phosphorus concentrations, the slow recovery of root biomass after Hurricane Hugo, and the fact that most data on roots come from the wet tropical Luquillo Experimental Forest, causing other habitat types to be underrepresented. This review also shows the gaps in knowledge about fine roots in the island's ecosystems, which should be used to promote and guide future studies. Abstract in Spanish is available with online material.

Intense hurricanes disturb many tropical forests, but the key mechanisms driving post-hurricane forest changes are not fully understood. In Puerto Rico, we used a replicated factorial experiment to determine the mechanisms of forest change associated with canopy openness and organic matter (debris) addition. Cascading effects from canopy openness accounted for most of the shifts in the forest biota and biotic processes, which included increased plant recruitment and richness, as well as the decreased abundance and diversity of several animal groups. Canopy opening decreased litterfall and litter moisture, thereby inhibiting lignin-degrading fungi, which slowed decomposition. Debris addition temporarily increased tree basal area. Elevated soil solution nitrate was a dominant response after past hurricanes; this effect only occurred in our experiment with simultaneous canopy-opening and debris treatments. Although debris is an important carbon and nutrient source, short-term responses to cyclonic storms appear to be largely driven by canopy opening.

The Anthropocene is a time of rapid change induced by human activities, including pulse and press disturbances that affect the species composition of local communities and connectivity among them, giving rise to spatiotemporal dynamics at multiple scales. We evaluate effects of global warming and repeated intense hurricanes on gastropod metacommunities in montane tropical rainforests of Puerto Rico for each of 28 consecutive years. Specifically, we quantified metacommunity structure each year; assessed effects of global warming, hurricane-induced disturbance, and secondary succession on interannual variation in metacommunity structure; and evaluated legacies of previous land use on metacommunity structure. Gastropods were sampled annually during a 28-year period characterized by disturbance and succession associated with 3 major hurricanes (Hurricanes Hugo, Georges, and Maria). For each year, we evaluated coherence (the extent to which the environmental distributions of species are uninterrupted along a common latent environmental gradient), species range turnover, and species range boundary clumping; and conducted co-occurrence analyses for each pair of species. We used generalized linear mixed-effects model to evaluate long-term responses of the metacommunity to aspects of global warming and disturbance. Metacommunity structure was remarkably stable, with consistent patterns of species co-occurrence. Disturbance, warming, and successional stage had little effect on metacommunity structure. Despite great temporal variation in environmental conditions, groups of species tracked their niche through space and time to maintain the same general structure. Consequently, metacommunity structure was highly resistant and resilient to multiple disturbances, even those that greatly altered forest structure.

The Anthropocene is a time of unprecedented and accelerating rates of environmental change that includes press (e.g., climate change) and pulse disturbances (e.g., cyclonic storms, land use change) that interact to affect spatiotemporal dynamics in the density, distribution, and biodiversity of organisms. We leverage three decades of spatially explicit data on the density of a tropical folivore (Lamponius portoricensis [Insecta, Phasmida]) in a hurricane-mediated ecosystem (montane rainforest of Puerto Rico), along with associated environmental attributes, to disentangle the effects of interacting disturbances at multiple spatial scales. Spatiotemporal variation in density at a small spatial scale is affected by disturbance-related characteristics (hurricane severity, time after most recent major hurricane, ambient temperature, and understory temperature), legacies of previous land use, and understory habitat structure. Nonetheless, only a small proportion of spatiotemporal variation in density was related to those characteristics. In contrast, the majority of interannual variation in mean density at a larger scale was related to disturbance characteristics and understory habitat structure. These factors combine to affect a weak and declining trend in the density of L. portoricensis over time. The low resistance of L. portoricensis to Hurricane Hugo, as compared to Hurricanes Georges and Maria, likely arose because a drought followed Hurricane Hugo. The disturbance regime of the region is predicted to include increases in ambient temperatures, frequency of high-intensity storms, and frequency of droughts. Such trends may combine to threaten the conservation status of L. portoricensis, and other species with which it shares similar life history characteristics.

The impact of Hurricane Maria on the U.S. Caribbean was used to study the causes of remotely-sensed spatial variation in the effects of (1) vegetation index loss and (2) landslide occurrence. The vegetation index is a measure of canopy ‘greenness’, a combination of leaf chlorophyll, leaf area, canopy cover and structure. A generalized linear model was made for each kind of effect, using idealized maps of the hurricane forces, along with three landscape characteristics that were significantly associated. In each model, one of these characteristics was forest fragmentation, and another was a measure of disturbance-propensity. For the greenness loss model, the hurricane force was wind, the disturbance-propensity measure was initial greenness, and the third landscape characteristic was fraction forest cover. For the landslide occurrence model, the hurricane force was rain, the disturbance-propensity measure was amount of land slope, and the third landscape characteristic was soil clay content. The model of greenness loss had a pseudo R2 of 0.73 and showed the U.S. Caribbean lost 31% of its initial greenness from the hurricane, with 51% lost from the initial in the Luquillo Experimental Forest (LEF) from Hurricane Maria along with Hurricane Irma. More greenness disturbance was seen in areas with less wind sheltering, higher elevation and topographic sides. The model of landslide occurrence had a pseudo R2 of 0.53 and showed the U.S. Caribbean had 34% of its area and 52% of the LEF area with a landslide density of at least one in 1 km2 from Hurricane Maria. Four experiments with parameters from previous storms of wind speed, storm duration, rainfall, and forest structure over the same storm path and topographic landscape were run as examples of possible future scenarios. While intensity of the storm makes by far the largest scenario difference, forest fragmentation makes a sizable difference especially in vulnerable areas of high clay content or high wind susceptibility. This study showed the utility of simple hurricane force calculations connected with landscape characteristics and remote-sensing data to determine forest susceptibility to hurricane effects.

After hurricane disturbances in tropical forests, the size structure and species composition are affected by immediate mortality, and subsequent recruitment and individual growth. Often, immediate post-disturbance stand-level data are presented but understanding of the components that affect changes in growth and longer-term responses to forest structure and composition are lacking. To answer questions about how mortality, recruitment, and growth change among successional Plant Functional Types (PFT) through time after a hurricane disturbance, we use long-term census data (1989–2014) collected in the Luquillo Experimental Forest, Puerto Rico. We developed an algorithm to fill missing diameter data from the long-term data set that was collected three months after Hurricane Hugo; and subsequently at five-year intervals. Both the immediate hurricane-induced mortality and subsequent mortality were lower in stems with larger diameters, but varied among successional PFTs Early, Mid, Late, and Palm. Tree growth rates were observed to decrease with time since the hurricane disturbance. Five years after the hurricane, mortality was minimal but then increased gradually with time. In contrast, recruitment was highest five years after the hurricane and then decreased with time. The palm Prestoea montana became the most abundant species in the forest after the hurricane, as it had the lowest immediate hurricane-induced and subsequent mortality, and the highest recruitment. Twenty-five years after the hurricane, the palm and the Late PFT dominate the forest after shifting species composition from pre-hurricane conditions.

Fine roots play an important role in plant nutrition, as well as in carbon, water, and nutrient cycling. Fine roots account for a third of terrestrial net primary production (NPP), and inclusion of their structure and function in global carbon models should improve predictions of ecosystem responses to climate change. However, studies focusing on underground plant components are much less frequent than those on aboveground structure. This is more marked in the tropics, where one-third of the planet's terrestrial NPP is produced. Some tropical forests have been more represented in the literature than others, as demonstrated in the collective studies in Puerto Rico. This Caribbean island's biodiversity, frequency of natural disturbances, ease of access to forests, and long-term plots have created an ideal place for the study of tropical ecological processes. This literature review emphasizes 50 years of root research and patterns revealed around Puerto Rico. The data in this review were compiled from scientific publications, conference reports, symposiums, and raw data shared by some researches. Emergent patterns include the shallow distribution of fine roots, the great variation in root biomass among different forest types, little variation in root phosphorus concentrations, the slow recovery of root biomass after Hurricane Hugo, and the fact that most data on roots come from the wet tropical Luquillo Experimental Forest, causing other habitat types to be underrepresented. This review also shows the gaps in knowledge about fine roots in the island's ecosystems, which should be used to promote and guide future studies.

Hurricanes cause dramatic changes to forests by opening the canopy and depositing debris onto the forest floor. How invasive rodent populations respond to hurricanes is not well understood, but shifts in rodent abundance and foraging may result from scarce fruit and seed resources that follow hurricanes. We conducted studies in a wet tropical forest in Puerto Rico to better understand how experimental (canopy trimming experiment) and natural (Hurricane Maria) hurricane effects alter populations of invasive rodents (Rattus rattus [rats] and Mus musculus [mice]) and their foraging behaviors. To monitor rodent populations, we used tracking tunnels (inked and baited cards inside tunnels enabling identification of animal visitors' footprints) within experimental hurricane plots (arborist trimmed in 2014) and reference plots (closed canopy forest). To assess shifts in rodent foraging, we compared seed removal of two tree species (Guarea guidonia and Prestoea acuminata) between vertebrate‐excluded and free‐access treatments in the same experimental and reference plots, and did so 3 months before and 9 months after Hurricane Maria (2017). Trail cameras were used to identify animals responsible for seed removal. Rat incidences generated from tracking tunnel surveys indicated that rat populations were not significantly affected by experimental or natural hurricanes. Before Hurricane Maria there were no mice in the forest interior, yet mice were present in forest plots closest to the road after the hurricane, and their forest invasion coincided with increased grass cover resulting from open forest canopy. Seed removal of Guarea and Prestoea across all plots was rat dominated (75%–100% rat‐removed) and was significantly less after than before Hurricane Maria. However, following Hurricane Maria, the experimental hurricane treatment plots of 2014 had 3.6 times greater seed removal by invasive rats than did the reference plots, which may have resulted from rats selecting post‐hurricane forest patches with greater understory cover for foraging. Invasive rodents are resistant to hurricane disturbance in this forest. Predictions of increased hurricane frequency from expected climate change should result in forest with more frequent periods of grassy understories and mouse presence, as well as with heightened rat foraging for fruit and seed in preexisting areas of disturbance.