A framework for optimizing hydrologic performance of green roof media.

One of the primary functions of green roofs in urban areas is to moderate rainwater runoff, and one of the major impediments to the survival of plants on an extensive green roof (EGR) is a lack of available water during dry periods. Runoff moderation and water storage are both influenced by the composition of the growing media. Here we present a framework for evaluating the hydrologic performance of EGR growing media and also provide hydrologic attribute data for several commonly used EGR media constituents. In this three-phase study, we: 1) measured hydrologic attributes of individual EGR media constituents, 2) predicted attributes of media mixtures using individual constituent data, and 3) tested the seven top-ranking mixtures to evaluate hydrologic performance. Hydrologic attributes included wet weight and water held at maximum retentive capacity, long-term water retention, and hydraulic conductivity. Because perlite was light in weight yet held the greatest amount of water both at its maximum retentive capacity and in the long term, media mixtures dominated by perlite were predicted to have the best overall hydrologic performance. Mixtures dominated by pumice were also predicted to perform relatively well but were heavier. Despite the slightly greater weight and slightly lower performance, pumice may be a preferred alternative to perlite because perlite is a processed constituent with greater estimated embodied energy. Results indicate that performance of mixtures can be adequately predicted using performance of individual constituents for wet weight, water held, and long-term water retention. Hydraulic conductivity was less predictable because the pore volume in mixtures can be unrelated to the pore volume of the individual constituents. The framework presented here can be used to evaluate the performance of other EGR media, and the media attribute data can be used in formulating EGR media mixtures for specific applications. In addition, the attribute data can serve as a benchmark for evaluating other EGR media. Our results underscore the need for standardization of methods for more effective comparisons of EGR substrates, and also reinforce the need to evaluate EGR components using real-world scenarios.

The effects of long-term management on patterns of carbon storage in a northern highbush blueberry production system.

Perennial crops potentially provide a sink for atmospheric carbon. However, there is a poor understanding of how perennial crops differ in their carbon allocation patterns, and few studies have tested how agronomic practices such as fertilization influence long-term patterns of carbon allocation in actual production systems. In this study, we report results of a long-term field experiment that tested the individual and combined effects of organic matter incorporation and nitrogen fertilization on carbon allocation. The mature (nine-year-old) blueberry plants in this study had an average standing carbon stock of 1147gCm-2 and average annual Net Primary Production (NPP) of 523gCm-2yr-1, values that are similar to those reported for other woody crops. Forty-four percent of blueberry annual NPP was sequestered in persistent biomass, 19% was exported as harvested fruit, and 37% entered the detrital pathway. Nitrogen applied at rates typical for blueberry production throughout the span of the study had no significant effect on total plant or soil C. However, pre-planting organic matter incorporation and periodic mulching with sawdust significantly increased both soil organic matter and soil C. Pre-planting organic matter incorporation also increased total standing plant C nine years later at maturity. At the field scale, we estimate that fields receiving pre-planting organic matter incorporation would have 4.8% (4.5Mgha-1) more standing C relative to non-amended fields, although the difference is within the range of uncertainty of the estimated values. These results suggest that blueberry production can provide a valuable medium-term carbon store that is comparable in magnitude to that of temperate tree crops, but overall carbon budgets are influenced by management practices over the first decade after planting.

Plant species' origin predicts dominance and response to nutrient enrichment and herbivores in global grasslands.

Exotic species dominate many communities; however the functional significance of species' biogeographic origin remains highly contentious. This debate is fuelled in part by the lack of globally replicated, systematic data assessing the relationship between species provenance, function and response to perturbations. We examined the abundance of native and exotic plant species at 64 grasslands in 13 countries, and at a subset of the sites we experimentally tested native and exotic species responses to two fundamental drivers of invasion, mineral nutrient supplies and vertebrate herbivory. Exotic species are six times more likely to dominate communities than native species. Furthermore, while experimental nutrient addition increases the cover and richness of exotic species, nutrients decrease native diversity and cover. Native and exotic species also differ in their response to vertebrate consumer exclusion. These results suggest that species origin has functional significance, and that eutrophication will lead to increased exotic dominance in grasslands.

Predicting invasion in grassland ecosystems: is exotic dominance the real embarrassment of richness?

Invasions have increased the size of regional species pools, but are typically assumed to reduce native diversity. However, global-scale tests of this assumption have been elusive because of the focus on exotic species richness, rather than relative abundance. This is problematic because low invader richness can indicate invasion resistance by the native community or, alternatively, dominance by a single exotic species. Here, we used a globally replicated study to quantify relationships between exotic richness and abundance in grass-dominated ecosystems in 13 countries on six continents, ranging from salt marshes to alpine tundra. We tested effects of human land use, native community diversity, herbivore pressure, and nutrient limitation on exotic plant dominance. Despite its widespread use, exotic richness was a poor proxy for exotic dominance at low exotic richness, because sites that contained few exotic species ranged from relatively pristine (low exotic richness and cover) to almost completely exotic-dominated ones (low exotic richness but high exotic cover). Both exotic cover and richness were predicted by native plant diversity (native grass richness) and land use (distance to cultivation). Although climate was important for predicting both exotic cover and richness, climatic factors predicting cover (precipitation variability) differed from those predicting richness (maximum temperature and mean temperature in the wettest quarter). Herbivory and nutrient limitation did not predict exotic richness or cover. Exotic dominance was greatest in areas with low native grass richness at the site- or regional-scale. Although this could reflect native grass displacement, a lack of biotic resistance is a more likely explanation, given that grasses comprise the most aggressive invaders. These findings underscore the need to move beyond richness as a surrogate for the extent of invasion, because this metric confounds monodominance with invasion resistance. Monitoring species' relative abundance will more rapidly advance our understanding of invasions.

Productivity is a poor predictor of plant species richness.

For more than 30 years, the relationship between net primary productivity and species richness has generated intense debate in ecology about the processes regulating local diversity. The original view, which is still widely accepted, holds that the relationship is hump-shaped, with richness first rising and then declining with increasing productivity. Although recent meta-analyses questioned the generality of hump-shaped patterns, these syntheses have been criticized for failing to account for methodological differences among studies. We addressed such concerns by conducting standardized sampling in 48 herbaceous-dominated plant communities on five continents. We found no clear relationship between productivity and fine-scale (meters(-2)) richness within sites, within regions, or across the globe. Ecologists should focus on fresh, mechanistic approaches to understanding the multivariate links between productivity and richness.

Nitrogen inputs promote the spread of an invasive marsh grass.

Excess nutrient loading and large-scale invasion by nonnatives are two of the most pervasive and damaging threats to the biotic and economic integrity of our estuaries. Individually, these are potent forces, but it is important to consider their interactive impacts as well. In this study we investigated the potential limitation of a nonnative intertidal grass, Spartina alterniflora, by nitrogen (N) in estuaries of the western United States. Nitrogen fertilization experiments were conducted in three mud-flat habitats invaded by S. alterniflora in Willapa Bay, Washington, USA, that differed in sediment N. We carried out parallel experiments in San Francisco Bay, California, USA, in three habitats invaded by hybrid Spartina (S. alterniflora x S. foliosa), in previously unvegetated mud flat, and in native S. foliosa or Salicornia virginica marshes. We found similar aboveground biomass and growth rates between habitats and estuaries, but end-of-season belowground biomass was nearly five times greater in San Francisco Bay than in Willapa Bay. In Willapa Bay, aboveground biomass was significantly correlated with sediment N content. Addition of N significantly increased aboveground biomass, stem density, and the rate of spread into uninvaded habitat (as new stems per day) in virtually all habitats in both estuaries. Belowground biomass increased in Willapa Bay only, suggesting that belowground biomass is not N limited in San Francisco Bay due to species differences, N availability, or a latitudinal difference in the response of Spartina to N additions. The relative impact of added N was greater in Willapa Bay, the estuary with lower N inputs from the watershed, than in San Francisco Bay, a highly eutrophic estuary. Nitrogen fertilization also altered the competitive interaction between hybrid Spartina and Salicornia virginica in San Francisco Bay by increasing the density and biomass of the invader and decreasing the density of the native. There was no significant effect of N on the native, Spartina foliosa. Our results indicate that excess N loading to these ecosystems enhances the vulnerability of intertidal habitats to rapid invasion by nonnative Spartina sp.

Invasion in a heterogeneous world: resistance, coexistence or hostile takeover?

We review and synthesize recent developments in the study of the invasion of communities in heterogeneous environments, considering both the invasibility of the community and impacts to the community. We consider both empirical and theoretical studies. For each of three major kinds of environmental heterogeneity (temporal, spatial and invader-driven), we find evidence that heterogeneity is critical to the invasibility of the community, the rate of spread, and the impacts on the community following invasion. We propose an environmental heterogeneity hypothesis of invasions, whereby heterogeneity both increases invasion success and reduces the impact to native species in the community, because it promotes invasion and coexistence mechanisms that are not possible in homogeneous environments. This hypothesis could help to explain recent findings that diversity is often increased as a result of biological invasions. It could also explain the scale dependence of the diversity-invasibility relationship. Despite the undoubted importance of heterogeneity to the invasion of communities, it has been studied remarkably little and new research is needed that simultaneously considers invasion, environmental heterogeneity and community characteristics. As a young field, there is an unrivalled opportunity for theoreticians and experimenters to work together to build a tractable theory informed by data.

Ecosystem engineering in space and time.

The ecosystem engineering concept focuses on how organisms physically change the abiotic environment and how this feeds back to the biota. While the concept was formally introduced a little more than 10 years ago, the underpinning of the concept can be traced back to more than a century to the early work of Darwin. The formal application of the idea is yielding new insights into the role of species in ecosystems and many other areas of basic and applied ecology. Here we focus on how temporal, spatial and organizational scales usefully inform the roles played by ecosystem engineers and their incorporation into broader ecological contexts. Two particular, distinguishing features of ecosystem engineers are that they affect the physical space in which other species live and their direct effects can last longer than the lifetime of the organism--engineering can in essence outlive the engineer. Together, these factors identify critical considerations that need to be included in models, experimental and observational work. The ecosystem engineering concept holds particular promise in the area of ecological applications, where influence over abiotic variables and their consequent effects on biotic communities may facilitate ecological restoration and counterbalance anthropogenic influences.

Using ecosystem engineers to restore ecological systems.

Ecosystem engineers affect other organisms by creating, modifying, maintaining or destroying habitats. Despite widespread recognition of these often important effects, the ecosystem engineering concept has yet to be widely used in ecological applications. Here, we present a conceptual framework that shows how consideration of ecosystem engineers can be used to assess the likelihood of restoration of a system to a desired state, the type of changes necessary for successful restoration and how restoration efforts can be most effectively partitioned between direct human intervention and natural ecosystem engineers.

Spatially variable propagule pressure and herbivory influence invasion of chaparral shrubland by an exotic grass.

Although numerous studies have identified mechanisms that either resist or facilitate biological invasions, few studies have explicitly tested how resisting and facilitating mechanisms interact to drive invasion success. In California, USA, undisturbed Mediterranean-type shrublands have resisted invasion by the perennial tussock grass Cortaderia jubata. In some cases, however, this resistance has been spectacularly breached even in the absence of large-scale disturbance. I tested the hypothesis that these invasions are facilitated by local reductions in the strength of biotic resistance. I evaluated invasive success using C. jubata seed and seedling additions at different microhabitats: the edge of a chaparral stand, under shrub canopy at different distances from the stand edge, and in canopy gaps within the stand. When left exposed to mammalian herbivores, seedling survivorship decreased sharply from nearly 40% on the stand edge to zero just 10 m into the stand. When transplants were protected from herbivory, however, distance from the edge had no significant influence on transplant survivorship. Seedling emergence was also greater on the edge and in canopy gaps than under the canopy, but these differences were not caused by differences in herbivory. The flux of invasive propagules reaching the soil surface was immense and greater along the edge and within gaps than under the stand canopy. Mirroring these patterns, naturally occurring seedling abundance declined dramatically with distance from the stand edge, and seedlings were far more common within stand gaps than would be expected given gap frequency within the stand. Despite strong biotic resistance to invasion within the stand, the cover of C. jubata has increased 20% over the last 9 years. These results suggest that the relative amount of susceptible edge habitat and the supply of invasive propagules can facilitate invasion even in the face of strong local biotic resistance.

Pollen limitation causes an Allee effect in a wind-pollinated invasive grass (Spartina alterniflora).

It is usually assumed that pollen availability does not limit reproduction in wind-pollinated plants. Little evidence either supporting or contradicting this assumption exists, despite the importance of seed production to population persistence and growth. We investigated the role of pollen limitation in an invasive estuarine grass (Spartina alterniflora), with a manipulative pollen supplementation and exclusion experiment in areas of high population density and at the low-density leading edge of the invasion. We also quantified pollen deposition rates on stigmas and pollen traps along a windward to leeward gradient. We found pollen impoverishment at the low-density leading edge of a large invasion, causing an 8-fold reduction in seed set. We found 9-fold more pollen on stigmas of high-density plants than on those of low-density plants. Pollen deposition rates on stigmas and traps did not increase downwind of low-density plants but did increase downwind of high-density plants and dropped off precipitously across a gap that lacked pollen donors. The delay of appreciable numbers of seed caused by pollen limitation persists for decades until vegetative growth coalesces plants into continuous meadows, and this Allee effect has slowed the rate of spread of the invasion.