Jeremy T. Lundholm

Green Roofs as Urban Ecosystems: Ecological Structures, Functions, and Services

ABSTRACT Green roofs (roofs with a vegetated surface and substrate) provide ecosystem services in urban areas, including improved storm-water management, better regulation of building temperatures, reduced urban heat-island effects, and increased urban wildlife habitat. This article reviews the evidence for these benefits and examines the biotic and abiotic components that contribute to overall ecosystem services. We emphasize the potential for improving green-roof function by understanding the interactions between its ecosystem elements, especially the relationships among growing media, soil biota, and vegetation, and the interactions between community structure and ecosystem functioning. Further research into green-roof technology should assess the efficacy of green roofs compared to other technologies with similar ends, and ultimately focus on estimates of aggregate benefits at landscape scales and on more holistic cost-benefit analyses.

Plant species and functional group combinations affect green roof ecosystem functions.

Background Green roofs perform ecosystem services such as summer roof temperature reduction and stormwater capture that directly contribute to lower building energy use and potential economic savings. These services are in turn related to ecosystem functions performed by the vegetation layer such as radiation reflection and transpiration, but little work has examined the role of plant species composition and diversity in improving these functions. Methodology/Principal Findings We used a replicated modular extensive (shallow growing- medium) green roof system planted with monocultures or mixtures containing one, three or five life-forms, to quantify two ecosystem services: summer roof cooling and water capture. We also measured the related ecosystem properties/processes of albedo, evapotranspiration, and the mean and temporal variability of aboveground biomass over four months. Mixtures containing three or five life-form groups, simultaneously optimized several green roof ecosystem functions, outperforming monocultures and single life-form groups, but there was much variation in performance depending on which life-forms were present in the three life-form mixtures. Some mixtures outperformed the best monocultures for water capture, evapotranspiration, and an index combining both water capture and temperature reductions. Combinations of tall forbs, grasses and succulents simultaneously optimized a range of ecosystem performance measures, thus the main benefit of including all three groups was not to maximize any single process but to perform a variety of functions well. Conclusions/Significance Ecosystem services from green roofs can be improved by planting certain life-form groups in combination, directly contributing to climate change mitigation and adaptation strategies. The strong performance by certain mixtures of life-forms, especially tall forbs, grasses and succulents, warrants further investigation into niche complementarity or facilitation as mechanisms governing biodiversity-ecosystem functioning relationships in green roof ecosystems.

Insect species composition and diversity on intensive green roofs and adjacent level-ground habitats

While it is expected that green roofs support a wider variety of insects compared with conventional roof surfaces, few studies have quantified insect diversity on green roofs. Even fewer have attempted to determine whether green roofs can support insect communities comparable to level-ground urban habitats. In this study, insect richness, abundance and diversity indices were compared between five pairs of intensive green roofs and adjacent ground-level habitat patches in downtown Halifax, Nova Scotia. Pitfall traps were set at each site, collected bi-weekly between May-October 2009 and captured insects were identified to morphospecies (except where taxonomic expertise was available). No significant differences in richness, abundance or any of the indices (S, H’, Evar) were detected in analysis, which included plant species richness, site area and sampling effort as covariables. However, richness and abundance tended to be greater at ground level for all orders (except Heteroptera), and diversity appeared to increase away from the downtown core. Insect composition differed slightly between green roof and ground-level sites; only 17 species were collected from a single site type in numbers greater than five specimens. Nevertheless, a wide variety of insects, including many uncommon species were collected from green roofs, supporting the idea that these habitats can contribute to sustaining biodiversity in cities.

FORUM: Do green roofs help urban biodiversity conservation?

Summary Green roofs are novel ecosystems that are increasingly common in cities. While their hydrologic and energy saving benefits are well-established, green roofs have also been proposed as having significant value for conserving biodiversity. We evaluate six hypotheses that describe the purported biodiversity conservation benefits of green roofs. Green roofs largely support generalist species particularly insects, but their conservation value for rare taxa, and other taxonomic groups especially vertebrates, is poorly documented. Further, their ability to replicate biotic communities in the context of ecological restoration is largely untested, as is their potential to connect ground-level habitats. Synthesis and applications. Given the evidence, green roof proponents should use restraint in claiming conservation benefits and it is premature for policymakers to consider green roofs equivalent to ground-level urban habitats. Ecologists need to work with the industry to evaluate green roof biodiversity and help design green roofs based on ecological principles to maximize biodiversity gains.

Habitat origins and microhabitat preferences of urban plant species

Urban vegetation is commonly described as dominated by weedy species that are adapted to human disturbance. In this study, we determined the original (pre-agriculture) habitats of urban plant species sampled quantitatively in the spontaneous vegetation of a university campus in Halifax, Nova Scotia (eastern Canada). We sampled 11 distinct patch types corresponding to different built forms. Differences in vegetation among patch types were related in part to environmental variables such as soil moisture and light availability. The urban vegetation was dominated by species from rocky habitats such as cliffs and talus slopes, with lesser representation from grassland and floodplain habitats. When compared to a null model of species origins based on the global area and species richness of different original habitats, species from rock outcrops and grassland habitats were overrepresented in the urban vegetation. These results contrast with the received view that cities represented highly “unnatural” ecosystems: built form appears to replicate the habitat templates required by rock outcrop species. Urban ecological theory should incorporate the replication of habitat analogs by built forms in addition to the creation of ecologically novel habitats.

Green roof plant species diversity improves ecosystem multifunctionality

Summary Constructed ecosystems such as green roofs often contain monocultures or low-diversity plant communities, but adding more plant species to these systems can increase ecosystem service provisioning. Mixture advantage, when species-rich treatments outperform the best monocultures, is desirable in constructed ecosystems due to the cost of increasing diversity. However, there have not been any studies in constructed ecosystems that have quantitatively compared mixtures with the best monocultures for multifunctionality, and there have been few studies that have examined how provision of ecosystem services changes over time as plant communities develop. In a green roof system, I predicted (i) that the mixture advantage would be stronger for ecosystem multifunctionality than for single ecosystem functions and (ii) that ecosystem service provisioning and complementarity in above-ground biomass would increase over time. Fifteen monocultures of plant species from five life-form groups (succulents, tall forbs, dwarf shrubs, creeping forbs, grasses) were compared with three-species mixtures of the same life-form and mixtures of species from three and five different life-forms in a modular green roof system. Indicators of ecosystem services including above-ground production, thermal regulation, stormwater retention, nutrient uptake and carbon sequestration and two indices of ecosystem multifunctionality were compared. Canopy density increased over time while substrate temperature decreased, suggesting higher provisioning of valuable ecosystem services. For single services, the positive relationship between planted species richness and ecosystem service grew stronger over time, but was consistently strong over time for multifunctionality. Quantile regression indicated a weak mixture advantage for several services including both multifunctionality indices. While the effects were small, different species optimized different functions, thus multifunctioning is enhanced in more diverse mixtures by combining species that maximize different functions. Tripartite partitioning of canopy density showed that overyielding and trait-independent complementarity fluctuated between years in response to shifts in species abundances, but dominance and trait-dependent complementarity increased over time. Synthesis and applications. This study provides the first evidence in a constructed ecosystem that mixtures can outperform the best monocultures for multiple ecosystem services. Mixtures of plant life-forms can improve green roof performance. The biodiversity–ecosystem function relationships observed in natural ecosystems can also occur in novel and highly simplified engineered ecosystems.

Performance of dryland and wetland plant species on extensive green roofs

BACKGROUND AND AIMS Green roofs are constructed ecosystems where plants perform valuable services, ameliorating the urban environment through roof temperature reductions and stormwater interception. Plant species differ in functional characteristics that alter ecosystem properties. Plant performance research on extensive green roofs has so far indicated that species adapted to dry conditions perform optimally. However, in moist, humid climates, species typical of wetter soils might have advantages over dryland species. In this study, survival, growth and the performance of thermal and stormwater capture functions of three pairs of dryland and wetland plant species were quantified using an extensive modular green roof system. METHODS Seedlings of all six species were germinated in a greenhouse and planted into green roof modules with 6 cm of growing medium. There were 34 treatments consisting of each species in monoculture and all combinations of wet- and dryland species in a randomized block design. Performance measures were survival, vegetation cover and roof surface temperature recorded for each module over two growing seasons, water loss (an estimate of evapotranspiration) in 2007, and albedo and water capture in 2008. KEY RESULTS Over two seasons, dryland plants performed better than wetland plants, and increasing the number of dryland species in mixtures tended to improve functioning, although there was no clear effect of species or habitat group diversity. All species had survival rates >75 % after the first winter; however, dryland species had much greater cover, an important indicator of green roof performance. Sibbaldiopsis tridentata was the top performing species in monoculture, and was included in the best treatments. CONCLUSIONS Although dryland species outperformed wetland species, planting extensive green roofs with both groups decreased performance only slightly, while increasing diversity and possibly habitat value. This study provides further evidence that plant composition and diversity can influence green roof functions.

Ecosystem services provided by urban spontaneous vegetation

Spontaneous vegetation colonizes large areas in and around cities. These unmanaged areas are considered to have low economic value or indicate dereliction, but recent research suggests that these can contribute valuable ecosystem services. This study evaluates indicators of ecosystem services in three habitats: urban spontaneous vegetation (USV), managed lawns, and semi-natural urban forest, in Halifax, Nova Scotia. USV had higher indicator values for habitat provisioning (plant species diversity, invertebrate abundance and taxonomic diversity) than the other habitats. Indicators of climatic regulatory services (albedo and leaf area index) in USV were similar to those in lawn habitats. Organic carbon content of the soils, an indicator of carbon storage, was lowest in USV but only marginally lower than in lawns. Standing biomass, an indicator of production services, was lowest in USV but lawn production may have been overestimated. While USV sites are usually transitory components of the urban landscape, they deserve further consideration due to their provision of ecosystem services, in some cases to a greater extent than conventionally valued urban habitats.

Species richness, abundance, rarity and environmental gradients in coastal barren vegetation

Coastal barrens in Nova Scotia are heathlands characterised by short, predominantly ericaceous vegetation, sparse tree cover, exposed bedrock, pockets of Sphagnum bog, and stressful climatic conditions. Although coastal barrens are prominent in the physical and cultural landscape, they are largely unprotected. We selected six barrens along the Atlantic coast, and surveyed 20 1-m2 plots at each barren for vascular plants, macrolichens, mosses and environmental factors. We recorded 173 species (105 vascular, 41 macrolichen, 27 moss), including six provincially rare vascular species found predominantly in nearshore areas with high levels of substrate salt and nutrients, variable substrate depth, and short vegetation. Although vascular plant and moss richness were similarly correlated with vegetation height, substrate depth, organic matter content, and rock exposure, there were no clear correlations between vascular plant, macrolichen and moss richness across all sites. Vascular plant rarity and species richness were not correlated, but had inverse relationships with key environmental gradients. Tailoring conservation efforts to protect areas of high richness may thus mean that rare species are missed, and vice versa. Ordination and ANOSIM show that barrens vegetation differs widely among sites; therefore, protecting any singular coastal barren will not protect the entire range of vegetation communities and species in this heathland type. Conservation planning should emphasize protecting environmental gradients correlated with richness, rarity and plant community structure, including substrate depth and moisture, and vegetation height. Additionally, protected areas should include a coastal-inland gradient and a diversity of substrate types, including exposed rock and trees.

Experimental separation of resource quantity from temporal variability: seedling responses to water pulses

We tested the hypothesis that higher temporal variability in water supply will promote higher species richness of germinating and surviving seedlings using assemblages of 70 species of herbaceous plants from limestone pavement habitats. In a two-factor greenhouse experiment, doubling the total volume of water added led to greater germination (measured as number of germinated seeds and species) and establishment (survival and biomass) but the effects of temporal variability depended on the response variable considered. Low pulse frequencies of water addition with total volume added held constant resulted in greater temporal variability in soil moisture concentration that in turn promoted higher density and richness of germinated seedlings. Low pulse frequencies caused an eight-fold greater mortality in the low total volume treatment and biomass production to decline by one-third in the high total volume treatment. The effects of increasing temporal variability in water supply during recruitment stages can thus be opposite on different components of plant fitness and may also depend on total resource quantity. While greater species richness in more temporally variable soil moisture conditions was attributable to sampling effects rather than species-specific responses to the water treatments, species relative abundances did vary significantly with temporal variability. Changes in the amplitude or frequency of resource fluctuations may alter recruitment patterns, and could have severe and relatively rapid effects on community structure in unproductive ecosystems.