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Building Height Matters: Nesting Activity of Bees and Wasps on Vegetated Roofs J This article was downloaded by: [173.32.110.191] On: 19 June 2015, At: 12:23 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK Israel Journal of Ecology & Evolution Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/tiee20 Click for updates Building height matters: nesting activity of bees and wasps on vegetated roofs J. Scott MacIvora a Biology Department, York University, Toronto, Canada Published online: 16 Jun 2015. To cite this article: J. Scott MacIvor (2015): Building height matters: nesting activity of bees and wasps on vegetated roofs, Israel Journal of Ecology & Evolution, DOI: 10.1080/15659801.2015.1052635 To link to this article: http://dx.doi.org/10.1080/15659801.2015.1052635 PLEASE SCROLL DOWN FOR ARTICLE Taylor & Francis makes every effort to ensure the accuracy of all the information (the “Content”) contained in the publications on our platform. However, Taylor & Francis, our agents, and our licensors make no representations or warranties whatsoever as to the accuracy, completeness, or suitability for any purpose of the Content. Any opinions and views expressed in this publication are the opinions and views of the authors, and are not the views of or endorsed by Taylor & Francis. The accuracy of the Content should not be relied upon and should be independently verified with primary sources of information. Taylor and Francis shall not be liable for any losses, actions, claims, proceedings, demands, costs, expenses, damages, and other liabilities whatsoever or howsoever caused arising directly or indirectly in connection with, in relation to or arising out of the use of the Content. This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. Terms & Conditions of access and use can be found at http:// www.tandfonline.com/page/terms-and-conditions Israel Journal of Ecology & Evolution, 2015 http://dx.doi.org/10.1080/15659801.2015.1052635 Building height matters: nesting activity of bees and wasps on vegetated roofs J. Scott MacIvor* Biology Department, York University, Toronto, Canada (Received 11 October 2014; accepted 12 May 2015) 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). Keywords: urban; biodiversity; green roofs; cavity-nesting; trap nest; Megachile Introduction facilitate À as well as study À both positive and negative Studies investigating local and landscape impacts on novel responses of biodiversity to urban and environmen- insect populations are increasingly carried out in and tal changes (McIntyre 2000; Felson & Pickett 2005; around cities (Blair 1999; Hostetler & McIntyre 2001; Goddard et al. 2010). Cane et al. 2006; Matteson et al. 2008; Sattler et al. 2010; Many species in a wide range of taxonomic groups Bates et al. 2012; Banaszak-Cibicka & Zmihorski_ 2012; have been identified from surveys on vegetated roofs (e.g. Geslin et al. 2013; Braaker et al. 2014). City landscapes Jones 2002; Millet 2004; Kadas 2006; Pearce & Walters are usually more strongly three-dimensional and complex 2012; Madre et al. 2013, MacIvor, Ruttan, & Salehi than surrounding natural landscapes (Allen 1998). In 2015; reviewed in MacIvor & Ksiazek 2015); however, many cases, city buildings create even more complexity adjacent or nearby ground level reference habitat yields Downloaded by [173.32.110.191] at 12:23 19 June 2015 than that on cliff faces or other mountainous habitat higher diversity (Colla et al. 2009; MacIvor & Lundholm (Larson et al. 2004). Studying how taxa interact with this 2011; Tonietto et al. 2011). There is some quantitative complex environment may provide new insight into their data on the reproductive success of birds (Baumann 2006) ecology and conservation (Dearborn & Kark 2010). but there is little data on whether vegetated roofs can act Green infrastructure elevated above ground in three- as a source or sink for species seeking nesting habitat dimensional space (including vegetated roof and walls, (MacIvor & Ksiazek 2015). Cavity-nesting bees and gardened terraces, balconies, and garages) increases green wasps in particular may benefit from vegetated roofs as space vertically where space at ground is developed, pro- nesting habitat, as they regularly use cracks, crevices, viding new urban habitat opportunities (Pickett et al. nail- and drill holes in human-made infrastructure, and 2013). The contribution of vegetated roofs is perhaps the other cavities as nesting habitat in place of natural ones best studied among them, with some suggesting they in wood and plant stems (Krombein 1967; Cane could aid in the conservation of rare species, or increase et al. 2007; MacIvor, Cabral, & Packer 2014). These connectivity among fragmented habitat patches (reviewed include those that are accidentally or intentionally added in Williams et al. 2014). However, vegetated roofs like to vegetated roofs and other forms of green infrastructure. some ground level habitat might act as dispersal platforms Pollination by bees and predation on pest insects by soli- for exotic or undesirable species that undermine native tary wasps are desirable ecosystem functions carried out biodiversity conservation. It is important to consider habi- by cavity-nesting species that should be encouraged in tat conditions in the design of green infrastructure to both natural and human-dominated landscapes *Email: [email protected] Ó 2015 Taylor & Francis 2 J.S. MacIvor (Tscharntke et al. 1998). However, it is not clear which, if correlated with reduced observations of wild bees on flow- any species of cavity-nesting bees and wasps will search ers (Jauker et al. 2009; Garibaldi et al. 2011). Moreover, for nesting habitat on roofs of buildings, especially when increasing horizontal distance from floral resources has situated dozens of meters above ground. resulted in fewer offspring provisioned in cavity-nesting Nest site locating behaviors of cavity-nesting bees and bees (Peterson & Roitberg 2006; Schuepp€ et al. 2011). In wasps often involve vertical movement as the insects this study, I hypothesized building height, which reflects search for holes in dead wood in trees (Wcislo 1996). vertical isolation from ground level habitat opportunities, Thus, cavity-nesting bees and wasps might have some would result in decreasing colonization of trap nests by pre-adapted traits for searching for nest sites at the heights bees and wasps. Further, I hypothesized that as the propor- of vegetated roofs. The use of trap nests (Krombein 1967) tion of green space at ground level increases there would can provide such suitable nesting sites and many cavity- be an increase in colonization of trap nests, as studies nesting species readily adopt them (Tscharntke et al. sampling bees on vegetated roofs have determined ground 1998). Trap nests have been used in many ecological and level green space to be a predictor of greater abundance conservation studies (Gathmann et al. 1994; Steffan-Dew- and diversity (Tonietto et al. 2011; Braaker et al. 2014). enter et al. 2002; Tylianakis et al. 2006; Praz et al. 2008; MacIvor, Cabral, & Packer 2014) and they have been shown to reflect overall bee diversity (Westphal et al. Methods 2008). However, few studies have used trap nests to sur- Bees and wasps were sampled from May to October over vey cavity-nesting bees and wasps in urban landscapes three years (2011À2013) using trap nests set up on 29 (Alves-dos-Santos, 2003; Gaston et al. 2005; Loyola & rooftops (one per roof) each separated by a minimum Martins 2006; Everaars et al. 2011; Pereira-Peixoto et al. 250 m throughout the city of Toronto (Figure 1). The 2014; MacIvor & Packer 2015). roofs were at varying heights from one (approximately In this study I assess the impact of building height, 3.3 m from ground level) to nine building levels (approxi- number of buildings, and the proportion of local ground mately 29.7 m) and were classified into three types: (1) level green space upon the species richness and abun- intensively planted vegetated roofs with numerous wild dance of bees and wasps colonizing trap nests. Horizontal flowers, some shrubs and with growing media deeper than landscape isolation from florally diverse habitats has been 15 cm; (2) extensive Sedum- or grass-dominant vegetated Downloaded by [173.32.110.191] at 12:23 19 June 2015 Figure 1.
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