pH selects for distinct N2O-reducing microbiomes in tropical soil microcosms.

Nitrous oxide (N2O), a greenhouse gas with ozone destruction potential, is mitigated by the microbial reduction to dinitrogen catalyzed by N2O reductase (NosZ). Bacteria with NosZ activity have been studied at circumneutral pH but the microbiology of low pH N2O reduction has remained elusive. Acidic (pH < 5) tropical forest soils were collected in the Luquillo Experimental Forest in Puerto Rico, and microcosms maintained with low (0.02 mM) and high (2 mM) N2O assessed N2O reduction at pH 4.5 and 7.3. All microcosms consumed N2O, with lag times of up to 7 months observed in microcosms with 2 mM N2O. Comparative metagenome analysis revealed that Rhodocyclaceae dominated in circumneutral microcosms under both N2O feeding regimes. At pH 4.5, Peptococcaceae dominated in high-N2O, and Hyphomicrobiaceae in low-N2O microcosms. Seventeen high-quality metagenome-assembled genomes (MAGs) recovered from the N2O-reducing microcosms harbored nos operons, with all eight MAGs derived from acidic microcosms carrying the Clade II type nosZ and lacking nitrite reductase genes (nirS/K). Five of the eight MAGs recovered from pH 4.5 microcosms represent novel taxa indicating an unexplored N2O-reducing diversity exists in acidic tropical soils. A survey of pH 3.5-5.7 soil metagenome datasets revealed that nosZ genes commonly occur, suggesting broad distribution of N2O reduction potential in acidic soils.

Termite sensitivity to temperature affects global wood decay rates.

Deadwood is a large global carbon store with its store size partially determined by biotic decay. Microbial wood decay rates are known to respond to changing temperature and precipitation. Termites are also important decomposers in the tropics but are less well studied. An understanding of their climate sensitivities is needed to estimate climate change effects on wood carbon pools. Using data from 133 sites spanning six continents, we found that termite wood discovery and consumption were highly sensitive to temperature (with decay increasing >6.8 times per 10°C increase in temperature)-even more so than microbes. Termite decay effects were greatest in tropical seasonal forests, tropical savannas, and subtropical deserts. With tropicalization (i.e., warming shifts to tropical climates), termite wood decay will likely increase as termites access more of Earth's surface.

The contribution of insects to global forest deadwood decomposition.

The amount of carbon stored in deadwood is equivalent to about 8 per cent of the global forest carbon stocks1. The decomposition of deadwood is largely governed by climate2-5 with decomposer groups-such as microorganisms and insects-contributing to variations in the decomposition rates2,6,7. At the global scale, the contribution of insects to the decomposition of deadwood and carbon release remains poorly understood7. Here we present a field experiment of wood decomposition across 55 forest sites and 6 continents. We find that the deadwood decomposition rates increase with temperature, and the strongest temperature effect is found at high precipitation levels. Precipitation affects the decomposition rates negatively at low temperatures and positively at high temperatures. As a net effect-including the direct consumption by insects and indirect effects through interactions with microorganisms-insects accelerate the decomposition in tropical forests (3.9% median mass loss per year). In temperate and boreal forests, we find weak positive and negative effects with a median mass loss of 0.9 per cent and -0.1 per cent per year, respectively. Furthermore, we apply the experimentally derived decomposition function to a global map of deadwood carbon synthesized from empirical and remote-sensing data, obtaining an estimate of 10.9 ± 3.2 petagram of carbon per year released from deadwood globally, with 93 per cent originating from tropical forests. Globally, the net effect of insects may account for 29 per cent of the carbon flux from deadwood, which suggests a functional importance of insects in the decomposition of deadwood and the carbon cycle.

Habitability Models for Astrobiology.

Habitability has been generally defined as the capability of an environment to support life. Ecologists have been using Habitat Suitability Models (HSMs) for more than four decades to study the habitability of Earth from local to global scales. Astrobiologists have been proposing different habitability models for some time, with little integration and consistency among them, being different in function to those used by ecologists. Habitability models are not only used to determine whether environments are habitable, but they also are used to characterize what key factors are responsible for the gradual transition from low to high habitability states. Here we review and compare some of the different models used by ecologists and astrobiologists and suggest how they could be integrated into new habitability standards. Such standards will help improve the comparison and characterization of potentially habitable environments, prioritize target selections, and study correlations between habitability and biosignatures. Habitability models are the foundation of planetary habitability science, and the synergy between ecologists and astrobiologists is necessary to expand our understanding of the habitability of Earth, the Solar System, and extrasolar planets.

Metagenomic Characterization of Soil Microbial Communities in the Luquillo Experimental Forest (Puerto Rico) and Implications for Nitrogen Cycling.

The phylogenetic and functional diversities of microbial communities in tropical rainforests and how these differ from those of temperate communities remain poorly described but are directly related to the increased fluxes of greenhouse gases such as nitrous oxide (N2O) from the tropics. Toward closing these knowledge gaps, we analyzed replicated shotgun metagenomes representing distinct life zones and an elevation gradient from four locations in the Luquillo Experimental Forest (LEF), Puerto Rico. These soils had a distinct microbial community composition and lower species diversity compared to those of temperate grasslands or agricultural soils. In contrast to the overall distinct community composition, the relative abundances and nucleotide sequences of N2O reductases (nosZ) were highly similar between tropical forest and temperate soils. However, respiratory NO reductase (norB) was 2-fold more abundant in the tropical soils, which might be relatable to their greater N2O emissions. Nitrogen fixation (nifH) also showed higher relative abundance in rainforest than in temperate soils, i.e., 20% versus 0.1 to 0.3% of bacterial genomes in each soil type harbored the gene, respectively. Finally, unlike temperate soils, LEF soils showed little stratification with depth in the first 0 to 30 cm, with ∼45% of community composition differences explained solely by location. Collectively, these results advance our understanding of spatial diversity and metabolic repertoire of tropical rainforest soil communities and should facilitate future ecological studies of these ecosystems. IMPORTANCE Tropical rainforests are the largest terrestrial sinks of atmospheric CO2 and the largest natural source of N2O emissions, two greenhouse gases that are critical for the climate. The microbial communities of rainforest soils that directly or indirectly, through affecting plant growth, contribute to these fluxes remain poorly described by cultured-independent methods. To close this knowledge gap, the present study applied shotgun metagenomics to samples selected from three distinct life zones within the Puerto Rico rainforest. The results advance our understanding of microbial community diversity in rainforest soils and should facilitate future studies of natural or manipulated perturbations of these critical ecosystems.

A new Caribbean species of Hylaeanura Arlé, 1966 (Collembola, Neanuridae, Pseudachorutinae).

We here describe a new Collembola species, Hylaeanura emiliae sp. nov., from the Luquillo Experimental Forest in Puerto Rico. We describe H. emiliae sp. nov. as a distinct species based on the enlarged sensilla s3 in antennal segment IV, the absence of modified sensorial setae in abdominal segment IV and the presence of four setae on each dens. An updated key with illustrations for the identification of worldwide species of the genus is included.

Green roofs in the tropics: design considerations and vegetation dynamics.

Green roofs (GR) have been proposed as a possible solution for urban stressors that, integrated with other remediation and mitigation actions, can lead the way to a more sustainable society. Even when some aspects of green roof design are well established and known (i.e. depth arrangements, materials, structural components, etc.) there is a need for further development on ecological attributes. This study is a descriptive analysis of suitable plant species for their possible incorporation in green roof designs with tropical climate conditions. Green roof research has been mostly led by temperate climate countries and has neglected to address tropical areas; this study aims to move research towards this knowledge gap. The evaluation of the vegetation dynamics in these novel ecosystems was done through a case study in the renovated facilities of the International Institute of Tropical Forestry in Río Piedras, Puerto Rico, which incorporated a set of green roofs in their infrastructure. We also sampled an older green roof built in the Social Sciences Faculty at the University of Puerto Rico at Río Piedras. A three-dimensional approach, the Point-Intercept Method, was taken in the vegetation surveys to capture as much as possible the green infrastructure of the roofs. Most of the originally planted species did not appear in these surveys. On the contrary, mainly new species dominated the areas. Along with the findings of these surveys and those in other tropical countries, a list of suitable species for green roofs in Puerto Rico is suggested, and some general recommendations are made for the better management of green roofs in tropical zones.

Checklist and distribution of Collembola from Greater Puerto Rico.

BACKGROUND: Springtails (Arthropoda, Hexapoda, Collembola) are small arthropods commonly found in soil, litter and other habitats all around the Globe. More than 9,000 species have been described worldwide, but knowledge about their diversity and distribution remains far from complete. Reports of springtail diversity in the Antilles are uneven, some islands are relatively well known, whereas others have not been explored at all. The fauna of Puerto Rico is reasonably well known, but many recent reports are scattered in published literature and unpublished theses. NEW INFORMATION: Here, we present a summary of all springtail species identified from the Bank of Puerto Rico, including unpublished records. As a result, we list 146 species including 43 unnamed, included in 65 genera and 17 families. Most species, 33, belong to Entomobryidae, but this possibly reflects the taxonomic expertise of specialists working in Puerto Rico rather than a real bias in the distribution of higher taxa in the islands. In addition to the new records, the database provides information on the world and local distribution of species listed. The dataset, presented here, is work in progress and will be updated as ongoing taxonomic inventories are completed.

New species of Furculanurida (Collembola, Neanuridae, Pseudachorutinae) from the Luquillo Mountains, Puerto Rico.

A new species of Furculanurida is described and illustrated. Furculanurida bistribus sp. nov. differs from other species of the genus by the presence of three eyes, three setae on the dens, and the white and purple coloration pattern. A key for identification of the world species of the genus is included.

A dataset for the effect of earthworm abundance and functional group diversity on plant litter decay and soil organic carbon level.

This paper describes data of earthworm abundance and functional group diversity regulate plant litter decay and soil organic carbon (SOC) level in global terrestrial ecosystems. The data also describes the potential effect of vegetation types, litter quality, litterbag mesh size, soil C/N, soil aggregate size, experimental types and length of experimental time on earthworm induced plant litter and SOC decay. The data were collected from 69 studies published between 1985 and 2018, covering 340 observations. This data article is related to the paper "Earthworm Abundance and Functional Group Diversity Regulate Plant Litter Decay and Soil Organic Carbon Level: A Global Meta-analysis" [1].

Tropical herbivorous phasmids, but not litter snails, alter decomposition rates by modifying litter bacteria.

Consumers can alter decomposition rates through both feces and selective feeding in many ecosystems, but these combined effects have seldom been examined in tropical ecosystems. Members of the detrital food web (litter-feeders or microbivores) should presumably have greater effects on decomposition than herbivores, members of the green food web. Using litterbag experiments within a field enclosure experiment, we determined the relative effects of common litter snails (Megalomastoma croceum) and herbivorous walking sticks (Lamponius portoricensis) on litter composition, decomposition rates, and microbes in a Puerto Rican rainforest, and whether consumer effects were altered by canopy cover presence. Although canopy presence did not alter consumers' effects, focal organisms had unexpected influences on decomposition. Decomposition was not altered by litter snails, but herbivorous walking sticks reduced leaf decomposition by about 50% through reductions in high quality litter abundance and, consequently, lower bacterial richness and abundance. This relatively unexplored but potentially important link between tropical herbivores, detritus, and litter microbes in this forest demonstrates the need to consider autotrophic influences when examining rainforest ecosystem processes.

Infrared heater system for warming tropical forest understory plants and soils.

The response of tropical forests to global warming is one of the largest uncertainties in predicting the future carbon balance of Earth. To determine the likely effects of elevated temperatures on tropical forest understory plants and soils, as well as other ecosystems, an infrared (IR) heater system was developed to provide in situ warming for the Tropical Responses to Altered Climate Experiment (TRACE) in the Luquillo Experimental Forest in Puerto Rico. Three replicate heated 4-m-diameter plots were warmed to maintain a 4°C increase in understory vegetation compared to three unheated control plots, as sensed by IR thermometers. The equipment was larger than any used previously and was subjected to challenges different from those of many temperate ecosystem warming systems, including frequent power surges and outages, high humidity, heavy rains, hurricanes, saturated clayey soils, and steep slopes. The system was able to maintain the target 4.0°C increase in hourly average vegetation temperatures to within ± 0.1°C. The vegetation was heterogeneous and on a 21° slope, which decreased uniformity of the warming treatment on the plots; yet, the green leaves were fairly uniformly warmed, and there was little difference among 0-10 cm depth soil temperatures at the plot centers, edges, and midway between. Soil temperatures at the 40-50 cm depth increased about 3°C compared to the controls after a month of warming. As expected, the soil in the heated plots dried faster than that of the control plots, but the average soil moisture remained adequate for the plants. The TRACE heating system produced an adequately uniform warming precisely controlled down to at least 50-cm soil depth, thereby creating a treatment that allows for assessing mechanistic responses of tropical plants and soil to warming, with applicability to other ecosystems. No physical obstacles to scaling the approach to taller vegetation (i.e., trees) and larger plots were observed.

Improving predictions of tropical forest response to climate change through integration of field studies and ecosystem modeling.

Tropical forests play a critical role in carbon and water cycles at a global scale. Rapid climate change is anticipated in tropical regions over the coming decades and, under a warmer and drier climate, tropical forests are likely to be net sources of carbon rather than sinks. However, our understanding of tropical forest response and feedback to climate change is very limited. Efforts to model climate change impacts on carbon fluxes in tropical forests have not reached a consensus. Here, we use the Ecosystem Demography model (ED2) to predict carbon fluxes of a Puerto Rican tropical forest under realistic climate change scenarios. We parameterized ED2 with species-specific tree physiological data using the Predictive Ecosystem Analyzer workflow and projected the fate of this ecosystem under five future climate scenarios. The model successfully captured interannual variability in the dynamics of this tropical forest. Model predictions closely followed observed values across a wide range of metrics including aboveground biomass, tree diameter growth, tree size class distributions, and leaf area index. Under a future warming and drying climate scenario, the model predicted reductions in carbon storage and tree growth, together with large shifts in forest community composition and structure. Such rapid changes in climate led the forest to transition from a sink to a source of carbon. Growth respiration and root allocation parameters were responsible for the highest fraction of predictive uncertainty in modeled biomass, highlighting the need to target these processes in future data collection. Our study is the first effort to rely on Bayesian model calibration and synthesis to elucidate the key physiological parameters that drive uncertainty in tropical forests responses to climatic change. We propose a new path forward for model-data synthesis that can substantially reduce uncertainty in our ability to model tropical forest responses to future climate.

Reassessing rainfall in the Luquillo Mountains, Puerto Rico: Local and global ecohydrological implications.

Mountains receive a greater proportion of precipitation than other environments, and thus make a disproportionate contribution to the world's water supply. The Luquillo Mountains receive the highest rainfall on the island of Puerto Rico and serve as a critical source of water to surrounding communities. The area's role as a long-term research site has generated numerous hydrological, ecological, and geological investigations that have been included in regional and global overviews that compare tropical forests to other ecosystems. Most of the forest- and watershed-wide estimates of precipitation (and evapotranspiration, as inferred by a water balance) have assumed that precipitation increases consistently with elevation. However, in this new analysis of all known current and historical rain gages in the region, we find that similar to other mountainous islands in the trade wind latitudes, leeward (western) watersheds in the Luquillo Mountains receive lower mean annual precipitation than windward (eastern) watersheds. Previous studies in the Luquillo Mountains have therefore overestimated precipitation in leeward watersheds by up to 40%. The Icacos watershed, however, despite being located at elevations 200-400 m below the tallest peaks and to the lee of the first major orographic barrier, receives some of the highest precipitation. Such lee-side enhancement has been observed in other island mountains of similar height and width, and may be caused by several mechanisms. Thus, the long-reported discrepancy of unrealistically low rates of evapotranspiration in the Icacos watershed is likely caused by previous underestimation of precipitation, perhaps by as much as 20%. Rainfall/runoff ratios in several previous studies suggested either runoff excess or runoff deficiency in Luquillo watersheds, but this analysis suggests that in fact they are similar to other tropical watersheds. Because the Luquillo Mountains often serve as a wet tropical archetype in global assessments of basic ecohydrological processes, these revised estimates are relevant to regional and global assessments of runoff efficiency, hydrologic effects of reforestation, geomorphic processes, and climate change.

A soil burn severity index for understanding soil-fire relations in tropical forests.

Methods for evaluating the impact of fires within tropical forests are needed as fires become more frequent and human populations and demands on forests increase. Short- and long-term fire effects on soils are determined by the prefire, fire, and postfire environments. We placed these components within a fire-disturbance continuum to guide our literature synthesis and develop an integrated soil burn severity index. The soil burn severity index provides a set of indicators that reflect the range of conditions present after a fire. The index consists of seven levels, an unburned level and six other levels that describe a range of postfire soil conditions. We view this index as a tool for understanding the effects of fires on the forest floor, with the realization that as new information is gained, the index may be modified as warranted.

Characterizing forest fragments in boreal, temperate, and tropical ecosystems.

An increased ability to analyze landscapes in a spatial manner through the use of remote sensing leads to improved capabilities for quantifying human-induced forest fragmentation. Developments of spatially explicit methods in landscape analyses are emerging. In this paper, the image delineation software program eCognition and the spatial pattern analysis program FRAGSTATS were used to quantify patterns of forest fragments on six landscapes across three different climatic regions characterized by different moisture regimes and different influences of human pressure. Our results support the idea that landscapes with higher road and population density are more fragmented; however, there are other, equally influential factors contributing to fragmentation, such as moisture regime, historic land use, and fire dynamics. Our method provided an objective means to characterize landscapes and assess patterns of forest fragments across different forested ecosystems by addressing the limitations of pixel-based classification and incorporating image objects.

Forest structure and downed woody debris in boreal, temperate, and tropical forest fragments.

Forest fragmentation affects the heterogeneity of accumulated fuels by increasing the diversity of forest types and by increasing forest edges. This heterogeneity has implications in how we manage fuels, fire, and forests. Understanding the relative importance of fragmentation on woody biomass within a single climatic regime, and along climatic gradients, will improve our ability to manage forest fuels and predict fire behavior. In this study we assessed forest fuel characteristics in stands of differing moisture, i.e., dry and moist forests, structure, i.e., open canopy (typically younger) vs. closed canopy (typically older) stands, and size, i.e., small (10-14 ha), medium (33 to 60 ha), and large (100-240 ha) along a climatic gradient of boreal, temperate, and tropical forests. We measured duff, litter, fine and coarse woody debris, standing dead, and live biomass in a series of plots along a transect from outside the forest edge to the fragment interior. The goal was to determine how forest structure and fuel characteristics varied along this transect and whether this variation differed with temperature, moisture, structure, and fragment size. We found nonlinear relationships of coarse woody debris, fine woody debris, standing dead and live tree biomass with mean annual median temperature. Biomass for these variables was greatest in temperate sites. Forest floor fuels (duff and litter) had a linear relationship with temperature and biomass was greatest in boreal sites. In a five-way multivariate analysis of variance we found that temperature, moisture, and age/structure had significant effects on forest floor fuels, downed woody debris, and live tree biomass. Fragment size had an effect on forest floor fuels and live tree biomass. Distance from forest edge had significant effects for only a few subgroups sampled. With some exceptions edges were not distinguishable from interiors in terms of fuels.

Decay of aspen (Populus tremuloides Michx.) wood in moist and dry boreal, temperate, and tropical forest fragments.

In this study, we set up a wood decomposition experiment to i) quantify the percent of mass remaining, decay constant and performance strength of aspen stakes (Populus tremuloides) in dry and moist boreal (Alaska and Minnesota, USA), temperate (Washington and Idaho, USA), and tropical (Puerto Rico) forest types, and ii) determine the effects of fragmentation on wood decomposition rates as related to fragment size, forest age (and/or structure) and climate at the macro- and meso-scales. Fragment sizes represented the landscape variability within a climatic region. Overall, the mean small fragments area ranged from 10-14 ha, medium-sized fragments 33 to 60 ha, and large fragments 100-240 ha. We found that: i) aspen stakes decayed fastest in the tropical sites, and the slowest in the temperate forest fragments, ii) the percent of mass remaining was significantly greater in dry than in moist forests in boreal and temperate fragments, while the opposite was true for the tropical forest fragments, iii) no effect of fragment size on the percent of mass remaining of aspen stakes in the boreal sites, temperate dry, and tropical moist forests, and iv) no significant differences of aspen wood decay between forest edges and interior forest in boreal, temperate and tropical fragments. We conclude that: i) moisture condition is an important control over wood decomposition over broad climate gradients; and that such relationship can be non linear, and ii) the presence of a particular group of organism (termites) can significantly alter the decay rates of wood more than what might be predicted based on climatic factors alone. Biotic controls on wood decay might be more important predictors of wood decay in tropical regions, while abiotic constraints seems to be important determinants of decay in cold forested fragments.

Soil ecological interactions: comparisons between tropical and subalpine forests.

Soil fauna can influence soil processes through interactions with the microbial community. Due to the complexity of the functional roles of fauna and their effects on microbes, little consensus has been reached on the extent to which soil fauna can regulate microbial activities. We quantified soil microbial biomass and maximum growth rates in control and fauna-excluded treatments in dry and wet tropical forests and north- and south-facing subalpine forests to test whether soil fauna effects on microbes are different in tropical and subalpine forests. Exclusion of fauna was established by physically removing the soil macrofauna and applying naphthalene. The effect of naphthalene application on the biomass of microbes that mineralize salicylate was quantified using the substrate induced growth response method. We found that: (1) the exclusion of soil fauna resulted in a higher total microbial biomass and lower maximum growth rate in the subalpine forests, (2) soil fauna exclusion did not affect the microbial biomass and growth rate in the tropical forests, and (3) the microbial biomass of salicylate mineralizers was significantly enhanced in the fauna-exclusion treatment in the tropical wet and the south-facing subalpine forests. We conclude that non-target effects of naphthalene on the microbial community alone cannot explain the large differences in total microbial biomass found between control and fauna-excluded treatments in the subalpine forests. Soil fauna have relatively larger effects on the microbial activities in the subalpine forests than in tropical dry and wet forests.