Structural diversity is better associated with forest productivity than species or functional diversity.

Understanding the relationship between biodiversity and productivity can be advanced by improving metrics used to quantify biodiversity. Structural diversity, that is, variation of size and form of plant organs, is an emerging biodiversity metric. However, compared with the other biodiversity metrics, its relative importance in specific components of forest productivity, for example, recruitment of new individuals, biomass net change after accounting for mortality, is largely unknown, particularly across a large spatial scale with multiple influential gradients. To address the knowledge gap, we used USDA Forest Service Forest Inventory and Analysis (FIA) data across the southcentral USA from 2008 to 2017. We calculated forest biomass increments due to recruitment and growth and net change in biomass. Then, we quantified the effects of a range of abiotic and biotic variables on the biomass increments and net change. Our results showed that (1) Structural diversity was negatively associated with the two biomass increments and net change in biomass. The negative effects were supported by increased occurrences of insects and diseases with greater structural diversity. (2) Compared with species and functional diversity, structural diversity showed a better association with biomass increments and net change, suggested by its larger absolute values of standardized coefficients, and the effects of structural diversity were negative in contrast to species diversity. (3) The effects of structural diversity, stand age, and elevation differed between natural and planted forests that may stem from the differences in stand development and species composition between the two forest types. Together, structural diversity may represent an important dimension of biodiversity impacts on plant productivity, which could be related to the exacerbated disturbances with greater structural diversity.

Trade-off of ecosystem productivity and water use related to afforestation in southcentral USA under climate change.

The increase of tree canopy cover due to woody plant encroachment and tree plantations modifies both carbon and water dynamics. The tradeoffs between ecosystem net primary productivity (NPP) and water use with increasing tree cover in different climate conditions, particularly under future climate scenarios, are not well understood. Within the climate transition zone of the southern Great Plains, USA, we used the Soil and Water Assessment Tool+ (SWAT+) to investigate the combined impacts of increasing tree cover and climate change on carbon and water dynamics in three watersheds representing semiarid, subhumid, and humid climates. Model simulations incorporated two land use modifications (Baseline: existing tree cover; Forest +: increasing evergreen tree cover), in conjunction with two climate change projections (the RCP45 and the RCP85), spanning two time periods (historic: 1991-2020; future: 2070-2099). With climate change, the subhumid and humid watersheds exhibited a greater increase in evapotranspiration (ET) and a corresponding reduction in runoff compared to the semi-arid watershed, while the semi-arid and subhumid watersheds encountered pronounced losses in water availability for streams (>200 mm/year) due to increasing tree cover and climate change. With every 1 % increase in tree cover, both NPP and water use efficiency were projected to increase in all three watersheds under both climate change scenarios, with the subhumid watershed demonstrating the largest increases (>0.16 Mg/ha/year and 170 %, respectively). Increasing tree cover within grasslands, either through woody plant expansion or afforestation, boosts ecosystem NPP, particularly in subhumid regions. Nevertheless, this comes with a notable decrease in water resources, a concern made worse by future climate change. While afforestation offers the potential for greater NPP, it also brings heightened water scarcity concerns, highlighting the importance of tailoring carbon sequestration strategies within specific regions to mitigate unintended repercussions on water availability.

Escherichia coli efflux from rangeland ecosystems in the southcentral Great Plains of the United States.

Bacterial contamination of surface water is a public health concern. To quantify the efflux of Escherichia coli into ephemeral and intermittent streams and assess its numbers in relation to secondary body contact standards, we monitored runoff and measured E. coli numbers from 10 experimental watersheds that differed in vegetation cover and cattle access in north-central Oklahoma. Escherichia coli numbers were not significantly different among the watersheds, with one exception; the grazed prairie watershed (GP1) had greater numbers compared to one ungrazed prairie watershed (UP2). Median E. coli numbers in runoff from ungrazed watersheds ranged from 260 to 1482 MPN/100 mL in comparison with grazed watersheds that ranged from 320 to 8878 MPN/100 mL. In the GP1 watershed, higher cattle stocking rates during pre- and post-calving (February-May) resulted in significantly greater bacterial numbers and event loading compared to periods with lower stocking rates. The lack of significance among watersheds is likely due to the grazed sites being rotationally (and lightly) grazed, data variability, and wildlife contributions. To address wildlife sources, we used camera trap data to assess the usage in the watersheds; however, the average number of animals in a 24-h period did not correlate with observed median E. coli numbers. Because of its impacts on E. coli numbers in water, grazing management (stocking rate, rotation, and timing) should be considered for improving water quality in streams and reservoirs.

Economic returns and the perceived obstacles to adopting active management in the forest-grassland transition ecoregion in south-central USA.

Forest-grassland ecotones are a mosaic of grassland, savanna, and upland forest. As such, landowners may have opportunities to choose to manage their lands for multiple objectives. We estimated the economic returns from managing forest and rangeland in southeastern Oklahoma, USA to produce different combinations of timber, cattle forage, and white-tailed deer (Odocoileus virginianus Zimmermann) browse for a 40-year period. We further conducted a survey to understand landowner perceptions of obstacles to adopting active management that involve timber harvest and prescribed fire. The highest net return was obtained from the treatment with harvested timber that was burned every four years (uneven-aged woodland/forest) because it had the greatest gross return from a combination of timber (46%), cattle forage (42%), and deer browse (11%). The return from this treatment was greater than that for managed for timber only (closed-canopy forest) or prioritizing cattle and deer (savanna). Survey results suggested that landowners were aware of the benefits of active management but that the majority (66%) considered cost a major obstacle in the management of their forest or rangeland. In particular, women forestland owners and older landowners considered cost an obstacle. Our findings advocate integrated timber, cattle, and deer management as the best economic strategy within the forest-grassland ecotone and for targeted outreach and landowner education related to the benefits of active management.

Intentions of Landowners towards Active Management of Ecosystem for Deer Habitat.

Active management such as prescribed fire and thinning can restore savanna and prairie ecosystem to maintain a full suite of ecosystem services and create suitable habitat for wildlife species such as white-tailed deer (Odocoileus virginianus). Active management comes with the cost of management and acceptance of management tools. The south-central transitional ecoregion of the USA, which otherwise was a mixture of forest, savanna, and tallgrass prairie, is increasing in woody plant dominance due to the exclusion of fire and other anthropogenic factors. Deer hunting is a vital source of revenue generation to offset the landowner's management cost in the region. We studied Oklahoma landowners' perceptions regarding active and sustainable management of forest and rangeland for deer habitat using two established theories of reasoned action and planned behavior as well as expanded theories adding moral norms. We analyzed mailed survey data using structural equation modeling. We found that subjective norms and perceived behavior control significantly affected deer hunting intention when moral norms were introduced into the model. Attitudes independently significantly affected intentions of deer hunting but have negative relations with the intentions. The study suggested that landowners have positive social pressure and were interested in active management but associated financial burden and risk could be shaping negative attitudes.

Spatial variability in tree-ring carbon isotope discrimination in response to local drought across the entire loblolly pine natural range.

Considering the temporal responses of carbon isotope discrimination (Δ13C) to local water availability in the spatial analysis of Δ13C is essential for evaluating the contribution of environmental and genetic facets of plant Δ13C. Using tree-ring Δ13C from years with contrasting water availability at 76 locations across the natural range of loblolly pine, we decomposed site-level Δ13C signals to maximum Δ13C in well-watered conditions (Δ13Cmax) and isotopic drought sensitivity (m) as a change in Δ13C per unit change of Palmer's Drought Severity Index (PDSI). Site water status, especially the tree lifetime average PDSI, was the primary factor affecting Δ13Cmax. The strong spatial correlation exhibited by m was related to both genetic and environmental factors. The long-term average water availability during the period relevant to trees as indicated by lifetime average PDSI correlated with Δ13Cmax, suggesting acclimation in tree gas-exchange traits, independent of incident water availability. The positive correlation between lifetime average PDSI and m indicated that loblolly pines were more sensitive to drought at mesic than xeric sites. The m was found to relate to a plant's stomatal control and may be employed as a genetic indicator of efficient water use strategies. Partitioning Δ13C to Δ13Cmax and m provided a new angle for understanding sources of variation in plant Δ13C, with several fundamental and applied implications.

Drought Tolerance and Competition in Eastern Redcedar (Juniperus virginiana) Encroachment of the Oak-Dominated Cross Timbers.

On the dry, western edge of the eastern deciduous forest of the USA (Cross Timbers), the drought-tolerant, evergreen eastern redcedar (Juniperus virginiana) is encroaching into post oak- (Quercus stellata) dominated woodlands. The overall goal of this study was to examine whether the drought tolerance strategies of eastern redcedar provide it a competitive advantage over post oak and whether this is a key attribute facilitating its successful establishment in the Cross Timbers. Specifically, we assessed xylem water potential and leaf gas exchange of these two species growing in single-species stands and in a mixed-species stand. We found that both species exhibit a similar degree of isohydry and close their stomates to the same extent in response to declining xylem water potentials. Both species had similar relative reductions in gas exchange in response to drought, despite differences in xylem anatomy. However, post oak had leaf-level gas exchange rates approximately 5× greater than eastern redcedar during periods of high moisture availability. Therefore, it did not appear that eastern redcedar encroachment into an oak-dominated forest is facilitated by growing season differences in carbon gain, although evergreen eastern redcedar can conduct gas exchange year-round when conditions are favorable while post oak is deciduous. We found that volumetric soil water content (0-45 cm) was lower in the pure eastern redcedar stand than the mixed-species or pure post oak stand which may indicate that eastern redcedar may experience favorable soil moisture conditions when encroaching into open oak woodlands. Moreover, water potentials in eastern redcedar tended to be more negative in pure stands compared to the mixed stand. Our results suggest the two species may be using water from different depths, reducing competition. Overall, our findings indicate that eastern redcedar encroachment into formerly oak-dominated Cross Timbers forests likely will continue under moderate drought, in the absence of fire, with consequences for water budgets, carbon cycling, grazing forage, wildlife habitat, and wildfire risk.

The ecological and economic determinants of eastern redcedar (Juniperus virginiana) encroachment in grassland and forested ecosystems: A case study from Oklahoma.

Frequent fires were used as a management tool to maintain prairies, savannas, and woodlands in the southern Great Plains of the United States. However, fire exclusion beginning in the mid-1900s allowed for the establishment and growth of fire-intolerant species such as eastern redcedar (Juniperus virginiana: ERC) beyond their naturally occurring habitats. Apart from the reduction in burning, wide soil and climate adaptability, and seed dispersal by birds have facilitated the expansion of ERC in the southern Great Plains. The encroachment of ERC has caused heavy ecological and economic losses to Oklahoma and thus has been a major management concern for the past few decades. This study utilized count data modeling to analyze USDA Forest Service's (USFS) Forest Inventory and Analysis (FIA) data to investigate the relationship between available ecological and economic factors and the abundance of ERC in grassland and forested ecosystems of Oklahoma. The results suggested that low site productivity, high basal area, dense canopy, and silt loam soil texture significantly increase the abundance of ERC on a given site. The results also indicated that the rate of ERC encroachment is 3.3% higher in the softwood and 2.0% higher in the miscellaneous forests, compared to the hardwood forests. However, the economic variables of the study such as ownership type, adoption of active management, and proximity to a metropolitan area did not show a significant relationship to the abundance of ERC.

Dead or dying? Quantifying the point of no return from hydraulic failure in drought-induced tree mortality.

Determining physiological mechanisms and thresholds for climate-driven tree die-off could help improve global predictions of future terrestrial carbon sinks. We directly tested for the lethal threshold in hydraulic failure - an inability to move water due to drought-induced xylem embolism - in a pine sapling experiment. In a glasshouse experiment, we exposed loblolly pine (Pinus taeda) saplings (n = 83) to drought-induced water stress ranging from mild to lethal. Before rewatering to relieve drought stress, we measured native hydraulic conductivity and foliar color change. We monitored all measured individuals for survival or mortality. We found a lethal threshold at 80% loss of hydraulic conductivity - a point of hydraulic failure beyond which it is more likely trees will die, than survive, and describe mortality risk across all levels of water stress. Foliar color changes lagged behind hydraulic failure - best predicting when trees had been dead for some time, rather than when they were dying. Our direct measurement of native conductivity, while monitoring the same individuals for survival or mortality, quantifies a continuous probability of mortality risk from hydraulic failure. Predicting tree die-off events and understanding the mechanism involved requires knowledge not only of when trees are dead, but when they begin dying - having passed the point of no return.

Vegetation Controls on the Spatio-Temporal Heterogeneity of Deep Moisture in the Unsaturated Zone: A Hydrogeophysical Evaluation.

Information on the spatio-temporal variability of soil moisture in the vadose zone is important to assess groundwater recharge and solute transport in unconsolidated substrate as influenced by biological processes. Time-lapse electrical resistivity imaging (ERI) was used to monitor soil moisture dynamics to a depth of 9 m in a grassland, a grassland encroached by a juniper species (eastern redcedar, Juniperus virginiana), a juniper woodland and an oak forest in the south-central Great Plains, Oklahoma, USA. A site-specific relationship between moisture content and electrical conductivity data was developed for the soil zone, and a perched water zone was monitored at two of the sites. Results showed that (a) change in soil moisture content was linearly correlated to change in electric conductivity in the soil zone; (b) vegetation cover type induced differences in vertical bulk electrical resistivity (ER) profiles and influenced the temporal evolution of soil moisture profiles; and (c) juniper encroachment lowered the water level in the perched groundwater aquifer. Our results suggest land use and vegetation cover type, as opposed to rock properties, controls deep water drainage for the vegetation transition zone. Methods used to measure hydrogeophysical changes, such as ERI, can be used for broader understanding of geological, physical, and biological processes and their links in Earth's critical zones.

Canopy Interception for a Tallgrass Prairie under Juniper Encroachment.

Rainfall partitioning and redistribution by canopies are important ecohydrological processes underlying ecosystem dynamics. We quantified and contrasted spatial and temporal variations of rainfall redistribution for a juniper (Juniperus virginiana, redcedar) woodland and a tallgrass prairie in the south-central Great Plains, USA. Our results showed that redcedar trees had high canopy storage capacity (S) ranging from 2.14 mm for open stands to 3.44 mm for closed stands. The canopy funneling ratios (F) of redcedar trees varied substantially among stand type and tree size. The open stands and smaller trees usually had higher F values and were more efficient in partitioning rainfall into stemflow. Larger trees were more effective in partitioning rainfall into throughfall and no significant changes in the total interception ratios among canopy types and tree size were found. The S values were highly variable for tallgrass prairie, ranging from 0.27 mm at early growing season to 3.86 mm at senescence. As a result, the rainfall interception by tallgrass prairie was characterized by high temporal instability. On an annual basis, our results showed no significant difference in total rainfall loss to canopy interception between redcedar trees and tallgrass prairie. Increasing structural complexity associated with redcedar encroachment into tallgrass prairie changes the rainfall redistribution and partitioning pattern at both the temporal and spatial scales, but does not change the overall canopy interception ratios compared with unburned and ungrazed tallgrass prairie. Our findings support the idea of convergence in interception ratio for different canopy structures under the same precipitation regime. The temporal change in rainfall interception loss from redcedar encroachment is important to understand how juniper encroachment will interact with changing rainfall regime and potentially alter regional streamflow under climate change.

Increased vapor pressure deficit due to higher temperature leads to greater transpiration and faster mortality during drought for tree seedlings common to the forest-grassland ecotone.

Tree species growing along the forest-grassland ecotone are near the moisture limit of their range. Small increases in temperature can increase vapor pressure deficit (VPD) which may increase tree water use and potentially hasten mortality during severe drought. We tested a 40% increase in VPD due to an increase in growing temperature from 30 to 33°C (constant dewpoint 21°C) on seedlings of 10 tree species common to the forest-grassland ecotone in the southern Great Plains, USA. Measurement at 33 vs 30°C during reciprocal leaf gas exchange measurements, that is, measurement of all seedlings at both growing temperatures, increased transpiration for seedlings grown at 30°C by 40% and 20% for seedlings grown at 33°C. Higher initial transpiration of seedlings in the 33°C growing temperature treatment resulted in more negative xylem water potentials and fewer days until transpiration decreased after watering was withheld. The seedlings grown at 33°C died 13% (average 2 d) sooner than seedlings grown at 30°C during terminal drought. If temperature and severity of droughts increase in the future, the forest-grassland ecotone could shift because low seedling survival rate may not sufficiently support forest regeneration and migration.

Forest floor depth mediates understory vigor in xeric Pinus palustris ecosystems.

Longleaf pine (Pinus palustris) woodlands and savannas are among the most frequently burned ecosystems in the world with fire return intervals of 1-10 years. This fire regime has maintained high levels of biodiversity in terms of both species richness and endemism. Land use changes have reduced the area of this ecosystem by >95%, and inadequate fire frequencies threaten many of the remnants today. In the absence of frequent fire, rapid colonization of hardwoods and shrubs occurs, and a broad-leaved midstory develops. This midstory encroachment has been the focus of much research and management concern, largely based on the assumption that the midstory reduces understory plant diversity through direction competition via light interception. The general application of this mechanism of degradation is questionable, however, because midstory density, leaf area, and hardwood species composition vary substantially along a soil moisture gradient from mesic to extremely xeric sites. Reanalysis of recently reported data from xeric longleaf pine communities suggests that the development of the forest floor, a less conspicuous change in forest structure, might cause a decline in plant biodiversity when forests remain unburned. We report here a test of the interactions among fire, litter accumulation, forest floor development, and midstory canopy density on understory plant diversity. Structural equation modeling showed that within xeric sites, forest floor development was the primary factor explaining decreased biodiversity. The only effects of midstory development on biodiversity were those mediated through forest floor development. Boundary line analysis of functional guilds of understory plants showed sensitivity to even minor development of the forest floor in the absence of fire. These results challenge the prevailing management paradigm and suggest that within xeric longleaf pine communities, the primary focus of managed fire regime should be directed toward the restoration of forest floor characteristics rather than the introduction of high-intensity fires used to regulate midstory structure.

Radiation-use efficiency and gas exchange responses to water and nutrient availability in irrigated and fertilized stands of sweetgum and sycamore.

We investigated how water and nutrient availability affect radiation-use efficiency (epsilon) and assessed leaf gas exchange as a possible mechanism for shifts in epsilon. We measured aboveground net primary production (ANPP) and annual photosynthetically active radiation (PAR) capture to calculate epsilon as well as leaf-level physiological variables (light-saturated net photosynthesis, Asat; stomatal conductance, gs; leaf internal CO2 concentration, Ci; foliar nitrogen concentration, foliar [N]; and midday leaf water potential, Psileaf) during the second (2001) and third (2002) growing seasons in sweetgum (Liquidambar styraciflua L.) and sycamore (Platanus occidentalis L.) stands receiving a factorial combination of irrigation and fertilization at the Savannah River Site, South Carolina. Irrigation and fertilization increased PAR capture (maximum increase 60%) in 2001 and 2002 for both species and annual PAR capture was well correlated with ANPP (mean r2 = 0.77). A decreasing trend in epsilon was observed in non-irrigated stands for sweetgum in 2001 and for sycamore in both years, although this was only significant for sycamore in 2002. Irrigated stands maintained higher gas exchange rates than non-irrigated stands for sweetgum in 2001 and for sycamore in both years, although foliar [N] and Psileaf were generally unaffected. Because Ci decreased in proportion to gs in non-irrigated stands, it appeared that greater stomatal limitation of photosynthesis was associated with decreased Asat. On several measurement dates for sweetgum in 2001 and for sycamore in both years, epsilon was positively correlated with gas exchange variables (Asat, gs, Ci) (r ranged from 0.600 to 0.857). These results indicate that PAR capture is well correlated with ANPP and that gas exchange rates modified by irrigation can influence the conversion of captured light energy to biomass.

Diurnal and seasonal variability in the radial distribution of sap flow: predicting total stem flow in Pinus taeda trees.

We monitored the radial distribution of sap flux density (v; g H2O m(-2) s(-1)) in the sapwood of six plantation-grown Pinus taeda L. trees during wet and dry soil periods. Mean basal diameter of the 32-year-old trees was 33.3 cm. For all trees, the radial distribution of sap flow in the base of the stem (i.e., radial profile) was Gaussian in shape. Sap flow occurred maximally in the outer 4 cm of sapwood, comprising 50-60% of total stem flow (F), and decreased toward the center, with the innermost 4 cm of sapwood (11-15 cm) comprising less than 10% of F. The percent of flow occurring in the outer 4 cm of sapwood was stable with time (average CV < 10%); however, the percentage of flow occurring in the remaining sapwood was more variable over time (average CV > 40%). Diurnally, the radial profile changed predictably with time and with total stem flow. Seasonally, the radial profile became less steep as the soil water content (theta) declined from 0.38 to 0.21. Throughout the season, daytime sap flow also decreased as theta decreased; however, nighttime sap flow (an estimate of stored water use) remained relatively constant. As a result, the percentage of stored water use increased as theta declined. Time series analysis of 15-min values of F, theta, photosynthetically active radiation (PAR) and vapor pressure deficit (D) showed that F lagged behind D by 0-15 min and behind PAR by 15-30 min. Diurnally, the relationship between F and D was much stronger than the relationship between F and PAR, whereas no relationship was found between F and theta. An autoregressive moving average (ARIMA) model estimated that 97% of the variability in F could be predicted by D alone. Although total sap flow in all trees responded similarly to D, we show that the radial distribution of sap flow comprising total flow could change temporally, both on daily and seasonal scales.

Acclimation of loblolly pine (Pinus taeda) seedlings to high temperatures.

To determine the extent to which loblolly pine seedlings (Pinus taeda L.) acclimate to high temperatures, seedlings from three provenances-southeastern Texas (mean annual temperature 20.3 degrees C), southwestern Arkansas (mean annual temperature 16.2 degrees C) and Chesapeake, Maryland (mean annual temperature 12.8 degrees C)-were grown at constant temperatures of 25, 30, 35 or 40 degrees C in growth chambers. After two months, only 14% of the seedlings in the 40 degrees C treatment survived, so the treatment was dropped from the experiment. Provenance and family differences were not significant for most measured variables. Total biomass was similar in the 25 and 30 degrees C treatments, and less in the 35 degrees C treatment. Foliage biomass was higher, and root biomass lower, in the 30 degrees C treatment compared with the 25 degrees C treatment. Net photosynthesis and dark respiration of all seedlings were measured at 25, 30 and 35 degrees C. Both net photosynthesis and dark respiration exhibited acclimation to the temperature at which the seedlings were grown. For each temperature treatment, the highest rate of net photosynthesis was measured at the growth temperature. Dark respiration rates increased with increasing measurement temperature, but the basal rate of respiration, measured at 25 degrees C, decreased from 0.617 &mgr;mol m(-2) s(-1) in the 25 degrees C treatment to 0.348 &mgr;mol m(-2) s(-1) in the 35 degrees C treatment, resulting in less carbon loss in the higher temperature treatments than if the seedlings had not acclimated to the growth conditions. Temperature acclimation, particularly of dark respiration, may explain why total biomass of seedlings grown at 30 degrees C was similar to that of seedlings grown at 25 degrees C.