Trait-based filtering mediates the effects of realistic biodiversity losses on ecosystem functioning.

Biodiversity losses are a major driver of global changes in ecosystem functioning. While most studies of the relationship between biodiversity and ecosystem functioning have examined randomized species losses, trait-based filtering associated with species-specific vulnerability to drivers of diversity loss can strongly influence how ecosystem functioning responds to declining biodiversity. Moreover, the responses of ecosystem functioning to diversity loss may be mediated by environmental variability interacting with the suite of traits remaining in depauperate communities. We do not yet understand how communities resulting from realistic diversity losses (filtered by response traits) influence ecosystem functioning (via effect traits of the remaining community), especially under variable environmental conditions. Here, we directly test how realistic and randomized plant diversity losses influence productivity and invasion resistance across multiple years in a California grassland. Compared with communities based on randomized diversity losses, communities resulting from realistic (drought-driven) species losses had higher invasion resistance under climatic conditions that matched the trait-based filtering they experienced. However, productivity declined more with realistic than with randomized species losses across all years, regardless of climatic conditions. Functional response traits aligned with effect traits for productivity but not for invasion resistance. Our findings illustrate that the effects of biodiversity losses depend not only on the identities of lost species but also on how the traits of remaining species interact with varying environmental conditions. Understanding the consequences of biodiversity change requires studies that evaluate trait-mediated effects of species losses and incorporate the increasingly variable climatic conditions that future communities are expected to experience.

Will there be water? Climate change, housing needs, and future water demand in California.

Climate change in California is expected to alter future water availability, impacting water supplies needed to support future housing growth and agriculture demand. In groundwater-dependent regions like California's Central Coast, new land-use related water demand and decreasing recharge is already stressing depleted groundwater basins. We developed a spatially explicit state-and-transition simulation model that integrates climate, land-use change, water demand, and groundwater gain-loss to examine the impact of future climate and land use change on groundwater balance and water demand in five counties along the Central Coast from 2010 to 2060. The model incorporated downscaled groundwater recharge projections based on a Warm/Wet and a Hot/Dry climate future from a spatially explicit hydrological process-based model. Two urbanization projections from a parcel-based, regional urban growth model representing 1) recent historical and 2) state-mandated housing growth projections were used as alternative spatial targets for future urban growth. Agricultural projections were based on recent historical trends from remote sensing data. Annual projected changes in groundwater balance were calculated as the difference between land-use related water demand, based on historical estimates, and climate-driven recharge plus agriculture return flows. Results indicate that future changes in climate-driven groundwater recharge, coupled with cumulative increases in agricultural water demand, result in overall declines in future groundwater balance, with a Hot/Dry future resulting in cumulative groundwater decline in all but Santa Cruz County. Cumulative declines by 2060 are especially prominent in San Luis Obispo (-2.9 to -5.1 Bm3) and Monterey counties (-6.5 to -8.7 Bm3), despite limited changes in agricultural water demand over the model period. These two counties show declining groundwater reserves in a Warm/Wet future as well, while San Benito and Santa Barbara County barely reach equilibrium. These results suggest future groundwater supplies may not be able to keep pace with regional demand and declining climate-driven recharge, resulting in a potential reduction in water security in the region. However, our county-scale projections showed new housing and associated water demand does not conflict with California's groundwater sustainability goals. Rather, future climate coupled with increasing agricultural groundwater demand may reduce water security in some counties, potentially limiting available groundwater supplies for new housing.

Flowering phenology shifts in response to biodiversity loss.

Observational studies and experimental evidence agree that rising global temperatures have altered plant phenology-the timing of life events, such as flowering, germination, and leaf-out. Other large-scale global environmental changes, such as nitrogen deposition and altered precipitation regimes, have also been linked to changes in flowering times. Despite our increased understanding of how abiotic factors influence plant phenology, we know very little about how biotic interactions can affect flowering times, a significant knowledge gap given ongoing human-caused alteration of biodiversity and plant community structure at the global scale. We experimentally manipulated plant diversity in a California serpentine grassland and found that many plant species flowered earlier in response to reductions in diversity, with peak flowering date advancing an average of 0.6 days per species lost. These changes in phenology were mediated by the effects of plant diversity on soil surface temperature, available soil N, and soil moisture. Peak flowering dates were also more dispersed among species in high-diversity plots than expected based on monocultures. Our findings illustrate that shifts in plant species composition and diversity can alter the timing and distribution of flowering events, and that these changes to phenology are similar in magnitude to effects induced by climate change. Declining diversity could thus contribute to or exacerbate phenological changes attributed to rising global temperatures.

Effects of 21st-century climate, land use, and disturbances on ecosystem carbon balance in California.

Terrestrial ecosystems are an important sink for atmospheric carbon dioxide (CO2 ), sequestering ~30% of annual anthropogenic emissions and slowing the rise of atmospheric CO2 . However, the future direction and magnitude of the land sink is highly uncertain. We examined how historical and projected changes in climate, land use, and ecosystem disturbances affect the carbon balance of terrestrial ecosystems in California over the period 2001-2100. We modeled 32 unique scenarios, spanning 4 land use and 2 radiative forcing scenarios as simulated by four global climate models. Between 2001 and 2015, carbon storage in California's terrestrial ecosystems declined by -188.4 Tg C, with a mean annual flux ranging from a source of -89.8 Tg C/year to a sink of 60.1 Tg C/year. The large variability in the magnitude of the state's carbon source/sink was primarily attributable to interannual variability in weather and climate, which affected the rate of carbon uptake in vegetation and the rate of ecosystem respiration. Under nearly all future scenarios, carbon storage in terrestrial ecosystems was projected to decline, with an average loss of -9.4% (-432.3 Tg C) by the year 2100 from current stocks. However, uncertainty in the magnitude of carbon loss was high, with individual scenario projections ranging from -916.2 to 121.2 Tg C and was largely driven by differences in future climate conditions projected by climate models. Moving from a high to a low radiative forcing scenario reduced net ecosystem carbon loss by 21% and when combined with reductions in land-use change (i.e., moving from a high to a low land-use scenario), net carbon losses were reduced by 55% on average. However, reconciling large uncertainties associated with the effect of increasing atmospheric CO2 is needed to better constrain models used to establish baseline conditions from which ecosystem-based climate mitigation strategies can be evaluated.

Realistic diversity loss and variation in soil depth independently affect community-level plant nitrogen use.

Numerous experiments have demonstrated that diverse plant communities use nitrogen (N) more completely and efficiently, with implications for how species conservation efforts might influence N cycling and retention in terrestrial ecosystems. However, most such experiments have randomly manipulated species richness and minimized environmental heterogeneity, two design aspects that may reduce applicability to real ecosystems. Here we present results from an experiment directly comparing how realistic and randomized plant species losses affect plant N use across a gradient of soil depth in a native-dominated serpentine grassland in California. We found that the strength of the species richness effect on plant N use did not increase with soil depth in either the realistic or randomized species loss scenarios, indicating that the increased vertical heterogeneity conferred by deeper soils did not lead to greater complementarity among species in this ecosystem. Realistic species losses significantly reduced plant N uptake and altered N-use efficiency, while randomized species losses had no effect on plant N use. Increasing soil depth positively affected plant N uptake in both loss order scenarios but had a weaker effect on plant N use than did realistic species losses. Our results illustrate that realistic species losses can have functional consequences that differ distinctly from randomized losses, and that species diversity effects can be independent of and outweigh those of environmental heterogeneity on ecosystem functioning. Our findings also support the value of conservation efforts aimed at maintaining biodiversity to help buffer ecosystems against increasing anthropogenic N loading.

Ecosystem carbon storage does not vary with mean annual temperature in Hawaiian tropical montane wet forests.

Theory and experiment agree that climate warming will increase carbon fluxes between terrestrial ecosystems and the atmosphere. The effect of this increased exchange on terrestrial carbon storage is less predictable, with important implications for potential feedbacks to the climate system. We quantified how increased mean annual temperature (MAT) affects ecosystem carbon storage in above- and belowground live biomass and detritus across a well-constrained 5.2 °C MAT gradient in tropical montane wet forests on the Island of Hawaii. This gradient does not systematically vary in biotic or abiotic factors other than MAT (i.e. dominant vegetation, substrate type and age, soil water balance, and disturbance history), allowing us to isolate the impact of MAT on ecosystem carbon storage. Live biomass carbon did not vary predictably as a function of MAT, while detrital carbon declined by ~14 Mg of carbon ha(-1) for each 1 °C rise in temperature - a trend driven entirely by coarse woody debris and litter. The largest detrital pool, soil organic carbon, was the most stable with MAT and averaged 48% of total ecosystem carbon across the MAT gradient. Total ecosystem carbon did not vary significantly with MAT, and the distribution of ecosystem carbon between live biomass and detritus remained relatively constant across the MAT gradient at ~44% and ~56%, respectively. These findings suggest that in the absence of alterations to precipitation or disturbance regimes, the size and distribution of carbon pools in tropical montane wet forests will be less sensitive to rising MAT than predicted by ecosystem models. This article also provides needed detail on how individual carbon pools and ecosystem-level carbon storage will respond to future warming.

Phosphorus and soil development: does the Walker and Syers model apply to semiarid ecosystems?

The Walker and Syers model of phosphorus (P) transformations during pedogenesis is widely accepted for the development of humid ecosystems, but long-term P dynamics of more arid ecosystems remain poorly understood. We tested the Walker and Syers model in semiarid piñon-juniper woodlands by measuring soil P fractions under tree canopies and in intercanopy spaces along a well-constrained, approximately 3000 ka (1 ka = 1000 years) volcanic substrate age gradient in northern Arizona, USA. The various pools of soil P behaved largely as predicted; total soil P and primary mineral P declined consistently with substrate age, labile inorganic P increased early in soil development and then declined at later stages, and organic phosphorus increased consistently across the chronosequence. Within each site, soils under tree canopies tended to have higher concentrations of labile and intermediately available P fractions compared to intercanopy soils. However, the degree of spatial heterogeneity conferred by tree islands was moderated by the stage of soil development. In contrast, tree islands had no influence on within-site distribution of more recalcitrant soil P pools, which appear to be controlled solely by the stage of pedogenesis. Coincident with declines in total P, primary mineral P, and labile inorganic P, we found that phosphatase enzyme activity increased with substrate age; a result consistent with greater ecosystem-level P demand on older, more highly weathered substrates. Our results suggest that, compared to humid climates, reduced inputs of water, energy, and acidity to semiarid ecosystems slow the rate of change in P fractions during pedogenesis, but the overall pattern remains consistent with the Walker and Syers model. Furthermore, our data imply that pedogenic change may be an important factor controlling the spatial distribution of labile P pools in semiarid ecosystems. Taken together, these data should both broaden and unify terrestrial ecosystem development theory.

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.

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.].

Soil responses to management, increased precipitation, and added nitrogen in ponderosa pine forests.

Forest management, climatic change, and atmospheric N deposition can affect soil biogeochemistry, but their combined effects are not well understood. We examined the effects of water and N amendments and forest thinning and burning on soil N pools and fluxes in ponderosa pine forests near Flagstaff, Arizona (USA). Using a 15N-depleted fertilizer, we also documented the distribution of added N into soil N pools. Because thinning and burning can increase soil water content and N availability, we hypothesized that these changes would alleviate water and N limitation of soil processes, causing smaller responses to added N and water in the restored stand. We found little support for this hypothesis. Responses of fine root biomass, potential net N mineralization, and the soil microbial N to water and N amendments were mostly unaffected by stand management. Most of the soil processes we examined were limited by N and water, and the increased N and soil water availability caused by forest restoration was insufficient to alleviate these limitations. For example, N addition caused a larger increase in potential net nitrification in the restored stand, and at a given level of soil N availability, N addition had a larger effect on soil microbial N in the restored stand. Possibly, forest restoration increased the availability of some other limiting resource, amplifying responses to added N and water. Tracer N recoveries in roots and in the forest floor were lower in the restored stand. Natural abundance delta15N of labile soil N pools were higher in the restored stand, consistent with a more open N cycle. We conclude that thinning and burning open up the N cycle, at least in the short-term, and that these changes are amplified by enhanced precipitation and N additions. Our results suggest that thinning and burning in ponderosa pine forests will not increase their resistance to changes in soil N dynamics resulting from increased atmospheric N deposition or increased precipitation due to climatic change. Restoration plans should consider the potential impact on long-term forest productivity of greater N losses from a more open N cycle, especially during the period immediately after thinning and burning.

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.

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.

Leaf litter decomposition rates increase with rising mean annual temperature in Hawaiian tropical montane wet forests.

Decomposing litter in forest ecosystems supplies nutrients to plants, carbon to heterotrophic soil microorganisms and is a large source of CO2 to the atmosphere. Despite its essential role in carbon and nutrient cycling, the temperature sensitivity of leaf litter decay in tropical forest ecosystems remains poorly resolved, especially in tropical montane wet forests where the warming trend may be amplified compared to tropical wet forests at lower elevations. We quantified leaf litter decomposition rates along a highly constrained 5.2 °C mean annual temperature (MAT) gradient in tropical montane wet forests on the Island of Hawaii. Dominant vegetation, substrate type and age, soil moisture, and disturbance history are all nearly constant across this gradient, allowing us to isolate the effect of rising MAT on leaf litter decomposition and nutrient release. Leaf litter decomposition rates were a positive linear function of MAT, causing the residence time of leaf litter on the forest floor to decline by ∼31 days for each 1 °C increase in MAT. Our estimate of the Q 10 temperature coefficient for leaf litter decomposition was 2.17, within the commonly reported range for heterotrophic organic matter decomposition (1.5-2.5) across a broad range of ecosystems. The percentage of leaf litter nitrogen (N) remaining after six months declined linearly with increasing MAT from ∼88% of initial N at the coolest site to ∼74% at the warmest site. The lack of net N immobilization during all three litter collection periods at all MAT plots indicates that N was not limiting to leaf litter decomposition, regardless of temperature. These results suggest that leaf litter decay in tropical montane wet forests may be more sensitive to rising MAT than in tropical lowland wet forests, and that increased rates of N release from decomposing litter could delay or prevent progressive N limitation to net primary productivity with climate warming.

Does dissolved organic carbon regulate biological methane oxidation in semiarid soils?

In humid ecosystems, the rate of methane (CH4 ) oxidation by soil-dwelling methane-oxidizing bacteria (MOB) is controlled by soil texture and soil water holding capacity, both of which limit the diffusion of atmospheric CH4 into the soil. However, it remains unclear whether these same mechanisms control CH4 oxidation in more arid soils. This study was designed to measure the proximate controls of potential CH4 oxidation in semiarid soils during different seasons. Using a unique and well-constrained 3-million-year-old semiarid substrate age gradient, we were able to hold state factors constant while exploring the relationship between seasonal potential CH4 oxidation rates and soil texture, soil water holding capacity, and dissolved organic carbon (DOC). We measured unexpectedly higher rates of potential CH4 oxidation in the wet season than the dry season. Although other studies have attributed low CH4 oxidation rates in dry soils to desiccation of MOB, we present several lines of evidence that this may be inaccurate. We found that soil DOC concentration explained CH4 oxidation rates better than soil physical factors that regulate the diffusion of CH4 from the atmosphere into the soil. We show evidence that MOB facultatively incorporated isotopically labeled glucose into their cells, and MOB utilized glucose in a pattern among our study sites that was similar to wet-season CH4 oxidation rates. This evidence suggests that DOC, which is utilized by MOB in other environments with varying effects on CH4 oxidation rates, may be an important regulator of CH4 oxidation rates in semiarid soils. Our collective understanding of the facultative use of DOC by MOB is still in its infancy, but our results suggest it may be an important factor controlling CH4 oxidation in soils from dry ecosystems.