An Ecohydrological Cellular Automata Model Investigation of Juniper Tree Encroachment in a Western North American Landscape

Woody plant encroachment over the past 140 years has substantially changed grasslands in western North American. We studied encroachment of western juniper (Juniperus occidentalis var. occidentalis) into a previously mixed shrub–grassland site in central Oregon (USA) using a modified version of Cellular Automata Tree–Grass–Shrub Simulator (CATGraSS) ecohydrological model. We developed simple algorithms to simulate three encroachment factors (grazing, fire frequency reduction, and seed dispersal by herbivores) in CATGraSS. Local ecohydrological dynamics represented by the model were first evaluated using satellite-derived leaf area index and measured evapotranspiration data. Reconstructed pre-encroachment vegetation cover percentages and the National Land Cover Database (NLCD 2006) vegetation map were used to estimate parameters for encroachment factors to represent juniper encroachment in CATGraSS. Model sensitivity experiments examined the influence of each encroachment factor and their combinations on trajectories of modeled percent cover of each plant functional type and emergent spatial vegetation patterns in the modeled domain. Simulation results identified grazing as the key factor leading to juniper encroachment, by reducing shrub and grass cover and promoting the formation of juniper tree clusters. Reduced fire frequency and increased seed dispersal by grazers further amplified juniper encroachment into grassland patches between clusters of juniper trees. Each encroachment factor showed different consequences on modeled vegetation patterns. Time series of modeled plant cover and spatial patterns of plant functional types were found to be consistent with an existing conceptual model described in the literature. The proposed model provides a tool that can be used to improve our understanding of the drivers and processes of woody plant encroachment and vegetation response to global change.     

Epiphytic lichens on Juniperus communis – an unexplored component of biodiversity in threatened alvar grassland

Semi‐natural grasslands, among them thin‐soil calcareous grasslands (alvars), have great conservation value but have become increasingly rare in Europe. The main threat to alvar grasslands is the encroachment by juniper Juniperus communis and therefore it is usually removed during the restoration practice. Juniper can also be a host plant for many epiphytic lichens, but its role as a phorophyte is poorly known. We studied epiphytic lichen diversity on 126 junipers in 17 sites in western Estonia and found 140 lichenized taxa including several rare and red‐listed species. Using indirect and direct multivariate analyses (DCA, pCCA) and general linear models we revealed that both habitat and phorophyte properties affect lichen assemblies on juniper. Lichen species richness per site showed a unimodal relationship with compound factors of site productivity and juniper characteristics (stem circumference and juniper width). Lichen species richness per phorophyte was increasing with its size and with the proportion of dead branches, and was twice higher in plate alvars than in ryhk alvars. Also, the species composition in plate alvars differed from ryhk alvars by having 42 characteristic lichen species in plate alvars vs three indicators of ryhk alvars. The composition of lichens was significantly influenced by encroachment of alvars, e.g. by high juniper cover and shrub layer height, as well as by the proportion of dead branches and stem circumference of juniper. We conclude that the epiphytic lichen assemblies on junipers are threatened by grassland encroachment similarly to ground layer lichen assemblies. We suggest that some old and scencent junipers should be preserved during the restoration of alvar grasslands.     

Diverse responses across soil parent materials during ecological restoration

A major challenge to advancing the science and practice of ecological restoration is working across large landscapes containing diverse sites that may respond differently to restoration. We conducted a 5‐year restoration experiment, replicated across 9 sites spanning 3 soil parent material types within a 9,000‐ha Pinus ponderosa forest landscape. We evaluated plant community response to restoration Pinus thinning, grazing, and aqueous smoke application. We measured vegetation before (2003) and 3 (2006) and 5 (2008) years after treatment. Plant community responses of species richness, cover, and composition were diverse, ranging from increases, decreases, or no change depending on soil parent material, tree thinning, and presence or exclusion of grazing. Restoration outcomes were under hierarchical control: soil parent material constrained response to Pinus thinning, which in turn influenced grazing effects. On limestone‐derived soil, responses included no change in species richness but increased plant cover with Pinus thinning. Both plant richness and cover increased on benmorite soil after thinning, and cover generally increased more without grazing. On rocky, basalt soil, plant richness increased but cover did not after any treatment. Diversity of responses to restoration has implications for: (1) setting goals or monitoring indicators tailored to inherent soil capability; (2) identifying where grazing most affects restoration outcomes; and (3) forecasting responses to restoration across landscapes. Diverse responses to restoration along physiographic gradients such as soil parent material warrant consideration when developing restoration across degraded landscapes.     

Influence of Colophospermum mopane canopy cover on litter decomposition and nutrient dynamics in a semi‐arid African savannah

The objective of this study was to investigate the influence of mopane canopy cover on litter decomposition in a semi‐arid African savannah. We used a randomized block design with five blocks of 100 × 100 m demarcated in a 10‐ha pocket of open mopane woodland. Litterbags were placed beneath large (8.3 m crown diameter) and small mopane trees (2.7 m crown diameter) and in the intercanopy area. Decomposition was fastest in the intercanopy area exposed to solar radiation (k = 0.35 year^?1), intermediate beneath small trees (k = 0.28 year^?1) and slowest beneath large trees (k = 0.23 year^?1). Soil temperatures beneath small and large trees were 3–5 and 6–9°C lower than in the intercanopy area, respectively. Bacterial and fungal counts were significantly higher (P < 0.05) beneath large than small trees and in the intercanopy area. The amount of N and P released did not vary significantly among sampling sites. Soil moisture in the dry season was similar among sampling sites but rainy‐season soil moisture was significantly greater (P < 0.05) beneath large than small trees and in the intecanopy area. Mopane canopy cover retarded litter decomposition suggesting that photodegradation could be an important factor controlling carbon turnover in semi‐arid African savannahs.     

Effects of young Artemisia rothrockii shrubs on soil moisture,soil nitrogen cycling,and resident herbs

Questions: How do young sagebrush shrubs (Artemisia rothrockii, Asteraceae) affect soil moisture availability? How do young sagebrush shrubs affect soil nitrogen cycling? How does the resident herb community respond to shrub removal in the early stages of sagebrush encroachment? Location: Mulkey and Bullfrog Meadows on the Kern Plateau in the Golden Trout Wilderness, Sierra Nevada Mountains, Inyo National Forest, Inyo County, California, USA. Methods: We removed young encroaching sagebrush shrubs from 3.5 m × 3.5 m plots and compared soil moisture, net mineralization, net nitrification, and herb cover with paired control plots over four growing seasons. Results: On average throughout the experiment, the difference between removal plots and control plots in soil moisture was small. Removal plots were wetter by 1.3 ± 2.0% at 0–30 cm depth, 2.1 ± 3.1% at 30–60 cm depth and 3.1 ± 5.8% at 60–90 cm depth. By contrast, after four years, net mineralization was 32 ± 26% (mean ± 95% CI) lower in sagebrush removal plots, suggesting that sagebrush encroachment increases rates of N‐cycling. Total herb cover was 13.0 ± 6.4% (mean ± 95% CI) higher in plots where young sagebrush shrubs were removed. This difference in cover appeared during the first season in which sagebrush shrubs were removed. Conclusions: Our results suggest that while young sagebrush shrubs do not contribute substantially to meadow drying, they alter N cycling rates, and may indirectly increase the rate of their own encroachment by competitively reducing resident herbs.     

Topographic control of vegetation in a mountain big sagebrush steppe

Mountain big sagebrush steppes in Wyoming have strong spatial patterning associated with topography. We describe the spatial variability of vegetation in a sagebrush steppe, and test the relationship between topography and vegetation using canonical correlation. Results of the analysis suggest that the main control over vegetation distribution in this system is wind exposure. Exposed sites are characterized by cushion plant communities and Artemisia nova, and less exposed sites by the taller sagebrush species Artemisia tridentata ssp. vaseyana. Topographic depressions and leeward slopes are characterized by aspen stands and nivation hollows. Measurements of soil microclimate suggest that a major influence of topographic position on vegetation is snow redistribution and its effect on soil moisture and temperature.Abbreviations ARNO Artemisia nova - ARTRW Artemisia tridentata ssp. wyomingensis - ARTRV Artemisia tridentata ssp. vaseyana - PUTR Purshia tridentata - RIP riparian community - POTR Populus tremuloides - NIV nivation hollow community     

To burn or not to burn: Comparing reintroducing fire with cutting an encroaching conifer for conservation of an imperiled shrub‐steppe

Woody vegetation has increased on rangelands worldwide for the past 100–200 years, often because of reduced fire frequency. However, there is a general aversion to reintroducing fire, and therefore, fire surrogates are often used in its place to reverse woody plant encroachment. Determining the conservation effectiveness of reintroducing fire compared with fire surrogates over different time scales is needed to improve conservation efforts. We evaluated the conservation effectiveness of reintroducing fire with a fire surrogate (cutting) applied over the last ~30 years to control juniper (Juniperus occidentalis Hook.) encroachment on 77 sagebrush‐steppe sites. Critical to conservation of this imperiled ecosystem is to limit juniper, not encourage exotic annual grasses, and promote sagebrush dominance of the overstory. Reintroducing fire was more effective than cutting at reducing juniper abundance and extending the period of time that juniper was not dominating the plant community. Sagebrush was reduced more with burning than cutting. Sagebrush, however, was predicted to be a substantial component of the overstory longer in burned than cut areas because of more effective juniper control. Variation in exotic annual grass cover was explained by environmental variables and perennial grass abundance, but not treatment, with annual grasses being problematic on hotter and drier sites with less perennial grass. This suggests that ecological memory varies along an environmental gradient. Reintroducing fire was more effective than cutting at conserving sagebrush‐steppe encroached by juniper over extended time frames; however, cutting was more effective for short‐term conservation. This suggests fire and fire surrogates both have critical roles in conservation of imperiled ecosystems.     

Inter- and under-canopy soil water, leaf-level and whole-plant gas exchange dynamics of a semi-arid perennial C4 grass

It is not clear if tree canopies in savanna ecosystems exert positive or negative effects on soil moisture, and how these might affect understory plant carbon balance. To address this, we quantified rooting-zone volumetric soil moisture (??_25?cm), plant size, leaf-level and whole-plant gas exchange of the bunchgrass, bush muhly (Muhlenbergia porteri), growing under and between mesquite (Prosopis velutina) in a southwestern US savanna. Across two contrasting monsoon seasons, bare soil ??_25?cm was 1.0?C2.5% lower in understory than in the intercanopy, and was consistently higher than in soils under grasses, where ??_25?cm was similar between locations. Understory plants had smaller canopy areas and volumes with larger basal diameters than intercanopy plants. During an above-average monsoon, intercanopy and understory plants had similar seasonal light-saturated leaf-level photosynthesis (A _net-sat), stomatal conductance (g _s-sat), and whole-plant aboveground respiration (R _auto), but with higher whole-plant photosynthesis (GEP_plant) and transpiration (T _plant) in intercanopy plants. During a below-average monsoon, intercanopy plants had higher diurnally integrated GEP_plant, R _auto, and T _plant. These findings showed little evidence of strong, direct positive canopy effects to soil moisture and attendant plant performance. Rather, it seems understory conditions foster competitive dominance by drought-tolerant species, and that positive and negative canopy effects on soil moisture and community and ecosystem processes depends on a suite of interacting biotic and abiotic factors.     

Habitat degradation,topography and rainfall variability interact to determine seed distribution and recruitment in a sand dune grassland

Question: Understanding the mechanisms underlying how habitat degradation, topography and rainfall variability interactively affect seed distribution and seedling recruitment is crucial for explaining plant community patterns and dynamics. Interactions between these major factors were studied together in a semiarid sand dune grassland. Location: Eastern Inner Mongolia, China. Methods: The study system used four sites of fixed, semifixed, semishifting and shifting sand dune grasslands, representing a gradient of habitat degradation. We investigated the density of germinable seeds deposited in the top 5 cm of soil and in situ seedling emergence (number of seedlings emerging early in the growing season) and establishment (number of plants recruited at the end of the growing season) at three topographic positions (dune top, windward and leeward sides) within each site over 2 years that differed in rainfall. Habitat characteristics (i.e. vegetation cover, plant species composition and diversity, soil moisture and nutrient availability and soil erodibility) of the four sites were also measured. Results: Habitat degradation (i.e. decreased vegetation cover and enhanced wind erosion rate) significantly reduced the size of the germinable soil seed bank. On average, germinable seed number from the high‐vegetation cover fixed dune was 36‐fold larger than the low‐vegetation cover shifting dune, and eight‐ and two‐fold larger, respectively, than the semishifting and semifixed dunes with intermediate vegetation cover. We observed within‐habitat variability in seed distribution, but among‐topographic position variation differed among habitats. Seedling recruitment showed large between‐year, and among‐ and within‐habitat variability, but these variations varied significantly depending on the response variables evaluated (i.e. initial seedling density, final plant density, emergence rate and recruitment rate). Path analysis revealed complex density‐dependent positive and negative, direct and indirect effects of germinable seed density and initial seedling density on recruitment, but the relative importance of these density‐dependent effects varied depending on habitat type and rainfall availability. Conclusion: Our results suggest that habitat degradation, microtopography and rainfall availability interact in shaping sand dune seed bank and plant community recruitment patterns and dynamics. Their effects were mainly mediated through changes in both the biotic and abiotic environment during the process of habitat deterioration.     

Predicting the post-fire establishment and persistence of an invasive tree species across a complex landscape

The reintroduction of pre-European fire regimes has allowed the entry of many invasive plant species into fire-dependant ecosystems of North America. However, the environmental factors that favor the post-fire establishment of these species across complex landscapes are not well understood and the initial establishment of invasive species does not necessarily result in long-term persistence. To evaluate the post-fire establishment and persistence of disturbance-dependent invasive plants, we studied the invasion of Paulownia tomentosa (princess tree, an early-successional species introduced from Asia) across three burns in the southern Appalachian Mountains. Based upon classification tree analysis, the presence/absence of P. tomentosa 2 years after burning was most strongly related to the cover of residual vegetation, topographic shading, and moisture availability. Spatial application of classification tree models to repeated survey data showed that P. tomentosa established across a wide range of microsites 2 years after burning. However, predicted habitat for P. tomentosa decreased by 63% 4 years after fire and by 73% 6 years after fire. Following its initial widespread establishment, P. tomentosa only persisted on xeric and exposed topographic positions that experienced high intensity burning. However, the sites where it persisted include rare community types that contain two endangered plant species that depend upon fire for successful reproduction. The control of P. tomentosa on these ecologically important sites may require special attention from land managers.     

Spatially explicit patterns in a dryland's soil respiration and relationships with climate,whole plant photosynthesis and soil fertility

Arid and semiarid ecosystems play a significant role in regulating global carbon cycling, yet our understanding of the controls over the dominant pathways of dryland CO₂ exchange remains poor. Substantial amounts of dryland soil are not covered by vascular plants and this patchiness in cover has important implications for spatial patterns and controls of carbon cycling. Spatial variation in soil respiration has been attributed to variation in soil moisture, temperature, nutrients and rhizodeposition, while seasonal patterns have been attributed to changes in moisture, temperature and photosynthetic inputs belowground. To characterize how controls over respiration vary spatially and temporally in a dryland ecosystem and to concurrently explore multiple potential controls, we estimated whole plant net photosynthesis (A_net) and soil respiration at four distances from the plant base, as well as corresponding fine root biomass and soil carbon and nitrogen pools, four times during a growing season. To determine if the controls vary between different plant functional types for Colorado Plateau species, measurements were made on the C₄ shrub, Atriplex confertifolia, and C₃ grass, Achnatherum hymenoides. Soil respiration declined throughout the growing season and diminished with distance from the plant base, though variations in both were much smaller than expected. The strongest relationship was between soil respiration and soil moisture. Soil respiration was correlated with whole plant A_net, although the relationship varied between species and distance from plant base. In the especially dry year of this study we did not observe any consistent correlations between soil respiration and soil carbon or nitrogen pools. Our findings suggest that abiotic factors, especially soil moisture, strongly regulate the response of soil respiration to biotic factors and soil carbon and nitrogen pools in dryland communities and, at least in dry years, may override expected spatial and seasonal patterns.     

Decadal‐scale aspen changes: evidence in remote sensing and tree ring data

Question: (1) Which remote sensing classification most successfully identify aspen using multitemporal Landsat 5 TM images and airborne lidar data? (2) How has aspen distribution changed in southwestern Idaho? (3) Are topographic variables and conifer encroachment correlated with aspen changes? Location: Reynolds Creek Experimental Watershed in southwestern Idaho, USA. Methods: Multi‐temporal Landsat 5 TM and lidar data were used individually and fused together. The best classification model was compared with a 1965 aspen map and tree ring data. Conifer encroachment was examined via image‐based change detection and field mapping. Lidar‐derived topographic variables were correlated with aspen change patterns using quantile regression models. Results: The best Landsat 5 TM classification was a normalized difference vegetation index (NDVI)‐based approach with 92% overall accuracy. The lidar classification of tree presence/absence performed with 100% overall accuracy. Fusing the lidar classification with various Landsat 5 TM classifications improved overall accuracies 3 to 6%. Among the fusion models, the NDVI‐lidar fusion performed best with 96% overall accuracy. Change detection indicated 69% decline in aspen cover, but 179% increase in aspen cover in other areas of the watershed. Conifers have completely replaced 17% of the aspen, while 93% of the remaining aspen stands have young Douglas‐fir and western juniper trees underneath the aspen canopy. Aspen significantly decreased (P‐values <0.05) with increasing elevation (up to 2150 m) and decreasing slope. Conclusions: Landsat 5 TM data used with a NDVI‐based approach provide an accurate method to classify aspen distribution. Landsat 5 TM classifications can be further improved via fusion with lidar data. Aspen change patterns are spatially variable: while aspen is drastically declining in some parts of this watershed, aspen is increasing in other areas.     

Ontogenetic shifts in interactions of two dominant shrub species in a semi‐arid coastal sand dune system

Question: How does the interaction between two dominant shrub species in a coastal sand dune community change during their life history? Does this interaction influence their population dynamics? Location: A semiarid coastal sand dune system in southeast Spain. Methods: For 3 years we monitored physiological status, growth and reproductive effort of Juniperus phoenicea and Pistacia lentiscus, the dominant shrub species, growing either alone or in close spatial association. We also recorded adult mortality patterns and characterized seedling survival, soil properties and microclimate conditions beneath canopies and in bare ground. Results and conclusions: There was a strong bi‐directional interaction between the two studied species, with a net balance that changed in sign with increasing plant development. While mature individuals facilitated the establishment of seedlings of both species, adult mortality patterns suggested asymmetric competition at later life stages. The interaction with Pistacia negatively affected growth of juniper and contributed to its high mortality rates, while juniper had almost no effect on mature Pistacia individuals. Physiological data suggested that Pistacia had a competitive advantage over juniper, most likely because of differences in rooting patterns and tolerance to salinity, which may determine the source of water available for each species. Community dynamics are governed by facilitation at the seedling stage and shaped by differences in physiological traits in adult plants. Plant‐plant interactions, which are strongly affected by environmental gradients, are important drivers of community dynamics in this system.     

Spatial and temporal changes in ecosystem carbon pools following juniper encroachment and removal

Proliferation of woody plants is a predominant global land cover change of the past century, particularly in dryland ecosystems. Woody encroachment and its potential impacts (e.g., livestock forage, wildlife habitat, hydrological cycling) have led to widespread brush management. Although woody plants may have substantial impacts on soils, uncertainty remains regarding woody encroachment and brush management influences on carbon (C) pools. Surface C pools (shallow soils and litter) may be particularly dynamic in response to encroachment and brush management. However, we have limited understanding of spatiotemporal patterns of surface C responses or how surface pools respond relative to aboveground C, litter, roots, and deep soil organic C. Spatial variability and lack of basic ecological data in woody-encroached dryland ecosystems present challenges to filling this data gap. We assessed the impact of western juniper (Juniperus occidentalis) encroachment and removal on C pools in a semi-arid sagebrush ecosystem. We used spatially-intensive sampling to create sub-canopy estimates of surface soil C (0–10 cm depth) and litter C pools that consider variation in tree size/age and sub-canopy location for live juniper and around stumps that were cut 7 years prior to sampling. We coupled the present size distribution of junipers with extensive existing allometric information about juniper in this region to estimate how landscape-level C pools would change through time under future management and land cover scenarios. Juniper encroachment and removal leads to substantial changes in C pools. Best-fit models for surface soil and litter C included positive responses to shrub basal diameter and negative responses to increasing relative distance from the bole to dripline. Juniper removal led to a net loss of surface C as a function of large decreases in litter C and small increases in surface soil C. At the landscape scale, deep soil C was the largest C pool (77 Mg C ha^?1), with an apparent lack of sensitivity to management. Overall, encroachment led to substantial increases in C storage over time as juniper size increased (excluding deep soil C, ecosystem C pools increased from 13.5 to 30.2 Mg C ha^?1 with transition from sagebrush-dominated to present encroachment levels). The largest pool of accumulation was juniper aboveground C, with important other pools including juniper roots, litter, and surface soil C. Woody encroachment and subsequent brush management can have substantive impacts on ecosystem C pools, although our data suggest the spatiotemporal patterns of surface C pools need to be properly accounted for to capture C pool responses. Our approach of coupling spatially-intensive surface C information with shrub distribution and allometric data is an effective method for characterizing ecosystem C pools, offering an opportunity for filling in knowledge gaps regarding woody encroachment and brush management impacts on local-to-regional ecosystem C pools.     

Soil C and N responses to woody plant expansion in a mesic grassland

Changes in land management and reductions in fire frequency have contributed to increased cover of woody species in grasslands worldwide. These shifts in plant community composition have the potential to alter ecosystem function, particularly through changes in soil processes and properties. In semi-arid grasslands, the invasion of shrubs and trees is often accompanied by increases in soil resources and more rapid N and C cycling. We assessed the effects of shrub encroachment in a mesic grassland in Kansas (USA) on soil CO₂ flux, extractable inorganic N, and N mineralization beneath shrub communities (Cornus drummondii) and surrounding undisturbed grassland sites. In this study, a shift in plant community composition from grassland to shrubland resulted in a 16% decrease in annual soil CO₂ flux(4.78 kg CO₂ m^–2 year^–1 for shrub dominated sites versus 5.84 kg CO₂ m^–2 year^–1 for grassland sites) with no differences in total soil C or N or inorganic N. There was considerable variability in N mineralization rates within sites, which resulted in no overall difference in cumulative N mineralized during this study (4.09 g N m^–2 for grassland sites and 3.03 g N m^–2 for shrub islands). These results indicate that shrub encroachment into mesic grasslands does not significantly alter N availability (at least initially), but does alter C cycling by decreasing soil CO₂ flux.     

Linking small-scale topography with microclimate,plant species diversity and intra-specific trait variation in an alpine landscape

Background: Small-scale topographic complexity is a characteristic feature of alpine landscapes, with important effects on alpine plant distribution.

Aims: We investigated the links between small-scale topographic complexity and resultant microclimatic heterogeneity, vascular-plant species richness and beta diversity, and realised niche width and trait variation of some target species.

Methods: We recorded temperature and soil moisture within 10 sites (40 m × 40 m) of differing topographic complexity in alpine terrain at Finse, Norway (N 60° 36?, E 7° 33?). Plant species occurrence and traits of target species were recorded in 16 sample plots at each site.

Results: Sites differed significantly in microclimatic heterogeneity, and topographically rough sites were always more heterogeneous than flatter ones. Greater species richness and turnover was associated with greater microclimatic heterogeneity, and rough sites contained 15–55% more species than flatter ones. Plant species had on average wider realised niches when growing at rough sites. Individuals of Bistorta vivipara, but not those of Luzula spicata, tended to exhibit greater phenotypic variation at rough sites.

Conclusions: Rough alpine terrains create small-scale variation in microclimate, promoting species richness and beta diversity. In the event of climate change, small-scale microclimatic heterogeneity might allow plant species to escape from regional climate change by short-distance migration to local micro-refugia. This study suggests that the opportunity for such responses would be greater in topographically complex terrains.     

Woody plant encroachment reduces species richness of herb‐rich woodlands in southern Australia

Abstract Woody plants have been increasing in many woodland and savanna ecosystems owing to land use changes in recent decades. We examined the effects of encroachment by the indigenous shrub Leptospermum scoparium (Myrtaceae) on herb‐rich Eucalyptus camaldulensis woodlands in southern Australia. Species richness and compositional patterns were examined under the canopy of L. scoparium and in surrounding open areas to determine the species most susceptible to structural changes. Richness was significantly lower in areas of moderate to high L. scoparium cover (>15%), suggesting that a threshold shrub cover caused major change in this ecosystem. Shrubs were associated with a significant reduction in above‐ground biomass of the ground‐layer flora and a significant shift in community composition. The few species that were positively associated with high L. scoparium cover were also common in the woodland flora; no new species were recorded under the shrub canopy. Important environmental changes associated with L. scoparium cover were decreased light availability and increased litter cover, which were likely a consequence of encroachment. Leptospermum scoparium cover was also associated with greater surface soil moisture, which may be a consequence of increased shading under the shrub canopy or indicate favourable soil conditions for L. scoparium establishment. Reductions in species richness and abundance of the germinable seed bank were found in soil samples taken from under L. scoparium. With ongoing recruitment of L. scoparium and consequent increases in shrub cover, ground‐layer diversity in these species‐rich woodlands should continue to decline over time.     

Physiological and growth responses of switchgrass (Panicum virgatum L.) in native stands under passive air temperature manipulation

In the central Great Plains of North America, climate change predictions include increases in mean annual temperature of 1.5–5.5 °C by 2100. Ecosystem responses to increased temperatures are likely to be regulated by dominant plant species, such as the potential biofuel species Panicum virgatum (switchgrass) in the tallgrass prairie. To describe the potential physiological and whole‐plant responses of this species to future changes in air temperatures, we used louvered open‐sided chambers (louvered OSC; 1 × 1 m, adjustable height) to passively alter canopy temperature in native stands of P. virgatum growing in tallgrass prairie at varying topographic positions (upland/lowland). The altered temperature treatment decreased daily mean temperatures by 1 °C and maximum temperatures by 4 °C in May and June, lowered daytime stomatal conductance and transpiration, decreased tiller density, increased specific leaf area, and delayed flowering. Among topographic contrasts, aboveground biomass, flowering tiller density, and tiller weight were greater in lowland sites compared to upland sites, with no temperature treatment interactions. Differences in biomass production responded more to topography than the altered temperature treatment, as soil water status varied considerably between topographic positions. These results indicate that while water availability as a function of topography was a strong driver of plant biomass, many leaf‐level physiological processes were responsive to the small decreases in daily mean and maximum temperature, irrespective of landscape position. The varying responses of leaf‐level gas exchange and whole‐plant growth of P. virgatum in native stands to altered air temperature or topographic position illustrate that accurately forecasting yields for P. virgatum in mixed communities will require greater integration of physiological responses to simulated climate change (increased temperature) and resource availability over natural environmental gradients (soil moisture).     

Environmental gradients,plant distribution,and species richness in arctic salt marsh near Prudhoe Bay,Alaska

Spatial patterns of plant cover and species composition in arctic salt marsh and salt affected tundra near Prudhoe Bay, Alaska reflect gradients in elevation, soil conductivity, and soil concentrations of the ions prevalent in seawater. Soil conductivity and soil concentrations of Ca²⁺, Mg²⁺, Na⁺, K⁺, SO₄ ⁼ and Cl^– were significantly related to site elevation, decreasing as elevation increased. Vascular plant species richness increased significantly as soil conductivity and soil ion concentrations decreased, and site elevation increased. Puccinellia phryganodes was the only species present in low elevation sites with low plant cover, high soil conductivity and high soil concentrations of Ca²⁺, Mg²⁺, Na⁺, K⁺, SO₄ ⁼ and Cl^–. Mid-gradient sites were dominated by Carex subspathaceae. Plant cover at these sites was greater than at lower elevation sites, but bare ground was still present. Higher elevation sites had the lowest concentrations of soil ions and the lowest soil conductivities. These sites had little bare ground, contained as many as 16 species, and were dominated by Dupontia fischeri and Eriophorum angustifolium. Ordinations indicated that a complex topographic gradient related most closely to elevation and site distance from the coast best explains variation in the vegetation cover. Irregular deposition along the coastline partially or completely buried three sites in peat or sand up to 20 cm deep. Such rapid changes in plant cover and species composition contributes to the community patch mosaic typical of these marshes. Results suggest an individualistic response of plant species to the environmental gradients in salt marsh and salt affected tundra and are indicative of successional models developed in other marginal arctic environments.     

Carbon dioxide fluxes in a spatially and temporally heterogeneous temperate grassland

Landscape position, grazing, and seasonal variation in precipitation and temperature create spatial and temporal variability in soil processes, and plant biomass and composition in grasslands. However, it is unclear how this variation in plant and soil properties affects carbon dioxide (CO₂) fluxes. The aim of this study is to explore the effect of grazing, topographic position, and seasonal variation in soil moisture and temperature on plant assimilation, shoot and soil respiration, and net ecosystem CO₂ exchange (NEE). Carbon dioxide fluxes, vegetation, and environmental variables were measured once a month inside and outside long-term ungulate exclosures in hilltop (dry) to slope bottom (mesic) grassland throughout the 2004 growing season in Yellowstone National Park. There was no difference in vegetation properties and CO₂ fluxes between the grazed and the ungrazed sites. The spatial and temporal variability in CO₂ fluxes were related to differences in aboveground biomass and total shoot nitrogen content, which were both related to variability in soil moisture. All sites were CO₂ sinks (NEE>0) for all our measurments taken throughout the growing season; but CO₂ fluxes were four- to fivefold higher at sites supporting the most aboveground biomass located at slope bottoms, compared to the sites with low biomass located at hilltops or slopes. The dry sites assimilated more CO₂ per gram aboveground biomass and stored proportionally more of the gross-assimilated CO₂ in the soil, compared to wet sites. These results indicate large spatio-temporal variability of CO₂ fluxes and suggest factors that control the variability in Yellowstone National Park.