Jonathan D. Bakker

Contingency of grassland restoration on year, site, and competition from introduced grasses

Semiarid ecosystems such as grasslands are characterized by high temporal variability in abiotic factors, which has led to suggestions that management actions may be more effective in some years than others. Here we examine this hypothesis in the context of grassland restoration, which faces two major obstacles: the contingency of native grass establishment on unpredictable precipitation, and competition from introduced species. We established replicated restoration experiments over three years at two sites in the northern Great Plains in order to examine the extent to which the success of several restoration strategies varied between sites and among years. We worked in 50-yr-old stands of crested wheatgrass (Agropyron cristatum), an introduced perennial grass that has been planted on >10 × 106 ha in western North America. Establishment of native grasses was highly contingent on local conditions, varying fourfold among years and threefold between sites. Survivorship also varied greatly and increased signi...

Integrative modelling reveals mechanisms linking productivity and plant species richness

How ecosystem productivity and species richness are interrelated is one of the most debated subjects in the history of ecology. Decades of intensive study have yet to discern the actual mechanisms behind observed global patterns. Here, by integrating the predictions from multiple theories into a single model and using data from 1,126 grassland plots spanning five continents, we detect the clear signals of numerous underlying mechanisms linking productivity and richness. We find that an integrative model has substantially higher explanatory power than traditional bivariate analyses. In addition, the specific results unveil several surprising findings that conflict with classical models. These include the isolation of a strong and consistent enhancement of productivity by richness, an effect in striking contrast with superficial data patterns. Also revealed is a consistent importance of competition across the full range of productivity values, in direct conflict with some (but not all) proposed models. The promotion of local richness by macroecological gradients in climatic favourability, generally seen as a competing hypothesis, is also found to be important in our analysis. The results demonstrate that an integrative modelling approach leads to a major advance in our ability to discern the underlying processes operating in ecological systems.

Herbaceous Vegetation Responses (1992–2004) to Restoration Treatments in a Ponderosa Pine Forest

Abstract Ecological restoration treatments are widely applied in southwestern ponderosa pine forests to convert them to an open canopy structure similar to that found at the time of Euro-American settlement. An experiment was initiated in northern Arizona in 1994 to evaluate long-term ecosystem responses to 3 restoration treatments: 1) thinning from below (thinning), 2) thinning from below plus forest floor manipulation with periodic prescribed burning (composite), and 3) an untreated control. Results focus on total herbaceous and functional-group standing crop response to these restoration treatments. Pretreatment data were collected in 1992 and posttreatment responses were measured from 1994 through 2004. Total herbaceous standing crop was significantly higher on the 2 treated areas than on the control over the entire posttreatment period, but did not differ between the thinning and composite treatments. Plant functional groups responded differently to treatments and to drought. In general, the graminoid standing crop responded within several years after the initial treatments and continued to increase through time, until a series of severe droughts reduced standing crop to pretreatment levels. C3 graminoids dominated the standing-crop response, of which bottlebrush squirreltail (Elymus elymoides (Raf.) Swezey ssp. elymoides) was the primary contributor. C4 graminoids had a minimal response to restoration treatments, possibly because they were less abundant before the experiment began or because they were adversely affected by autumn burning. Legumes and forbs exhibited a 4–5 year lag before responding to the thinning and composite treatments. Annual and biennial plants showed a large biomass increase approximately 5 years after implementation of the composite treatment. The restoration goal of optimizing herbaceous standing crop must be weighed against the competing goals of increasing the abundance of specific functional groups, increasing biodiversity or rare plants, and managing invasive plant species.

Regional Contingencies in the Relationship between Aboveground Biomass and Litter in the World's Grasslands

Based on regional-scale studies, aboveground production and litter decomposition are thought to positively covary, because they are driven by shared biotic and climatic factors. Until now we have been unable to test whether production and decomposition are generally coupled across climatically dissimilar regions, because we lacked replicated data collected within a single vegetation type across multiple regions, obfuscating the drivers and generality of the association between production and decomposition. Furthermore, our understanding of the relationships between production and decomposition rests heavily on separate meta-analyses of each response, because no studies have simultaneously measured production and the accumulation or decomposition of litter using consistent methods at globally relevant scales. Here, we use a multi-country grassland dataset collected using a standardized protocol to show that live plant biomass (an estimate of aboveground net primary production) and litter disappearance (represented by mass loss of aboveground litter) do not strongly covary. Live biomass and litter disappearance varied at different spatial scales. There was substantial variation in live biomass among continents, sites and plots whereas among continent differences accounted for most of the variation in litter disappearance rates. Although there were strong associations among aboveground biomass, litter disappearance and climatic factors in some regions (e.g. U.S. Great Plains), these relationships were inconsistent within and among the regions represented by this study. These results highlight the importance of replication among regions and continents when characterizing the correlations between ecosystem processes and interpreting their global-scale implications for carbon flux. We must exercise caution in parameterizing litter decomposition and aboveground production in future regional and global carbon models as their relationship is complex.

Grassland productivity limited by multiple nutrients

Terrestrial ecosystem productivity is widely accepted to be nutrient limited1. Although nitrogen (N) is deemed a key determinant of aboveground net primary production (ANPP)2,3, the prevalence of co-limitation by N and phosphorus (P) is increasingly recognized4–8. However, the extent to which terrestrial productivity is co-limited by nutrients other than N and P has remained unclear. Here, we report results from a standardized factorial nutrient addition experiment, in which we added N, P and potassium (K) combined with a selection of micronutrients (K+μ), alone or in concert, to 42 grassland sites spanning five continents, and monitored ANPP. Nutrient availability limited productivity at 31 of the 42 grassland sites. And pairwise combinations of N, P, and K+μ co-limited ANPP at 29 of the sites. Nitrogen limitation peaked in cool, high latitude sites. Our findings highlight the importance of less studied nutrients, such as K and micronutrients, for grassland productivity, and point to significant variations in the type and degree of nutrient limitation. We suggest that multiple-nutrient constraints must be considered when assessing the ecosystem-scale consequences of nutrient enrichment.

Competitive abilities of introduced and native grasses

Differencesin competitive ability may explain the maintenance of existing plantpopulationsand the invasion of new areas by plant species. We used field experiments toexamine the competitive responses of Agropyron cristatum(L.) Gaertn., an introduced C3 grass, and Boutelouagracilis (HBK.) Lag., a native C4 grass, and thecompetitive effects of Agropyron-dominated vegetation andsuccessional prairie. We also tested whether the outcome of competitiveinteractions varied with water availability. In each vegetation type,transplants of each species were grown under two levels of competition(presenceor absence of neighboring vegetation) and three levels of water availability(high, medium, or low). Transplant survival, growth, and biomass allocationpatterns were measured. Water availability had no effect on the measuredvariables, suggesting that both species were limited by another resource.Growthrates were affected more by competition, while survival and root: shoot ratiowere affected more by transplant species identity. In the successional prairie,neighboring vegetation suppressed the growth of Agropyrontransplants less than that of Bouteloua transplants,suggesting that Agropyron has a stronger ability to resistcompetitive suppression in that vegetation type. The spread ofAgropyron into surrounding vegetation may relate to itsability to resist competitive suppression. In theAgropyron-dominated vegetation, neighboring vegetationsuppressed the growth of both species by the same extent. However, competitionaccounted for more variation in transplant growth inAgropyron-dominated vegetation than in successionalprairie, suggesting that Agropyron has strong competitiveeffects which hinder plant growth and prevent other species from establishinginAgropyron fields.

Life-history constraints in grassland plant species: a growth-defence trade-off is the norm

Plant growth can be limited by resource acquisition and defence against consumers, leading to contrasting trade-off possibilities. The competition-defence hypothesis posits a trade-off between competitive ability and defence against enemies (e.g. herbivores and pathogens). The growth-defence hypothesis suggests that strong competitors for nutrients are also defended against enemies, at a cost to growth rate. We tested these hypotheses using observations of 706 plant populations of over 500 species before and following identical fertilisation and fencing treatments at 39 grassland sites worldwide. Strong positive covariance in species responses to both treatments provided support for a growth-defence trade-off: populations that increased with the removal of nutrient limitation (poor competitors) also increased following removal of consumers. This result held globally across 4 years within plant life-history groups and within the majority of individual sites. Thus, a growth-defence trade-off appears to be the norm, and mechanisms maintaining grassland biodiversity may operate within this constraint.

Examining conifer canopy structural complexity across forest ages and elevations with LiDAR data

LiDAR measurements of canopy structure can be used to classify forest stands into structural stages to study spatial patterns of canopy structure, identify habitat, or plan management actions. A key assumption in this process is that differences in canopy structure based on forest age and elevation are consistent with predictions from models of stand de- velopment. Three LiDAR metrics (95th percentile height, rumple, and canopy density) were computed for 59 secondary and 35 primary forest plots in the Pacific Northwest, USA. Hierarchical clustering identified two precanopy closure classes, two low-complexity postcanopy closure classes, and four high-complexity postcanopy closure classes. Forest de- velopment models suggest that secondary plots should be characterized by low-complexity classes and primary plots char- acterized by high-complexity classes. While the most and least complex classes largely confirmed this relationship, intermediate-complexity classes were unexpectedly composed of both secondary and primary forest types. Complexity classes were not associated with elevation, except that primary Tsuga mertensiana (Bong.) Carriere (mountain hemlock) plots were complex. These results suggest that canopy structure does not develop in a linear fashion and emphasize the im- portance of measuring structural conditions rather than relying on development models to estimate structural complexity across forested landscapes.

CONTROLS ON VEGETATION STRUCTURE IN SOUTHWESTERN PONDEROSA PINE FORESTS, 1941 AND 2004

Long-term studies can broaden our ecological understanding and are particularly important when examining contingent effects that involve changes to dominance by long-lived species. Such a change occurred during the last century in Southwestern (USA) ponderosa pine (Pinus ponderosa) forests. We used five livestock grazing exclosures established in 1912 to quantify vegetation structure in 1941 and 2004. Our objectives were to (1) assess the effects of historical livestock grazing on overstory structure and age distribution, (2) assess the effects of recent livestock grazing and overstory on understory vegetation, and (3) quantify and explain changes in understory vegetation between 1941 and 2004. In 1941, canopy cover of tree regeneration was significantly higher inside exclosures. In 2004, total tree canopy cover was twice as high, density was three times higher, trees were smaller, and total basal area was 40% higher inside exclosures. Understory species density, herbaceous plant density, and herbaceous cover were negatively correlated with overstory vegetation in both years. Most understory variables did not differ between grazing treatments in 1941 but were lower inside exclosures in 2004. Differences between grazing treatments disappeared once overstory effects were accounted for, indicating that they were due to the differential overstory response to historical livestock grazing practices. Between 1941 and 2004, species density declined by 34%, herbaceous plant density by 37%, shrub cover by 69%, total herbaceous cover by 59%, graminoid cover by 39%, and forb cover by 82%. However, these variables did not differ between grazing treatments or years once overstory effects were accounted for, indicating that the declines were driven by the increased dominance of the overstory during this period. Our results demonstrate that historical livestock grazing practices are an aspect of land-use history that can affect ecosystem development. Grazing history must be considered when extrapolating results from one site to another. In addition, the understory vegetation was more strongly controlled by the ponderosa pine overstory than by recent livestock grazing or by temporal dynamics, indicating that overstory effects must be accounted for when examining understory responses in this ecosystem.

SEMIARID OLD‐FIELD RESTORATION: IS NEIGHBOR CONTROL NEEDED?

Restoration practice suggests that neighbor control is essential in semiarid grasslands, but ecological theory predicts that neighbor effects are relatively small in young fields. We investigated the effectiveness of neighbor control (mowing and herbicide) for establishing native grasses in a recently abandoned field in southwestern Saskatchewan, Canada. We also examined its interactions with common restoration techniques, such as mulching (straw and sawdust) and contrasting sowing methods (drilling, and broadcasting cleaned seeds, cleaning remainders, and native hay). The experiment was repeated over three years to examine the effect of weather. Neighbor control had no effect on establishment and rarely any effect on first-year survival. This contrasted with significant effects of neighbor control on community and ecosystem-level variables (species richness, water, N). The lack of neighbor effects is concordant with theory which predicts low competition intensity from ruderal annuals. Establishment in seeded plots varied two-fold among years in drilled plots and 50-fold in broadcast plots, and it was lowest during a cool, dry summer. Thus, variables beyond human control are a major factor determining restoration success. Overall, broadcasting was as effective as drilling. The highest long-term establishment of native species was produced by broadcasting cleaning remainders. Almost no seedlings emerged from plots supplied with native hay. Straw mulch increased soil moisture and available N, and sawdust decreased N, but neither had any long-term effect on native grasses. Our results suggest that restorations of semiarid old fields should focus less on neighbor control and more on strategies for exploiting suitable years for germination, either by monitoring soil moisture or through repeated seeding.