The importance of heterogeneity revisited from a multiscale and multitaxa approach Martin M. Gossner a,⇑ , Stephan Getzin b,1 , Markus Lange c,2 , Esther Pašalic ´ c , Manfred Türke a , Kerstin Wiegand b , Wolfgang W. Weisser a a Terrestrial Ecology Research Group, Department of Ecology and Ecosystem Management, Center for Food and Life Sciences Weihenstephan, Technische Universität München, Hans-Carl-von-Carlowitz-Platz 2, 85354 Freising-Weihenstephan, Germany b Department of Ecosystem Modelling, Büsgen-Institut, Georg-August-University of Göttingen, Büsgenweg 4, 37077 Göttingen, Germany c Institute of Ecology, Friedrich-Schiller-University, Dornburger Str. 159, 07743 Jena, Germany article info Article history: Received 4 January 2013 Received in revised form 16 June 2013 Accepted 26 June 2013 Keywords: Beetles Ecosystem function Multiplicative diversity partitioning q-Metric Spatial scale Species turnover abstract The importance of spatial scale for b-diversity has been shown in several studies, but it is unclear how spatial diversity patterns correlate among different organismic groups. We studied spatial diversity orga- nization of plants and several trophic guilds of beetles in beech-dominated forests in two regions of Ger- many to test whether different trophic guilds are organized independently in space. We applied multiplicative diversity partitioning using a nested hierarchical design of four increasingly broader spa- tial levels (subplot, plot, forest class, region) and tested for correlations among trophic guilds by using Pearson product moment correlations and Mantel-tests. We observed similar general diversity patterns at different trophic guilds showing a high contribution of b-diversity to total c-diversity and found b-diversity to be higher at different spatial scales and a-diversity to be lower than expected by random distributions of individuals. Results, however, partly depended on the weighting of rare and abundant species. Beta-diversity in our study was caused mainly by species spatial turnover rather than by nested- ness. Correlations of a-diversity between trophic guilds were low whereas correlations of b-diversity above subplot level were high. Importantly, more strongly connected trophic guilds revealed not gener- ally stronger relationships than less strongly connected guilds. Three important implications for conser- vation can be deduced from our results: (1) heterogeneity of beech forests at different spatial scales should be supported in conservation strategies to enhance biodiversity and related functions; (2) the observed high importance of spatial turnover in relation to nestedness implies a concentration of conser- vation efforts to a large number of not necessarily the richest sites, and (3) recommendation for particular conservation strategies (e.g. selection of priority sites for conservation at regional scale) based on single indicator taxa or functional guild is difficult because of the varied response of the species in our study. Ó 2013 Elsevier Ltd. All rights reserved. 1. Introduction Conserving biodiversity requires detailed knowledge of how diversity is distributed within and between habitats. Starting from Whittaker (1960), an increasing number of studies has emphasized the importance of compositional heterogeneity between places, or beta-diversity, for total biodiversity (gamma-diversity) in a region (Gossner and Müller, 2011; Hirao et al., 2007; Müller and Gossner, 2010; Summerville et al., 2003). Beta-diversity has been shown to be important for understanding broad bio-geographical diversity patterns such as elevational, latitudinal and longitudinal gradients (Kraft et al., 2011; Qian et al., 2005). If b-diversity is high, site selec- tion for conservation presents a formidable challenge. For example, for beech forests and for headwater streams it is crucial to consider complementarity in species composition in the selection of conser- vation target sites as b-diversity contributes greatly to overall diversity (Clarke et al., 2010; Müller et al., 2013). Clough et al. (2007) showed that conservation-orientated evaluation of man- agement schemes in agricultural landscapes needs to include b-diversity, because of its contribution to total diversity at the landscape scale. Importantly, however, measures of species diver- sity including b-diversity are dependent on the spatial scale considered (Gabriel et al., 2006; Gossner and Müller, 2011; 0006-3207/$ - see front matter Ó 2013 Elsevier Ltd. All rights reserved. http://dx.doi.org/10.1016/j.biocon.2013.06.033 ⇑ Corresponding author. Tel.: +49 8161 71 3713; fax: +49 8161 71 4577. E-mail addresses: martin.gossner@tum.de (M.M. Gossner), stephan.getzin@ ufz.de (S. Getzin), mlange@bgc-jena.mpg.de (M. Lange), estherpasalic@googlemail. com (E. Pašalic ´), manfred.tuerke@tum.de (M. Türke), kerstin.wiegand@uni- goettingen.de (K. Wiegand), wolfgang.weisser@tum.de (W.W. Weisser). 1 Present address: UFZ Helmholtz Centre for Environmental Research-UFZ, Depart- ment of Ecological Modelling, PF 500136, D-04301 Leipzig, Germany. 2 Present address: Max Planck Institute for Biogeochemistry, Hans Knöll Str. 10, 07745 Jena, Germany. Biological Conservation 166 (2013) 212–220 Contents lists available at SciVerse ScienceDirect Biological Conservation journal homepage: www.elsevier.com/locate/biocon