Ecological Consequenses of Rapid Urban Expansion

China has been undergoing a period of economic reformand expansion since the late 1970s, accompanied by

rapid and widespread urbanization. During the early 1980s,18% of China’s population lived in cities, but this rose to39% by 2003, while the number of cities increased from 190to 660 (including about 170 cities with a population greaterthan 1 million) over the same time period. From 1980 to2003, the contribution of Chinese cities to gross nationalproduct (GNP) increased from 69.9% to 85.9% (ChineseStatistical Bureau 2004). However, urbanization has alsoled to serious environmental and ecological problems, bothin urban and surrounding areas, including increased air andwater pollution (Briant and Guo 2000; Liu and Diamond

2005; Shao et al. 2006), local climate alteration (Zhou et al.2004), and a major reduction in natural vegetation coverand production (Fang et al. 2003). Urban residents haveexperienced an increase in levels of cholesterol-related dis-eases (Lee 2004) and an overall decline in quality of life.

As the largest and most modern city, Shanghai hasexperienced extensive urban expansion over the pastthree decades (Figure 1a). Between 1975 and 2003, thecity’s population increased from 10.8 million to 13.4 mil-lion (Shanghai Statistic Bureau 2004). The resulting eco-logical consequences of urban sprawl have caused consid-erable concern among scientists and policy makers.Several studies have documented some environmentalimpacts of Shanghai’s increasing urbanization; for exam-ple, an analysis of meteorological data from both the city’s

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RESEARCH COMMUNICATIONS RESEARCH COMMUNICATIONS

Ecological consequences of rapid urbanexpansion: Shanghai, China

Shuqing Zhao1, Liangjun Da2, Zhiyao Tang1, Hejun Fang2, Kun Song2, and Jingyun Fang1*

Since China’s economic reform in the late 1970s, Shanghai, the country’s largest and most modern city, hasexperienced rapid expansion and urbanization. Here, we explore its land-use and land-cover changes, focusingon the impacts of the urbanization process on air and water quality, local climate, and biodiversity. Over thepast 30 years, Shanghai’s urban area and green land (eg urban parks, street trees, lawns) have increased dramat-ically, at the expense of cropland. Concentrations of major air pollutants (eg SO2, NOx, and total suspended par-ticles) were higher in urban areas than in suburban and rural areas. Overall, however, concentrations havedecreased (with the exception of NOx), due primarily to a decline in coal consumption by industry and in pri-vate households. Increased NOx pollution was mainly attributed to the huge increase in the number of vehicleson the roads. Water quality changes showed a pattern similar to that of air quality, with the most severe pollu-tion occurring in urban areas. Differences in mean air temperatures between urban and rural areas alsoincreased, in line with the rapid pace of urban expansion, indicating an accelerating “urban heat island” effect.Urban expansion also led to a decrease in native plant species. Despite its severe environmental problems,Shanghai has also seen major economic development. Managing the tradeoffs between urbanization and envi-ronmental protection will be a major challenge for Chinese policy makers.

Front Ecol Environ 2006; 4(7): 341–346

Authors’ contact details are on p346)

urban and adjacent rural areas has revealed the presenceof a “heat island” effect (higher air temperatures in urbanareas than in the surrounding areas; Chow 1992; Chen etal. 2003). Water quality in the city’s central business dis-trict has been seriously degraded (Ren et al. 2003), and soilin the Baoshan District is often contaminated with lead,zinc, and cadmium (Hu et al. 2004). Ye et al. (2000) havesuggested that the huge increase in the number of motorvehicles associated with the urbanization process has ledto serious human health risks in the city.

To date, however, long-term (spanning both pre- andpost-urbanization periods) and spatially explicit monitor-ing of the urbanization of the entire Shanghai area,together with comprehensive studies of the ecologicalconsequences, have not been conducted. Here, we docu-ment the processes of urbanization in Shanghai between1975 and 2005, through the analyses of satellite-deriveddata regarding land-use and land-cover changes. Weexplore the changes in air and water quality, the urbanheat island effect, and the biodiversity loss, and examinetheir possible associations with urban expansion.

� Data and methods

Site description

Shanghai is located on the coast of the East China Sea, at theestuary of the Yangtze River. The climate is subtropical, withan annual precipitation of 1200 mm and a mean annual tem-perature of 16˚ C. The total area covered by the city is 8010km2 (latitude: 30˚ 40’ to 31˚ 55’ N; longitude: 120˚ 50’ to121˚ 55’ E). In order to examine the ecological consequences

Urbanization in Shanghai SQ Zhao et al.

of urbanization, we divided Shanghaiinto three parts, according to the levelof urbanization: urban, suburban, andrural (Figure 1b).

Remote sensing data and dataprocessing

Between 1975 and 2005, data on landuse and land cover in Shanghai wasobtained using cloud-free LandsatMultispectral Scanner (MSS) andThematic Mapper (TM) remote sens-ing images. The data, which spannedthree decades, covered seven periods:1975, 1981, 1987, 1990, 1995, 2000,and 2005 (mostly from April to June).The first two sets of satellite images(1975 and 1981) were obtained fromthe MSS; for all other years, the TMwas used. Landsat data were inter-preted using ERDAS 8.4 image-pro-cessing software. Land-cover data weredivided into four types: urban land(urbanized area), green land (ie urban

parks, street trees, lawns, etc), cropland, and water body. Fordetails on the data processing, see Fang et al. (2005).

Air and water quality data

Systematic monitoring data on air and water pollution inShanghai are available from 1983 onwards (ShanghaiEnvironmental Protection Bureau 1983–2004). To mea-sure air quality, data were collected on sulfur dioxide(SO2), acid rain frequency, total suspended particles(TSP), and nitrous oxides (NOx); for water quality, thedata covered dissolved oxygen (DO), chemical oxygendemand (COD), biochemical oxygen demand (BOD),Kjeldahl nitrogen (KN), total phosphorus (TP), oils,volatile phenol (VP), and total mercury (THg). We ana-lyzed variations in the concentrations of these componentsin urban, suburban, and rural areas between 1983 and2004. With the exception of DO, all these measures of airand water quality are inverse indices; that is, the higher thevalues, the worse the air or water quality.

Climatic data

Urbanization leads to alterations of the local climate, andin particular creates a significant heat island effect(Kalnay and Cai 2003; Zhou et al. 2004). We collectedmonthly mean air temperatures at 10 meteorological sta-tions for the period between January 1975 and December2004. Four of these stations were located in urban areas(Xujiahui [Shanghai proper], Minhang, Jiading, andBaoshan), with the remainder in rural areas (Nanhui,Fengxian, Jinshan, Qingpu, Chongming, and Songjiang;

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Figure 1. The city of Shanghai. (a) An aerial view of the downtown area. (b and inset)Map showing the location of Shanghai, at the estuary of the Yangtze River, on the coast ofthe East China Sea. For the purposes of the current study, Shanghai was divided into threeareas: (A) urban; (B) suburban; and (C) rural.

SQ Zhao et al. Urbanization in Shanghai

Figure 1b). We analyzed the differences in mean annualtemperature (MAT), monthly mean maximum tempera-ture (MTmax), and monthly mean minimum temperature(MTmin) between the urban and rural areas. In order tostudy the urban heat island effect, we correlated the areaof urban land with the differences in mean temperaturesbetween the urban and rural areas from 1975 to 2005.

Biodiversity data

Urbanization exerts a substantial effect on biodiversity, result-ing in the loss of native species and the introduction of non-native species (Blair 1999; McKinney 2000). To explore theimpact of urbanization on biodiversity, we collected speciesrichness data for flora and fauna from various literaturesources (Xu et al. 1999; Yang et al. 2002; Shanghai Agricultureand Forestry Bureau 2004). We also carried out fieldwork toinvestigate the numbers of native and non-native woodyplant species in different green areas (Da et al. 2005).

� Results and discussion

Land-use and land-cover change

Land use and land cover in Shanghai have been greatlyaltered over the past three decades, as a result of the rapidexpansion of urban areas (Figure 2). The area of urbanland increased from 159.1 km2 in 1975 to 1179.3 km2 in2005 (Figure 2i). There were distinct phases in the urban-ization process. The slowest rate of urbanization, 17.7 km2

yr–1, occurred between 1975 and 1981 (Figures 2a and2b), increasing to 52.4 km2 yr–1 between 1990 and 1995and 54.9 km2 yr–1 from 2000 to 2005 (Figures 2d to 2g).This is consistent with China’s economic policies, sincethe country began its economic reform in 1978, andaccelerated the process in 1992 (Lin 1999).

Urbanization is generally associated with an increase inmanaged green areas, such as street trees, lawns, and parksfor urban recreation; these improve both the visual appealof a city and environmental quality (Attwell 2000). In par-allel with its urban expansion, Shanghai’s green areas havecontinued to increase in size, from 8.7 km2 in 1975 to 252.9km2 in 2005. In contrast, agricultural land area has fallenrapidly, from 6030.7 km2 in 1975 to 4743.1 km2 in 2005. Inaddition, the area covered by water has shown small fluctu-ations over the past three decades (Figure 2i).

Air and water quality changes

Urbanization places a heavy burden on local air and waterquality. Air quality monitoring data obtained from differentobservatories showed that concentrations of SO2, TSP, andNOx, along with acid rain frequency, varied considerably indifferent areas along the urbanization gradient.Concentrations of air pollutants were consistently highestin urban areas, lower in suburban areas, and lowest in ruralareas over the study period, indicating a strongly negative

influence of major urban development on air quality(Figure 3). The temporal variations in air pollutants suggestthat SO2, TSP, and acid rain were largely the result of coalcombustion, since levels of these pollutants decreased from1983 to 2004 in all three areas, while NOx concentrationincreased (Figure 3). The falling concentrations weremainly related to the decline in the use of coal, both byindustry and in private households (Yuan and James 2002).The increase in green lands may also have helped to miti-gate air pollution. The rising concentrations of NOx areattributed primarily to a rapidly increasing number of motorvehicles in recent years; for example, vehicles in Shanghainumbered less than 100 000 in the early 1980s, butrose to more than 2 million by 2004 (www.shtaq.com/

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Figure 2. Changes in land-cover types in Shanghai. (a–g)Spatial distribution of land-cover types for seven time periods:1975, 1981, 1987, 1990, 1995, 2000, and 2005. (h) Overlapof urban lands from 1975–2005. (i) The changes in area of fourland-cover types: urban land, green land, cropland, and water,from 1975–2005.

(a) 1975 (b) 1981 (c) 1987

(d) 1990 (e) 1995 (f) 2000

(g) 2005 (h) Urban expansion 1975–2005

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Legends19751975–19811981–19871987–19901990–19951995–20002000–2005

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showcontent/contentdetail_old.asp?id=39703). A similarresult was seen in Beijing (He et al. 2002).

There have been two exceptions to these general trends.First, as shown in Figures 3a and 3b, SO2 pollution andacid rain have tended to increase in the suburbs in recentyears, which may be attributed to a growing number of fac-tories in suburban areas. In the late 1990s, the municipalGovernment of Shanghai initiated an environmental

improvement program, aimed at movingindustrial units from urban centers tosuburban areas in order to improve urbanair quality (Shanghai EnvironmentalProtection Bureau 2001).

The second exception is that NOx pol-lution has decreased in urban areas, sincethe late 1990s, a result of the CleanVehicle Program which was imple-mented in 1999 as a means of controllingair pollution due to vehicle emissions(He et al. 2002).

Water pollution levels show similarpatterns, being consistently more severein the central urban areas than in subur-ban and rural areas (Figure 4). During thestudy period, different pollutants variedover time in the central urban areas; con-centrations of COD, BOD, KN, and

volatile phenols increased from the early 1980s to theearly 1990s, and then began to decrease, while DO, totalphosphorus, oils, and total mercury did not exhibit signifi-cant changes over the same period. This suggests thatwater quality in the city center has been improving sincethe 1990s. However, water quality in the suburban andrural areas has deteriorated over the past decade, althoughgenerally much better than in the central urban area. Forexample, BOD, KN, and TP pollution has increased inrecent years, probably as a result of the transfer of factoriesfrom the city center to suburban and rural areas (ShanghaiEnvironmental Protection Bureau 2001).

Local climate change and the heat island effect

As illustrated in Figure 5, a substantial urban heat islandeffect was found in Shanghai. The difference in MATbetween urban and rural areas increased from 0.1 ˚C inthe late 1970s (average for 1975–1979; a mean MAT of15.7˚C vs 15.6˚C at urban and rural stations) to 0.7˚C inthe early 2000s (average for 2000–2004; 17.3˚C vs16.6˚C), with an increase of 0.24˚C per decade (Figure5a). Similarly, MTmax and MTmin did not show a differ-ence between urban and rural areas in the late 1970s(28.2˚C vs 3.4˚C for MTmax and MTmin, respectively),although the differences increased to 0.7˚C (29.2˚C vs28.5˚C for the urban and rural area) and 0.5˚C (5.0˚C vs4.5˚C) in the early 2000s, respectively, with a decadalincrease of 0.26˚C and 0.21˚C (Figures 5b and 5c).

The correlation analysis of the relationship betweenthe differences in mean temperatures in urban vs ruralareas, and the amount of urban land, indicate that thedifferences in temperature between urban and rural areasincreased substantially as the city expanded, and that thisincrease was faster for MTmax than for MTmin (Figure 5d).This strongly suggests that rapid urbanization canincrease temperatures considerably in the city and adja-cent areas (Zhou et al. 2004).

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Figure 3. Changes in major components of air quality between 1983 and 2004 inthe different areas of Shanghai: (a) sulfur dioxide (SO2); (b) acid rain frequency;(c) total suspended particles (TSP); and (d) nitrous oxides (NOx).

Figure 4. Changes in major components of water qualitybetween 1983 and 2004 in the different areas of Shanghai: (a)dissolved oxygen (DO); (b) chemical oxygen demand (COD);(c) biochemical oxygen demand (BOD); (d) Kjeldahl nitrogen(KN); (e) total phosphorus (TP); (f) oils; (g) volatile phenols(VP); and (h) total mercury (THg).

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SQ Zhao et al. Urbanization in Shanghai

Changes in biodiversity

The area around Shanghai is rich inbiodiversity. There are 1904 spermato-phyte plants, representing 981 generaand 168 families. However, there aremore non-native species than native(968 and 936, respectively; Xu et al.1999). Seven hundred and sixty verte-brate species are found in Shanghai, ofwhich 40 are mammals, 424 birds, 32reptiles, 14 amphibians, and 250 fish(Shanghai Agriculture and ForestryBureau 2004).

Urban expansion and the associatedincrease in human activities have led toa considerable loss of biodiversity in thestudy area. The number of native plantspecies has fallen rapidly in the rela-tively wild regions; for example, thenumber of plant species in the Sheshanarea fell from 535 in the 1980s to 254 bythe end of the 1990s (Xu et al. 1999),while on Dajinshan Island plant speciesdeclined from 254 in the 1980s to 145in 2000 (Yang et al. 2002). At the same time, the numberof non-native plant species has increased greatly due tothe introduction of many species into the managed greenspaces; about 300 alien plant species were introduced tothe Shanghai area between 1980 and 2005, for instance.In 2004, a comprehensive investigation by the authorsrecorded a total of 206 woody plant species within thegreen spaces in the city, of which only 64 (31.1%) werenative to Shanghai and adjacent areas (Da et al. 2005).The proportion of native species is even lower in recentlyestablished green areas within the city. For example, inthe newly created Yanzhong green land, only 26 (18.4%)of 142 woody plants are native, while a much higher pro-portion (28 out of 69 species) of woody plant species werefound in the green spaces established in the 1950s on thecampus of East China Normal University (Table 1).

� Conclusions

Shanghai has experiencedrapid urbanization over thepast three decades, accom-panied by large-scale eco-nomic development. How-ever, this urban growth hascaused a number of ecolog-ical problems, includingdegradation of air andwater quality, alteration ofthe local climate, a declinein native species, and anincrease in numbers ofalien species. This, to-

gether with an improvement in public awareness of envi-ronmental issues, has been the basis for attempts to achieveboth socioeconomic and environmental sustainability inthe urbanization process. In recent years, the ShanghaiGovernment has implemented a series of measures specifi-cally designed to protect the local environment. First, anumber of policies have been developed which are aimed atpromoting clean energy use to reduce water and air pollu-tion; over the past 10 years, these have involved improve-ment of the sewage treatment infrastructure, removal ofexhaust emission sources, improvement of transportationsystems, and control of vehicle density. As a result, air andwater quality have begun to improve, beginning in the1990s (Figures 3 and 4). Second, afforestation and theestablishment of parks within Shanghai have been incorpo-rated into city planning, leading to an increase in vegeta-tion coverage. For example, the area occupied by greenland in Shanghai has risen from 870 ha in 1975 to 25 300 ha

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Figure 5. Changes in temperature differences between urbanized and rural areas inthe years 1975–2004. (a) Difference in mean annual temperature (dMAT); (b)difference in monthly mean maximum temperature (dMTmax); (c) difference inmonthly mean minimum temperature (dMTmin); and (d) relationships between thetemperature differences and degree of urbanization.

Table 1. Comparison of the number and proportion of native woody plant species inthree different green lands in Shanghai

No. of No. (%) of species No. (%) of species native No. (%) ofSite woody plants native to Shanghai to the adjacent areas native species

All green lands in the urban center 206 43 (20.9%) 21 (10.2%) 64 (31.1%)

Newly established Yanzhong green land 142 11 (7.8%) 15 (10.6%) 26 (18.4%)

Green land on the East China Normal University campus, established inthe 1950s 69 16 (23.2%) 12 (17.4%) 28 (40.6%)Based on Da et al. (2005)

(a) (b)

(c) (d)


in 2005 (Figure 2). Third, nature reserves and forested parkshave been established to protect biodiversity and providerecreation sites. The first nature reserve in Shanghai wasestablished in the early 1990s, followed by four more reservesand a national forest park. Finally, in 2003, ChongmingIsland, the third largest island in China and the largest allu-vial island in the world, was declared an “ecological island”by the Shanghai Government. Chongming Island, situatedat the estuary of the Yangtze River in northern Shanghai,covers an area of 1100 km2 and includes a large coastal wet-land and tidal flats that provide habitat for a wide variety ofspecies. All development activities that conflict with theprotection of the environment will be prohibited on theisland. At the same time, the Shanghai Government pro-vides monetary compensation to local communities andencourages local resident participation in the protection ofthe island (Yuan et al. 2003).

In summary, China is facing many challenges as itsurban growth rate continues to accelerate. Rapid urban-ization has greatly accelerated economic and social devel-opment, but has also created severe environmental prob-lems. Maintaining a balance between environmentalsustainability and the continuing process of urbanizationis a major issue facing the Chinese Government. Anational strategy for sustainable development – China’sAgenda 21 (Department of Planning Committee of China1994) – has been established, aimed at reducing the nega-tive environmental impacts of economic developmentwhile at the same time maintaining economic and socialbenefits. This involves land-use management strategiesand the development of greener cities, thereby providing ahealthier environment for both humans and wildlife.

� Acknowledgment

We thank LN Li, NN Liu, XY Wang, JQ Qian, L Li, JYWang, CX Wang, and KK Shang from East China NormalUniversity for their help in data collection. This study wasfunded by the State Key Basic Research and DevelopmentPlan of China (#G2000046801), the National NaturalScience Foundation of China (#40571150 and#40024101), and Special Research Fund for the DoctoralProgramme of Higher Education (#20050269015).

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1Department of Ecology, College of Environmental Sciences, and KeyLaboratory for Earth Surface Processes of the Ministry of Education,Peking University, Beijing, 100871, China *([email protected]);2Department of Environmental Sciences, and Shanghai Key Laboratory forEcology of Urbanization Process and Ecorestoration, East China NormalUniversity, Shanghai, 200062, China

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China spans a huge geographical area, ranging from trop-ical to boreal zones and from very low altitudes (156 m

below sea level in the Turpan Desert, Xinjiang) to theworld’s highest mountain, Mount Qomolangma (Everest),on the border of Tibet and Nepal. Almost all of the differ-ent types of biomes found on Earth, from rainforests todeserts, are found in China. This geographic diversity pro-vides abundant habitats for plants and animals. Home tomore than 30 000 vascular plants (surpassed only by Braziland Colombia) and 6300 vertebrates, China is one of theworld’s “megabiodiversity countries” (McNeely et al. 1990).However, this biodiversity has suffered severe degradationdue to the density of the human population, a long historyof cultivation, and an increase in the intensity and extent ofhuman disturbances in the plains and lowland areas.

The mountain regions of China provide rich habitatsfor much of the country’s remaining biodiversity, owingto the heterogeneity of climates and soils, rapid eleva-tional changes, varying aspects of slope direction, abun-dant microhabitats, and limited suitability for cultivation

(Körner and Spehn 2002). Thus, China’s mountains arelikely to be especially important for preserving its remain-ing biodiversity (Chen 1998).

In this study, we used well-developed county level dis-tribution data for plants and animals to explore the pat-terns of richness of plant genera and terrestrial mammals,endemic plant genera and mammal species, and endan-gered plant and vertebrate species in China. Our goalswere to identify hotspots for three different aspects of bio-diversity (overall, endemic, and endangered) and toinvestigate the potential role of the mountain regions inconserving China’s biodiversity.

� Data sources

There are 2383 counties in China. A county level mapwas digitized and overlaid with a Digital Elevation Model(DEM) with 1 km x 1 km resolution, obtained from theUnited States Geological Survey (USGS), to documentthe topographic attributes of each county.

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Biodiversity in China’s mountains

Zhiyao Tang, Zhiheng Wang, Chengyang Zheng, and Jingyun Fang*

China, one of the world’s “megabiodiversity countries”, is home to more than 30 000 vascular plant and 6300vertebrate species. Over thousands of years, however, cultivation has led to the disappearance of many of thesespecies from the plains and lowland areas. The mountain regions still harbor large numbers of species, becausethere have been fewer human and natural disturbances and there are more diverse habitats. We used countylevel species distribution databases to explore patterns of biodiversity and to identify biodiversity hotspotswithin China. Ten hotspot ecoregions were identified, containing 3110 plant genera (92.0% of the country’stotal), 220 (90.5%) endemic plant genera, 366 (94.3%) endangered plants, and 254 (72.2%) endangered verte-brates, 427 (91.0%) terrestrial mammal species, and 65 (85.5%) endemic mammals. All 10 hotspot ecoregionsare located in the mountainous areas of China. Although high richness of overall, endangered, and endemicplants and animals co-occurred in many of the same hotspot ecoregions, they often occurred in different coun-ties within these ecoregions and showed low spatial congruence. In conclusion, China’s mountain regions arecritical for protecting biodiversity and should be made conservation priorities in the future.

Front Ecol Environ 2006; 4(7): 347–352

The overall numbers of plant genera and mammalspecies, endemic plant genera and mammal species, andendangered plant and vertebrate species for each countywere derived from the following resources: “Seed plants ofChina” (Wu and Ding 1999), “Mammal species inChina” (Zhang 1997), “China red data book of endan-gered plants” (Fu 1992) and “China red data book ofendangered animals” (Wang 1998).

The “Seed plants of China” database (Wu and Ding1999) is the main product of a national project spanning10 years (1990–1999) that described county-level distrib-utions for each of the 32 308 native and cultivated seedplants, representing 363 families and 3428 genera (includ-ing 243 genera endemic to China). For each genus, thegeographic distribution was resolved to the county level,and habitat type, elevational range, and longitudinal andlatitudinal range were documented based on nationalflora, local reports, and other scientific literature. Becausethis database is derived primarily from plant specimenrecords, it is potentially biased by regional differences inlevels of investigation. In the study presented here, genera(or family) richness was used instead of species richness tomeasure plant diversity, because it is less likely that thesehigher taxa would be overlooked in surveys (Qian 1998).To help reduce the possibility of bias in the records as aresult of regional differences in investigation intensity, weexplored the patterns of seed plant density at the genus, asopposed to species, level in this study.

“Mammal species in China” (Zhang 1997) was compiledbased on the scientific literature as well as observations oflocal fauna over the past 50 years. This book includes infor-mation on the county level distribution and habitat type foreach of the 537 mammals in China. In our study, 68 marinemammals whose distributions are not clear were excludedfrom the analysis. We therefore included only the data for469 terrestrial mammal species (of which 77 are endemic toChina) to investigate patterns of mammal diversity.

The “China red data book of endangered plants” (Fu1992) and the “China red data book of endangered ani-mals” (Wang 1998) provided information on the distrib-utions of endangered plant and vertebrate species.These two datasets record the geographic distribution,habitat type, and elevational range for each of China’s388 endangered plant species and 352 endangered ver-tebrate species (including 134 mammals, 95 reptiles, 92fishes, and 31 amphibians). Endangered birds were alsonot included in this study as most avian species aremigratory, thus making their distributions unclear.

�Methods

We first calculated the animal and plant richness per countyfor the three aspects of biodiversity mentioned above (over-all, endemic, and endangered). Geographic information sys-tem (GIS) software (ArcView GIS 3.2) was used to calculatethe area covered by each county. The administrative divi-sion atlas of China (Editorial Committee for the

Biodiversity in China’s mountains ZY Tang et al.

Administrative Division Atlas of China 2002) was digitizedto obtain information on county boundaries. The area cov-ered by the different counties in China varies greatly; weused the following area-adjusted species (genera for overalland endemic plants) density as a measure of diversity, toeliminate the influence of county size (Qian 1998):

D = S/ln (A)

where D is the species density for a county, S is the num-ber of species, and A is the area of the county.

We then used ArcView GIS 3.2 to map different aspectsof biodiversity in order to identify biodiversity hotspotsacross the country. Here, we defined hotspot counties asthe top 5% of land area with the highest density of each ofthe three aspects of biodiversity. We first selected thecounty that contained the highest density of species, thenadded the county with the second highest density, thenthe third, and so on, until the total area of selected coun-ties equaled 5% of China’s total land area. All of the coun-ties ranked within the top 5% for each aspect of biodiver-sity were designated as the hotspot counties. Becauseecoregions can be used as conservation units at regionalscales (Olson and Dinerstein 1998), we assigned all thecounties to different ecoregions, according to their physi-cal geography (Editorial Committee for China’s PhysicalGeography 1985). We then calculated the percentage ofthe area covered by hotspot counties within each ecore-gion. An ecoregion was designated as a hotspot if 5% of itstotal area was covered by hotspot counties.

The topographic attributes (maximum and minimumelevation, average elevation, elevational range, andslope) of each county were derived from the DEM byusing ArcView GIS 3.2. The UNEP-WCMC (2002) cri-terion was used to identify mountainous areas versusplains. According to the criterion, areas that fit any of thefollowing four descriptions are defined as mountainous:(1) an elevation of 300–1000 m and a local elevationalrange of > 300 m; (2) an elevation of 1000–1500 m abovesea level (asl), and a slope of > 5˚ or a local elevationalrange of > 300 m asl; (3) an elevation of 1500–2500 m asl,and a slope of > 2˚; or (4) an elevation of > 2500 m asl.

� Results and discussion

Macrotopography in China

According to the UNEP-WCMC (2002) definition,mountains cover a total area of approximately 4.6 millionkm2 in China (48% of the total land area). Figure 1 showsthe macrotopography and a frequency distribution of themean elevation per county, together with major geo-graphic regions. Low-lands with an elevation of < 500 masl covered 2.63 million km2 (27.4% of the total landarea), while areas with an altitude of > 1000 m asl encom-passed an area of 5.42 million km2 (56.5% of the totalland area). Another 1.93 million km2 (20.1% of the

348


ZY Tang et al. Biodiversity in China’s mountains

country’s total area) was covered by alti-tudes of > 4000 m asl.

Patterns of overall plant genera andmammal species densities

The density of plant genera was higher ineastern compared to western China, higherin southern compared to northern China,and also in the mountainous areas com-pared to the plains and plateaus (Figure2a). One hundred and seventy-three coun-ties were assigned as hotspot counties forplant genera; they contained densities of> 65 genera ln–1 (km2). In total, 2948 gen-era of seed plants (86.0% of China’s total)grow in these counties. The counties werelocated mostly within eight regions: theHengduan Mountains, the Xishuangbannaarea, the upper reaches of the HongshuiRiver (the Wumeng Mountains), theNanling Mountains, the WulingMountains, the East China Mountains,Hainan Island, and Taiwan Island.Compared to the mountainous areas listedabove, counties in the North China Plain, Loess Plateau,Sichuan Basin, and Qinghai-Xizang Plateau containedvery low densities of plant genera (< 20 genera ln–1 [km2]).

Mammal species densities showed a slightly different pat-tern from that of plant genera. One hundred and fifty-fourcounties were designated as hotspot counties, with mam-mal species densities of > 5.8 species ln–1 (km2). Thesecounties contained 384 mammals (81.9% of China’s total)and were mostly located in the following eight regions: theHengduan Mountains, the Nanling Mountains, theXishuangbanna Area, the Qinling Mountains, the upperreaches of the Hongshui River, the East China Mountains,Hainan Island, and Taiwan Island. Densities of mammalspecies were lower in the North China Plain, the CentralYangtze Plain, the Loess Plateau, the western part of theQinhai-Xizang Plateau, the Northeast China Plain, andthe Sichuan Basin (Figure 2b).

The distribution of overall richness is positively associ-ated with habitat diversity (Orme et al. 2005). In the pre-sent study, we used the elevation range (ie the differencebetween the highest and the lowestelevation in a county) as a surrogate ofhabitat heterogeneity, and found posi-tive relationships between elevationrange and the densities of both plantgenera (Pearson correlation coeffi-cient, r = 0.32, P < 0.01), and mammalspecies (r = 0.36, P < 0.01). Statisticalanalysis also suggested a positive rela-tionship between the density of over-all plant genera and mammal species(r = 0.36, P < 0.01; Table 1).

Patterns of endemic plant genera and mammalspecies densities

The density of endemic plant genera was also higher inmountainous areas than in other areas. Altogether, 161counties were assigned as hotspots for endemic plants(> 0.85 genera ln–1 [km2]). These counties contained 204endemic plant genera (84.0% of the total), with mostlocated in the following seven regions: the HengduanMountains, the Wuling Mountains, the NanlingMountains, the Qinling Mountains, the East ChinaMountains, Hainan Island, and Taiwan Island (Figure 2c).On the other hand, the 123 hotspot counties for endemicmammals (a density of > 0.75 species ln–1 [km2]) contained56 endemic mammals (73.7% of the total), which werelocated primarily in the Hengduan Mountains, the QinlingMountains, and the eastern part of the Qinghai-XizangPlateau (Figure 2d). These mammal hotspot counties werelocated either in the ecotones between different biomes orin areas with high environmental heterogeneity, and havenot suffered the influences of the last period of glaciation

349


Figure 1. Macrotopographic pattern in China, together with locations of majormountains. The inset figure illustrates the elevation–area frequency distribution.

Table 1. Correlation between densities of overall and endemic plant genera,overall and endemic mammal species, and endangered plant and vertebratespecies in different counties of China

Endemic Endangered Overall Endemic EndangeredAspect of diversity plant genera plants mammals mammals vertebrates

Overall plant genera 0.41* 0.42* 0.36* 0.07 0.41*Endemic plant genera 0.44* 0.31* 0.39* 0.34*Endangered plants 0.38* 0.26 0.45*Overall mammals 0.57* 0.54*Endemic mammals 0.31**P < 0.01. For all correlations, n = 2383 counties.

Elevation (m)<200200–500500–10001000–20002000–30003000–40004000–50005000–6000<6000

Are

a (x

106

km2 ) 3

2

1

0

>5

5–10

10–1

5

15–2

0

20–2

5

25–3

0

30–3

5

35–4

0

40–4

5

45–5

0

50–5

5

55–6

0

>60

Elevation (x100 m)


(eg the Hengduan Mountains, the Qinling Mountains) orare areas with high rates of speciation (eg the TibetanPlateau; Chen 1998).

Patterns of endangered plant and vertebratedensities

Hotspots for endangered plants included 185 counties(> 1.1 species ln–1 [km2]). These counties contained a totalof 333 endangered plants (85.8% of the total) and werelocated in the following five regions: the HengduanMountains, the Xishuangbanna Area, upper reaches of theHongshui River area, Hainan Island, and Taiwan Island.Compared with these mountainous areas, the counties inthe North China Plain, Northeast China Plain, CentralYangtze Plain, Sichuan Basin, Loess Plateau, InnerMongolia Plateau, and Qinghai-Xizang Plateau possessvery low densities of endangered plant species (Figure 2e).

The density patterns of endangered vertebrates (exclud-

ing birds) are similar to that of endan-gered plants. For these species, 143 coun-ties were assigned as hotspot counties(> 2.2 species ln–1 [km2]); they contained225 endangered vertebrates (63.9% of thetotal). These counties were located in thefollowing eight regions: the HengduanMountains, upper reaches of the HongheRiver area, the Xishuangbanna area, theQinling Mountains, the NanlingMountains, the East China Mountains,Hainan Island, and the east part of theQinghai-Xizang Plateau. Low densities ofendangered vertebrates were found in thenorthern and northeast plains andplateaus in China (Figure 2f).

Endangered species richness is widelyused as a proxy for conservation priorityidentification because of the significantand positive relationship between endan-gered species and overall species richness(Dobson et al. 1997). As in previous stud-ies (eg Dobson et al. 1997), we observed apositive correlation between densities ofoverall mammals and endangered verte-brates (r = 0.54, P < 0.01), and betweendensities of overall plant genera andendangered plants (r = 0.42, P < 0.01;Table 1). Further, positive correlationswere found between the densities ofendangered plants and endemic plantgenera (r = 0.44, P < 0.01) and betweenendangered and endemic animals (r =0.31, P < 0.05; Table 1), as most of theendemic plants and vertebrates in Chinaare endangered because of their limitedgeographic distributions (Wang 1998).We also found that the densities of

endangered plant and vertebrate species were positivelycorrelated (r = 0.45, P < 0.01; Table 1).

Richness of endangered species is shaped by the inter-action between the biological mechanisms promotingspecies diversity and the anthropogenic mechanismseroding that diversity (Orme et al. 2005). To explore theinfluence of human threat on the distribution of endan-gered species richness, we calculated the ratio of endan-gered compared to overall densities (endangered plants tooverall plant genera and endangered vertebrates to over-all mammals). This ratio was high in the North ChinaPlain for plants, and in the East China Mountains, theWuling Mountains, and the Nanling Mountains for ani-mals. The North China Plain, the east (including theEast China Mountains), the central (where the WulingMountains are located), and the southern (including theNanling Mountain) regions of China have high humanpopulation densities, suggesting the influence of humanson species endangerment (Dobson et al. 1997).

350


Figure 2. Distribution patterns for the county level density of: (a) seed plant genera; (b)mammal species; (c) endemic plant genera; (d) endemic mammal species; (e) endangeredplant species; and (f) endangered vertebrate species (excluding birds) in China. Hotspotcounties are shown in dark orange and brown.

(a) (b)

(c) (d)

(e) (f)

Plants (ln–1 [km2])0–2020–4040–6565–100>100

Mammals (ln–1 [km2])0–11–33–5.85.8–8>8

Endemic plants (ln–1 [km2])0–0.20.2–0.50.5–0.850.85–2>2

Endemic mammals (ln–1 [km2])0–0.20.2–0.40.4–0.750.75–1.25>1.25

Endangered plants (ln–1 [km2])0–11–1.51.5–2.052.05–3>3

Endangered vertebrates (ln–1 [km2])0–0.50.5–11–2.22.2–4.5>4.5

ZY Tang et al. Biodiversity in China’s mountains

Comparison between the different aspects ofbiodiversity hotspots

The various types of biodiversity hotspot counties (thosebased on the density of all, endemic, and endangered taxa)tended to be clustered in a few key regions such as theHengduan Mountains, the upper reaches of the HongshuiRiver, the East China Mountains, Hainan Island, andTaiwan Island (Figure 2). However, there remains somespatial dissimilarity among the different types of hotspotcounties: for example, although the co-occurrence of highrichness of overall, endangered, and endemic plants andanimals was apparent in many of the same ecoregions,there was little plant–animal richness overlap, as they wereoften found in different counties within these ecoregions.

The three aspects of plant hotspot counties occupied atotal area of 977 450 km2, of which only 11.5% (112 281km2) was common to all three measures (Figure 3a). Anadditional 21.1% (206 025 km2) was shared between pairsof aspects. However, 22% (215 497 km2), 23.7% (231 801km2), and 21.7% (211 846 km2) were idiosyncratic toindividual aspects of overall, endemic, and endangeredhotspots, respectively (Figure 3a).

Similarly, the animal hotspot counties also showed lowspatial congruence between different aspects. The cumula-tive area of the three types of animal hotspot counties was1 001 709 km2, of which only 8.2% (82 080 km2) was com-mon to all three aspects (Figure 3b). The idiosyncratichotspot counties occupied a total of 67.5% (675 977 km2),with 20.0% (200 328 km2), 25.5% (254 852 km2), and 22%(220 797 km2) for the overall, endemic, and endangeredanimal hotspots, respectively. The remaining 24.3%(243 652 km2) was shared between pairs of the differentaspects (Figure 3b). Similarly low spatial congruenciesbetween different aspects of hotspots were observed withthe global pattern of avian richness (Orme et al. 2005).

Ten hotspot ecoregions in major mountain ranges

High densities of plant genera and mammal species,endemic plant genera and mammal species, and endan-

gered plant and vertebrate species appearedin different counties, which were aggregatedin several ecoregions (Editorial Committeefor China’s Physical Geography 1985). Theseencompassed the Hengduan Mountains, theWuling Mountains, the Nanling Mountains,upper reaches of the Hongshui River area, theXishuangbanna area, the Qinling Mountains,the East China Mountains, the Qinghai-Xizang Plateau, Hainan Island, and TaiwanIsland. According to the criteria of UNEP-WCMC (2001), all of these ecoregions arelocated in mountainous areas and should beconsidered as the hotspot ecoregions (with a“hotspot ecoregion” determined by an areahaving at least 5% of its land area composedof hotspot counties) for biodiversity in

China. These 10 biodiversity hotspot ecoregions cover anarea of ca 2.3 million km2, or 24.2 % of China’s total landarea. Table 2 illustrates the plant diversity of these areas.In total, they contained 3110 plant genera (92.0% of thecountry’s total), 220 (90.5%) endemic plant genera, 366(94.3%) endangered plants, 427 (91.0%) terrestrial mam-mal species, 65 (85.5%) endemic mammals, and 254(72.2%) endangered vertebrates. It is worth noting thatthe alpine areas of the Tibetan Plateau contain globallyunique ecosystems and their species are highly endemic,despite low overall taxonomic richness (Table 2).

These hotspot ecoregions are generally the same asthose proposed by Chen (1998) and Olson and Dinerstein(1998), who used relatively descriptive and approximatespecies information. Chen (1998), using estimated speciesrichness and number of endemics, proposed 11 terrestrialcritical regions for the conservation of China’s biodiver-sity, eight of which were located within the hotspot ecore-gions identified in this study. Of the WWF’s Global 200most critical and endangered ecoregions, 17 are located inor around China, with 11 consistent with the priorityareas identified in this study. Several of the others areeither located in aquatic ecoregions or along China’s bor-ders (Olson and Dinerstein 1998).

� Conclusions

The patterns of biodiversity, as well as the mechanismsthat cause these patterns, are a primary focus of biodiver-sity research (Gaston 2000). We explored the distribu-tion patterns of different aspects of biodiversity at acounty level, and identified ten biodiversity hotspotecoregions that should serve as conservation priorities inChina. All of the identified hotspot ecoregions for over-all, endemic, and endangered diversity for plants and ani-mals in China are located in the mountainous areas.These mountainous areas are therefore critical for pro-tecting China’s biodiversity.

By the end of 2004, China had established more than2000 national and local nature reserves, which collectively

351


Figure 3. Venn diagrams illustrating the spatial congruence between different aspects(overall, endemic, and endangered) of biodiversity hotspots for (a) plants and (b) animals.For each of the aspects, the hotspots are the top 5% land areas with the highest density.

(a) PlantsOverall

215 497(22.0%)

80 832(8.3%)

59 037(6.0%)

112 281(11.5%)

66 156(6.8%)

231 801(23.7%)

Endemic

211 846(21.7%)

Endangered

(b) AnimalsOverall

200 328(20.0%)

111 507(11.1%)

77 879(7.8%)

82 080(8.2%) 254 852

(25.5%)

Endemic

220 797(22.0%)

Endangered54 266(5.4%)


cover approximately 1.7 million km2, or 17.8% of the coun-try’s total land area (Chinese Species Information Service2005). Of these, 1498 (or 73.8% of all of the protectedareas) are located in mountainous regions, covering a totalarea of 1.5 million km2 (83.0% of all protected areas).Although these reserves play a key role in China’s plan forbiodiversity conservation, most are located in remote andpoorer areas and are often viewed as obstacles to humanwelfare and economic development (Liu et al. 2003).Therefore, maintaining the functionality of these reservesis, and will continue to be, a critical but difficult issue.Increasing financial support to reserves and local residents,and assisting local communities in sharing the benefitsderived from tourism and natural resources are importantfactors in addressing this problem (Liu et al. 2003).

� Acknowledgement

We are grateful to XP Wang, YH Chen, X Lin, W Xu, andY Chen for their assistance in data collection. This studyhas been funded by the National Natural ScienceFoundation of China (#49971002, #40501025,#40024101, and #90211016) and the State Key BasicResearch and Development Plan (#G2000046801).

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Department of Ecology, College of Environmental Sciences, and KeyLaboratory for Earth Surface Processes of the Ministry of Education,Peking University, Beijing, China, 100871 *([email protected]).

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Table 2. Plant species richness of China’s major mountain ranges and the Qinghai-Xizang (Tibetan) Plateau

Area Area (103 km2) of Summit Seed plants Genera/species Reference*Mountain range (103 km2) hotpot counties (%) (m asl) (family/genus/species) endemic to China (local)

Hengduan Mts ~ 500 409 (81.7%) 7556 175/1348/7962 56/5079 (12/2988) Wang 1994

Upper reaches of Hongshui River (Wumeng Mts) 110 76 (69.1%) 2900 178/1293/5061 55/2555 (8/468) Wu 1996

East China Mts ~ 600 213 (35.5%) 2158 174/1180/4259 54/2147 (5/425) Zhang and Lai 1993

Wuling Mts ~ 100 97 (96.8%) 2570 20/1005/4119 65/2682 (0/126) Chen et al. 2002

Nanling Mts 45 44 (98.0%) 2142 200/1045/3854 41/na (2/na) Chen and Zhang 1994

Taiwan Island 35 35 (100%) 3997 186/1201/3656 17/1275 (4/1070) Ying and Hsu 2002

Qinling Mts 70 67 (96.1%) 3767 198/1007/3446 46/1428 (0/192) Ying 1994

Xishuangbanna 20 20 (100%) 2500 197/1140/3336 8(0)/na (121) Zhu et al. 2001

Hainan Island 35 35 (100%) 1867 204/1206/3315 19/1110 (8/505) Xing et al. 1995

Alpine Qinghai -Xizang Plateau (> 4200 m) ~ 800 255 (31.9%) 8848 67/339/1816 14/636 (10/604) Wu et al. 1995

*for detailed data source, see Web Only material; na = not available

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