This is a repository copy of Land-use simplification weakens the association between terrestrial producer and consumer diversity in Europe. White Rose Research Online URL for this paper: http://eprints.whiterose.ac.uk/108853/ Version: Accepted Version Article: Dainese, M, Isaac, NJB, Powney, GD et al. (11 more authors) (2017) Land-use simplification weakens the association between terrestrial producer and consumer diversity in Europe. Global Change Biology, 23 (8). pp. 3040-3051. ISSN 1354-1013 https://doi.org/10.1111/gcb.13601 This article is protected by copyright. All rights reserved. This is the peer reviewed version of the following article: Dainese, M., Isaac, N. J. B., Powney, G. D., Bommarco, R., Öckinger, E., Kuussaari, M., Pöyry, J., Benton, T. G., Gabriel, D., Hodgson, J. A., Kunin, W. E., Lindborg, R., Sait, S. M. and Marini, L. (2016), Landscape simplification weakens the association between terrestrial producer and consumer diversity in Europe. 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[email protected] https://eprints.whiterose.ac.uk/ 1 Land-use simplification weakens the association between terrestrial producer 2 and consumer diversity in Europe 3 4 Running head: Diversity associations and land-use change 5 6 Matteo Dainese1,2,*, NickF J. B. Isaac3, Gary D. Powney3, Riccardo Bommarco4, Erik Öckinger4, 7 Mikko Kuussaari5, Juha P,yryo 5, Tim G. Benton6, Doreen Ga.riel7, Jenny 0. Hodgson2, William 2 E. Kunin6, Reginar 4indborg5, Steven M. Sait6, 4oren7o Marini1 5 Re 10 1D08NAE, University of Padova, Viale dell’9niversit=v 16, 35020 4egnaro, Padova, Italy 2 e 11 Department of Animal Ecology and Tropical Biology,w Biocenter, University of [email protected], 12 0m Hubland, 97074 [email protected], Germany 3 O 13 Natural Environment Research Council (NERAC Centre for Ecologn y and 1ydrology, Benson 14 4ane, Crowmarsh Gifford, OEfordshire, OF10 8BB, UK l 15 4Department of Ecology, Swedish University of 0gricultural Sciences,y SE750 07 Uppsala, 16 Sweden 17 5Natural Environment Centre, Finnish Environment Institute, PO BoE 140, FI-00251 Helsinki, 12 8inland 6 15 School of Biology, Faculty of Biological Sciences, 9niversity of 4eeds, Leeds, 4S2 9JT, UK 1 20 7Institute of Crop and Soil Science, Julius K@hn-Institut (JKIC, 8ederal Research Centre for 21 Aultivated Plants, Bundesallee 50, D-32116 Braunschweig, Germany 22 2Department of Evolution, Ecology and Behaviour, Biosciences Building, Arown Street, 23 9niversity of 4iverpool, 9K 5 24 Department of Physical Geography and Quaternary Geology, Stockholm University, SE106 91 25 Stockholm, Sweden 26 *Corresponding author:For Matteo ReviewDainese, Tel.: +45 (0C531 Only 31-22733, Email: 27 matteo.daineseIuni-wuer7.urg.de 22 25 eywords: biodiversity loss, co-evolution, her.ivory, ecosystem resilience, functional and 30 phylogenetic diversity, ha.itat loss, host speciali7ation, land-use change, plant-insect diversity, 31 trophic associations 32 33 Paper type: Primary Research 0rticles 2 34 Abstract 35 4and-use change is one of the primary drivers of species loss, yet little is known a.out its effect 36 on other components of biodiversity that may be at risk, such as local associations between 37 trophic levels. Here, we ask whether, and to what eEtent, landscape simplification, measured as 32 the percentage of ara.le land in the landscape, disrupts the functional and phylogenetic 35 association between plants and primary consumers. Across seven European regions, we inferred 40 the potential associationsFor (functional Review and phylogeneticC betweenOnly host plants and butterflies in 41 561 semi-natural grasslands. 4ocal plant diversity showed a strong bottom-up effect on butterfly 42 diversity in the most compleE landscapes, but this effect disappeared in simple landscapes. The 43 functional associations between plant and butterflies are, therefore, the results of processes that 44 act not only locally but are also dependent on the surrounding landscape conteEt. Similarly, 45 landscape simplification reduced the phylogenetic congruence among host plants and butterflies 46 indicating that closely related butterfly species are more generalist in the potential resource 47 lineages used. These processes occurred without any detecta.le change in species richness of 42 plants or butterflies along the gradient of ara.le land. The structural properties of ecosystems are 45 eEperiencing substantial erosion, with potentially pervasive effects on ecosystem functions and 50 future evolutionary traJectories. 4oss of interacting species might trigger cascading eEtinction 51 events and reduce the sta.ility of trophic interactions, as well as influence the longer-term 52 resilience of ecosystem functions. This underscores a growing reali7ation that species richness is 53 a crude and insensitive metric and that both functional and phylogenetic associations, measured 54 across multiple trophic levels, are likely to provide far deeper insights into the resilience of 55 ecosystems, and the functions they provide 3 56 Introduction 57 4and-use simplification has emerged as one of the fundamental components of global change 52 (8oley et al., 2005K Turner II et al., 2007K Ver.urg et al., 2011K Allan et al., 2015K New.old et 55 al., 2015C. Ecology has provided ample scientific evidence that decreasing ha.itat heterogeneity 60 and increasing fragmentation, e.g. through agricultural eEpansion and intensification (a process 61 often termed Llandscape simplificationMC (Meehan et al., 2011C, are main anthropogenic drivers 62 of biodiversity loss (TscharntkeFor etReview al., 2012C. However, biodiversityOnly science has largely focused 63 on species richness loss, underplaying other components of biodiversity that may be at risk of 64 landscape simplification (Valiente-Banuet et al., 2015C. Traditionally, studies have focused on a 65 single trophic level, when instead the biodiversity loss at a given trophic level may also affect 66 other levels, and, hence the associated diversity relationships (Duffy et al., 2007K Scher.er et al., 67 2010C. 0ssociations between trophic levels can have a large impact on community responses to 62 global change (Duffy, 2002K Cardinale et al., 2012K Oliver et al., 2015C. 4oss of interacting 65 species can trigger cascading eEtinction events and reduce the sta.ility of trophic interactions 70 (Dunne et al., 2002K Haddad et al., 2011C, as well as influence the longer-term resilience of 71 ecosystem functions (Oliver et al., 2015C 72 In many human-managed landscapes that are characteri7ed by fragmented ha.itats, the 73 resource base for consumers is often scattered across space (Tscharntke & Brandl, 2004K Winfree 74 et al., 2011C. Because consumer insects are generally highly mobile and affected by land use 75 change, landscape simplification can also alter relationships between the diversity of different 76 taEa (Tscharntke et al., 2012K Weiner et al., 2014C. Our understanding of these associations is 77 mainly based on analyses of manipulative eEperiments (e g. Haddad et al., 2005K Scher.er et al., 72 2010C or studies at the local scale (e g. Manning et al., 2015C, while empirical data considering 4 75 the effect of land-use change at larger spatial scales are largely missing. For instance, it remains 20 less clear how local associations between producer and consumer diversity are affected by 21 landscape simplification Nevertheless, focusing on the conservation status of local scale trophic 22 associations can provide early diagnosis of the functional conseOuences of .iodiversity loss due 23 to global scale change (Valiente-Banuet et al., 2015K Harvey et al., 2016C 24 For Review Only 25 26 Figure 1. Schematic representation of potential associations .etween plants and butterflies and the 27 expected landscape effect on these associations. (aC Butterflies have distinctive functional links with 22 plants: they feed on plant tissues as larvae and on nectar as adults. As adult butterflies show low 25 speciali7ation with flower resources (Rosas-Guerrero et al., 2014C we hypothesi7ed that butterfly 50 a.undance depends on the species richness of flowering plants and their functional trait composition. The 51 diet breadth of butterfly larvae is more restricted than that of adults due to (.C co-evolution .etween host 52 and consumer (phylogenetically closely related butterflies often prefer to feed on phylogenetically closely 53 related host plantsC. Such functional and phylogenetic associations determine the .ottom-up effect of host 54 plant diversity on butterfly evenness and species richness. (cC As losses of producer-consumer diversity 55 associations may freOuently precede the loss of species, we hypothesi7e a stronger negative effect of 56 landscape simplification