Alien Plant Species Do Have a Clear Preference for Different Land Uses Within Urban Environments
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Urban Ecosystems (2018) 21:1189–1198 https://doi.org/10.1007/s11252-018-0792-4 Alien plant species do have a clear preference for different land uses within urban environments Sandrine Godefroid1,2,3 & Carlo Ricotta4 Published online: 16 August 2018 # Springer Science+Business Media, LLC, part of Springer Nature 2018 Abstract Since neophytes can become invasive in the future, untangling their ecological preferences is of paramount importance, espe- cially in urban areas where they represent a substantial proportion of the local flora. Studies exploring alien species assemblages in urban environments are however scarce. This study aims to unravel alien plant species preferences for five urban land uses (densely built-up areas, open built-up areas, industrial areas, broadleaved urban forests, and agricultural areas and small land- scape elements). We took the city of Brussels as a model, in which we recorded all vascular species growing spontaneously in grid cells of 1 km2. We tested two different ways of classifying the 1-km2 cells: (1) We simply associated each cell with its dominant land cover; (2) We used a fuzzy approach for which the degree of association of a given cell to a given land cover depended on the proportion of that land cover within the cell. For both classification methods, we calculated the indicator species of the resulting land cover types based on alien species only. The crisp and fuzzy classifications identified 33 and 49 species, respectively, with a clear preference for some urban land use types (from a total of 129 alien plant species analyzed). Results showed that urban land use types having apparently similar environmental conditions can actually harbor different neophyte assemblages. Fine-tuning the categorization of urban environments in future ecological studies is therefore important for under- standing spatial patterns of alien species occurrence. Keywords Exotic plants . Neophytes . Indicator species . Brussels . Land use classification . Ellenberg’sindicators Introduction ecosystems through their impacts on resident plant and animal communities, soil biota and nutrient concentrations (e.g. The number of alien species increased over recent decades in Vanderhoeven et al. 2005; Morales and Traveset 2009; Europe (Butchart et al. 2010), especially in urban areas where Maurel et al. 2010;Pyšek et al. 2012; Downey and they represent on average 28% of the local flora (Aronson et Richardson 2016). They may also be responsible for species al. 2014). Alien species are able to influence the functioning of extinction. For instance, five plant species recorded as extinct on the IUCN red list have exotic plant species listed as one of the causes of their extinction (IUCN 2015). * Sandrine Godefroid [email protected] Cities impose strong environmental filters on the urban flora (Ricotta et al. 2009; Williams et al. 2009;Aronsonet al. 2016). In spite of this, it has been demonstrated on several 1 Research Department, Botanic Garden Meise, Nieuwelaan 38, occasions that cities are species-rich (e.g. Kühn et al. 2004a). 1860 Meise, Belgium The affinity of several alien plant species towards urban areas 2 ’ Service général de l Enseignement supérieur et de la Recherche has been shown previously. For instance, an urbanity scale scientifique, Fédération Wallonie-Bruxelles, rue A. Lavallée 1, 1080 Brussels, Belgium ranging from urbanophilic to urbanophobic depending on the species’ presence or absence in cities has been developed 3 Laboratory of Plant Ecology and Biogeochemistry, Université libre de Bruxelles, CP 244, Boulevard du Triomphe, for 3659 taxa of the European flora (Klotz et al. 2002;Kühnet 1050 Brussels, Belgium al. 2004b). However, if we want to improve our knowledge of 4 Department of Environmental Biology, University of Rome ‘La the ecology of urbanophilous species we cannot consider the Sapienza’, Piazzale Aldo Moro 5, 00185 Rome, Italy urban environment as a homogeneous entity but rather as a set 1190 Urban Ecosyst (2018) 21:1189–1198 of more or less distinct habitats, e.g. housing areas of different Materials and methods types and densities, transport infrastructures, industrial areas, cemeteries, parks, etc. Study area Very little published work on the distribution of aliens in cities has taken into account sub-categories of the urban envi- The area considered in the present study is the city of Brussels ronment. When considering several urban land-use types in a (50°48’N – 004°21′E; 161 km2; 1.2 million inhabitants). It is city of southern Korea, Zerbe et al. (2004)showedthatrural characterized by a temperate climate with a mean annual tem- residential areas and industrial areas harbor the highest pro- perature of 10.5 °C and a mean annual rainfall of 852 mm portions of non-native species. Preliminary work on the urban (data provided for the period 1981–2010 by the Belgian flora of Brussels city showed the positive relationship between Royal Meteorological Institute). The mean annual tempera- densely built-up areas and the presence of aliens (Godefroid ture of the city has risen by 1.5 °C in the last 40 years and Koedam 2007; Ricotta et al. 2010a). In a comparative (Godefroid 2011). From a structural point of view, the city study across 32 cities in central Europe, Lososová et al. appears as a succession of four concentric zones, from its (2012) found that the highest proportion of alien species was business and historical center to the outlying suburbs (De found in historical squares and boulevards, whereas very few Bruyn and Lannoy 1991 in IBGE-BIM 1995): (1) the core is aliens had been identified in parks. Similarly, in 45 Central dominated by commercial and administrative activities with a European settlements, Čeplová et al. (2017) highlighted that the limited residential function; (2) the districts, constructed in the number of neophytes was higher in residential areas compared to last century, are densely built-up; (3) the periphery, less dense- settlement centers and older successional sites. These studies did ly built-up; and (4) the suburbs which can be considered as the not, however, focus on the requirements of individual species. maximum (or peak) demographic growth zone. This part of This is unfortunate since a better understanding of the ecology of the city still keeps green areas. This structure is in line with the alien species can help to assess the risks to native biodiversity and concentric zone model presented by Burgess (1925)introduc- implement mitigation measures (Polce et al. 2011). ing the complexity of urban land use. If the ecological niche of some alien species is quite well described (e.g. Tiley et al. 1996;Phartyaletal.2009; Flora census Godefroid and Koedam 2010;Chmuraetal.2013), it is be- cause these are among the most invasive plants on earth, The city was divided in 189 grid cells of 1 km2. However, in which calls for urgent research given the economic impact this study, only 159 cells which are included in the adminis- that these species have on their environment. The situation is trative limits of the city for at least 75% of their area were quite different for those alien species which are not (yet) re- considered. Within each of these 1 km2-cells, we recorded all corded as invasive. These can however become invasive after spontaneous (growing naturally) species of the vascular flora staying for decades in relatively low numbers (Crooks 2005) between March and October during 3 years (1992–1994). or under future climate warming scenarios (Ricotta et al. With the exception of private gardens, the whole city area 2010b). It is therefore imperative to try to understand the eco- has been surveyed, including managed areas (e.g., road logical needs of as many exotic species as possible. In doing verges, parks, lawns), sometimes requiring a special permit this, we cannot rely on the ecological data available for the (e.g. in the case of train stations or properties of the royal area of origin of the species since they may behave differently family). In order to avoid under-sampling because of the in their native vs. introduced areas (Kudoh et al. 2007;Hierro seasonal variation, each 1 km2-cell was surveyed twice along et al. 2009;Beckmannetal.2011). the growing season (early spring and summer or early au- This paper aims to fine-tune our knowledge of alien species tumn). Upon each visit, the field survey was long enough to preferences for different land uses within urban environments. reach the point where it was difficult to add further species to To this end, we took the city of Brussels as a model, in which the list. we recorded the flora in grid cells of 1 km2.Wetestedtwo Each species was then classified as alien or native, where different ways of classifying the 1-km2 cells in different urban alien species are defined according to Pyšek et al. (2004)as environments: (1) We simply associated each cell with its those introduced to the region as a result of human activities dominant land cover; (2) We used a fuzzy approach for which and successfully naturalized. Only alien species introduced the degree of association of a given cell to a given land cover after AD1500 were considered in this study. This date rep- depended on the proportion of that land cover within the cell. resents the discovery of the New World and the start of a Next, for both classification methods we calculated the indi- period of radical change in human movement, commerce cator species of the resulting land cover types based on alien and industry. species only. We believe that the proposed approach could Soil nutrient availability (N), soil reaction (R), soil mois- help to identify indicator species of various urban environ- ture (F) and light (L) in each cell were estimated using ments among the alien flora.