Dispersal capacity and diet breadth modify the response of wild bees to habitat loss

● Drakare, S., Lennon, J. J. & Hillebrand, H. 2006 The imprint of the geographical, evolutionary and ecological context on species–area relationships. Ecol. Lett. 9, 215–227. (doi:10.1111/j.1461-0248.2005.00848.x) ● Ewers, R. M. & Didham, R. K. 2006 Confounding factors in the detection of species responses to habitat fragmentation. Biol. Rev. 81, 117–142. (doi:10.1017/S1464793105006949) ● Fahrig, L. 2003 Effects of habitat fragmentation on biodiversity. Ann. Rev. Ecol. Syst. 34, 487–515. (doi:10.1146/annurev.ecolsys.34.011802.132419) ● Fortuna, M. A. & Bascompte, J. 2006 Habitat loss and the structure of plant–animal mutualistic networks. Ecol. Lett. 9, 278–283. (doi:10.1111/j.1461-0248.2005.00868.x) ● Franze´n, M. & Nilsson, S. G. 2010 Both population size and patch quality affect local extinctions and colonizations. Proc. R. Soc. B 277, 78–85. (doi:10.1098/rspb.2009.1584) ● Gaston, K. J. 1988 Patterns in local and regional dynamics of moth populations. Oikos 53, 49–57. (doi:10.2307/ 3565662) ● Greenleaf, S. S., Williams, N. M., Winfree, R. & Kremen, C. 2007 Bee foraging ranges and their relationship to body size. Oecologia 153, 589–596. (doi:10.1007/s00442-007- 0752-9) ● Hanski, I. & Ovaskainen, O. 2000 The metapopulation capacity of a fragmented landscape. Nature 404, 755– 758. (doi:10.1038/35008063) ● Heinrich, B. 1979 Bumblebee economics. Cambridge, MA: Harvard University Press. Henle, K., Davies, K. F., Kleyer, M., Margules, C. & Settele, J. 2004 Predictors of species sensitivity to fragmentation. Biodiv. Cons. 13, 207–251. (doi:10.1023/B:BIOC.0000004319.91643.9e) ● Hodgson, J. G. 1993 Commonness and rarity in British butterflies. J. Appl. Ecol. 30, 407–427. (doi:10.2307/ 2404182) ● Holzschuh, A., Steffan-Dewenter, I., Kleijn, D. & Tscharntke, T. 2007 Diversity of flower-visiting bees in cereal fields: effects of farming system, landscape composition and regional context. J. Appl. Ecol. 44, 41–49. (doi:10.1111/j.1365-2664.2006.01259.x) ● Johnson, S. D. & Steiner, K. E. 2000 Generalization versus specialization in plant pollination systems. TREE 15, 140–143. (doi:10.1016/S0169-5347(99)01811-X) ● Kiviniemi, K. & Eriksson, O. 2002 Size-related deterioration of semi-natural grassland fragments in Sweden. Div. Dist. 8, 21–29. (doi:10.1046/j.1366-9516.2001. 00125.x) ● Kleijn, D. & Raemakers, I. 2008 A retrospective analysis of pollen host plant use by stable and declining bumble bee species. Ecology 89, 1811–1823. (doi:10.1890/07- 1275.1) ● Klein, A.-M., Steffan-Dewenter, I. & Tscharntke, T. 2003 Pollination of Coffea canephora in relation to local and regional agroforestry management. J. Appl. Ecol. 40, 837–845. (doi:10.1046/j.1365-2664.2003.00847.x) ● Klein, A.-M., Vaissie`re, B. E., Cane, J. H., Steffan- Dewenter, I., Cunningham, S. A., Kremen, C. & Tscharntke, T. 2007 Importance of pollinators in changing landscapes for world crops. Proc. R. Soc. B. 274, 303–313. (doi:10.1098/rspb.2006.3721) ● Kotiaho, J. S., Kaitala, V., Komonen, A. & Pa¨ivinen, J. 2005 Predicting the risk of extinction from shared ecological characteristics. Proc. Natl Acad. Sci. USA 102, 1963–1967. (doi:10.1073/pnas.0406718102) ● Krauss, J., Steffan-Dewenter, I. & Tscharntke, T. 2003 How does landscape context contribute to effects of habitat fragmentation on diversity and population density of butterflies? J. Biogeogr. 30, 889–900. (doi:10.1046/j. 1365-2699.2003.00878.x) ● Krauss, J., Alfert, T. & Steffan-Dewenter, I. 2009 Habitat area but not habitat age determines wild bee richness in limestone quarries. J. Appl. Ecol. 46, 194–202. (doi:10. 1111/j.1365-2664.2008.01582.x) ● Littell, R. C., Milliken, G. A., Stroup, W. W., Wolfinger, R. D. & Schabenberger, O. 2006 SAS for mixed models, 2nd edn. Cary, NC: SAS Institute Inc. Lomolino, M. V. 1984 Mammalian island biogeography: effects of area, isolation and vagility. Oecologia 61, 376– 382. (doi:10.1007/BF00379638) ● McKinney, M. L. & Lockwood, J. L. 1999 Biotic homogenization: a few winners replacing many losers in the next mass extinction. TREE 14, 450–453. (doi:10.1016/ S0169-5347(99)01679-1) ● Michener, Ch. D. 2000 The bees of the world. Baltimore, MD: Johns Hopkins University Press. Montoya, D., Zavala, M. A., Rodriguez, M. A. & Purves, D. W. 2008 Animal versus wind dispersal and the robustness of tree species to deforestation. Science 320, 1502–1504. (doi:10.1126/science.1158404) ● Moretti, M., de Bello, F., Roberts, S. P. M. & Potts, S. G. 2009 Taxonomical vs. functional responses of bee communities to fire in two contrasting climatic regions. J. Anim. Ecol. 78, 98–108. (doi:10.1111/j.1365-2656. 2008.01462.x) ● Öckinger, E. & Smith, H. G. 2006 Landscape composition and habitat area affect butterfly species richness. Oecologia 149, 526–534. (doi:10.1007/s00442-006-0464-6) ● Öckinger, E. & Smith, H. G. 2007 Semi-natural grasslands as population sources for pollinating insects in agricultural landscapes. J. Appl. Ecol. 44, 50–59. (doi:10.1111/j.1365-2664.2006.01250.x) ● Öckinger, E., Franze´n, M., Rundlo¨ f, M. & Smith, H. G. 2009 Mobility-dependent effects on species richness in fragmented landscapes. Basic Appl. Ecol. 10, 573–578. (doi:10.1016/j.baae.2008.12.002) ● Pöyry, J., Paukkunen, J., Helio¨ la¨, J. & Kuussaari, M. 2009 Relative contributions of local and regional factors to species richness and total density of butterflies and moths in semi-natural grasslands. Oecologia 160, 577– 587. (doi:10.1007/s00442-009-1328-7) ● Prugh, L. R., Hodges, K. E., Sinclair, A. R. E. & Brashares, J. S. 2008 Effect of habitat area and isolation on fragmented animal populations. Proc. Natl Acad. Sci. USA 105, 20 770–20 775. (doi:10.1073/pnas.0806080105) ● Rand, T. A., Tylianakis, J. M. & Tscharntke, T. 2006 Spillover edge effects: the dispersal of agriculturally subsidized insect natural enemies into adjacent natural habitats. Ecol. Lett. 9, 603–614. (doi:10.1111/j.1461- 0248.2006.00911.x) ● Roland, J. & Taylor, P. D. 1997 Insect parasitoid species respond to forest structure at different spatial scales. Nature 386, 710–713. (doi:10.1038/386710a0) ● Rosenzweig, M. L. 1995 Species diversity in space and time. Cambridge, MA: Cambridge University Press. Roy, D. B., Quinn, R. M. & Gaston, K. J. 1998 Coincidence in the distribution of butterflies and their food plants. Ecography 21, 279–289. (doi:10.1111/j.1600-0587.1998. tb00565.x) ● Rundlöf, M., Nilsson, H. & Smith, H. G. 2008 Interacting effects of farming practice and landscape context on bumble-bees. Biol. Cons. 141, 417–426. SAS. 2009 SAS Institute. Cary, NC, USA. Steffan-Dewenter, I. & Tscharntke, T. 2000 Butterfly community structure in fragmented habitats. Ecol. Lett. 3, 449–456. (doi:10.1111/j.1461-0248.2000.00175.x) ● Steffan-Dewenter, I. & Tscharntke, T. 2002 Insect communities and biotic interaction on fragmented calcareous Wild bee responses to habitat loss R. Bommarco et al. 7 Proc. R. Soc. B Downloaded from rspb.royalsocietypublishing.org on March 10, 2010 grasslands—a mini review. Biol. Cons. 104, 275–284. (doi:10.1016/S0006-3207(01)00192-6) ● Steffan-Dewenter, I.,Mu¨nzenberg, U., Bu¨ rger, C., Thies, C. & Tscharntke, T. 2002 Scale-dependent effects of landscape context on three pollinator guilds. Ecology 83, 1421–1432. (doi:10.1890/0012-9658(2002)083[1421:SDEOLC]2.0. CO;2) ● Steffan-Dewenter, I., Klein, A. M., Gaebele, V., Alfert, T. & Tscharntke, T. 2006 Bee diversity and plant–pollinator interactions in fragmented landscapes. In Plant–pollinator interactions: from specialization to generalization (eds N. M. Waser & J. Ollerton), pp. 387–407. Chicago, IL, USA: University of Chicago Press. Swihart, R. K., Gehring, T. M., Kolozsvary, M. B. & Nupp, T. E. 2003 Responses of ‘resistant’ vertebrates to habitat loss and fragmentation: the importance of niche breadth and range boundaries. Div. Dist. 9, 1–18. (doi:10.1046/ j.1472-4642.2003.00158.x) ● Thomas, C. D. 2000 Dispersal and extinction in fragmented landscapes. Proc. R. Soc. Lond. B. 267, 139–145. (doi:10. 1098/rspb.2000.0978) ● Thomas,C.D. & Hanski, I. 2004Metapopulation dynamics in changing environments: butterfly responses to habitat and climate change. In Ecology, genetics and evolution of metapopulations. (eds I. Hanski & O. E. Gaggiotti), pp. 489–514. Amsterdam, The Netherlands: Elsevier Academic Press. Thomas, J. A., Bourn, N. A. D., Clarke, R. T., Stewart, K. E., Simcox, D. J., Pearman, G. S., Curtis, R. & Goodger, B. 2001 The quality and isolation of habitat patches both determine where butterflies persist in fragmented landscapes. Proc. R. Soc. Lond. B. 268, 1791–1796. (doi:10. 1098/rspb.2001.1693) ● Westphal, C., Steffan-Dewenter, I. & Tscharntke, T. 2003 Mass flowering crops enhance pollinator densities at a landscape scale. Ecol. Lett. 6, 961–965. (doi:10.1046/j. 1461-0248.2003.00523.x) ● Westrich, P. 1990 Die Wildbienen Baden-Wu¨ rttembergs. Allgemeiner Teil. Stuttgart, Germany: Ulmer Verlag. Winfree, R., Aguilar, R., Va´zquez, D. P., LeBuhn, G. & Aizen, M. A. 2009 A meta-analysis of bees’ responses to anthropogenic disturbance. Ecology 90, 2068–2076. (doi:10.1890/08-1245.1)