Shauna M. Uselman

Effects of increased atmospheric CO2, temperature, and soil N availability on root exudation of dissolved organic carbon by a N-fixing tree (Robinia pseudoacacia L.)

Root exudation has been hypothesized as one possible mechanism that may lead to increased inputs of organic C into the soil under elevated atmospheric CO2, which could lead to greater long-term soil C storage. In this study, we analyzed exudation of dissolved organic C from the roots of seedlings of the N-fixing tree Robinia pseudoacacia L. in a full factorial design with 2 CO2 (35.0 and 70.0 Pa) × 2 temperature (26° and 30 °C during the day) × 2 N fertilizer (0 and 10.0 mM N concentration) levels. We also analyzed the decomposition rates of root exudate to estimate gross rates of exudation. Elevated CO2 did not affect root exudation of organic C. A 4 °C increase in temperature and N fertilization did, however, significantly increase organic C exudation rates. Approximately 60% of the exudate decomposed relatively rapidly, with a turnover rate of less than one day, while the remaining 40% decomposed more slowly. These results suggest that warmer climates, as predicted for the next century, may accelerate root exudation of organic C, which will probably stimulate rapid C cycling and may make a minor contribution to intermediate to more long-term soil C storage. However, as these losses to root exudation did not exceed 1.2% of the net C fixed by Robinia pseudoacacia, root exudation of organic C appears to have little potential to contribute to long-term soil C sequestration.

Tamarisk biocontrol in the western United States: ecological and societal implications

Tamarisk species (genus Tamarix), also commonly known as saltcedar, are among the most successful plant invaders in the western United States. At the same time, tamarisk has been cited as having enormous economic costs. Accordingly, local, state, and federal agencies have undertaken considerable efforts to eradicate this invasive plant and restore riparian habitats to pre-invasion status. Traditional eradication methods, including herbicide treatments, are now considered undesirable, because they are costly and often have unintended negative impacts on native species. A new biological control agent, the saltcedar leaf beetle (Diorhabda elongata), has been released along many watersheds in the western US, to reduce the extent of tamarisk cover in riparian areas. However, the use of this insect as a biological control agent may have unintended ecological, hydrological, and socioeconomic consequences that need to be anticipated by land managers and stakeholders undertaking restoration efforts. Here, we examine the possible ramifications of tamarisk control and offer recommendations to reduce potential negative impacts on valued riparian systems in the western US.

Soil formation and organic matter accretion in a young andesitic chronosequence at Mt. Shasta, California

The objectives of this work were to study rates of increase in allophane concentration, specific surface area changes, and organic matter accretion in a young andesitic chronosequence. We sampled the 0–10-, 10–20-, 30–40-, 70–80-, and 140–150-cm depths of 77-, 255-, 616-, and approximately 1200+-year-old soils and analyzed them for allophane, ferrihydrite, specific surface area, cation exchange capacity, soil pH, and C and N concentrations. Allophane concentrations increased at rates up to a maximum of 0.14 g kg−1 year−1, and concentrations are up to 68 times higher in the oldest than in the youngest soil. During the same time ferrihydrite concentrations increased only by a factor of 2.3. The specific surface area that could be attributed to allophane was only 2–4% in the youngest soil but was 41–97% in the oldest soil. Carbon and N stocks increased linearly with soil age over the first ∼600 years with rates of 139 kg C ha−1 year−1 and 5.3 kg N ha−1 year−1, respectively. After about 600 years, accretion rates were lower. Increases in allophane concentrations lead to increased cation exchange capacity in the soil. Our results indicate that the ability of the soil to retain nutrients improved with soil development.

A test of a potential short cut in the nitrogen cycle: The role of exudation of symbiotically fixed nitrogen from the roots of a N-fixing tree and the effects of increased atmospheric CO2 and temperature

N-fixing trees facilitate the growth of neighboring trees of other species. These neighboring species benefit from the simple presence of the N fixation symbiosis in their surroundings. Because of this phenomenon, it has been hypothesized that a change in atmospheric CO2 concentration may alter the role of N-fixing trees in their environment. It is thought that the role of N-fixing trees in ecosystems of the future may be more important since they may help sustain growth increases due to increased CO2 concentration in nitrogen limited forests. We examined: (1) whether symbiotically fixed N was exuded from roots, (2) whether a doubled atmospheric CO2 concentration would result in increased organic N exudation from roots, and (3) whether increased temperature or N availability affected N exudation from roots. This study analyzed exudation of dissolved organic N from the roots of seedlings of the N-fixing tree Robinia pseudoacacia L. in a full factorial design with 2 CO2 (35.0 and 70.0 Pa) × 2 temperature (26 or 30 °C during the day) × 2 N fertilizer (0 and 10.0 mM N concentration) levels. Trees with no other source of N except N fixation exuded about 1% to 2% of the fixed N through their roots as dissolved organic N. Increased atmospheric CO2 concentrations did not, however, increase N exudation rates on a per gram belowground biomass basis. A 4 °C increase in temperature and N fertilization did, however, significantly increase N exudation rates. These results suggest that exudation of dissolved organic N from roots or nodules of N-fixing trees could be a significant, but minor, pathway of transferring N to neighboring plants in a much more rapid and direct way than cycling through death, decomposition and mineralization of plant residues. And, while exudation rates of dissolved organic N from roots were not significantly affected by atmospheric CO2 concentration, the previously observed ‘CO2 fertilization effect’ on N-fixing trees suggests that N exudation from roots could play a significant but minor role in sustaining increases in forest growth, and thus C storage, in a CO2 enriched atmosphere.

Ecophysiological responses of salt cedar (Tamarix spp. L.) to the northern tamarisk beetle (Diorhabda carinulata Desbrochers) in a controlled environment.

The northern tamarisk beetle (Diorhabda carinulata Desbrochers) was released in several western states as a biocontrol agent to suppress Tamarix spp. L. which has invaded riparian ecosystems; however, effects of beetle herbivory on Tamarix physiology are largely undocumented and may have ecosystem ramifications. Herbivory by this insect produces discoloration of leaves and premature leaf drop in these ecosystems, yet the cause of premature leaf drop and the effects of this leaf drop are still unknown. Insect herbivory may change leaf photosynthesis and respiration and may affect a plant’s ability to regulate water loss and increase water stress. Premature leaf drop may affect plant tissue chemistry and belowground carbon allocation. We conducted a greenhouse experiment to understand how Tamarix responds physiologically to adult beetle and larvae herbivory and to determine the proximate cause of premature leaf drop. We hypothesized that plants experiencing beetle herbivory would have greater leaf and root respiration rates, greater photosynthesis, increased water stress, inefficient leaf nitrogen retranslocation, lower root biomass and lower total non-structural carbohydrates in roots. Insect herbivory reduced photosynthesis rates, minimally affected respiration rates, but significantly increased water loss during daytime and nighttime hours and this produced increased water stress. The proximate cause for premature leaf drop appears to be desiccation. Plants exposed to herbivory were inefficient in their retranslocation of nitrogen before premature leaf drop. Root biomass showed a decreasing trend in plants subjected to herbivory. Stress induced by herbivory may render these trees less competitive in future growing seasons.

First-Year Establishment, Biomass and Seed Production of Early vs. Late Seral Natives in Two Medusahead (Taeniatherum caput-medusae) Invaded Soils

Abstract Re-seeding efforts to restore or rehabilitate Great Basin rangelands invaded by exotic annual grasses are expensive and have generally achieved limited success. There is a need to identify new strategies to improve restoration outcomes. We tested the performance of a native early seral seed mix (annual forbs, early seral grasses and shrubs) with that of a native late seral mix representative of species commonly used in restoration when growing with medusahead in soils of contrasting texture (sandy loam and clay loam) through the first growing season after seeding. Natives were also seeded without medusahead. We found that the grasses and forbs in the early seral mix established significantly better than those in the late seral mix, and the early seral mix significantly reduced aboveground biomass and seed production of medusahead by 16 and 17% respectively, likely because of competition with the early seral native forb, bristly fiddleneck. Medusahead performance was reduced in both soil types, suggesting utility of bristly fiddleneck in restoration is not limited to only one soil type. In contrast, the late seral mix did not suppress medusahead establishment, aboveground biomass or seed production. Although the native perennial grasses, particularly early seral species, were able to establish with medusahead, these grasses did not appear to have a suppressive effect on medusahead during the first growing season. Medusahead was able to establish and produce seeds on both soil types, demonstrating an ability to expand its current range in the Intermountain West, though aboveground biomass and seed production was higher in the clay loam. Our results suggest that certain species may play a key role in restoration, and that targeting early seral species in particular to find additional native species with the ability to suppress exotic annual grasses is an important next step in improving restoration outcomes in desert ecosystems. Nomenclature: Medusahead (Taeniatherum caput-medusae (L.) Nevski), bristly fiddleneck (Amsinckia tessellata A. Gray). Management Implications: Medusahead is an exotic annual grass that that has invaded into the Intermountain West of the U.S., reducing native species biodiversity and increasing fire frequency. In a study of native and medusahead performance, we found that the early seral native annual forb, bristly fiddleneck, was an effective competitor with medusahead in two soil types, significantly reducing biomass and seed production by 16 to 17%. Given that this effect was relatively small, further research to examine whether the use of increased seeding density of bristly fiddleneck and/or whether greater diversity of species in the seeding mix would enhance exotic suppression is warranted. Native perennial grasses, particularly early seral grasses, established in higher numbers than native forbs and shrubs, demonstrating their importance in restoration seedings. Although they did not appear to have a suppressive effect on medusahead during their first growing season, perennial grasses have been found to be effective competitors with exotic annual grasses once mature. Our findings suggest that efforts to find additional novel candidate species for seed mixtures may be best focused on early successional species, similar to bristly fiddleneck, to improve restoration/rehabilitation outcomes in disturbed rangeland ecosystems.

Restoring dryland old fields with native shrubs and grasses: Does facilitation and seed source matter?

Restoration of agricultural fields is challenging, especially in arid and semi-arid ecosystems. We conducted experiments in two fields in the Great Basin, USA, which differed in cultivation history and fertility. We tested the effects of different levels of functional diversity (planting grasses and shrubs together, vs. planting shrubs alone), seed source (cultivars, local or distant wild-collections), and irrigation regime (spring or fall and spring) on restoration outcomes. We sowed either: 1) grasses and shrubs in year one, 2) shrubs only, in year one, 3) grasses in year one with herbicide, shrubs in year two, or 4) shrubs alone in year two, after a year of herbicide. We irrigated for two years and monitored for three years. Shrub emergence was highest in the lower fertility field, where increasing functional diversity by seeding grasses had a neutral or facilitative effect on shrub emergence. In the higher fertility field, increasing functional diversity appeared to have a neutral to competitive effect. After declines in shrub densities after irrigation ceased, these effects did not persist. Grasses initially suppressed or had a neutral effect on weeds relative to an unseeded control, but had neutral or facilitative effects on weeds relative to shrub-only seeding. Initially, commercial grasses were either equivalent to or outperformed wild-collected grasses, but after irrigation ceased, commercial grasses were outperformed by wild-collected grasses in the higher fertility field. Local shrubs initially outperformed distant shrubs, but this effect did not persist. Fall and spring irrigation combined with local shrubs and wild-collected grasses was the most successful strategy in the higher fertility field, while in the lower fertility field, irrigation timing had fewer effects. Superior shrub emergence and higher grass persistence indicated that the use of wild and local seed sources is generally warranted, whereas the effects of functional diversity and irrigation regime were context-dependent. A bet-hedging approach that uses a variety of strategies may maximize the chances of restoration success.