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Testing the Relative Importance of Local Resources and Landscape Connectivity on Bombus Impatiens (Hymenoptera, Apidae) Colonies

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Testing the Relative Importance of Local Resources and Landscape Connectivity on Bombus Impatiens (Hymenoptera, Apidae) Colonies Apidologie Original article * INRA, DIB and Springer-Verlag France, 2017 DOI: 10.1007/s13592-017-0499-1 Testing the relative importance of local resources and landscape connectivity on Bombus impatiens (Hymenoptera, Apidae) colonies 1,2,3 4 5 John D. HERRMANN , Nick M. HADDAD , Douglas J. LEVEY 1Department of Biology, University of Washington, Seattle, WA 98195-1800, USA 2Department of Plant Biology, Michigan State University, East Lansing, MI 48824, USA 3Department of Landscape Ecology, University of Kiel, Olshausenstr, 75, 24118, Kiel, Germany 4Department of Applied Ecology, North Carolina State University, Raleigh, NC 27695-7617, USA 5Division of Environmental Biology, National Science Foundation, Arlington, VA 22230, USA Received 19 November 2015 – Revised 24 November 2016 – Accepted 18 January 2017 Abstract – Bee populations are decreasing worldwide. The underlying causes are likely determined by factors at different scales. We tested the relative importance of local resources and landscape connectivity on 64 bumblebee (Bombus impatiens ) colonies in experimentally isolated and connected habitat fragments. We used colony mass, no. of workers, and no. of gynes to estimate colony performance. Landscape connectivity did not significantly affect colony performance, but local floral resources had a significantly positive effect, especially in isolated fragments. These results suggest that bumblebee colonies encountered sufficient floral resources within the local 1.4 ha habitat fragments to support colony growth, making long-distance foraging trips to neighboring fragments unnecessary. From a conservation perspective, we suggest that efforts to improve colony performance should prioritize boosting local floral resources over manipulation of large-scale landscape features. agri-environment scheme / foraging distance / landscape connectivity / habitat fragmentation / floral resources 1. INTRODUCTION on the Status of Pollinators in North America 2007, Daily 1997,Kremenetal.2002). A first step is to Insect pollinators are suffering sharp declines in identify and assess potential causes of population abundance and richness, worldwide (Potts et al. declines. 2010). These declines have led to a simultaneous Habitat loss has been identified as one of the most decrease of ecosystem services that pollinators pro- important drivers of pollinator declines (Potts et al. vide (Cunningham 2000) and may have triggered a 2010). Decreasing habitat connectivity, which typi- global pollination crisis (Steffan-Dewenter et al. cally accompanies habitat loss, is also thought to 2005). Reduced pollination is a concern because of negatively affect pollinator populations, but positive its importance for conservation of global biodiversi- as well as negative effects of connectivity on polli- ty (Bodin et al. 2006) and human food production nator populations have been observed (e.g., Schüepp (Potts et al. 2010). Conservation strategies are ur- et al. 2011, Steffan-Dewenter & Tscharntke 1999). gently needed to counteract ongoing pollinator de- The contrasting conclusions reached by differ- clines and sustain pollination services (Committee ent studies on connectivity effects may have two main causes. First, results might be artifacts of study design, particularly when some designs fail Corresponding author: J. Herrmann, to separate effects of habitat loss from those of [email protected] habitat connectivity (Hadley & Betts 2012). This Handling editor: Yves Le Conte almost inevitable link between habitat loss and J.D. Herrmann et al. connectivity continues to generate controversy spatial scales, such as flower density in a habitat among landscape ecologists regarding their inde- fragment (Rundlof et al. 2008). Offspring produc- pendent effects and the importance of separating tion of solitary bees nesting in high-quality habi- them (Bailey et al. 2010, Doerr et al. 2011, Fahrig tat, for instance, was buffered against negative 2013, Hodgson et al. 2009, 2011). Second, struc- isolation effects by switching to local floral re- tural connectivity (i.e., physical contiguity of sources when other resources were too distant landscape elements) and functional connectivity (Williams & Kremen 2007). The importance of (i.e., behavioral responses that vary among spe- local resources, however, might be contingent cies and result in different probabilities of travel upon accessibility and amount of additional re- between landscape elements) may not be synony- sources in the surrounding landscape. mous for some organisms. This distinction be- Within a large-scale landscape experiment comes important when an organism’s movement composed of seminatural savanna fragments, we is not confined to its preferred habitat, which is tested the effects of landscape connectivity and common (Kuefler et al. 2010,Haddad& local floral resources on bumblebee (Bombus im- Tewksbury 2005, Tischendorf & Fahrig 2000). patiens ) colonies. We established bumblebee col- Structural connectivity can be increased through onies in fragments, which were either isolated or landscape corridors, strips of habitat that connect connected to neighboring fragments via a corri- otherwise isolated fragments of the same habitat dor. By simultaneously measuring corridor and (Hilty et al. 2006). Increased structural connectivity local resource effects on different parameters of may lead to increased functional connectivity by colony growth, we avoided pitfalls associated facilitating movement between otherwise isolated with visual surveys of pollinators and gained di- habitat fragments (Haddad et al. 2015). Insect pol- rect estimates of colony performance (Heard et al. linators have been observed to use habitat strips 2007). Measuring multiple metrics of colony per- and landscape features (e.g., hedgerows) as corri- formance is important, as conservation measures dors (Cranmer et al. 2012), and it has been exper- need to be based on both short-term fitness pa- imentally demonstrated that corridors increase pol- rameters (e.g., colony mass and number of len transfer between habitat fragments that they workers) and long-term fitness parameters (e.g., connect (Townsend & Levey 2005), leading to number of virgin queens produced to establish increased reproductive success of plants (Cranmer new colonies). We hypothesized that (i) colony et al. 2012,Tewksburyetal.2002). However, this performance is higher in connected than in uncon- observed increase in pollinator movement via cor- nected fragments, (ii) local floral resources in- ridors does not provide evidence that habitat con- crease colony performance, and (iii) colony per- nectivity also benefits the pollinators (Hogdson formance depends more on local floral resources et al. 2011). Although it is thought that pollinator in unconnected than in connected fragments. fitness will increase with structural connectivity (Hopfenmüller et al. 2014), it remains untested 2. MATERIAL AND METHODS whether increased structural connectivity increases pollinator fitness through functional connectivity 2.1. Study sites (Hadley & Betts 2012). In addition to large-scale effects such as con- We tested our hypotheses in a large experimen- nectivity between fragments, population dynam- tal landscape at the Savannah River Site (SRS) in ics of many species are determined by processes South Carolina, USA (Figure 1a). Eight replicate occurring at smaller scales, for example, in a landscapes (hereafter Bblocks^)consistingoffrag- single fragment (McGarigal and Cushman ments and corridors of open habitat were created 2002). Even though many pollinators are regarded in 1999/2000 (six blocks) and 2007 (two blocks) as highly mobile foragers that perceive resources by clearing mature pine forest. The cleared areas at large spatial scales (e.g., bumblebees; Westphal are being actively restored to longleaf pine savan- et al. 2006), survival of pollinators may most na through implementation of prescribed fires on strongly depend on resource quality at small a2–3-year rotation, planting of longleaf pine Impact of local and landscape factors on bumblebees a 5 3 South Carolina 4 SRS 3 Latitude 3 3 2 050100 km 3 -83 -82 -81 -80 -79 Longitude b Wi Re Ce Co 0100 m Wi Figure 1. a Locations of the eight experimental blocks (black circles ) at the Savannah River Site (SRS ) in South Carolina (USA). b One experimental block with one center fragment (Ce ) and four peripheral fragments: One peripheral fragment is connected by a 150 × 25 m corridor (Co ), whereas the rectangular (Re ) and winged (Wi ) fragments are isolated by mature pine forest. All blocks contain one duplicate unconnected fragment type. Two Bombus impatiens colonies (red box ) were established within protective setups in each peripheral fragment seedlings (Pinus palustris ), and removal of hard- rectangular (four blocks) or winged fragment woods. Each block consists of one center frag- (four blocks). Each block was surrounded by a ment (100 m x 100 m) surrounded by four periph- forested buffer zone with limited management eral fragments, each 150 m from the center frag- activities (400 m radius around the block’s center ment (Figure 1b). One of the peripheral fragments point). The experimental landscapes have been (Bconnected fragment^) in each block is connect- used to investigate effects of fragment shape and ed to the central fragment by a 25 m wide and connectivity on dispersal of various organisms, 150 m long corridor. The other three peripheral including plants, butterflies, and birds fragments are isolated from the central
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