Plant cover and biomass change on extensive green roofs over a decade and ten lessons learned.

Green roofs are well studied for the environmental, social, and economic services these provide. As a result, green roofs are widespread and within the common vernacular of city residents. Green roof bylaws and construction standards are present in many cities in North America, rooting the presence of this green infrastructure within urban landscapes. Although examples of green roofs constructed decades ago exist, rarely are green roofs monitored over such long periods, and in ways that allow for experimentation, analysis, and conclusions about performance or function. In this study we present findings on plant cover and biomass from a green roof testing facility in Toronto, Canada that was monitored for over a decade. We examine the contributions of growing media, planting, and irrigation in the first seven years (2011-2021) of the eleven-year monitoring period. We found that during this maintenance phase period (2011-2017), plant cover and biomass was highest in modules planted with Sedum, included organic media, and were irrigated, whereas non-irrigated modules planted with forbs and grasses had the poorest performance regardless of media type. Following the stoppage of irrigation, and the post maintenance phase (2017-2021), modules initially planted with Sedum continued to sustain cover and biomass whereas planted forbs and grasses mostly disappeared, and these treatments were overtaken by Sedum. Our findings demonstrate that with irrigation, plantings of forbs and grasses can sustain plant cover and biomass. However, Sedum buffers against major changes to environmental conditions or abrupt changes to maintenance, adding insurance against failure of extensive green roofs.

Establishment of the non-native horned-face bee Osmia cornifrons and the taurus mason bee Osmia taurus (Hymenoptera: Megachilidae) in Canada.

Established populations of the non-native horned-face bee, Osmia cornifrons (Radoszkowski, 1887), and the taurus mason bee, Osmia taurus Smith, 1873 (Hymenoptera: Megachilidae), have been identified from Canada for the first time. In the US, the importation of O. cornifrons, beginning in the 1970s, led to its release for agricultural crop pollination and spread across the country. In this article, we report on O. cornifrons captured while sampling wild bees in Toronto, Ontario using hand nets, bug vacuums, and vane traps, as well as established populations in trap nests, from 2017-2020. The morphologically similar O. taurus, which was accidentally introduced to the US with shipments of imported O. cornifrons, was also recorded in our samples. Recently, a few individual O. taurus specimens have been identified from Ontario and Quebec; however, the extent of our sampling included nests, indicating it is also established in Canada. Others have shown its population growth to have been associated with concordant declines in abundances of native mason bee species in the US, and similar impacts are possible in Canada if action is not taken. We propose three non-mutually exclusive possible pathways for the arrival of O. cornifrons, as well as O. taurus, in Canada: (1) natural migration northward from non-native populations in the US, (2) international importation in the 1980s-2000s to support agricultural research programs, and (3) unintentional release of mason bee cocoons purchased from non-local vendors. We argue that a focus on enhancing populations of locally occurring native bees and stronger policy on the importation and sale of non-native bees are needed.

Reproductive trait differences drive offspring production in urban cavity-nesting bees and wasps.

The contrasting and idiosyncratic changes in biodiversity that have been documented across urbanization gradients call for a more mechanistic understanding of urban community assembly. The reproductive success of organisms in cities should underpin their population persistence and the maintenance of biodiversity in urban landscapes. We propose that exploring individual-level reproductive traits and environmental drivers of reproductive success could provide the necessary links between environmental conditions, offspring production, and biodiversity in urban areas. For 3 years, we studied cavity-nesting solitary bees and wasps in four urban green space types across Toronto, Canada. We measured three reproductive traits of each nest: the total number of brood cells, the proportion of parasite-free cells, and the proportion of non-emerged brood cells that were parasite-free. We determined (a) how reproductive traits, trait diversity and offspring production respond to multiple environmental variables and (b) how well reproductive trait variation explains the offspring production of single nests, by reflecting the different ways organisms navigate trade-offs between gathering of resources and exposure to parasites. Our results showed that environmental variables were poor predictors of mean reproductive trait values, trait diversity, and offspring production. However, offspring production was highly positively correlated with reproductive trait evenness and negatively correlated with trait richness and divergence. This suggests that a narrow range of reproductive traits are optimal for reproduction, and the even distribution of individual reproductive traits across those optimal phenotypes is consistent with the idea that selection could favor diverse reproductive strategies to reduce competition. This study is novel in its exploration of individual-level reproductive traits and its consideration of multiple axes of urbanization. Reproductive trait variation did not follow previously reported biodiversity-urbanization patterns; the insensitivity to urbanization gradients raise questions about the role of the spatial mosaic of habitats in cities and the disconnections between different metrics of biodiversity.

The direct and indirect effects of extreme climate events on insects.

Extreme climate events are predicted to increase in the future, which will have significant effects on insect biodiversity. Research into this area has been rapidly expanding, but knowledge gaps still exist. We conducted a review of the literature to provide a synthesis of extreme climate events on insects and identify future areas of research. In our review, we asked the following questions: 1) What are the direct and indirect mechanisms that extreme climate events affect individual insects? 2) What are the effects of extreme climate events on insect populations and demography? 3) What are the implications of the extreme climate events effects on insect communities? Drought was among the most frequently described type of extreme climate event affecting insects, as well as the effects of temperature extremes and extreme temperature variation. Our review explores the factors that determine the sensitivity or resilience to climate extremes for individuals, populations, and communities. We also identify areas of future research to better understand the role of extreme climate events on insects including effects on non-trophic interactions, alteration of population dynamics, and mediation of the functional the trait set of communities. Many insect species are under threat from global change and extreme climate events are a contributing factor. Biologists and policy makers should consider the role of extreme events in their work to mitigate the loss of biodiversity and delivery of ecosystem services by insects.

Urbanization and plant invasion alter the structure of litter microarthropod communities.

Anthropogenic activity underpins the creation of urban ecosystems, often with introduced or invasive species playing a large role in structuring ecological communities. While the effects of urbanization on charismatic taxa such as birds, bees or butterflies have received much attention, the impacts on small and inconspicuous organisms remain poorly understood. Here, we assess how the community structure of leaf litter-inhabiting microarthropods in city parks varies along an urbanization gradient in Toronto, Canada. At each park, we established paired forest understorey plots which were either dominated by native vegetation or dog-strangling vine Vincetoxicum rossicum, an invasive species that is spreading throughout northeastern North America and abundant in urban areas. We compared microarthropod richness, abundance and diversity in ecological traits between invaded and non-invaded plots as well as compositional dissimilarities among plots across the urbanization gradient. We recorded 123 genera and found (a) there was a negative effect of urbanization on microarthropod richness and abundance but only in invaded plots; (b) richness and abundance increased continuously with urbanization in non-invaded plots, but peaked at intermediate urbanization levels in invaded plots and (c) there was significant turnover with increasing urbanization, with distinct communities represented in highly urbanized areas compared to less urbanized areas, regardless of whether invaded. We also found litter microarthropod richness and abundance increased with soil ammonium and decreased with nitrate. These trends were especially strong for fungivorous microarthropods; however, there was no relationship between soil nutrients and urbanization or invasion. Urbanization and biological invasion drive biodiversity change, and there is a need to disentangle these effects on ecological communities and related ecosystem processes. We show microarthropod communities change with urbanization, with the effects of invasion most prominent in non-urban areas. Here, there is high richness and abundance but low ecological trait diversity, possibly because certain feeding traits are excluded and others overrepresented. Understanding of urban ecological systems must include knowledge of the microarthropods that interact widely across food webs, form distinct communities in highly urban areas and drive many of the important ecological functions upon which people in cities depend.

A roadmap for urban evolutionary ecology.

Urban ecosystems are rapidly expanding throughout the world, but how urban growth affects the evolutionary ecology of species living in urban areas remains largely unknown. Urban ecology has advanced our understanding of how the development of cities and towns change environmental conditions and alter ecological processes and patterns. However, despite decades of research in urban ecology, the extent to which urbanization influences evolutionary and eco-evolutionary change has received little attention. The nascent field of urban evolutionary ecology seeks to understand how urbanization affects the evolution of populations, and how those evolutionary changes in turn influence the ecological dynamics of populations, communities, and ecosystems. Following a brief history of this emerging field, this Perspective article provides a research agenda and roadmap for future research aimed at advancing our understanding of the interplay between ecology and evolution of urban-dwelling organisms. We identify six key questions that, if addressed, would significantly increase our understanding of how urbanization influences evolutionary processes. These questions consider how urbanization affects nonadaptive evolution, natural selection, and convergent evolution, in addition to the role of urban environmental heterogeneity on species evolution, and the roles of phenotypic plasticity versus adaptation on species' abundance in cities. Our final question examines the impact of urbanization on evolutionary diversification. For each of these six questions, we suggest avenues for future research that will help advance the field of urban evolutionary ecology. Lastly, we highlight the importance of integrating urban evolutionary ecology into urban planning, conservation practice, pest management, and public engagement.

Manipulating plant phylogenetic diversity for green roof ecosystem service delivery.

Plant species and functional trait diversity have each been shown to improve green roof services. Species and trait differences that contribute to ecosystem services are the product of past evolutionary change and phylogenetic diversity (PD), which quantifies the relatedness among species within a community. In this study, we present an experimental framework to assess the contribution of plant community PD for green roof ecosystem service delivery, and data from one season that support our hypotheses that PD would be positively correlated with two services: building cooling and rainwater management. Using 28 plant species in 12 families, we created six community combinations with different levels of PD. Each of these communities was replicated at eight green roofs along an elevation gradient, as well as a ground level control. We found that the minimum and mean roof temperature decreased with increasing PD in the plant community. Increasing PD also led to an increase in the volume of rainwater captured, but not the proportion of water lost via evapotranspiration 48 hr following the rain event. Our findings suggest that considering these evolutionary relationships could improve functioning of green infrastructure and we recommend that understanding how to make PD (and other measures of diversity) serviceable for plant selection by practitioners will improve the effectiveness of design and ecosystem service delivery. Lastly, since no two green roof sites are the same and can vary tremendously in microclimate conditions, our study illustrates the importance of including multiple independent sites in studies of green roof performance.

The Necessity of Multitrophic Approaches in Community Ecology.

Trophic interactions are a fundamental part of ecosystems; yet, most ecological studies focus on single trophic levels and this hampers our ability to detect the underlying mechanisms structuring communities as well as the effects of environmental change. Here, we argue that the historical dominance of studying competition within trophic levels, and the focus on taxonomic groups without differentiating the trophic level, has led to the under-representation of multitrophic research in community ecology. There are many hurdles that challenge multitrophic approaches and we discuss solutions to overcome these. To advance our understanding of the fundamental drivers of community assembly and to provide the necessary guidance for managing and mitigating the effects of environmental change, we argue that ecologists should better align research with a trophically inclusive definition of a community.

The Bees among Us: Modelling Occupancy of Solitary Bees.

Occupancy modelling has received increasing attention as a tool for differentiating between true absence and non-detection in biodiversity data. This is thought to be particularly useful when a species of interest is spread out over a large area and sampling is constrained. We used occupancy modelling to estimate the probability of three phylogenetically independent pairs of native-introduced species [Megachile campanulae (Robertson)-Megachile rotundata (Fab.), Megachile pugnata Say-Megachile centuncularis (L.), Osmia pumila Cresson-Osmia caerulescens (L.)] (Apoidea: Megachilidae) being present when repeated sampling did not always find them. Our study occurred along a gradient of urbanization and used nest boxes (bee hotels) set up over three consecutive years. Occupancy modelling discovered different patterns to those obtained by species detection and abundance-based data alone. For example, it predicted that the species that was ranked 4th in terms of detection actually had the greatest occupancy among all six species. The native M. pugnata had decreased occupancy with increasing building footprint and a similar but not significant pattern was found for the native O. pumila. Two introduced bees (M. rotundata and M. centuncularis), and one native (M. campanulae) had modelled occupancy values that increased with increasing urbanization. Occupancy probability differed among urban green space types for three of six bee species, with values for two native species (M. campanulae and O. pumila) being highest in home gardens and that for the exotic O. caerulescens being highest in community gardens. The combination of occupancy modelling with analysis of habitat variables as an augmentation to detection and abundance-based sampling is suggested to be the best way to ensure that urban habitat management results in the desired outcomes.

DNA barcoding to identify leaf preference of leafcutting bees.

Leafcutting bees (Megachile: Megachilidae) cut leaves from various trees, shrubs, wildflowers and grasses to partition and encase brood cells in hollow plant stems, decaying logs or in the ground. The identification of preferred plant species via morphological characters of the leaf fragments is challenging and direct observation of bees cutting leaves from certain plant species are difficult. As such, data are poor on leaf preference of leafcutting bees. In this study, I use DNA barcoding of the rcbL and ITS2 regions to identify and compare leaf preference of three Megachile bee species widespread in Toronto, Canada. Nests were opened and one leaf piece from one cell per nest of the native M. pugnata Say (N=45 leaf pieces), and the introduced M. rotundata Fabricius (N=64) and M. centuncularis (L.) (N=65) were analysed. From 174 individual DNA sequences, 54 plant species were identified. Preference by M. rotundata was most diverse (36 leaf species, H'=3.08, phylogenetic diversity (pd)=2.97), followed by M. centuncularis (23 species, H'=2.38, pd=1.51) then M. pugnata (18 species, H'=1.87, pd=1.22). Cluster analysis revealed significant overlap in leaf choice of M. rotundata and M. centuncularis. There was no significant preference for native leaves, and only M. centuncularis showed preference for leaves of woody plants over perennials. Interestingly, antimicrobial properties were present in all but six plants collected; all these were exotic plants and none were collected by the native bee, M. pugnata. These missing details in interpreting what bees need offers valuable information for conservation by accounting for necessary (and potentially limiting) nesting materials.

'Bee hotels' as tools for native pollinator conservation: a premature verdict?

Society is increasingly concerned with declining wild bee populations. Although most bees nest in the ground, considerable effort has centered on installing 'bee hotels'--also known as nest boxes or trap nests--which artificially aggregate nest sites of above ground nesting bees. Campaigns to 'save the bees' often promote these devices despite the absence of data indicating they have a positive effect. From a survey of almost 600 bee hotels set up over a period of three years in Toronto, Canada, introduced bees nested at 32.9% of sites and represented 24.6% of more than 27,000 total bees and wasps recorded (47.1% of all bees recorded). Native bees were parasitized more than introduced bees and females of introduced bee species provisioned nests with significantly more female larva each year. Native wasps were significantly more abundant than both native and introduced bees and occupied almost 3/4 of all bee hotels each year; further, introduced wasps were the only group to significantly increase in relative abundance year over year. More research is needed to elucidate the potential pitfalls and benefits of using bee hotels in the conservation and population dynamics of wild native bees.

Bee species-specific nesting material attracts a generalist parasitoid: implications for co-occurring bees in nest box enhancements.

Artificial nests (e.g., nest boxes) for bees are increasingly being used to contribute to nesting habitat enhancement for bees that use preexisting cavities to provision brood. They usually incorporate additional nesting materials that vary by species. Cavity-nesting bees are susceptible to brood parasitoids that recognize their host(s) using visual and chemical cues. Understanding the range of cues that attract parasitoids to bee nests, including human-made analogues, is important if we wish to control parasitism and increase the potential value of artificial nests as habitat-enhancement strategies. In this study, we investigated the cues associated with the orientation of the generalist brood parasitoid Monodontomerus obscurus Westwood (Hymenoptera: Torymidae) to the nests of a common cavity-nesting resin bee Megachile campanulae (Robertson) (Megachilidae). The parasitoids were reared from previously infested M. campanulae brood cells and placed into choice trials where they were presented with pairs of different nest material cues. Among different materials tested, we found that Mo. obscurus was most attracted to fresh resin collected directly from Pinus strobus trees followed by previously used resin collected from the bee nest. The parasitoid also attacked other bee species in the same nest boxes, including those that do not use resin for nesting. Our findings suggest that M. campanulae could act as a magnet, drawing parasites away from other bee hosts co-occurring in nest boxes, or, as an attractant of Mo. obscurus to nest boxes, increasing attacks on co-occurring host bee species, potentially undermining bee diversity enhancement initiatives.

Decoupling factors affecting plant diversity and cover on extensive green roofs.

Supplemental irrigation systems are often specified on green roofs to ensure plant cover and growth, both important components of green roof performance and aesthetics. Properties of the growing media environment too can alter the assemblage of plant species able to thrive. In this study we determine how plant cover, above ground biomass and species diversity are influenced by irrigation and growing media. Grass and forb vegetative cover and biomass were significantly greater in organic based growing media but there was no effect of supplemental irrigation, with two warm season grasses dominating in those treatments receiving no supplemental irrigation. On the other hand, plant diversity declined without irrigation in organic media, and having no irrigation in inorganic growing media resulted in almost a complete loss of cover. Sedum biomass was less in inorganic growing media treatments and species dominance shifted when growing media organic content increased. Our results demonstrate that supplemental irrigation is required to maintain plant diversity on an extensive green roof, but not necessarily plant cover or biomass. These results provide evidence that planting extensive green roofs with a mix of plant species can ensure the survival of some species; maintaining cover and biomass when supplemental irrigation is turned off to conserve water, or during extreme drought.

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.

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.