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View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by White Rose Research Online promoting access to White Rose research papers Universities of Leeds, Sheffield and York http://eprints.whiterose.ac.uk/ This is the author’s pre-print version of an article published in Advances in Ecological Research White Rose Research Online URL for this paper: http://eprints.whiterose.ac.uk/id/eprint/76090 Published article: Hagen, M, Kissling, WD, Rasmussen, C, Carstensen, DW, Dupont, YL, Kaiser- Bunbury, CN, O'Gorman, EJ, Olesen, JM, De Aguiar, MAM, Brown, LE, Alves- Dos-Santos, I, Guimarães, PR, Maia, KP, Marquitti, FMD, Vidal, MM, Edwards, FK, Genini, J, Jenkins, GB, Trøjelsgaard, K, Woodward, G, Jordano, P, Ledger, ME, Mclaughlin, T, Morellato, LPC and Tylianakis, JM (2012) Biodiversity, Species Interactions and Ecological Networks in a Fragmented World. Advances in Ecological Research, 46. 89 - 120. ISSN 0065-2504 http://dx.doi.org/10.1016/B978-0-12-396992-7.00002... White Rose Research Online [email protected] 1 Biodiversity, species interactions and ecological networks 2 in a fragmented world 3 4 MELANIE HAGEN1, W. DANIEL KISSLING1, CLAUS RASMUSSEN1, MARCUS A. M. 5 DE AGUIAR2, LEE BROWN3, DANIEL W. CARSTENSEN1, ISABEL ALVES-DOS- 6 SANTOS4, YOKO L. DUPONT1, FRANCOIS K. EDWARDS5, JULIETA GENINI6, 7 PAULO R. GUIMARÃES JR.4, GARETH B. JENKINS7, PEDRO JORDANO8, 8 CHRISTOPHER N. KAISER-BUNBURY1, MARK LEDGER9, KATE P. MAIA4, FLAVIA 9 M. DARCIE MARQUITTI4, ÓRLA MCLAUGHLIN10,11, L. PATRICIA C. MORELLATO6, 10 EOIN J. O’GORMAN7, KRISTIAN TRØJELSGAARD1, JASON M. TYLIANAKIS12, 11 MARIANA MORAIS VIDAL4, GUY WOODWARD7 AND JENS M. OLESEN1* 12 13 Author affiliations 14 1 Department of Bioscience, Aarhus University, Ny Munkegade 114, 8000 Aarhus C, 15 Denmark. 16 2 Instituto de Física Gleb Wataghin, Universidade Estadual de Campinas, 13083-970, 17 Campinas, SP, Brazil. 18 3 School of Geography, University of Leeds, Woodhouse Lane, Leeds, LS2 9JT, UK. 19 4 Departamento de Ecologia, Instituto de Biociências, Universidade de São Paulo, 05508-900, 20 São Paulo, SP, Brazil. 21 5 Centre for Ecology and Hydrology, Maclean Building, Benson Lane, Crowmarsh Gifford, 22 Wallingford, OX10 8BB, UK. 23 6 Departamento de Botânica, Laboratório de Fenologia, UNESP Univ Estadual Paulista, Av. 24 24A, 1515, 13506900, Rio Claro, SP, Brazil. 1 1 7 School of Biological and Chemical Sciences, Queen Mary University of London, Mile End 2 Road, London E1 4NS, UK. 3 8 Integrative Ecology Group, Estación Biológica de Doñana, CSIC, Avda. Americo Vespucio 4 s/n, Isla de La Cartuja, E-41092 Sevilla, Spain 5 9 School of Geography, Earth and Environmental Sciences, University of Birmingham, 6 Edgbaston, Birmingham, B15 2TT, UK. 7 10 Environmental Research Institute, University College Cork, Lee Road, Cork, Ireland. 8 11 School of Biological, Earth, and Environmental Sciences, University College Cork, 9 Distillery Fields, North Mall, Cork, Ireland. 10 12 School of Biological Sciences, University of Canterbury, Private Bag 4800, Christchurch, 11 New Zealand. 12 13 *corresponding author: Jens. M. Olesen, Department of Bioscience, Ny Munkegade 114, DK- 14 8000 Aarhus C, Denmark, phone: +45 8715 6563, fax: +45 8715 4326, e-mail: 15 [email protected] 16 17 18 TOC detailed 19 I. Abstract................................................................................................................................5 20 II. Introduction.........................................................................................................................6 21 III. Networks............................................................................................................................9 22 A. Ecological networks........................................................................................................9 23 1. Properties of mutualistic and antagonistic networks ............................................1211 24 2. Body size as a driver of ecological network structure...............................................13 25 3. Species abundance as a driver of ecological network structure ...............................14 26 4. Functional groups in ecological networks.............................................................1514 2 1 B. Spatial networks........................................................................................................1615 2 C. Combining spatial and ecological networks.............................................................1716 3 IV. Habitat fragmentation..................................................................................................1918 4 A. General introduction.................................................................................................1918 5 B. Fragment characteristics ...............................................................................................20 6 C. Habitat edges.............................................................................................................2221 7 D. Matrix ...........................................................................................................................25 8 E. Spatial and temporal turnover of species and individuals.............................................27 9 F. Scales of habitat fragmentation.................................................................................2928 10 V. Habitat fragmentation and species traits ......................................................................3231 11 A. Plant traits.................................................................................................................3332 12 B. Animal traits ................................................................................................................3534 13 C. Species trait combinations...........................................................................................3736 14 VI. Habitat fragmentation and biotic interactions .............................................................3938 15 A. Mutualistic plant–pollinator interactions..................................................................3938 16 B. Mutualistic plant–frugivore interactions...................................................................4140 17 C. Mutualistic plant–ant interactions.............................................................................4342 18 D. Antagonistic interactions within food webs .................................................................44 19 E. Antagonistic host–parasitoid interactions .................................................................4645 20 F. Summary of fragmentation effects on mutualistic and antagonistic interactions .....4847 21 VII. Effects of habitat fragmentation on different kinds of networks................................5049 22 A. General introduction.................................................................................................5049 23 B. Mutualistic plant–pollinator networks ......................................................................5049 24 C. Mutualistic plant–frugivore networks...........................................................................59 25 D. Mutualistic plant–ant networks.....................................................................................62 26 E. Antagonistic food webs.................................................................................................63 27 F. Antagonistic host–parasitoid networks .........................................................................67 3 1 G. General effects of habitat fragmentation on network properties..................................68 2 VIII. Habitat fragmentation in a meta-network context.........................................................70 3 A. Meta-networks and dispersal....................................................................................7271 4 B. Meta-networks and extinction ......................................................................................73 5 C. Meta-networks and colonization...................................................................................74 6 IX. Effects of habitat fragmentation on the coevolutionary dynamics of networks..............76 7 A. The geographic mosaic theory of coevolution (GMTC) ..............................................76 8 B. Habitat fragmentation and its effects on basic components of GMTC.........................77 9 C. Habitat fragmentation and selection mosaics in ecological networks ..........................78 10 X. Applications in conservation and agriculture ...................................................................81 11 XI. Conclusions .....................................................................................................................84 12 Acknowledgements..............................................................................................................8786 13 References................................................................................................................................87 14 Glossary. ..........................................................................................................................155153 15 16 17 4 1 2 I. ABSTRACT 3 Biodiversity is organized into complex ecological networks of interacting species in local 4 ecosystems, but our knowledge about the effects of habitat fragmentation on such systems 5 remains limited. We consider the effects of this key driver of both local and global change on 6 both mutualistic