Distance Decay of Similarity Among European Urban Floras
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Global Ecology and Biogeography, (Global Ecol. Biogeogr.) (2008) 17, 363–371 Blackwell Publishing Ltd RESEARCH Distance decay of similarity among PAPER European urban floras: the impact of anthropogenic activities on β diversity Frank A. La Sorte1*, Michael L. McKinney2, Petr Pyßek3,4, Stefan Klotz5,6, G.L. Rapson7, Laura Celesti-Grapow8 and Ken Thompson9 1Division of Biological Sciences, University of ABSTRACT California, San Diego, La Jolla, CA 92093, USA, Aim We examine how two categories of non-native species (archaeophyte and 2Department of Earth and Planetary Sciences, neophyte, introduced before and after 1500, respectively) have had different University of Tennessee, Knoxville, TN 37996, β USA, 3Institute of Botany, Academy of Sciences impacts on diversity across European urban floras. Our goal is to use the unique of the Czech Republic, CZ-252 43 Pruhonice, biological perspective provided by urban areas, and the contrasting historical and Czech Republic, 4Department of Ecology, geographical perspectives provided by archaeophytes and neophytes, to infer how β Faculty of Science, Charles University, Vini4ná non-native species will impact upon diversity in the future. 7, CZ-128 01 Praha 2, Czech Republic, Location Twenty-two urban areas located in seven European countries. 5UFZ – Helmholtz Centre for Environmental Research, Department of Community Ecology Methods We used the β-sim dissimilarity index to estimate the level of β diversity (BZF), Theodor-Lieser-Strasse 4, D-06120 for 231 unique pair-wise combinations of 22 urban floras. We examined bivariate Halle, Germany, 6Virtual Institute for plots of dissimilarity by geographical separation of city centres to evaluate distance Macroecology, Theodor-Lieser-Strasse 4, decay of similarity for native species, archaeophytes and neophytes. D-06120 Halle, Germany, 7Ecology Group, Results Based on average percentages, 52.8% (SD = 8.2%) of species in the urban Institute of Natural Resources, Massey University, Palmerston North, New Zealand, floras were identified as non-native with 28.3% (SD = 6.9%) classified as neophytes 8Department of Plant Biology, Sapienza and 24.5% (SD = 4.9%) as archaeophytes. Relative to native species, across urban floras, University of Rome, Piazzale Aldo Moro 5, archaeophytes were associated with higher compositional similarity and weaker I-00185, Rome, Italy, 9Department of Animal distance decay patterns, whereas neophytes were associated with lower compositional and Plant Sciences, University of Sheffield, similarity and stronger distance decay patterns. Sheffield S10 2TN, UK Main conclusions Across European urban floras, archaeophytes and neophytes occurred in similar numbers but archaeophytes were consistently associated with lower β diversity and neophytes with higher β diversity. Thus, the impact of non-native species on β diversity can be determined, at least in part, through their historical and geographical associations with anthropogenic activities. If archaeophytes represent the long-term biogeographical outcome for human commensal species, neophytes could develop similar patterns. The consequences, however, are likely to be more substantial ecologically and geographically due to the increasing numbers of neophytes and their global anthropogenic associations. Nevertheless, at present, our findings suggest that, based on occurrence information, neophytes have not achieved this state with European urban floras retaining regionally distinct assemblages of neophytes. *Correspondence: Frank A. La Sorte, Ecology, Behavior and Evolution Section, Division of Keywords Biological Sciences, University of California, Anthropogenic activities, archaeophyte, beta diversity, biotic homogenization, San Diego, 9500 Gilman Drive, MC 0116, La Jolla, CA 92093, USA. compositional similarity, distance decay, Europe, neophyte, non-native species, E-mail: fl[email protected] urban floras. having consequences for the composition, structure and INTRODUCTION function of ecological communities (Vitousek et al., 1997; Mack Anthropogenic activities are increasingly disrupting biogeo- et al., 2000; Lockwood, 2004; Hobbs & Mooney, 2005; McNeely, graphical barriers to dispersal, resulting in the spread and 2005). Two outcomes of this process that are commonly investigated establishment of species beyond their historical ranges, thereby include changes in biological diversity within (species richness or © 2007 The Authors DOI: 10.1111/j.1466-8238.2007.00369.x Journal compilation © 2007 Blackwell Publishing Ltd www.blackwellpublishing.com/geb 363 F. A. La Sorte et al. α diversity) and among (species turnover or β diversity; in determining local patterns in urban floras (Celesti-Grapow Whittaker, 1972) species assemblages. Within assemblages at et al., 2006), habitat structure across cities tends to share many local and regional extents there is evidence that species richness commonalities that are distinct from the surrounding landscape. has increased for many taxonomic groups through the addition This island-like characteristic of urban areas makes them suitable of new species (Sax & Gaines, 2003; Sax et al., 2005). This for evaluating the broad-scale impact of the intentional and includes the addition of native and non-native species that, in unintentional translocation of species by humans. Therefore, by many cases, have benefited directly or indirectly from anthropogenic focusing on urban areas – particularly in Europe which contains activities (e.g. La Sorte & Boecklen, 2005). Among assemblages, some of the oldest and best studied examples of these environ- the outcome for β diversity is more complex and is dependent on ments (Sukopp, 2002; Antrop, 2004) – we can assess some of the a combination of factors, including the anthropogenic, biological long-term consequences of the globalization of the Earth’s biota and geographical characteristics of the region under consideration. (Vitousek et al., 1997) and the transformation of landscapes into This includes, for non-native species found within a region, their urban environments (Berry, 1990; Douglas, 1994). time of introduction and place of origin (McKinney, 2005; La In this study, we examine β diversity for native species, archaeo- Sorte & McKinney, 2006; La Sorte et al., 2007). An outcome of phytes and neophytes across 22 major urban areas located in particular interest is the broad-scale decline in β diversity associated seven European countries. We use distance decay of similarity with the loss of species with narrow distributions in conjunction (Nekola & White, 1999; McKinney, 2004; Soininen et al., 2007) with the acquisition of widely distributed species (biotic or to represent these patterns and we compare the outcome with an taxonomic homogenization; McKinney & Lockwood, 1999). analysis conducted on urban floras in the north-eastern United For vascular plants in European floras, there is evidence that States (La Sorte & McKinney, 2006). The goal of this study is to non-native species have had a different impact on β diversity use the unique biological perspective provided by European based on their time of introduction and place of origin. urban areas and the contrasting historical and geographical Specifically, across European floras, non-native species that were perspectives provided by archaeophytes and neophytes in Europe introduced before 1500 (archaeophytes) have lower β diversity, to infer how non-native species, in association with expanding and assemblages of non-native species that were introduced after anthropogenic activities, will impact upon β diversity or spatial 1500 (neophytes; see Pyßek et al., 2004a for definitions) have patterns of diversity in the future. higher β diversity (Kühn et al., 2003; Kühn & Klotz, 2006; La Sorte et al., 2007). The year 1500 signifies the initiation of MATERIALS AND METHODS European exploration of North and South America and the transition from regional (Mediterranean Basin and south- We assembled urban floras for 22 European cities located in eastern European steppes) to global (primarily North America seven countries (Fig. 1 & Table 1). Twenty-one of the cities were and Asia) origins of non-native species in Europe. Therefore, located between latitude 49° and 55° N. Rome, Italy was an contrasting patterns of β diversity for archaeophytes and outlier occurring at latitude 41° N (Fig. 1). The exclusion of neophytes reflect, at least in part, differences in their historical Rome from the analysis did not alter our conclusions, thus our and geographical associations with anthropogenic activities. analysis considered all 22 cities. For each flora, we combined all That is, archaeophytes have an ancient association functioning, varieties and subspecies into single species. We then standardized at least initially, at a regional extent, and neophytes have a modern the taxonomic nomenclature using the Missouri Botanical association functioning at a global extent. These contrasting Garden’s TROPICOS data base and the program TS, perspectives provide a unique opportunity to study the long- version 1.2 (Boyle, 2004) resulting in a data base of 4108 species. term and broad-scale impact of non-native species on β diversity. We then designated each species as native (indigenous, auto- The structure and composition of natural communities within chthonous) or non-native (non-indigenous, alien, exotic, urban areas represent the consequences of intensive anthropogenic allochthonous) within each flora (Pyßek et al., 2004a) using three activities. Specifically, urban areas act as the focal point for