N and P constrain C in ecosystems under climate change: Role of nutrient redistribution, accumulation, and stoichiometry.

We use the Multiple Element Limitation (MEL) model to examine responses of 12 ecosystems to elevated carbon dioxide (CO2 ), warming, and 20% decreases or increases in precipitation. Ecosystems respond synergistically to elevated CO2 , warming, and decreased precipitation combined because higher water-use efficiency with elevated CO2 and higher fertility with warming compensate for responses to drought. Response to elevated CO2 , warming, and increased precipitation combined is additive. We analyze changes in ecosystem carbon (C) based on four nitrogen (N) and four phosphorus (P) attribution factors: (1) changes in total ecosystem N and P, (2) changes in N and P distribution between vegetation and soil, (3) changes in vegetation C:N and C:P ratios, and (4) changes in soil C:N and C:P ratios. In the combined CO2 and climate change simulations, all ecosystems gain C. The contributions of these four attribution factors to changes in ecosystem C storage varies among ecosystems because of differences in the initial distributions of N and P between vegetation and soil and the openness of the ecosystem N and P cycles. The net transfer of N and P from soil to vegetation dominates the C response of forests. For tundra and grasslands, the C gain is also associated with increased soil C:N and C:P. In ecosystems with symbiotic N fixation, C gains resulted from N accumulation. Because of differences in N versus P cycle openness and the distribution of organic matter between vegetation and soil, changes in the N and P attribution factors do not always parallel one another. Differences among ecosystems in C-nutrient interactions and the amount of woody biomass interact to shape ecosystem C sequestration under simulated global change. We suggest that future studies quantify the openness of the N and P cycles and changes in the distribution of C, N, and P among ecosystem components, which currently limit understanding of nutrient effects on C sequestration and responses to elevated CO2 and climate change.

Light fuels while nitrogen suppresses symbiotic nitrogen fixation hotspots in neotropical canopy gap seedlings.

Mature neotropical lowland forests have relatively lower symbiotic nitrogen fixation (SNF) rates compared with secondary forests. Canopy gap formation may create transient SNF hotspots in mature forests that increase overall SNF rates in these ecosystems, as canopy gaps are pervasive across the landscape and increasing in frequency. However, what environmental conditions are driving SNF upregulation in canopy gaps is unknown. In a field experiment to test these potential environmental controls on SNF, we grew 540 neotropical nitrogen-fixing legume seedlings (Pentaclethra macroloba, Zygia longifolia, and Stryphnodendron microstachyum) under manipulated light and soil nitrogen availability in canopy gaps and intact forests at La Selva Biological Station, Costa Rica. Seedling biomass, nodule biomass, and SNF (g N seedling-1 h-1 ) were 4-, 17- and 42-fold higher, respectively, in canopy gaps than in the intact forest. Nitrogen additions decreased SNF, but light had a stronger positive effect. Upregulation of SNF in canopy gaps was driven by increased plant growth and not a disproportionate increased SNF allocation. These data provide evidence that canopy gap SNF hotspots are driven, in part, by light availability, demonstrating a potential driver of SNF spatial heterogeneity. This further suggests that canopy gap dynamics are important for understanding the biogeochemistry of neotropical forests.

A New Framework for Evaluating Estimates of Symbiotic Nitrogen Fixation in Forests.

Symbiotic nitrogen fixation (SNF) makes atmospheric nitrogen biologically available and regulates carbon storage in many terrestrial ecosystems. Despite its global importance, estimates of SNF rates are highly uncertain, particularly in tropical forests where rates are assumed to be high. Here we provide a framework for evaluating the uncertainty of sample-based SNF estimates and discuss its implications for quantifying SNF and thus understanding of forest function. We apply this framework to field data sets from six lowland tropical rainforests (mature and secondary) in Brazil and Costa Rica. We use this data set to estimate parameters influencing SNF estimation error, notably the root nodule abundance and variation in SNF rates among soil cores containing root nodules. We then use simulations to gauge the relationship between sampling effort and SNF estimation accuracy for a combination of parameters. Field data illuminate a highly right-skewed lognormal distribution of SNF rates among soil cores containing root nodules that were rare and spanned five orders of magnitude. Consequently, simulations demonstrated that sample sizes of hundreds to even thousands of soil cores are needed to obtain estimates of SNF that are within, for example, a factor of 2 of the actual rate with 75% probability. This represents sample sizes that are larger than most studies to date. As a result of this previously undescribed uncertainty, we suggest that current estimates of SNF in tropical forests are not sufficiently constrained to elucidate forest stand-level controls of SNF, which hinders our understanding of the impact of SNF on tropical forest ecosystem processes.

Remotely sensed canopy nitrogen correlates with nitrous oxide emissions in a lowland tropical rainforest.

Tropical forests exhibit significant heterogeneity in plant functional and chemical traits that may contribute to spatial patterns of key soil biogeochemical processes, such as carbon storage and greenhouse gas emissions. Although tropical forests are the largest ecosystem source of nitrous oxide (N2 O), drivers of spatial patterns within forests are poorly resolved. Here, we show that local variation in canopy foliar N, mapped by remote-sensing image spectroscopy, correlates with patterns of soil N2 O emission from a lowland tropical rainforest. We identified ten 0.25 ha plots (assemblages of 40-70 individual trees) in which average remotely-sensed canopy N fell above or below the regional mean. The plots were located on a single minimally-dissected terrace (<1 km2 ) where soil type, vegetation structure and climatic conditions were relatively constant. We measured N2 O fluxes monthly for 1 yr and found that high canopy N species assemblages had on average three-fold higher total mean N2 O fluxes than nearby lower canopy N areas. These differences are consistent with strong differences in litter stoichiometry, nitrification rates and soil nitrate concentrations. Canopy N status was also associated with microbial community characteristics: lower canopy N plots had two-fold greater soil fungal to bacterial ratios and a significantly lower abundance of ammonia-oxidizing archaea, although genes associated with denitrification (nirS, nirK, nosZ) showed no relationship with N2 O flux. Overall, landscape emissions from this ecosystem are at the lowest end of the spectrum reported for tropical forests, consist with multiple metrics indicating that these highly productive forests retain N tightly and have low plant-available losses. These data point to connections between canopy and soil processes that have largely been overlooked as a driver of denitrification. Defining relationships between remotely-sensed plant traits and soil processes offers the chance to map these processes at large scales, potentially increasing our ability to predict N2 O emissions in heterogeneous landscapes.

Chronosequence predictions are robust in a Neotropical secondary forest, but plots miss the mark.

Tropical secondary forests (TSF) are a global carbon sink of 1.6 Pg C/year. However, TSF carbon uptake is estimated using chronosequence studies that assume differently aged forests can be used to predict change in aboveground biomass density (AGBD) over time. We tested this assumption using two airborne lidar datasets separated by 11.5 years over a Neotropical landscape. Using data from 1998, we predicted canopy height and AGBD within 1.1 and 10.3% of observations in 2009, with higher accuracy for forest height than AGBD and for older TSFs in comparison to younger ones. This result indicates that the space-for-time assumption is robust at the landscape-scale. However, since lidar measurements of secondary tropical forest are rare, we used the 1998 lidar dataset to test how well plot-based studies quantify the mean TSF height and biomass in a landscape. We found that the sample area required to produce estimates of height or AGBD close to the landscape mean is larger than the typical area sampled in secondary forest chronosequence studies. For example, estimating AGBD within 10% of the landscape mean requires more than thirty 0.1 ha plots per age class, and more total area for larger plots. We conclude that under-sampling in ground-based studies may introduce error into estimations of the TSF carbon sink, and that this error can be reduced by more extensive use of lidar measurements.

Nutrient acquisition, soil phosphorus partitioning and competition among trees in a lowland tropical rain forest.

We hypothesized that dinitrogen (N2 )- and non-N2 -fixing tropical trees would have distinct phosphorus (P) acquisition strategies allowing them to exploit different P sources, reducing competition. We measured root phosphatase activity and arbuscular mycorrhizal (AM) colonization among two N2 - and two non-N2 -fixing seedlings, and grew them alone and in competition with different inorganic and organic P forms to assess potential P partitioning. We found an inverse relationship between root phosphatase activity and AM colonization in field-collected seedlings, indicative of a trade-off in P acquisition strategies. This correlated with the predominantly exploited P sources in the seedling experiment: the N2 fixer with high N2 fixation and root phosphatase activity grew best on organic P, whereas the poor N2 fixer and the two non-N2 fixers with high AM colonization grew best on inorganic P. When grown in competition, however, AM colonization, root phosphatase activity and N2 fixation increased in the N2 fixers, allowing them to outcompete the non-N2 fixers regardless of P source. Our results indicate that some tropical trees have the capacity to partition soil P, but this does not eliminate interspecific competition. Rather, enhanced P and N acquisition strategies may increase the competitive ability of N2 fixers relative to non-N2 fixers.

Nutrient limitation in tropical secondary forests following different management practices.

Secondary forests now make up more than one-half of all tropical forests, and constraints on their biomass accumulation will influence the strength of the terrestrial carbon (C) sink in the coming decades. However the variance in secondary tropical forest biomass for a given stand age and climate is high and our understanding of why is limited. We constructed a model of terrestrial C, nitrogen (N), and phosphorus (P) cycling to examine the influence of disturbance and management practices on nutrient limitation and biomass recovery in secondary tropical forests. The model predicted that N limited the rate of forest recovery in the first few decades following harvest, but that this limitation switched to P approximately 30-40 yr after abandonment, consistent with field data on N and P cycling from secondary tropical forest chronosequences. Simulated biomass recovery agreed well with field data of biomass accumulation following harvest (R2  = 0.80). Model results showed that if all biomass remained on site following a severe disturbance such as blowdown, regrowth approached pre-disturbance biomass in 80-90 yr, and recovery was faster following smaller disturbances such as selective logging. Field data from regrowth on abandoned pastures were consistent with simulated losses of nutrients in soil organic matter, particularly P. Following any forest disturbance that involved the removal of nutrients (i.e., except blowdown), forest regrowth produced reduced biomass relative to the initial state as a result of nutrient loss through harvest, leaching and/or sequestration by secondary minerals. Differences in nutrient availability accounted for 49-94% of the variance in secondary forest biomass C at a given stand age. Management lessons from this study are the importance of strategies that help retain nutrients on site, recognizing the role of coarse woody debris in immobilization and subsequent release of nutrients, and the potential for nutrient additions to enhance biomass growth and recovery in secondary tropical forests.

Solute and sediment export from Amazon forest and soybean headwater streams.

Intensive cropland agriculture commonly increases streamwater solute concentrations and export from small watersheds. In recent decades, the lowland tropics have become the world's largest and most important region of cropland expansion. Although the effects of intensive cropland agriculture on streamwater chemistry and watershed export have been widely studied in temperate regions, their effects in tropical regions are poorly understood. We sampled seven headwater streams draining watersheds in forest (n = 3) or soybeans (n = 4) to examine the effects of soybean cropping on stream solute concentrations and watershed export in a region of rapid soybean expansion in the Brazilian state of Mato Grosso. We measured stream flows and concentrations of NO3- , PO43- , SO42- , Cl- , NH4+ , Ca2+ , Mg2+ , Na+ , K+ , Al3+ , Fe3+ , and dissolved organic carbon (DOC) biweekly to monthly to determine solute export. We also measured stormflows and stormflow solute concentrations in a subset of watersheds (two forest, two soybean) during two/three storms, and solutes and δ18 O in groundwater, rainwater, and throughfall to characterize watershed flowpaths. Concentrations of all solutes except K+ varied seasonally in streamwater, but only Fe3+ concentrations differed between land uses. The highest streamwater and rainwater solute concentrations occurred during the peak season of wildfires in Mato Grosso, suggesting that regional changes in atmospheric composition and deposition influence seasonal stream solute concentrations. Despite no concentration differences between forest and soybean land uses, annual export of NH4+ , PO43- , Ca2+ , Fe3+ , Na+ , SO42- , DOC, and TSS were significantly higher from soybean than forest watersheds (5.6-fold mean increase). This increase largely reflected a 4.3-fold increase in water export from soybean watersheds. Despite this increase, total solute export per unit watershed area (i.e., yield) remained low for all watersheds (<1 kg NO3- N·ha-1 ·yr-1 , <2.1 kg NH4+ -N·ha-1 ·yr-1 , <0.2 kg PO43- -P·ha-1 ·yr-1 , <1.5 kg Ca2+ ·ha-1 ·yr-1 ). Responses of both streamflows and solute concentrations to crop agriculture appear to be controlled by high soil hydraulic conductivity, groundwater-dominated hydrologic flowpaths on deep soils, and the absence of nitrogen fertilization. To date, these factors have buffered streams from the large increases in solute concentrations that often accompany intensive croplands in other locations.

Ecological and evolutionary variation in community nitrogen use traits during tropical dry forest secondary succession.

We assessed the role of ecological and evolutionary processes in driving variation in leaf and litter traits related to nitrogen (N) use among tropical dry forest trees in old-growth and secondary stands in western Mexico. Our expectation was that legumes (Fabaceae), a dominant component of the regional flora, would have consistently high leaf N and therefore structure phylogenetic variation in N-related traits. We also expected ecological selection during succession for differences in nitrogen use strategies, and corresponding shifts in legume abundance. We used phylogenetic analyses to test for trait conservatism in foliar and litter N, C:N, and N resorption. We also evaluated differences in N-related traits between old-growth and secondary forests. We found a weak phylogenetic signal for all traits, partly explained by wide variation within legumes. Across taxa we observed a positive relationship between leaf and litter N, but no shift in resorption strategies along the successional gradient. Despite species turnover, N-resorption, and N-related traits showed little change across succession, suggesting that, at least for these traits, secondary forests rapidly recover ecosystem function. Collectively, our results also suggest that legumes should not be considered a single functional group from a biogeochemical perspective.

Environmental controls on canopy foliar nitrogen distributions in a Neotropical lowland forest.

Distributions of foliar nutrients across forest canopies can give insight into their plant functional diversity and improve our understanding of biogeochemical cycling. We used airborne remote sensing and partial least squares regression to quantify canopy foliar nitrogen (foliar N) across ~164 km2 of wet lowland tropical forest in the Osa Peninsula, Costa Rica. We determined the relative influence of climate and topography on the observed patterns of foliar N using a gradient boosting model technique. At a local scale, where climate and substrate were constant, we explored the influence of slope position on foliar N by quantifying foliar N on remnant terraces, their adjacent slopes, and knife-edged ridges. In addition, we climbed and sampled 540 trees and analyzed foliar N in order to quantify the role of species identity (phylogeny) and environmental factors in predicting foliar N. Observed foliar N heterogeneity reflected environmental factors working at multiple spatial scales. Across the larger landscape, elevation and precipitation had the highest relative influence on predicting foliar N (30% and 24%), followed by soils (15%), site exposure (9%), compound topographic index (8%), substrate (6%), and landscape dissection (6%). Phylogeny explained ~75% of the variation in the field collected foliar N data, suggesting that phylogeny largely underpins the response to the environmental factors. Taken together, these data suggest that a large fraction of the variance in foliar N across the landscape is proximately driven by species composition, though ultimately this is likely a response to abiotic factors such as climate and topography. Future work should focus on the mechanisms and feedbacks involved, and how shifts in climate may translate to changes in forest function.

Structure and composition of altered riparian forests in an agricultural Amazonian landscape.

Deforestation and fragmentation influence the microclimate, vegetation structure, and composition of remaining patches of tropical forest. In the southern Amazon, at the frontier of cropland expansion, forests are converted and fragmented in a pattern that leaves standing riparian forests whose dimensions are mandated by the Brazilian National Forest Code. These altered riparian forests share many characteristics of well-studied upland forest fragments, but differ because they remain connected to larger areas of forest downstream, and because they may experience wetter soil conditions because reduction of forest cover in the surrounding watershed raises groundwater levels and increases stream runoff. We compared forest regeneration, structure, composition, and diversity in four areas of intact riparian forest and four areas each of narrow, medium, and wide altered riparian forests that have been surrounded by agriculture since the early 1980s. We found that seedling abundance was reduced by as much as 64% and sapling abundance was reduced by as much as 67% in altered compared to intact riparian forests. The most pronounced differences between altered and intact forest occurred near forest edges and within the narrowest sections of altered riparian forests. Woody plant species composition differed and diversity was reduced in altered forests compared to intact riparian forests. However, despite being fragmented for several decades, large woody plant biomass and carbon storage, the number of live or dead large woody plants, mortality rates, and the size distribution of woody plants did not differ significantly between altered and intact riparian forests. Thus, even in these relatively narrow forests with high edge: area ratios, we saw no evidence of the increases in mortality and declines in biomass that have been found in other tropical forest fragment studies. However, because of the changes in both species community and reduced regeneration, it is unclear how long this relative lack of change will be sustained. Additionally, Brazil recently passed a law in their National Forest Code allowing narrower riparian buffers than those studied here in restored areas, which could affect their long-term sustainability.

Assessing nutrient limitation in complex forested ecosystems: alternatives to large-scale fertilization experiments.

Quantifying nutrient limitation of primary productivity is a fundamental task of terrestrial ecosystem ecology, but in a high carbon dioxide environment it is even more critical that we understand potential nutrient constraints on plant growth. Ecologists often manipulate nutrients with fertilizer to assess nutrient limitation, yet for a variety of reasons, nutrient fertilization experiments are either impractical or incapable of resolving ecosystem responses to some global changes. The challenges of conducting large, in situ fertilization experiments are magnified in forests, especially the high-diversity forests common throughout the lowland tropics. A number of methods, including fertilization experiments, could be seen as tools in a toolbox that ecologists may use to attempt to assess nutrient limitation, but there has been no compilation or synthetic discussion of those methods in the literature. Here, we group these methods into one of three categories (indicators of soil nutrient supply, organismal indicators of nutrient limitation, and lab-based experiments and nutrient depletions), and discuss some of the strengths and limitations of each. Next, using a case study, we compare nutrient limitation assessed using these methods to results obtained using large-scale fertilizations across the Hawaiian Archipelago. We then explore the application of these methods in high-diversity tropical forests. In the end, we suggest that, although no single method is likely to predict nutrient limitation in all ecosystems and at all scales, by simultaneously utilizing a number of the methods we describe, investigators may begin to understand nutrient limitation in complex and diverse ecosystems such as tropical forests. In combination, these methods represent our best hope for understanding nutrient constraints on the global carbon cycle, especially in tropical forest ecosystems.

Relationships among net primary productivity, nutrients and climate in tropical rain forest: a pan-tropical analysis.

Tropical rain forests play a dominant role in global biosphere-atmosphere CO(2) exchange. Although climate and nutrient availability regulate net primary production (NPP) and decomposition in all terrestrial ecosystems, the nature and extent of such controls in tropical forests remain poorly resolved. We conducted a meta-analysis of carbon-nutrient-climate relationships in 113 sites across the tropical forest biome. Our analyses showed that mean annual temperature was the strongest predictor of aboveground NPP (ANPP) across all tropical forests, but this relationship was driven by distinct temperature differences between upland and lowland forests. Within lowland forests (< 1000 m), a regression tree analysis revealed that foliar and soil-based measurements of phosphorus (P) were the only variables that explained a significant proportion of the variation in ANPP, although the relationships were weak. However, foliar P, foliar nitrogen (N), litter decomposition rate (k), soil N and soil respiration were all directly related with total surface (0-10 cm) soil P concentrations. Our analysis provides some evidence that P availability regulates NPP and other ecosystem processes in lowland tropical forests, but more importantly, underscores the need for a series of large-scale nutrient manipulations - especially in lowland forests - to elucidate the most important nutrient interactions and controls.

A framework for predicting global silicate weathering and CO2 drawdown rates over geologic time-scales.

Global silicate weathering drives long-time-scale fluctuations in atmospheric CO(2). While tectonics, climate, and rock-type influence silicate weathering, it is unclear how these factors combine to drive global rates. Here, we explore whether local erosion rates, GCM-derived dust fluxes, temperature, and water balance can capture global variation in silicate weathering. Our spatially explicit approach predicts 1.9-4.6 x 10(13) mols of Si weathered globally per year, within a factor of 4-10 of estimates of global silicate fluxes derived from riverine measurements. Similarly, our watershed-based estimates are within a factor of 4-18 (mean of 5.3) of the silica fluxes measured in the world's ten largest rivers. Eighty percent of total global silicate weathering product traveling as dissolved load occurs within a narrow range (0.01-0.5 mm/year) of erosion rates. Assuming each mol of Mg or Ca reacts with 1 mol of CO(2), 1.5-3.3 x 10(8) tons/year of CO(2) is consumed by silicate weathering, consistent with previously published estimates. Approximately 50% of this drawdown occurs in the world's active mountain belts, emphasizing the importance of tectonic regulation of global climate over geologic timescales.

Tripartite legume-rhizobia-mycorrhizae relationship is influenced by light and soil nitrogen in Neotropical canopy gaps.

Plants and their soil microbial symbionts influence ecosystem productivity and nutrient cycling, but the controls on these symbioses remain poorly understood. This is particularly true for plants in the Fabaceae family (hereafter legumes), which can associate with both arbuscular mycorrhizal fungi (AMF) and nitrogen (N) -fixing bacteria. Here we report results of the first manipulated field experiment to explore the abiotic and biotic controls of this tripartite symbiosis in Neotropical canopy gaps (hereafter gaps). We grew three species of Neotropical N-fixing legume seedlings under different light (gap-full light, gap-shadecloth, and understory) and soil nitrogen (20 g N·m-2 ·yr-1 vs. 0 g N·m-2 ·yr-1 ) conditions across a lowland tropical forest at La Selva Biological Station, Costa Rica. We harvested the seedlings after 4 months of growth in the field and measured percent AMF root colonization (%AMF), nodule and seeding biomass, and seedling aboveground:belowground biomass ratios. Our expectation was that seedlings in gaps would grow larger and, as a result of higher light, invest more carbon in both AMF and N-fixing bacteria. Indeed, seedlings in gaps had higher total biomass, nodule biomass (a proxy for N-fixing bacteria investment) and rates of AMF root colonization, and the three were significantly positively correlated. However, we only found a significant positive effect of light availability on %AMF when seedlings were fertilized with N. Furthermore, when we statistically controlled for treatment, species, and site effects, we found %AMF and seedling biomass had a negative relationship. This was likely driven by the fact that seedlings invested relatively less in AMF as they increased in biomass (lower %AMF per gram of seedling). Taken together, these results challenge the long-held assumption that high light conditions universally increase carbon investment in AMF and demonstrate that this tripartite symbiosis is influenced by soil nutrient and light conditions.

Terrestrial phosphorus limitation: mechanisms, implications, and nitrogen-phosphorus interactions.

Nutrient limitation to primary productivity and other biological processes is widespread in terrestrial ecosystems, and nitrogen (N) and phosphorus (P) are the most common limiting elements, both individually and in combination. Mechanisms that drive P limitation, and their interactions with the N cycle, have received less attention than mechanisms causing N limitation. We identify and discuss six mechanisms that could drive P limitation in terrestrial ecosystems. The best known of these is depletion-driven limitation, in which accumulated P losses during long-term soil and ecosystem development contribute to what Walker and Syers termed a "terminal steady state" of profound P depletion and limitation. The other mechanisms are soil barriers that prevent access to P; transactional limitation, in which weathering of P-containing minerals does not keep pace with the supply of other resources; low-P parent materials; P sinks; and anthropogenic changes that increase the supply of other resources (often N) relative to P. We distinguish proximate nutrient limitation (which occurs where additions of a nutrient stimulate biological processes, especially productivity) from ultimate nutrient limitation (where additions of a nutrient can transform ecosystems). Of the mechanisms that drive P limitation, we suggest that depletion, soil barriers, and low-P parent material often cause ultimate limitation because they control the ecosystem mass balance of P. Similarly, demand-independent losses and constraints to N fixation can control the ecosystem-level mass balance of N and cause it to be an ultimate limiting nutrient.

Climate and soil-age constraints on nutrient uplift and retention by plants.

Plants and soils represent coevolving components of ecosystems, and while the effects of soils (e.g., nutrient availability) on plants have been extensively documented, the effect of plants on soils has received less attention. Furthermore there has been no systematic investigation of how plant effects vary across important ecological gradients in climate or soil age, which leaves a substantial gap in our understanding of how plant-soil systems develop. In this context, we analyzed changes in nutrient availability and elemental losses from the entire weathering zone at 35 sites arrayed across climatic and soil-age gradients on the island of Hawai'i. The sites are located on three basaltic lava flows (ages 10, 170, and 350 kyr) each of which crosses a precipitation gradient from approximately 500 to 2500 mm/yr. By comparing the loss of nutrient (potassium, phosphorus) and non-nutrient (e.g., sodium) rock-derived elements, we identify a climatic zone at intermediate rainfall where the retention of plant nutrients in the upper soil is most pronounced. We further show that there are several abiotic constraints on plant-driven retention of nutrients. At the dry sites (< or = 750 mm/yr on all three flows), plants slow the loss of nutrients, but the effect (as measured by the difference between K and Na losses) is small, perhaps because of low plant cover and productivity. At intermediate rainfall (750-1400 mm/yr) but negative water balance, plants substantially enrich both nutrient cations and P relative to Na in the surface horizons, an effect that remains strong even after 350 kyr of soil development. In contrast, at high rainfall (> or = 1500 mm/yr) and positive water balance, the effect of plants on nutrient distributions diminishes with soil age as leaching losses overwhelm the uplift and retention of nutrients by plants after 350 kyr of soil development. The effect of plants on soil nutrient distributions can also be mediated by the movement of iron (Fe), and substantial Fe losses at high rainfall on the older flows are highly correlated with P losses. Thus redox-driven redistribution of Fe may place a further abiotic constraint on nutrient retention by plants. In combination, these data indicate that the effects of soil aging on plant uplift and retention of nutrients differ markedly with precipitation, and we view this as a potentially fruitful area for future research.

Sources of nutrients to windward agricultural systems in pre-contact Hawai'i.

Prior to European contact in 1778, Hawaiians developed intensive irrigated pondfield agricultural systems in windward Kohala, Hawai'i. We evaluated three potential sources of nutrients to windward systems that could have sustained intensive agriculture: (1) in situ weathering of primary and secondary minerals in upland soils; (2) rejuvenation of the supply of rock-derived nutrients on eroded slopes and in alluvium; and (3) transport of rock-derived nutrients to crops via irrigation water. Our results show that most windward soils are infertile and suggest that weathering of minerals within upland soils was insufficient to sustain intensive agriculture without substantial cultural inputs. Erosion enhances weathering and so increases nutrient supply, with soils of the largest alluvial valleys (>200 m deep) retaining 37% of calcium from parent material (compared to 2% in upland sites). However, soils of smaller valleys that also supported pre-contact agricultural systems are substantially less enriched. Isotopic 87Sr/86Sr analyses of stream water demonstrate that at low to moderate stream flow over 90% of dissolved strontium derives from weathering of basalt rather than deposition of atmospheric sources; most other dissolved cations also derive from basalt weathering. We calculate that irrigation water could have supplied approximately 200 kg ha(-1) yr(-1) of calcium to pondfield systems, nearly 100 times more than was supplied by weathering in soils on stable geomorphic surfaces. In effect, irrigation waters brought nutrients from rocks to the windward crops.