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Joel S. Brown, Keren Embar, Eric Hancock and Burt P. Kotler

Derring-do is how aggressive a predator is in stalking and capturing prey. We model predator–prey interactions in which prey adjust vigilance behavior to mitigate risk of predation and predators their derring-do to manage risk of injury from capturing prey. High derring-do increases a predator's likelihood of capturing prey, but at higher risk of injury to itself. For fixed predator derring-do, prey increase vigilance in response to predator abundance, predator lethality, and predator encounter probability with prey and decrease vigilance with their own feeding rate; there is a humped-shaped relationship between prey vigilance and effectiveness of vigilance. For fixed prey vigilance, predators increase derring-do with the abundance of prey and predator lethality and decrease it with benefit of vigilance to prey and level of prey vigilance. When both prey and predator are behaviorally flexible, a predator–prey foraging game ensues whose solution represents an evolutionarily stable strategy (ESS). At the ESS, prey provide themselves with a public good as their vigilance causes predators to decrease derring-do. Conversely, predators have negative indirect effects on themselves as their derring-do causes prey to be more vigilant. These behavioral feedbacks create negative intra-specific interaction coefficients. Increasing the population size of prey (or predators) now has a direct negative effect on the prey (or predators). Both effects help stabilize predator–prey dynamics. Besides highlighting a common way by which predators may experience a food-safety tradeoff via dangerous prey, the model suggests why natural selection favors even small defensive measures by prey and hulky predators.

Hannu Ylӧnen and Marko Haapakoski

Mating with close kin may cause inbreeding depression with negative consequences to offspring and local populations. There exist mechanisms like kin-recognition or sex-specific dispersal to avoid mating with kin. In fluctuating population densities, like in many small mammals, both very low and very high densities provide conditions for inbreeding, if kin males are prone to stay in their natal area. Females are choosy and male dominance is thought to be the key feature when selecting mating partners. The aim of this study was to test the possible discrepancy in mate choice and negative fitness effects of inbreeding in two experiments, one in the laboratory and one in field enclosures. We asked (1) how the quality of the potential mating partners affects female choice regardless of relatedness and (2) how inbreeding affects the field populations created either from inbred or outbred individuals. Our results show that primiparous females in post-partum oestrus mated preferably with a dominant male, measured with their urine-marking behaviour, regardless if the selected male was brother or not. Only if the two males offered were of same dominance rank, the female mated with the non-kin male. However, the field experiment verified a negative effect of inbreeding in the bank vole. Thus, there seems to be a mismatch between female mate choice when selecting for dominance among resident males and population viability through long-term inbreeding depression. The study suggests the high importance of sex-biased dispersal as a mechanism to avoid kin individuals to meet in mate choice situations.

Aziz Subach

Predators affect prey directly by predation and indirectly by triggering behavioral responses that aim at reducing predation risk. In this paper, I present a method for training an avian predator which can allow separating between its direct and indirect effects on prey in various experimental setups. Barn owls are found to be a valuable tool for empirically testing different hypotheses related to predator-prey interactions, population dynamics, and inter-specific competition, all performed in the field using authentic rodent prey and their natural predators. Barn owls are raised and trained to participate in field experiments using classical conditioning, and are trained either to catch rodents or only to fly above a certain area without making any attempt to attack the prey, simulating solely predation risk. Body mass is a crucial factor in the training procedure, and I thus define five body mass ranges that characterize different behavioral stages in the training of owls. A logistic model is used to calculate and to predict changes in the body mass during the growth and training periods of owls. Finally, I discuss several possible implications of the usage of trained barn owls in empirical studies.

Christine Thuring and Gary Grant

From its beginnings in Germany in the twentieth century, a thriving extensive green roof industry has become established in many countries in temperate climates. Based upon the success of the industry, and with an expectation that this technology will be adopted in other climates, this review of the ecological research of extensive green roofs aims to evaluate the application of this knowledge. The modern extensive green roof is the product of research in the 1970s by German green roof pioneers; the selection of suitable species from analogue habitats led to green roof vegetation dominated by drought tolerant taxa. The commercial success of extensive green roof systems can be attributed to engineering and horticultural research, to policy mechanisms in some places, and to a market that encourages innovation, and the origins in ecological design are now easily overlooked. Some of the work reviewed here, including the classification of spontaneous roof vegetation into plant communities, is not widely known due to its confinement to the German literature. By re-visiting the history of the extensive green roof and reviewing the ecological research that has contributed to our understanding of it, the intention is, for this paper, to inform those considering green roofs in other climatic regions, to apply an ecologically informed approach in using local knowledge for developing installations that are suited to the bioregion in which they occur. Finally the paper considers some future directions for research and practice.

Nicole L. Kinlocka, Bracha Y. Schindler and Jessica Gurevitch

Green roofs can mitigate a number of urban environmental problems when green roof plant communities provide ecosystem services. However, this perspective may fail to address ecological aspects of the plant community. In particular, it does not account for the potential for green roofs to facilitate biological invasions. We consider current research in green roof ecology in light of the literature on biological invasions, focusing on plant invasion. We evaluate the role of species composition and novel communities, species interactions, succession, and dispersal on the trajectory of green roof plant communities. Green roofs have the potential to introduce invasive species through initial plantings, to become dominated by invasive species, and to spread invasive species, and we provide recommendations for plant selection and maintenance to reduce the risks of facilitating plant invasions to surrounding communities.

J. Scott Maclvor

Vegetated, “green” infrastructure, including terraces, balconies, and vegetated roofs and walls are increasingly common in urban landscapes, elevating habitat into novel contexts above ground. Highly mobile species, like bees and wasps, are often seen foraging on green infrastructure, but whether nesting opportunities are facilitated is not known. Cavity-nesting bees and wasps that provision brood in human-made trap nests were monitored over three years on 29 vegetated and non-vegetated roofs in Toronto, Canada. The study identified 27 species nesting on rooftops but found that building height was negatively correlated with the abundance of brood cells provisioned in trap nests, and positively correlated with the number of unfinished nests. A decline in green space area within a 600 m radius around each rooftop resulted in decreasing species richness and abundance. Although the introduced bee, Megachile rotundata (Fabricius) occupied more sites than any other bee or wasp (27.6%) and was the most abundant species, amounting to half (48.9%) of all brood reared, native bees were 73% of all bee species reared. The most abundant wasp was the native spider-collecting Trypoxylon collinum Smith (11.4%), but the introduced aphid-collecting Psenulus pallipes (Panzer) occurred at more sites (24.1%). For the pollination and pest controlling services they provide, bees and wasps should be considered in the design of vegetated roofs. Evidence here suggests that building height and surrounding green space at ground level impact bee and wasp diversity on vegetated roofs. Efforts supporting their populations using trap nests should target low- and mid-rise buildings (<5 building levels).

O.S. Starry

This paper explores the application of an ecosystem ecology framework to greenroof systems. It investigates how aspects of greenroof design or structure relate to functions such as rates of nutrient and energy cycling. Three main sections include energy budgets, cycling of nutrients and water, and ecosystem response to disturbance. Comparisons between greenroofs and other systems indicated that, functionally, greenroofs may be very different from ecosystem analogs. A further assessment of the greenroof energy budget called into question how food webs are supported. An evaluation of factors predicting system response to disturbance identified ways in which greenroofs may be less resilient to disturbance. One challenge with the ecosystem approach is a lack of sufficient data for fully holistic models, especially with respect to management practices. Ecosystem ecology is nevertheless shown to be a valuable framework for integrating existing greenroof research as well as targeting areas for future research and model development.

Robert D. Holt

A “green roof” is a roof on a structure created by humans, which has a plant community (and various hangers-on) established on it. There are many potential benefits of green roofs, ranging from moderation of local microclimates to modulations of storm runoffs, and green roofs may serve the enhancement of biodiversity conservation, as well. Green roofs would seem to be quintessential examples of a “novel ecosystem”. Here, I first outline some dimensions of the novelty that warrant more research. Green roofs can provide many opportunities for creative ecological research in the assembly, dynamics and functioning of novel ecosystems. Then, I briefly discuss some potential biodiversity hazards that are created along with green roofs. Recognizing these potential “shadows” of a green roof by no means belies the strong and compelling rationale for promoting green roofs, in terms of sustainability and livability of human structures; instead the points I raise are simply issues which should be evaluated and quantified when promoting green roofs broadly as a design strategy in new buildings or retrofitted existing structures.

Michael L. Rosenzweig

Many contributions to the symposium seek to expand the role of green roofs in the conservation of biodiversity. Indeed, if green roofs can be harnessed for biodiversity, they will add area to that now available to nature. That would have the mass effect of increasing the sustainable number of species in simple conformity with the species--area relationship. Because all green roofs are novel ecosystems, all represent instances of reconciliation ecology, i.e., re-engineering human uses to permit simultaneous beneficial use by people and nature. Green roofs can provide a large number of experiments that might teach us how to improve their design. But those experiments, like any in science, must be overtly designed so that their hypotheses are clear and explicit, their methods repeatable, and their data appropriate for rigorous analysis. I present an embryonic example using native plant species growing at ground level in the urban environments of Tucson, AZ, USA. Steps include: (1) formulating a hypothesis; (2) developing a database of species' attributes to allow intelligent selection for hypothesis testing; (3) developing software to allow winnowing the list of species to sets with a good chance, according to the hypothesis, of growing together; (4) installing the sets of plants and measuring the results; (5) defining a continuous measure of conformity with the hypothesis; and (6) comparing results to hypothesis. If ecologists can successfully design reconciled ecosystems in urban settings – green roofs included – city people will be able to re-establish their everyday connection to nature.

Inga Dirks, Buzi Raviv, Oren Shelef, Amber Hill, Eppel Amir, Moses Kwame Aidoo, Brian Hoefgen, Tal Rapaport, Hila Gil, Endale Geta, Amnon Kochavi, Itay Cohen and Shimon Rachmilevitch

Green roofs in the Mediterranean region are often exposed to high levels of radiation, extreme temperatures, and an inconsistent water supply. To withstand these harsh conditions in shallow soils and poorly aerated growth media, plants must be armored with adaptations. Strategies that have evolved in desert plants can play significant roles in the use of plants for green covers. In the following, we will specifically focus on (1) heat and radiation, (2) drought, and (3) salinity. Further, we will discuss (4) interactions between neighboring plants. Finally, we will (5) propose a design for diverse green roofs that includes horticultural and medicinal products and provides diverse habitats. Many desert plants have developed morphological and anatomical features to avoid photo-inhibition, which can be advantageous for growth on green roofs. Plants exhibiting C4 photosynthesis or crassulacean acid metabolism (CAM) photosynthesis have a protected hydraulic system that enables growth under dry conditions. Furthermore, dew and high levels of relative humidity can provide reliable water sources under limited precipitation. Halophytes are protected against salinity, ionic specific stress, and nutritional imbalances, characteristics that can be advantageous for green roofs. Under limited space, competition for resources becomes increasingly relevant. Allelopathy can also induce the germination and growth inhibition of neighboring plants. Many desert plants, as a result of their exposure to environmental stress, have developed unique survival adaptations based on secondary metabolites that can be used as pharmaceuticals. A systematic survey of plant strategies to withstand these extreme conditions provides a basis for increasing the number of green roof candidates.