Timothy E. Crews

Changes in Soil Phosphorus Fractions and Ecosystem Dynamics across a Long Chronosequence in Hawaii

We tested the Walker and Syers (1976) conceptual model of soil development and its ecological implications by analyzing changes in soil P, vegetation, and other ecosystem properties on a soil chronosequence with six sites ranging in age from 300 yr to 4.1 x 10 6 yr. Climate, dominant vegetation, slope, and parent material of all of the sites were similar. As fractions of total P, the various pools of soil phosphorus behaved very much as predicted by Walker and Syers. HCl-extractable P (presumably primary mineral phosphates) comprised 82% of total P at the 300-yr-old site, and then decreased to 1% at the 20,000-yr-old site. Organic phosphorus increased from the youngest site to a maximum at the 150000 yr site, and then declined to the 4.1 x 10 6 yr site. Occluded (residual) P increased steadily with soil age. In contrast to the Walker and Syers model, we found the highest total P at the 150000-yr-old site, rather than at the onset of soil development, and we found that the non-occluded, inorganic P fraction persisted through to the oldest chronosequence site. Total soil N and C increased substantially from early to middle soil development in parallel with organic P, and then declined through to the oldest site. Readily available soil P, NH 4 + , and NO 3 - were measured using anion and cation exchange resin bags. P availability increased and decreased unimodally across the chronosequence. NH 4 + and NO 3 - pools increased through early soil development, but did not systematically decline late in soil development. In situ decomposition rates of Metrosideros polymorpha litter were highest at two intermediate-aged sites with high soil fertility (20000 yr and 150000 yr), and lowest at the less-fertile beginning (300 yr) and endpoint (4.1 x 10 6 yr) of the chronosequence. M. polymorpha leaves collected from these same four sites, and decomposed in a common site, suggested that leaves from intermediate-aged sites were inherently more decomposable than those from the youngest and oldest sites. Both litter tissue quality and the soil environment appeared to influence rates of decomposition directly. The highest mean soil N 2 O emissions (809 μg.m -2 .d -1 ) were measured at the 20 000-yr-old site, which also had the highest soil nitrogen fertility status. Plant communities at all six chronosequence sites were dominated primarily by M. polymorpha, and to a lesser extent by several other genera of trees and shrubs. There were, however, differences in overall vegetation community composition among the sites. Using a detrended correspondence analysis (DECORANA), we found that a high proportion of species variance among the sites (eigenvalue = 0.71) can be explained by site age and thus soil developmental stage. Overall, long-term soil development across the chronosequence largely coincides with the conceptual model of Walker and Syers (1976). How P is distributed among different organic and inorganic fractions at a given stage of soil development provides a useful context for evaluating contemporary cycling of P and other nutrients, and for determining how changes in P availability might affect diverse ecosystem processes.

Nutrient Imbalances in Agricultural Development

Nutrient additions to intensive agricultural systems range from inadequate to excessive—and both extremes have substantial human and environmental costs. Nutrient cycles link agricultural systems to their societies and surroundings; inputs of nitrogen and phosphorus in particular are essential for high crop yields, but downstream and downwind losses of these same nutrients diminish environmental quality and human well-being. Agricultural nutrient balances differ substantially with economic development, from inputs that are inadequate to maintain soil fertility in parts of many developing countries, particularly those of sub-Saharan Africa, to excessive and environmentally damaging surpluses in many developed and rapidly growing economies. National and/or regional policies contribute to patterns of nutrient use and their environmental consequences in all of these situations (1). Solutions to the nutrient challenges that face global agriculture can be informed by analyses of trajectories of change within, as well as across, agricultural systems.

Can the Synchrony of Nitrogen Supply and Crop Demand be Improved in Legume and Fertilizer-based Agroecosystems? A Review

Asynchrony between nitrogen (N) supply and crop demand is the source of many environmental hazards associated with excess N in the biosphere. In this review, we explore some of the complexity of the synchrony issue in agroecosystems that obtain N via legume rotations or synthetic fertilizers. Studies that have simultaneously compared the fate of both sources of N suggest that in rainfed agricultures, crops recover more N from fertilizer, but a higher proportion of the legume N is retained in the soil and N losses tend not to differ greatly from either source. However, investigations from irrigated cropping systems indicate that legume N is generally less susceptible to loss processes than fertilizers. Such general conclusions need to be qualified by acknowledging that not all comparative studies have used ȁ8best management practices’ when applying the fertilizer or legume residues. When information-intensive management approaches are used, fertilizer-based systems can potentially out-perform the synchrony achieved by legume-based rotations. We suggest that the inclusion of perennials in cropping systems may hold the greatest promise for decreasing the risk of N losses in future farming systems.

The presence of nitrogen fixing legumes in terrestrial communities: Evolutionary vs ecological considerations

Nitrogen is often a limiting factor to net primary productivity (NPP) and other processes in terrestrial ecosystems. In most temperate freshwater ecosystems, when nitrogen becomes limiting to NPP, populations of N-fixing cyanobacteria experience a competitive advantage, and begin to grow and fix nitrogen until the next most limiting resource is encountered; typically phosphorus or light. Why is it that N-fixing plants do not generally function to overcome N limitation in terrestrial ecosystems in the same way that cyanobacteria function in aquatic ecosystems? To address this question in a particular ecosystem, one must first know whether the flora includes a potential set of nitrogen fixers. I suggest that the presence or absence of N-fixing plant symbioses is foremost an evolutionary consideration, determined to a large extent by constraints on the geographical radiation of woody members of the family Fabaceae. Ecological factors such as competition, nutrient deficiencies, grazing and fire are useful to explain the success of N-fixing plants only when considered against the geographical distribution of potential N-fixers.

Both nitrogen and phosphorus limit plant production on young Hawaiian lava flows

We applied fertilizers in a 23complete factorial design to determine the effects of nutrient amendments on plant growth in Hawaiian montane forests growing on two different volcanic substrates: ‘a‘ā and pāhoehoe lava. Both sites were about 140 years old and their overstories were nearly monospecific stands of Metrosideros polymorpha. Fertilizer applications included N, P, a mixture of essential macro- and micronutrients excepting P and N, and all combinations thereof in each of four blocks. Additions of nutrients other than N or P had no significant effects on measured plant-growth variables. In contrast, additions of either N or P significantly increased tree height growth, diameter increments, biomass growth, and height growth of the understory fern Dicranopteris linearis in both sites. The effect of N was greater than that of P. Greatest growth rates occurred in plots receiving both N and P, and signficant N*P interactions occurred in several cases, suggesting a synergistic effect between these two elements. Plant growth on these young, poorly weathered, basaltic lavas is colimited by N and P availability. Growth in a similar-aged stand growing on a mixture of volcanic ash and cinders is N but not P limited, indicating that the texture of the parent material influences nutrient-availability patterns during early primary succession.

Changes in Asymbiotic, Heterotrophic Nitrogen Fixation on Leaf Litter of Metrosideros polymorpha with Long-Term Ecosystem Development in Hawaii

We measured nitrogenase activity (acetylene reduction) of asymbiotic, heterotrophic, nitrogen-fixing bacteria on leaf litter from the tree Metrosideros polymorpha collected from six sites on the Hawaiian archipelago. At all sites M. polymorpha was the dominant tree, and its litter was the most abundant on the forest floor. The sites spanned a soil chronosequence of 300 to 4.1 million y. We estimated potential nitrogen fixation associated with this leaf litter to be highest at the youngest site (1.25 kg ha-1 y-1), declining to between 0.05 and 0.22 kg ha-1 y-1 at the oldest four sites on the chronosequence. To investigate how the availability of weathered elements influences N fixation rates at different stages of soil development, we sampled M. polymorpha leaf litter from complete, factorial fertilization experiments located at the 300-y, 20,000-y and 4.1 million–y sites. At the youngest and oldest sites, nitrogenase activity on leaf litter increased significantly in the plots fertilized with phosphorus and “total” (all nutrients except N and P); no significant increases in nitrogenase activity were measured in leaf litter from treatments at the middle-aged site. The results suggest that the highest rates of N fixation are sustained during the “building” or early phase of ecosystem development when N is accumulating and inputs of geologically cycled (lithophilic) nutrients from weathering are substantial.

Predicting and mitigating weed invasions to restore natural post-fire succession in Mesa Verde National Park, Colorado, USA

Six large wildfires have burned in Mesa Verde National Park during the last 15 years, and extensive portions of burns were invaded by non-native plant species. The most threatening weed species include Carduus nutans, Cirsium arvense, and Bromus tectorum, and if untreated, they persist at least 13 years. We investigated patterns of weed distribution to identify plant communities most vulnerable to post-fire weed invasion and created a spatially explicit model to predict the most vulnerable sites. At the scale of the entire park, mature pinon–juniper woodlands growing on two soil series were most vulnerable to post-fire weed invasion; mountain shrublands were the least vulnerable. At a finer scale, greater richness of native species was correlated with greater numbers of non-native species, indicating that habitats with high native biodiversity are at the greatest risk of weed invasion. In unburned areas, weed density increased with greater soil nitrogen and phosphorus, and lower salinity. In burned areas weed density correlated with soil nitrogen status and textural class. We also evaluated the effectiveness of a variety of weed mitigation methods; aerial seeding of targeted high-risk areas with native grasses was the most effective treatment tested. We recommend a conservative mitigation plan using natives grass seed on only the most invasible sites.