Inertia in plant community structure: state changes after cessation of nutrient-enrichment stress

Includes bibliographical references (pages 457-458).Water, nitrogen, and water-plus-nitrogen at levels beyond the range normally experienced by shortgrass steppe communities were applied from 1971 through 1975, plant populations were sampled through 1977, and the results of the experiment were published. Upon revisiting the plots in 1982, we found it apparent that large changes had occurred since 1977. Sampling was re-established in 1982 to follow trajectories of recovery. Our purposes in this paper are to examine how conclusions from this study changed through time, and discuss implications of these changes for monitoring potentially stressed ecosystems. Although productivities increased, dissimilarities in plant species composition at the end of the 5 year of nutrient treatments were not significantly different from controls. Two years after cessation of the treatments exotic "weed" species were increasing in water plus-nitrogen treated communities, and community dissimilarities were diverging in water and water-plus-nitrogen treated communities. Seven years after cessation of treatments all communities were significantly different from controls. Exotics were more than ten times more abundant in water-plus-nitrogen and nitrogen treated communities than they had been2 year post-treatment. A consistent trend in recovery of all treated communities was evident over the next 5 yr. However, the trend towards recovery reversed over the next four consecutive years in the previously water-plus-nitrogen and water treated communities. The four-to-five year cycles in species composition and abundance of exotics towards, and then away from, conditions in undisturbed control communities were not related to weather, but large accumulations of litter suggested biotic regulation. Inertia of existing plant populations, or the tendency to continue to occupy a site when conditions become unfavorable, can mask both future deterioration in ecosystem condition and unstable behavior resulting from environmental stressors. Time lags in initial response means that an ecosystem can pass a threshold leading to transitions to alternate states before it is evident in structural characteristics such as species composition. Global climate change and sulfur and nitrogen oxide pollutants also have the potential to act as enrichment-stressors with initial time lags and/or positive effects and cumulative, subsequent negative effects, rather than as disturbance forces with immediate negative impacts. Sociopolitical systems, however, often require change in biological variables or negative impacts before acting to ameliorate environmental problems. The manner in which conclusions changed at various periods in time, and the potential for time lags in responses of species populations, raises questions about which variables are most useful for detection of stress and how long studies must last to be useful

Carbon dynamics and estimates of primary production by harvest, 14C dilution, and 14C turnover

Includes bibliographical references (page 607).Large plots of native shortgrass steppe were labeled with 14C to assess short-term patterns of carbon allocation and the long-term process of herbivory, death, and decomposition, and to compare estimates of net aboveground, crown, and root primary production using 14C dilution, 14C turnover, and traditional harvest methods. Stabilization of labile 14C via translocation, incorporation into structural tissue, and respiration and exudation required one growing season. Exudation was 17% of plant 14C after stabilization. Estimates of turnover time for leaves, crowns, and roots by 14C turnover were 3, 5, and 8 yr, respectively, yielding estimates of belowground production that were much lower than previously thought. Estimates of aboveground production by 14C turnover were close to those obtained by harvest of peak-standing crop, but lower than reported values obtained by harvest maxima-minima. Estimates of root production by harvest maxima-minima were zero in 2 of 4 yr. 14C turnover appeared to provide reliable estimates of aboveground, crown, and root production. In contrast to reliable estimates by 14C turnover, 14C dilution estimates of root production were anomalous. The anomalous estimates were attributed to a nonuniform labeling of tissue age classes resulting in differential decomposition/herbivory of 14C:12C through time, as well as movement and loss of labile 14C through the first growing season. Isotope-dilution methodologies may be unreliable for any estimate of pool turnover when the labeling period is not as long as pool-turnover time. Problems and biases associated with traditional harvest maxima-minima methods of estimating aboveground primary production are well known, but are greatly exacerbated when the method is used to estimate root production. Estimates of root production by 14C dilution were unrealistic. 14C turnover methodology provided reliable estimates of production in this community

Elevated CO\u3csub\u3e2\u3c/sub\u3e Enhances Productivity and the C/N Ratio of Grasses in the Colorado Shortgrass Steppe

Atmospheric CO2 concentrations have been increasing since the industrial revolution, and are projected to double within this century over today\u27s concentration of 360 µmol mol-1 . This study used six open-top chambers in the Colorado, USA shortgrass steppe to investigate how increasing CO2 will affect productivity and C and N status of indigenous perennial grasses and forbs. From March until October, chambers were placed on two plots in each of the three blocks. In each block, one chamber was assigned an ambient CO2 treatment (~360 µmol mol-1), the other an elevated CO2 treatment (~720 µmol mol-1). Each block also had an unchambered control plot. Growth under elevated CO2 increased above-ground phytomass an average 31% in 1997 and 47% in 1998, with no differences in relative growth responses of C3 and C4 grasses and forbs. Growth in chambers was greater than non-chambered control plots, presumably due to warmer temperatures in chambers and a longer growing season. Shoot N concentrations were reduced 21% and C/N ratios increased 23% in elevated compared to ambient chambers. Variation in aboveground phytomass due to year, CO2 and chamber effects correlated well to % shoot N and C/N ratios, although for both traits different regression lines were required for green plant material (harvested in July) and senescent plant material (harvested in October). Results suggest increased growth and reduced N concentrations in this mixed C3/C4 grassland in an elevated CO2 environment

Long-term ecological research on Colorado Shortgrass Steppe

The SGS-LTER research site was established in 1980 by researchers at Colorado State University as part of a network of long-term research sites within the US LTER Network, supported by the National Science Foundation. Scientists within the Natural Resource Ecology Lab, Department of Forest and Rangeland Stewardship, Department of Soil and Crop Sciences, and Biology Department at CSU, California State Fullerton, USDA Agricultural Research Service, University of Northern Colorado, and the University of Wyoming, among others, have contributed to our understanding of the structure and functions of the shortgrass steppe and other diverse ecosystems across the network while maintaining a common mission and sharing expertise, data and infrastructure.Poster presented at the LTER All Scientists Meeting held in Estes Park, CO on September 10-13, 2012

Nutrients cause grassland biomass to outpace herbivory

Human activities are transforming grassland biomass via changing climate, elemental nutrients, and herbivory. Theory predicts that food-limited herbivores will consume any additional biomass stimulated by nutrient inputs ('consumer-controlled'). Alternatively, nutrient supply is predicted to increase biomass where herbivores alter community composition or are limited by factors other than food ('resource-controlled'). Using an experiment replicated in 58 grasslands spanning six continents, we show that nutrient addition and vertebrate herbivore exclusion each caused sustained increases in aboveground live biomass over a decade, but consumer control was weak. However, at sites with high vertebrate grazing intensity or domestic livestock, herbivores consumed the additional fertilization-induced biomass, supporting the consumer-controlled prediction. Herbivores most effectively reduced the additional live biomass at sites with low precipitation or high ambient soil nitrogen. Overall, these experimental results suggest that grassland biomass will outstrip wild herbivore control as human activities increase elemental nutrient supply, with widespread consequences for grazing and fire risk

Growth and Competitive Effects of Centaurea stoebe Populations in Response to Simulated Nitrogen Deposition

Increased resource availability can promote invasion by exotic plants, raising concerns over the potential effects of global increases in the deposition of nitrogen (N). It is poorly understood why increased N favors exotics over natives. Fast growth may be a general trait of good invaders and these species may have exceptional abilities to increase growth rates in response to N deposition. Additionally, invaders commonly displace locals, and thus may have inherently greater competitive abilities. The mean growth response of Centaurea stoebe to two N levels was significantly greater than that of North American (NA) species. Growth responses to N did not vary among C. stoebe populations or NA species. Without supplemental N, NA species were better competitors than C. stoebe, and C. stoebe populations varied in competitive effects. The competitive effects of C. stoebe populations increased with N whereas the competitive effects of NA species decreased, eliminating the overall competitive advantage demonstrated by NA species in soil without N added. These results suggest that simulated N deposition may enhance C. stoebe invasion through increasing its growth and relative competitive advantage, and also indicate the possibility of local adaptation in competitive effects across the introduced range of an invader

Estimating fine-root production by tree species and understorey functional groups in two contrasting peatland forests

Background and aims Estimation of root-mediated carbon fluxes in forested peatlands is needed for understanding ecosystem functioning and supporting greenhouse gas inventories. Here, we aim to determine the optimal methodology for utilizing ingrowth cores in estimating annual fine-root production (FRP) and its vertical distribution in trees, shrubs and herbs. Methods We used 3-year data obtained with modified ingrowth core method and tested two calculation methods: 'ingrowth-dividing' and `ingrowth-subtracting'. Results The ingrowth-dividing method combined with a 2-year incubation of ingrowth cores can be used for the 'best estimate' of FRP. The FRP in the nutrient-rich fen forest (561 g m(-2)) was more than twice that in the nutrient-poor bog forest (244 g m(-2)). Most FRP occurred in the top 20-cm layer (76-82 %). Tree FRP accounted for 71 % of total FRP in the bog and 94 % in the fen forests, respectively, following the aboveground vegetation patterns; however, in fen forest the proportions of spruce and birch in FRP were higher than their proportions in stand basal area. Conclusions Our methodology may be used to study peatland FRP patterns more widely and will reduce the volume of labour-intensive work, but will benefit from verification with other methods, as is the case in all in situ FRP studies.Peer reviewe

Large grazers modify effects of aboveground–belowground interactions on small-scale plant community composition

Aboveground and belowground organisms influence plant community composition by local interactions, and their scale of impact may vary from millimeters belowground to kilometers aboveground. However, it still poorly understood how large grazers that select their forage on large spatial scales interact with small-scale aboveground–belowground interactions on plant community heterogeneity. Here, we investigate how cattle (Bos taurus) modify the effects of interactions between yellow meadow ants (Lasius flavus) and European brown hares (Lepus europaeus) on the formation of small-scale heterogeneity in vegetation composition. In the absence of cattle, hares selectively foraged on ant mounds, while under combined grazing by hares and cattle, vertebrate grazing pressure was similar on and off mounds. Ant mounds that were grazed by only hares had a different plant community composition compared to their surroundings: the cover of the grazing-intolerant grass Elytrigia atherica was reduced on ant mounds, whereas the relative cover of the more grazing-tolerant and palatable grass Festuca rubra was enhanced. Combined grazing by hares and cattle, resulted in homogenization of plant community composition on and off ant mounds, with high overall cover of F. rubra. We conclude that hares can respond to local ant–soil–vegetation interactions, because they are small, selective herbivores that make their foraging decisions on a local scale. This results in small-scale plant patches on mounds of yellow meadow ants. In the presence of cattle, which are less selective aboveground herbivores, local plant community patterns triggered by small-scale aboveground–belowground interactions can disappear. Therefore, cattle modify the consequences of aboveground–belowground interactions for small-scale plant community composition

Facilitation or Competition? Tree Effects on Grass Biomass across a Precipitation Gradient

Savanna ecosystems are dominated by two distinct plant life forms, grasses and trees, but the interactions between them are poorly understood. Here, we quantified the effects of isolated savanna trees on grass biomass as a function of distance from the base of the tree and tree height, across a precipitation gradient in the Kruger National Park, South Africa. Our results suggest that mean annual precipitation (MAP) mediates the nature of tree-grass interactions in these ecosystems, with the impact of trees on grass biomass shifting qualitatively between 550 and 737 mm MAP. Tree effects on grass biomass were facilitative in drier sites (MAP≤550 mm), with higher grass biomass observed beneath tree canopies than outside. In contrast, at the wettest site (MAP = 737 mm), grass biomass did not differ significantly beneath and outside tree canopies. Within this overall precipitation-driven pattern, tree height had positive effect on sub-canopy grass biomass at some sites, but these effects were weak and not consistent across the rainfall gradient. For a more synthetic understanding of tree-grass interactions in savannas, future studies should focus on isolating the different mechanisms by which trees influence grass biomass, both positively and negatively, and elucidate how their relative strengths change over broad environmental gradients. © 2013 Moustakas et al

Eaten out of house and home:impacts of grazing on ground-dwelling reptiles in Australian grasslands and grassy woodlands

Large mammalian grazers can alter the biotic and abiotic features of their environment through their impacts on vegetation. Grazing at moderate intensity has been recommended for biodiversity conservation. Few studies, however, have empirically tested the benefits of moderate grazing intensity in systems dominated by native grazers. Here we investigated the relationship between (1) density of native eastern grey kangaroos, Macropus giganteus, and grass structure, and (2) grass structure and reptiles (i.e. abundance, richness, diversity and occurrence) across 18 grassland and grassy Eucalyptus woodland properties in south-eastern Australia. There was a strong negative relationship between kangaroo density and grass structure after controlling for tree canopy cover. We therefore used grass structure as a surrogate for grazing intensity. Changes in grazing intensity (i.e. grass structure) significantly affected reptile abundance, reptile species richness, reptile species diversity, and the occurrence of several ground-dwelling reptiles. Reptile abundance, species richness and diversity were highest where grazing intensity was low. Importantly, no species of reptile was more likely to occur at high grazing intensities. Legless lizards (Delma impar, D. inornata) were more likely to be detected in areas subject to moderate grazing intensity, whereas one species (Hemiergis talbingoensis) was less likely to be detected in areas subject to intense grazing and three species (Menetia greyii, Morethia boulengeri, and Lampropholis delicata) did not appear to be affected by grazing intensity. Our data indicate that to maximize reptile abundance, species richness, species diversity, and occurrence of several individual species of reptile, managers will need to subject different areas of the landscape to moderate and low grazing intensities and limit the occurrence and extent of high grazing