Urbanization is one of the most environmentally damaging of human activities, producing large alterations in ecosystem structure, function, species composition and interactions. In this study, we performed a systematic investigation of the plant species richness and density in the city of Beijing, China. We also assessed which socio-economic factors have most influence on plant diversity. Within the city, we found 551 plant species of 313 genera and 103 families, of which 118 were trees, 99 shrubs, and 296 herbs. Nearly half (48.3 %) of the total plant species were aliens. Species richness and density were positively correlated both for tree/shrub and herb taxa, which indicate that although some species predominate in our study area, there is an important array of species in relation to their densities. As expected, most of the socio-economic variables studied showed to be related to at least one of the four plant diversity variables (i.e., herb richness, herb density, tree/shrub richness, and tree/shrub density). Land-use showed a significant relationship in all four cases, which generally had lower values in cultural and education areas (areas that in Beijing are generally characterized by large extensions of urbanized land). The year of establishment was also largely related to plant diversity, with higher values for recently developed areas. This was an expected result given the dynamics of the urban development of Beijing during the last 60 years, which consisted of intense urban sprawling, followed by more environmentally-friendly urbanization practices during the 2000s related to the greening of the city for the olympic games. This dynamics also explained the negative relationship found between the distance to the city center and both tree/shrub species richness and density.
Urbanization represents one of the most environmentally concerning of human activities (Grimm et al. 2008). Through urbanization original vegetation is generally removed and replaced by a different array of plants, mainly comprised by exotic species (Walker et al. 2009; Szlavecz et al. 2011). Although such a novel set of plants is not representative of the original vegetation, it still plays many important roles within urban areas, providing ecological, climatic, socio-economic, and health benefits (Dwyer et al. 1992; Solecki and Welch 1995; Melles et al. 2003; McPherson et al. 2005; Conway and Urbani 2007; Grimm et al. 2008; MacGregor-Fors et al. 2009; Cariñanos and Caseres-Porcel 2011; Kowarik et al. 2011).
The impact of global environmental changes has become a major focus in conservation biology (Vitousek et al. 1997a). The introduction and spread of non-native organisms has been recorded worldwide (Vitousek et al. 1997b; Meng et al. 2004; Liang et al. 2008; Zhao et al. 2010), with anthropogenic land-use change being emphasized as a major cause (Pyšek 1998; Deutschewitz et al. 2003). Although land-use and habitat variables are both crucial components of global environmental change, they are often studied independently (Heikkinen et al. 2006) and analyses considering their interactions are scarce (Jetz et al. 2007). Thus, our understanding of the possible effects of land-use types and socio-economic factors on plant diversity in urban ecosystem is still limited (Didham et al. 2007).
Many previous studies have confirmed a link between urban ecological processes and demographic, socio-economic, and spatial factors in urban areas (Pickett et al. 2008). For example, Iverson and Cook (2000) found a relationship between urban forest cover and household income and human density in Chicago (United States). Similarly, Hope et al. (2003) found a relationship between household wealth and plant diversity in another North American city (Phoenix). Hope et al. (2003) stated that plant diversity in urban areas depends on human preferences for particular landscapes, and the availability of sufficient financial resources to meet this preference, calling it the “luxury effect.” Grove et al. (2006) found that lifestyle behavior was the best predictor of vegetation cover in Baltimore (United States). Troy et al. (2007) found that community housing age, vacant housing rates, population density, and lifestyle were related in some way to urban vegetation cover. Luck et al. (2009) confirmed that human factors influence urban vegetation in their long-term extensive study of residential areas in southeastern Australia, where vegetation diversity was negatively correlated with housing density. Boone et al. (2010) also found that time matters when analyzing plant diversity in urban areas; socio-economic traits dating from almost four decades ago were often better predictors of current urban plant diversity patterns than those from 1 year before. Most recently, Ortega-Álvarez et al. (2011) found that tree species richness follows a hump-shaped pattern along a gradient of urbanization, while tree density is highest in green areas of Mexico City, and again, resident preferences and decisions seem to play a fundamental role in determining the vegetation component of urban systems. Interestingly, tree diversity patterns were very similar between native and exotic species in Mexico City, although the latter represented >60 % of the total recorded species richness.
In this study, we evaluated the role of urban traits in determining the plant diversity (species richness and density) of Beijing, China. We conducted a citywide survey of trees, shrubs, and herbaceous plants, and searched for relationships between plant diversity and several urban parameters, including geographic (i.e., latitude, longitude, distance to city center), land-use, demographic (i.e., human population density), temporal (i.e., year of establishment), and socio-economic (i.e., housing price) variables. As we selected the set of variables based on previous studies performed worldwide, we predict that all these variables would be related in some way to plant diversity in Beijing, with higher plant diversity values as sites are farther from the city center, in old areas and in wealthy locations, and lower plant diversity values in highly populated areas.
Materials and methods
Beijing is a very large urban area located in northeastern China (39°28′–41°25′N; 115°25′–117°130′E; Fig. 1). It has been the capital city of China for 800 years and has a three-millennium history as a human settlement. Beijing has a warm-temperate continental monsoon climate, with an average annual temperature of 11.7 °C and an annual average rainfall of 595 mm, most of which occurs during July and August. The municipality of Beijing covers an area of approximately 16,000 km2, and the city (the four urban core districts plus the four urban expansion districts; e.g., Montgomery 2008) represents about 8.3 % (ca. 1,370 km2) of the municipality’s territory (BISM 2005). As of 2008, the city of Beijing had ca. 589 km2 of green areas, of which approximately 24 % were public, for a total green area per capita of about 56 m2 (MCPRC 1998; BMAC 2006).
The municipality of Beijing is the politic, economic, educational, and cultural center of China, and it is composed by 16 districts and two counties. By the end of 2008, it held an estimated permanent population of 16.95 million inhabitants, whereas the city sheltered a population of 10.44 million residents, with a population density peaking in the Xuanwu district (28,000 people/km2). Such human population density decreases with the distance to the geographic center of the city (BMBS 2009).
Survey site selection and plant surveys
We conducted urban plant surveys within the most urbanized part of Beijing, which is an area encircled by the fifth ring road, covering an area of approximately 750 km2 (Fig. 1). Surveys were carried out in two different years: 2009 and 2011. Sampling sites were randomly selected and stratified by land-uses (see below for land-use definitions). The citywide survey comprised 428 tree and shrub sampling sites, and 1,242 herb sampling sites. We used quadrant surveys to record tree, shrub, and herb species richness and diversity. At each quadrant we recorded the correspondent plant individuals in nested patches (10 × 10 m for trees, 2 × 2 m for shrubs, 1 × 1 m for herbs). We distinguished trees and shrubs by the number of stems (trees = 1, shrubs > 1) and by their height (trees > 1.3 m, shrubs < 1.3 m). Infraspecific taxa (i.e., subspecies, varieties, forms, cultivars) were treated as equivalent to species for ease of analysis. For the identification of the studied plants, we used Flora of China (http://flora.huh.harvard.edu/china/) and Flora of Beijing (He 1992), which are the most authoritative taxonomic works covering our study area; other reliable sources such formerly published papers describing plants in Beijing (Meng et al. 2004; Zhao et al. 2010) were also used as supporting material.
Survey site characterization
We gathered data on spatial traits, land-use types and other factors, such as demographic, historical, and socio-economic variables for all of our sampling sites. Spatial variables included latitude, longitude, location in relation to the peri-urban belt, and distance to the city center. All geographic coordinates were taken from the centroid of each sampling site, from which the distance to the city center was calculated using GIS based on the center of Tian’anmen square (39°54′26.37″N, 116°23′29.22″E). The delimitation of the peri-urban area of the city of Beijing was calculated following a recently proposed method for urban ecology studies (MacGregor-Fors 2010). Based on such methodology, the length (width) of the peri-urban area of the city of Beijing was of 1.29 km.
We classified the land-uses of the city of Beijing following James and Bound’s (2009) categorization, which includes five main types: (1) public service area—including governmental agencies, bus and train stations, urban road trees, urban forests, and attached green spaces, (2) industrial and commercial areas—including enterprises, vegetable fields, plant nurseries, and public institutions, (3) cultural and education areas—including elementary and secondary schools, libraries, and universities, (4) residential areas—including villages, small towns, low-density homesteads (lower than six stories) and high-density homesteads (higher than six stories); and (5) recreation and leisure areas—including parks, public squares, sport centers, hilly areas, and wastelands.
We used data provided by the Beijing Municipal Bureau of Statistics (BMBS 2009) for population density values at each of our sampling sites. We collected data regarding the year of establishment of each sampling site in situ. At each site we interviewed at least 20 residents asking the year of establishment of the area when conducting fieldwork and used an average value. In all cases the variance was very low, reason why we consider our data to be highly reliable. Finally, housing price (yuan/m2) was obtained from China’s Internet Broker website (http://beijing.anjuke.com/), accessed during April and May of 2011.
In this study we considered trees and shrubs, and herbs as different groups, as differences in their nature and the way they were established in the city (e.g., planted, not planted) could obscure our results. Thus, we separated the studied plants in two groups for the analyses: (1) herbs, and (2) trees and shrubs. To seek for possible relationships between herb species richness and density, and between tree/shrub species richness and density, we conducted correlation analyses. We also used linear models to assess which of the measured city variables were related to herb and tree/shrub species richness and density in Beijing, using a statistical significance of P < 0.05. To avoid multicollinearity and to comply with linearity and normality assumptions, we transformed our numeric continuous data before the analyses, and checked for the existence of moderate-to-strong relationships between variables to determine their statistical independence. When two or more variables showed moderate-to-strong significant relationships (r > 0.5, P < 0.05; Peck et al. 2008), we only considered the variable with the highest variance for the analysis. Based on the correlation matrix, we removed two of the nine measured variables (location in relation to the peri-urban belt, and which ring road the sampling sites fall into) with seven remaining for input: (1) land-use, (2) latitude (3) longitude, (4) human population density, (5) year of establishment, (6) housing price, and (7) distance to the geographic center of the city. We performed post hoc range tests to discriminate among statistically different treatments of the categorical variable (i.e., land-use). All analyses were conducted using R (R Development Core Team 2010).
Flora features of Beijing
We recorded a total of 551 plant species belonging to 313 genera and 103 families (Appendix). Of the recorded species, 118 were trees, 99 shrubs, and 296 herbs, which accounted for 21.4, 18.0, and 53.7 % of the total recorded plant species, respectively. The remaining 38 species could not be classified into a single category, as 33 species can be either shrubs or trees, and 5 can be classified as herbs or shrubs (Appendix). Almost half of the total plant species recorded in this study were alien species (266 species, i.e., 48.3 %). The ratio native:alien differed between life-forms. Whereas native herbs predominated (68.9 %), shrubs and trees were mostly exotic (68.4 %). Phytogeographically, many of the plant genera recorded in this study belonged to the North temperate zone (26.5 %), followed by those of pantropical distribution (15.7 %), cosmopolitan (11.5 %), Old World temperate (10.5 %), and the East Asia-North America disjunct genera (7.3 %). Only 7.7 and 1.9 % of the plant genera found in Beijing urban areas were endemic to East Asia and to China, respectively.
The most abundant tree species in the city of Beijing were the Chinese White Poplar (Populus tomentosa Carrière var. tomentosa), Yulan Magnolia (Magnolia denudata Desr.), and Chinese Juniper (Juniperus chinensis L.), while the most abundant shrubs were the Gold-leaf Privet (Ligustrum lucidum W.T. Aiton), Rose (Rosa chinensis Jacq. var. chinensis), Japan Euonymus (Euonymus japonicus Thunb.), and Common Box (Buxus sinica (Rehder & E.H. Wilson) M. Cheng var. sinica). The most abundant herbs were the Common Meadow-grass (Poa pratensis subsp. pratensis), Common Yellow Oxalis (Oxalis stricta L.), Green Bristle Grass (Setaria viridis (L.) P. Beauv. subsp. viridis), and Indian Goosegrass (Eleusine indica (L.) Gaertn.).
Plant diversity drivers in Beijing
Tree/shrub species richness and density were weakly but significantly related (r = 0.30, P < 0.001), while herb species richness and density were highly and significantly related (r = 0.77, P < 0.001). Linear models showed that six of the seven independent variables were significantly related to one of the dependent variables (i.e., tree/shrub and herbaceous plant richness and density). Tree/shrub richness was significantly and negatively related to the distance to the city center, year of establishment, latitude, and longitude, with statistically lower values in public and cultural and education land-uses when compared to residential and recreation and leisure land-uses (F 4,1010 = 6.91, P < 0.001; Table 1a). Tree/shrub density was significantly and negatively related to the distance to the city center and longitude, with statistically higher values in residential areas when compared to the rest of the studied land-uses (F 4,1024 = 7.64, P < 0.001; Table 1b). Herbaceous plant species richness showed to be significantly and negatively related to the year of establishment and latitude, with statistically lower values in cultural and education land-uses (F 4,1020 = 7.68, P < 0.001; Table 1c). In this case, population density showed a positive non-significant trend (P = 0.06). Finally, herbaceous plant density showed to be significantly and negatively related to the year of establishment, latitude, and land-use, with statistically lower values in cultural and education land-uses (F 4,1017 = 4.57, P = 0.001; Table 1d). In this case, the distance to the center of the city showed a negative non-significant trend (P = 0.09).
The integration of social sciences and ecological approaches could enhance the understanding of human drivers that mould biodiversity patterns in urban areas (Cadenasso et al. 2006). Our results show that both social and geographic variables play a crucial role in explaining plant diversity patterns in the city of Beijing. One of the most evident human footprints on Beijing’s flora was the presence of an important group of alien species, which represents 48.3 % of the total recorded plant species. Among the studied variables, land-use was the only one that was significantly related to both herb and tree/shrub richness and density. Other variables with a large influence in urban plant diversity were latitude and year of establishment. Surprisingly, we found no relationship between human population density and the four studied plant diversity variables.
Finding a large proportion of alien species was not surprising, as high amounts of non-native plant species can be found in urban areas across the globe (Pyšek 1998; Xu 1999; Clemants and Moore 2003; Martínez-Carretero 2010; Ortega-Álvarez et al. 2011). In fact, three previous studies focused on the plant diversity of the city of Beijing have reported high proportions of alien plant species (Meng et al. 2004; Liang et al. 2008; Zhao et al. 2010). Although these studies and our work were conducted in different parts of the city and were aimed to answer contrastingly different questions, similar proportions of alien species have been found, with an average of 49.2 % (±SE 2.3). This finding is of particular environmental concern, as the occurrence and the potential invasiveness of alien plant species can be one of the most important factors influencing urban diversity and functioning (Mooney and Hobbs 2000; Kowarik 2008).
Our results show that species richness and density are positively correlated, both for tree/shrub and herb taxa, with a stronger effect for the latter. This correlation has often been found in nature, both in undisturbed and disturbed sites (e.g., Goldberg and Miller 1990; Denslow 1995; Pärtel and Zobel 1999; Stevens and Carson 1999; Bobo et al. 2006), although it seems to be a trade-off within the context of limiting resources and available space (e.g., Niklas et al. 2003) rather than a general rule. The positive significant relationship between plant species richness and density in urban Beijing is difficult to interpret given the high number of exotics present in the city (with most of them probably planted). We may suggest that humans have molded plant diversity in urban areas enhancing both density and species richness (as humans fancy a great spectrum of greening scenarios; Hope et al. 2003; Faeth et al. 2011; Table 2), perhaps in the same way to what occurs under natural conditions (but this should be tested in non-urbanized places around the city). In support of this argument, the stronger correlation for herbs would be due to the fact that most of them are native (nearly 70 %). Some herbaceous vegetation may still represent remnants of the original vegetation of Beijing (Kowarik 2011) and, for example, some urban parks such as Temple of Heaven still harbor some relic plants (Chen and Lin 2006); an alternative explanation is simply that native herbs are preferred to native trees and shrubs because the former are more cheap and their planting is more feasible (H.-F. Wang, pers. obs.).
As predicted, most of the response variables considered in this study were related to at least one of the four plant diversity variables (i.e., tree/shrub and herbaceous plant richness and density). Land-use showed a significant relationship in all four cases (Table 1). However, the differences among the different land-uses varied depending on the plant diversity variable. Both herb species richness and density were lower in cultural and education areas. These land-uses in Beijing are generally characterized by large extensions of urbanized land (H.-F. Wang, pers. obs.), leaving little space for vegetation and greening purposes. This is probably also the reason why tree/shrub species richness values are lower in cultural, education and public areas, and higher in residential and recreation and leisure areas. Interestingly, tree/shrub density values are also significantly higher in residential areas. These patterns may be explained by various factors, including: (1) real estate merchandising strategies aimed to furnish their surroundings, making recreation and leisure areas richer in plants, and (2) the residents’ preference for tree and shrubs, as reported by Iverson and Cook (2000), Hope et al. (2003), and Boone et al. (2010), and Ortega-Álvarez et al. (2011), and which may partly be due to traditional beliefs (Table 2). For example, Juniperus chinensis, Platycladus orientalis (L.) Franco, and Pinus tabuliformis Carrière var. tabuliformis are ubiquitous because these have been traditionally associated with longevity (Profous 1992).
In addition to land-use, two other anthropogenic variables are driving plant diversity in the city of Beijing: (1) year of establishment, and (2) population density. Year of establishment was significantly and negatively related to tree and shrub species richness and herb species richness and density, with higher values in newer areas compared to old ones. This result seems to be related to modern urbanization and landscape design practices, which tend to develop more diverse landscapes than just “green areas” (e.g., Hope et al. 2003). The dynamics of the urban development during the last 60 years (the year of establishment for most of the sampling sites are within this time frame) have also considerably changed in Beijing. In the 1960s, the city began to experience a rapid expansion that reached its zenith during the 1980s and the 1990s when entire residential districts were quickly built to accommodate the migrant population and the relocated people (Gaubatz 1995; Mu et al. 2007). During the last decade, in contrast, great attention has been paid to greening, mostly as a part of the preparation of the city for the 2008 Olympic Games (Wang et al. 2011). Human population density only showed a positive, non-significant trend with one variable, herb species richness. Population density is not always a good predictor of urban plant diversity, with sometimes no relationships between these two parameters although generally a negative correlation is shown (Conway and Urbani 2007). The weak positive relationship between population density and herb species richness in Beijing could be due to the preference of herbs for greening small areas in crowded places instead of tree and shrubs, which are more expensive (Hope et al. 2003; H.-F. Wang, pers. obs.).
The rest of the variables found to partially explain plant diversity in the city of Beijing were geographic (i.e., latitude, longitude, distance to the center of the city). In all cases, latitude and/or longitude showed negative relationships, indicating that both the southern and western part of the city of Beijing have higher plant diversity values. This pattern is completely contrary to the “luxury effect” proposed by Hope et al. (2003), as wealthier areas in Beijing are located in its northern section (a trend that can be traced back to the Qing Dynasty; Haw 2007). This result could be biased toward the presence of high portions of land designated to the cultural and educational land-use in the northern section of the city (almost all the university campuses are located there), where less space is considered for greening purposes when compared to other urban land-uses (e.g., public service, commercial and industrial, residential, recreation, and leisure). Finding a negative significant relationship between the distance to the city center and both tree/shrub species richness and density could also be related to growth history of the city. In the center of the city many tree and shrub species exists, as explained above, due to the traditional behavior of planting them in the yards of the traditional houses (siheyuan). However, modern urbanization strategies used in areas surrounding the old quarters of the city of Beijing left few space for greening purposes since the main purpose was housing the large population growth in the 1970–1990s (Liang et al. 2008; Tang and Kunzmann 2008).
In summary, this study shows how both socio-economic and spatial variables influence plant diversity in the city of Beijing. Based on our findings, we suggest two main urban management and planning activities that could allow new urbanization processes to be more environmentally friendly: (1) including a higher proportion of native plant species when establishing urban green areas, and (2) planning the urbanization of all land-use types considering greening strategies based on landscape ecological principles (e.g., isolation, connectivity).
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We are grateful to Mark Goddard for his valuable comments. We thank Zhi-Xin Zhu and Xiao-Ma Li for their help with the figure, Su Liu (Peking University) for the Appendix plant species checking and Juan-Juan Zhao, Xue-Song Meng and others for providing some original data. This study was funded by the National Natural Science Foundation of China (41030744) and it was supported by funding from the State Key Laboratory of Urban and Regional Ecology.
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Wang, HF., MacGregor-Fors, I. & López-Pujol, J. Warm-temperate, immense, and sprawling: plant diversity drivers in urban Beijing, China. Plant Ecol 213, 967–992 (2012). https://doi.org/10.1007/s11258-012-0058-9
- Urban ecology