Mycorrhizal feedbacks influence global forest structure and diversity.

One mechanism proposed to explain high species diversity in tropical systems is strong negative conspecific density dependence (CDD), which reduces recruitment of juveniles in proximity to conspecific adult plants. Although evidence shows that plant-specific soil pathogens can drive negative CDD, trees also form key mutualisms with mycorrhizal fungi, which may counteract these effects. Across 43 large-scale forest plots worldwide, we tested whether ectomycorrhizal tree species exhibit weaker negative CDD than arbuscular mycorrhizal tree species. We further tested for conmycorrhizal density dependence (CMDD) to test for benefit from shared mutualists. We found that the strength of CDD varies systematically with mycorrhizal type, with ectomycorrhizal tree species exhibiting higher sapling densities with increasing adult densities than arbuscular mycorrhizal tree species. Moreover, we found evidence of positive CMDD for tree species of both mycorrhizal types. Collectively, these findings indicate that mycorrhizal interactions likely play a foundational role in global forest diversity patterns and structure.

Impacts of insect frass and cadavers on soil surface litter decomposition along a tropical forest temperature gradient.

Insect herbivores play important roles in shaping many ecosystem processes, but how climate change will alter the effects of insect herbivory are poorly understood. To address this knowledge gap, we quantified for the first time how insect frass and cadavers affected leaf litter decomposition rates and nutrient release along a highly constrained 4.3°C mean annual temperature (MAT) gradient in a Hawaiian montane tropical wet forest. We constructed litterbags of standardized locally sourced leaf litter, with some amended with insect frass + cadavers to produce treatments designed to simulate ambient (Control = no amendment), moderate (Amended-Low = 2 × Control level), or severe (Amended-High = 11 × Control level) insect outbreak events. Multiple sets of these litterbags were deployed across the MAT gradient, with individual litterbags collected periodically over one year to assess how rising MAT altered the effects of insect deposits on litter decomposition rates and nitrogen (N) release. Increased MAT and insect inputs additively increased litter decomposition rates and N immobilization rates, with effects being stronger for Amended-High litterbags. However, the apparent temperature sensitivity (Q 10) of litter decomposition was not clearly affected by amendments. The effects of adding insect deposits in this study operated differently than the slower litter decomposition and greater N mobilization rates often observed in experiments which use chemical fertilizers (e.g., urea, ammonium nitrate). Further research is required to understand mechanistic differences between amendment types. Potential increases in outbreak-related herbivore deposits coupled with climate warming will accelerate litter decomposition and nutrient cycling rates with short-term consequences for nutrient cycling and carbon storage in tropical montane wet forests.

Distribution of biomass dynamics in relation to tree size in forests across the world.

Tree size shapes forest carbon dynamics and determines how trees interact with their environment, including a changing climate. Here, we conduct the first global analysis of among-site differences in how aboveground biomass stocks and fluxes are distributed with tree size. We analyzed repeat tree censuses from 25 large-scale (4-52 ha) forest plots spanning a broad climatic range over five continents to characterize how aboveground biomass, woody productivity, and woody mortality vary with tree diameter. We examined how the median, dispersion, and skewness of these size-related distributions vary with mean annual temperature and precipitation. In warmer forests, aboveground biomass, woody productivity, and woody mortality were more broadly distributed with respect to tree size. In warmer and wetter forests, aboveground biomass and woody productivity were more right skewed, with a long tail towards large trees. Small trees (1-10 cm diameter) contributed more to productivity and mortality than to biomass, highlighting the importance of including these trees in analyses of forest dynamics. Our findings provide an improved characterization of climate-driven forest differences in the size structure of aboveground biomass and dynamics of that biomass, as well as refined benchmarks for capturing climate influences in vegetation demographic models.

Optimizing invasive species management using mathematical programming to support stewardship of water and carbon-based ecosystem services.

Invasive species alter hydrologic processes at watershed scales, with impacts to biodiversity and the supporting ecosystem services. This effect is aggravated by climate change. Here, we integrated modelled hydrologic data, remote sensing products, climate data, and linear mixed integer optimization (MIP) to identify stewardship actions across space and time that can reduce the impact of invasive species. The study area is the windward coast of Hawai'i Island (USA) across which non-native strawberry guava occurrence varies from extremely dense stands in lower watershed reaches, to low densities in upper watershed forests. We focused on the removal of strawberry guava, an invader that exerts significant impacts on watershed condition. MIP analyses spatially optimized the assignment of effective management actions to increase water yield, generate revenue from enhanced freshwater services, and income from removed biomass. The hydrological benefit of removing guava, often marginal when considered in isolation, was financially quantified, and single- and multiobjective MIP formulations were then developed over a 10-year planning horizon. Optimization resulted in $2.27 million USD benefit over the planning horizon using a payment-for-ecosystem-services scheme. That value jumped to $4.67 million when allowing work schedules with overnight camping to reduce costs. Pareto frontiers of weighted pairs of management goals showed the benefit of clustering treatments over space and time to improve financial efficiency. Values of improved land-water natural capital using payment-for-ecosystem-services schemes are provided for several combinations of spatial, temporal, economical, and ecosystem services flows.

Intraspecific Variation Along an Elevational Gradient Alters Seed Scarification Responses in the Polymorphic Tree Species Acacia koa.

Physical dormancy in seeds can challenge restoration efforts where scarification conditions for optimal germination and seedling vigor are unknown. For species that occur along wide environmental gradients, optimal scarification conditions may also differ by seed source. We examined intraspecific variation in optimal scarification conditions for germination and seedling performance in koa (Acacia koa), which occurs across a wide range of environmental conditions. To evaluate scarification responses, we recorded imbibition percentage, germination percentage, germination time, seedling abnormalities, early mortality, seedling growth, and seedling survivorship. From these, we developed a scarification index (SI) that integrates these measures simultaneously. We hypothesized that seeds from lower elevation sources exposed to higher temperatures would have harder seed coats and would require more intense scarification treatments. To test this hypothesis, we repeatedly exposed seeds to hot water differing in temperature and time until seeds imbibed. Supporting the hypothesis, seeds from lower elevation sources generally required more intense scarification, although we found substantial variation among sources. Koa seeds germinated in about a week following imbibition. Boiling seeds (i.e., maintaining at 100°C) was effective for imbibing seeds but it also substantially reduced germination percentages. Repeated exposure to 90 to 100°C water did not reduce germination percentage but decreased seedling performance and increased early mortality. No seeds remained unimbibed after six attempts of boiling germinated whereas seeds remaining unimbibed after 15 attempts of exposure to 90 to 100°C water showed high germination percentages. Abnormalities in seedling development were rare but increased with treatment intensity. Exposure to 100°C water for 1 min overall generated the best SI values but the best treatment differed by elevation, and the treatment with the best SI was rarely predicted from the highest germination percentages. Seeds that imbibed without any treatment germinated at the same level as manually filed seeds but produced poor seedling quality. Variation in mother tree environments along an elevational gradient can lead to differences in seed coat characteristics, which may explain differing responses to treatments. Scarification treatments affected processes beyond imbibition and germination and using an index like SI may improve efficiency by identifying optimal scarification treatments while reducing seed waste.

Interannual variation in rainfall modulates temperature sensitivity of carbon allocation and flux in a tropical montane wet forest.

Tropical forests exert a disproportionately large influence on terrestrial carbon (C) balance but projecting the effects of climate change on C cycling in tropical forests remains uncertain. Reducing this uncertainty requires improved quantification of the independent and interactive effects of variable and changing temperature and precipitation regimes on C inputs to, cycling within and loss from tropical forests. Here, we quantified aboveground litterfall and soil-surface CO2 efflux ("soil respiration"; FS ) in nine plots organized across a highly constrained 5.2°C mean annual temperature (MAT) gradient in tropical montane wet forest. We used five consecutive years of these measurements, during which annual rainfall (AR) steadily increased, in order to: (a) estimate total belowground C flux (TBCF); (b) examine how interannual variation in AR alters the apparent temperature dependency (Q10 ) of above- and belowground C fluxes; and (c) quantify stand-level C allocation responses to MAT and AR. Averaged across all years, FS , litterfall, and TBCF increased positively and linearly with MAT, which accounted for 49, 47, and 46% of flux rate variation, respectively. Rising AR lowered TBCF and FS , but increased litterfall, with patterns representing interacting responses to declining light. The Q10 of FS , litterfall, and TBCF all decreased with increasing AR, with peak sensitivity to MAT in the driest year and lowest sensitivity in the wettest. These findings support the conclusion that for this tropical montane wet forest, variations in light, water, and nutrient availability interact to strongly influence productivity (litterfall+TBCF), the sensitivity of above- and belowground C fluxes to rising MAT (Q10 of FS , litterfall, and TBCF), and C allocation patterns (TBCF:[litterfall+TBCF]).

Arbuscular mycorrhizal trees influence the latitudinal beta-diversity gradient of tree communities in forests worldwide.

Arbuscular mycorrhizal (AM) and ectomycorrhizal (EcM) associations are critical for host-tree performance. However, how mycorrhizal associations correlate with the latitudinal tree beta-diversity remains untested. Using a global dataset of 45 forest plots representing 2,804,270 trees across 3840 species, we test how AM and EcM trees contribute to total beta-diversity and its components (turnover and nestedness) of all trees. We find AM rather than EcM trees predominantly contribute to decreasing total beta-diversity and turnover and increasing nestedness with increasing latitude, probably because wide distributions of EcM trees do not generate strong compositional differences among localities. Environmental variables, especially temperature and precipitation, are strongly correlated with beta-diversity patterns for both AM trees and all trees rather than EcM trees. Results support our hypotheses that latitudinal beta-diversity patterns and environmental effects on these patterns are highly dependent on mycorrhizal types. Our findings highlight the importance of AM-dominated forests for conserving global forest biodiversity.

Mean annual temperature influences local fine root proliferation and arbuscular mycorrhizal colonization in a tropical wet forest.

Mean annual temperature (MAT) is an influential climate factor affecting the bioavailability of growth-limiting nutrients nitrogen (N) and phosphorus (P). In tropical montane wet forests, warmer MAT drives higher N bioavailability, while patterns of P availability are inconsistent across MAT. Two important nutrient acquisition strategies, fine root proliferation into bulk soil and root association with arbuscular mycorrhizal fungi, are dependent on C availability to the plant via primary production. The case study presented here tests whether variation in bulk soil N bioavailability across a tropical montane wet forest elevation gradient (5.2°C MAT range) influences (a) morphology fine root proliferation into soil patches with elevated N, P, and N+P relative to background soil and (b) arbuscular mycorrhizal fungal (AMF) colonization of fine roots in patches. We created a fully factorial fertilized root ingrowth core design (N, P, N+P, unfertilized control) representing soil patches with elevated N and P bioavailability relative to background bulk soil. Our results show that percent AMF colonization of roots increased with MAT (r 2 = .19, p = .004), but did not respond to fertilization treatments. Fine root length (FRL), a proxy for root foraging, increased with MAT in N+P-fertilized patches only (p = .02), while other fine root morphological parameters did not respond to the gradient or fertilized patches. We conclude that in N-rich, fine root elongation into areas with elevated N and P declines while AMF abundance increases with MAT. These results indicate a tradeoff between P acquisition strategies occurring with changing N bioavailability, which may be influenced by higher C availability with warmer MAT.

Impact of Mean Annual Temperature on Nutrient Availability in a Tropical Montane Wet Forest.

Despite growing understanding of how rising temperatures affect carbon cycling, the impact of long-term and whole forest warming on the suite of essential and potentially limiting nutrients remains understudied, particularly for elements other than N and P. Whole ecosystem warming experiments are limited, environmental gradients are often confounded by variation in factors other than temperature, and few studies have been conducted in the tropics. We examined litterfall, live foliar nutrient content, foliar nutrient resorption efficiency (NRE), nutrient return, and foliar nutrient use efficiency (NUE) of total litterfall and live foliage of two dominant trees to test hypotheses about how increasing mean annual temperature (MAT) impacts the availability and ecological stoichiometry of C, N, P, K, Ca, Mg, Mn, Fe, Zn, and Cu in tropical montane wet forests located along a 5.2°C gradient in Hawaii. Live foliage responded to increasing MAT with increased N and K concentrations, decreased C and Mn concentrations, and no detectable change in P concentration or in foliar NRE. Increases in MAT increased nutrient return via litterfall for N, K, Mg, and Zn and foliar NUE for Mn and Cu, while decreasing nutrient return for Cu and foliar NUE for K. The N:P of litterfall and live foliage increased with MAT, while there was no detectable effect of MAT on C:P. The ratio of live foliar N or P to base cations and micronutrients was variable across elements and species. Increased MAT resulted in declining N:K and P:K for one species, while only P:K declined for the other. N:Ca and N:Mn increased with MAT for both species, while N:Mg increased for one and P:Mn increased for the other species. Overall, results from this study suggest that rising MAT in tropical montane wet forest: (i) increases plant productivity and the cycling and availability of N, K, Mg, and Zn; (ii) decreases the cycling and availability of Mn and Cu; (iii) has little direct effect on P, Ca or Fe; and (iv) affects ecological stoichiometry in ways that may exacerbate P-as well as other base cation and micronutrient - limitations to tropical montane forest productivity.

Changes in soil bacterial community diversity following the removal of invasive feral pigs from a Hawaiian tropical montane wet forest.

Nonnative, invasive feral pigs (Sus scrofa) modify habitats by disturbing soils and vegetation, which can alter biogeochemical processes. Soil microbial communities drive nutrient cycling and therefore also play important roles in shaping ecosystem structure and function, but the responses of soil microbes to nonnative ungulate removal remains poorly studied. We examined changes in the soil bacterial community over a ~25 year chronosequence of feral pig removal in tropical montane wet forests on the Island of Hawai'i. We extracted bacterial eDNA from soil samples collected inside and outside of ungulate exclosures along this chronosequence and sequenced the eDNA using the Illumina platform. We found that ungulate removal increased diversity of soil bacteria, with diversity scores positively correlated with time since removal. While functional and phylogenetic diversity were not significantly different between pig present and pig removed soils, soil bulk density, which decreases following the removal of feral pigs, was a useful predictor of dissimilarity among sites and correlated to changes in functional diversity. Additionally, increases in soil porosity, potassium, and calcium were correlated to increases in functional diversity. Finally, sites with greater mean annual temperatures were shown to have higher scores of both functional and phylogenetic diversity. As such, we conclude that feral pigs influence overall bacterial community diversity directly while influencing functional diversity indirectly through alterations to soil structure and nutrients. Comparatively, phylogenetic differences between communities are better explained by mean annual temperature as a climatic predictor of community dissimilarity.

OpenNahele: the open Hawaiian forest plot database.

BACKGROUND: This data paper provides a description of OpenNahele, the open Hawaiian forest plot database. OpenNahele includes 530 forest plots across the Hawaiian archipelago containing 43,590 individuals of 185 native and alien tree, shrub and tree fern species across six islands. We include estimates of maximum plant size (D950.1 and Dmax3) for 58 woody plant species, a key functional trait associated with dispersal distance and competition for light. OpenNahele can serve as a platform to test key ecological, evolutionary and conservation questions in a hotspot archipelago. NEW INFORMATION: OpenNahele is the first database that compiles data from a large number of forest plots across the Hawaiian archipelago to allow broad and high resolution studies of biodiversity patterns.Keywords: Hawaii, forests, islands, biodiversity, community ecology, evolutionary ecology.

Vertical foraging shifts in Hawaiian forest birds in response to invasive rat removal.

Worldwide, native species increasingly contend with the interacting stressors of habitat fragmentation and invasive species, yet their combined effects have rarely been examined. Direct negative effects of invasive omnivores are well documented, but the indirect effects of resource competition or those caused by predator avoidance are unknown. Here we isolated and examined the independent and interactive effects of invasive omnivorous Black rats (Rattus rattus) and forest fragment size on the interactions between avian predators and their arthropod prey. Our study examines whether invasive omnivores and ecosystem fragment size impact: 1) the vertical distribution of arthropod species composition and abundance, and 2) the vertical profile of foraging behaviors of five native and two non-native bird species found in our study system. We predicted that the reduced edge effects and greater structural complexity and canopy height of larger fragments would limit the total and proportional habitat space frequented by rats and thus limit their impact on both arthropod biomass and birds' foraging behavior. We experimentally removed invasive omnivorous Black rats across a 100-fold (0.1 to 12 ha) size gradient of forest fragments on Hawai'i Island, and paired foraging observations of forest passerines with arthropod sampling in the 16 rat-removed and 18 control fragments. Rat removal was associated with shifts in the vertical distribution of arthropod biomass, irrespective of fragment size. Bird foraging behavior mirrored this shift, and the impact of rat removal was greater for birds that primarily eat fruit and insects compared with those that consume nectar. Evidence from this model study system indicates that invasive rats indirectly alter the feeding behavior of native birds, and consequently impact multiple trophic levels. This study suggests that native species can modify their foraging behavior in response to invasive species removal and presumably arrival through behavioral plasticity.

Climate sensitive size-dependent survival in tropical trees.

Survival rates of large trees determine forest biomass dynamics. Survival rates of small trees have been linked to mechanisms that maintain biodiversity across tropical forests. How species survival rates change with size offers insight into the links between biodiversity and ecosystem function across tropical forests. We tested patterns of size-dependent tree survival across the tropics using data from 1,781 species and over 2 million individuals to assess whether tropical forests can be characterized by size-dependent life-history survival strategies. We found that species were classifiable into four 'survival modes' that explain life-history variation that shapes carbon cycling and the relative abundance within forests. Frequently collected functional traits, such as wood density, leaf mass per area and seed mass, were not generally predictive of the survival modes of species. Mean annual temperature and cumulative water deficit predicted the proportion of biomass of survival modes, indicating important links between evolutionary strategies, climate and carbon cycling. The application of survival modes in demographic simulations predicted biomass change across forest sites. Our results reveal globally identifiable size-dependent survival strategies that differ across diverse systems in a consistent way. The abundance of survival modes and interaction with climate ultimately determine forest structure, carbon storage in biomass and future forest trajectories.

Response to Comment on "Plant diversity increases with the strength of negative density dependence at the global scale".

Hülsmann and Hartig suggest that ecological mechanisms other than specialized natural enemies or intraspecific competition contribute to our estimates of conspecific negative density dependence (CNDD). To address their concern, we show that our results are not the result of a methodological artifact and present a null-model analysis that demonstrates that our original findings-(i) stronger CNDD at tropical relative to temperate latitudes and (ii) a latitudinal shift in the relationship between CNDD and species abundance-persist even after controlling for other processes that might influence spatial relationships between adults and recruits.

Response to Comment on "Plant diversity increases with the strength of negative density dependence at the global scale".

Chisholm and Fung claim that our method of estimating conspecific negative density dependence (CNDD) in recruitment is systematically biased, and present an alternative method that shows no latitudinal pattern in CNDD. We demonstrate that their approach produces strongly biased estimates of CNDD, explaining why they do not detect a latitudinal pattern. We also address their methodological concerns using an alternative distance-weighted approach, which supports our original findings of a latitudinal gradient in CNDD and a latitudinal shift in the relationship between CNDD and species abundance.

Correction: Evaluating the role of land cover and climate uncertainties in computing gross primary production in Hawaiian Island ecosystems.

[This corrects the article DOI: 10.1371/journal.pone.0184466.].

Evaluating the role of land cover and climate uncertainties in computing gross primary production in Hawaiian Island ecosystems.

Gross primary production (GPP) is the Earth's largest carbon flux into the terrestrial biosphere and plays a critical role in regulating atmospheric chemistry and global climate. The Moderate Resolution Imaging Spectrometer (MODIS)-MOD17 data product is a widely used remote sensing-based model that provides global estimates of spatiotemporal trends in GPP. When the MOD17 algorithm is applied to regional scale heterogeneous landscapes, input data from coarse resolution land cover and climate products may increase uncertainty in GPP estimates, especially in high productivity tropical ecosystems. We examined the influence of using locally specific land cover and high-resolution local climate input data on MOD17 estimates of GPP for the State of Hawaii, a heterogeneous and discontinuous tropical landscape. Replacing the global land cover data input product (MOD12Q1) with Hawaii-specific land cover data reduced statewide GPP estimates by ~8%, primarily because the Hawaii-specific land cover map had less vegetated land area compared to the global land cover product. Replacing coarse resolution GMAO climate data with Hawaii-specific high-resolution climate data also reduced statewide GPP estimates by ~8% because of the higher spatial variability of photosynthetically active radiation (PAR) in the Hawaii-specific climate data. The combined use of both Hawaii-specific land cover and high-resolution Hawaii climate data inputs reduced statewide GPP by ~16%, suggesting equal and independent influence on MOD17 GPP estimates. Our sensitivity analyses within a heterogeneous tropical landscape suggest that refined global land cover and climate data sets may contribute to an enhanced MOD17 product at a variety of spatial scales.

Plant diversity increases with the strength of negative density dependence at the global scale.

Theory predicts that higher biodiversity in the tropics is maintained by specialized interactions among plants and their natural enemies that result in conspecific negative density dependence (CNDD). By using more than 3000 species and nearly 2.4 million trees across 24 forest plots worldwide, we show that global patterns in tree species diversity reflect not only stronger CNDD at tropical versus temperate latitudes but also a latitudinal shift in the relationship between CNDD and species abundance. CNDD was stronger for rare species at tropical versus temperate latitudes, potentially causing the persistence of greater numbers of rare species in the tropics. Our study reveals fundamental differences in the nature of local-scale biotic interactions that contribute to the maintenance of species diversity across temperate and tropical communities.

Rapid forest carbon assessments of oceanic islands: a case study of the Hawaiian archipelago.

BACKGROUND: Spatially explicit forest carbon (C) monitoring aids conservation and climate change mitigation efforts, yet few approaches have been developed specifically for the highly heterogeneous landscapes of oceanic island chains that continue to undergo rapid and extensive forest C change. We developed an approach for rapid mapping of aboveground C density (ACD; units = Mg or metric tons C ha-1) on islands at a spatial resolution of 30 m (0.09 ha) using a combination of cost-effective airborne LiDAR data and full-coverage satellite data. We used the approach to map forest ACD across the main Hawaiian Islands, comparing C stocks within and among islands, in protected and unprotected areas, and among forests dominated by native and invasive species. RESULTS: Total forest aboveground C stock of the Hawaiian Islands was 36 Tg, and ACD distributions were extremely heterogeneous both within and across islands. Remotely sensed ACD was validated against U.S. Forest Service FIA plot inventory data (R2 = 0.67; RMSE = 30.4 Mg C ha-1). Geospatial analyses indicated the critical importance of forest type and canopy cover as predictors of mapped ACD patterns. Protection status was a strong determinant of forest C stock and density, but we found complex environmentally mediated responses of forest ACD to alien plant invasion. CONCLUSIONS: A combination of one-time airborne LiDAR data acquisition and satellite monitoring provides effective forest C mapping in the highly heterogeneous landscapes of the Hawaiian Islands. Our statistical approach yielded key insights into the drivers of ACD variation, and also makes possible future assessments of C storage change, derived on a repeat basis from free satellite data, without the need for additional LiDAR data. Changes in C stocks and densities of oceanic islands can thus be continually assessed in the face of rapid environmental changes such as biological invasions, drought, fire and land use. Such forest monitoring information can be used to promote sustainable forest use and conservation on islands in the future.

CTFS-ForestGEO: a worldwide network monitoring forests in an era of global change.

Global change is impacting forests worldwide, threatening biodiversity and ecosystem services including climate regulation. Understanding how forests respond is critical to forest conservation and climate protection. This review describes an international network of 59 long-term forest dynamics research sites (CTFS-ForestGEO) useful for characterizing forest responses to global change. Within very large plots (median size 25 ha), all stems ≥ 1 cm diameter are identified to species, mapped, and regularly recensused according to standardized protocols. CTFS-ForestGEO spans 25 °S-61 °N latitude, is generally representative of the range of bioclimatic, edaphic, and topographic conditions experienced by forests worldwide, and is the only forest monitoring network that applies a standardized protocol to each of the world's major forest biomes. Supplementary standardized measurements at subsets of the sites provide additional information on plants, animals, and ecosystem and environmental variables. CTFS-ForestGEO sites are experiencing multifaceted anthropogenic global change pressures including warming (average 0.61 °C), changes in precipitation (up to ± 30% change), atmospheric deposition of nitrogen and sulfur compounds (up to 3.8 g N m(-2) yr(-1) and 3.1 g S m(-2) yr(-1)), and forest fragmentation in the surrounding landscape (up to 88% reduced tree cover within 5 km). The broad suite of measurements made at CTFS-ForestGEO sites makes it possible to investigate the complex ways in which global change is impacting forest dynamics. Ongoing research across the CTFS-ForestGEO network is yielding insights into how and why the forests are changing, and continued monitoring will provide vital contributions to understanding worldwide forest diversity and dynamics in an era of global change.