Applied Geography 29 (2009) 577–587 Contents lists available at ScienceDirect Applied Geography journal homepage: www.elsevier.com/locate/apgeog Diversity and distribution of landscape trees in the compact Asian city of Taipei C.Y. Jim*, Wendy Y. Chen Department of Geography, The University of Hong Kong, Pokfulam Road, Hong Kong a b s t r a c t Keywords: Urban biodiversity Urban forest Urban tree communities Urban ecology Compact city Taipei The promotion and preservation of biodiversity in urban areas remains scant, especially in Asian cities. This study focuses on spatial pattern and diversity of landscape trees in compact Taipei. Aggregate species diversity of three urban habitats (streets, urban parks and riverside parks) exceeded the countryside’s secondary forests. Urban parks with site heterogeneity and multiple functions accommodate the highest richness, and streets with acute site limitations the poorest represented by popular native species. More affinities exist between urban and riverside parks. Low diversity in riverside parks echoes natural site constraints and primary use for river discharge and flood control. The compact urban form has not stifled species diversity and spatial variability of urban forests. Development history and park area have no significant relationship with species diversity. Understanding species composition in urban ecosystems could frame conservation strategies to augment species richness, appropriate site selection, habit preservation and wildlife recruitment. Ó 2009 Elsevier Ltd. All rights reserved. Introduction The world is heavily urbanized and 60% of humanity is projected to reside in cities by 2030 (United Nations, 2007). The considerable magnitude, extent and pace of urban expansion have brought drastic habitat degradation and biodiversity losses at different scales (Cilliers, Müller, & Drewes, 2004; Czech & Krausman, 1997; McKinney, 2006; Prasad & Badarinth, 2004). The protection of urban natural areas has become an important facet of nature conservation since the 1990s (Goode, 1993; Heywood, 1996; McKinney, 2006; Miller, 2005; Miller & Hobbs, 2002). A key endeavour is the protection of urban biodiversity and associated habitat assemblages. Biodiversity in cities provides social and biological functions to residents, including ecological balance, ecosystem services, environmental protection, outdoor recreation, aesthetic enjoyment, nature education, and nurturing grounds, shelters, refuges and dispersal centres for wildlife species (Box & Harrison, 1994; Cilliers et al., 2004; Reduron, 1996; Tsai, 2001). The sustainable development of human society could be achieved with the help of ecological sustainability, in which urban biodiversity conservation could play a useful role. Biodiversity conservation in Taiwan faces an uphill battle due to rapid development and urbanization (Tsai, 1999). Conservation efforts initiated since the 1980s have focused on natural ecosystems outside cities. The Cultural Heritage Preservation Act, enacted in 1981, mandated nature reserves and protected species. The Wildlife Conservation Act of 1989 permitted designation of 1955 species of rare fauna at three protection levels, namely endangered, rare, valuable and requiring conservation measures. The issue of global biodiversity conservation then attracted little attention from * Corresponding author. Tel.: þ852 2859 7020; fax: þ852 2559 8994. E-mail address: hragjcy@hkucc.hku.hk (C.Y. Jim). 0143-6228/$ – see front matter Ó 2009 Elsevier Ltd. All rights reserved. doi:10.1016/j.apgeog.2009.01.002 578 C.Y. Jim, W.Y. Chen / Applied Geography 29 (2009) 577–587 government agencies or the public in Taiwan. A special Committee for Global Environment Change was set up in the Executive Yuan (equivalent to the Cabinet), which established a Biodiversity Conservation Group in 1994. Lacking enabling policies and public awareness, related governmental agencies encountered difficulties in protecting biodiversity. In 2001, the Executive Yuan augmented nature conservation by formulating an action plan to pursue ‘‘the conservation of biological diversity, the sustainable use of its components, and the fair and equitable sharing of the benefits arising out of the use of genetic resources’’, modelled after the UN Convention on Biological Diversity. Such efforts thus far have designated six national parks aiming mainly at nature conservation. Meanwhile, biodiversity conservation and restoration in urban ecosystems and the adjoining rural envelope still receive little attention. Urban green spaces provide home and sustenance for many floral and faunal species. They contribute notably to urban diversity conservation (Box & Harrison, 1994; Smith, Thompson, Hodgson, Warren, & Gaston, 2006; Sukopp, 2002; Thompson et al., 2003). Many studies focused on plant diversity (e.g. Pyšek, 1989; Smith et al., 2006; Turner, Lefler, & Freedman, 2005; Zerbe, Maurer, Schmitz, & Sukopp, 2003), others investigated animals (e.g. Melles, Glenn, & Martin, 2003; Shochat, Stefanov, Whitehouse, & Faeth, 2004) and both (Angold et al., 2006). In Germany, habitat and species diversities were positively related to intensities of disturbance in the transition zone (between city centre and outskirts). The choice and combination of tree species influence the leaf area indices of urban forests, leading to differences in environmental and ecological benefits and facilitating smart growth (Gatrell & Jensen, 2002). Moreover, land use patterns and historical changes could affect floristic diversity in cities (Zerbe et al., 2003). The extensive introduction and invasion of alien species in urban areas, and heterogeneity of urban habitats (natural, disturbed and managed), could raise species richness above the surrounding natural or semi-natural habitats (Kowarik, 1990; Turner et al., 2005). The increasing cultural disturbance could reduce natural potential and the conservation and ecological values of flora (Gracı́a-Romero, 2001). The ‘‘Landscape Master Plan for Taipei City’’ proposed by the Taipei City Government in 2006 stipulated the need for biodiversity conservation. The enlightened policy could benefit from relevant research leading to practical plans and actions. This study aimed at: (1) to investigate the current status of species diversity in the typical compact Asian city of Taipei, which is the political, economic, cultural and transport centre of Taiwan; (2) to explore differentiation of the tree flora in the main urban habitats (streets, urban parks and riverside parks); (3) to examine the relationship between tree diversity and human disturbance; and (4) to deduce implications on urban green spaces management to achieve biodiversity and nature conservation in compact cities. Understanding plant diversity in urban green spaces could contribute to enhanced understanding of human and natural impacts on urban vegetation. The findings could inform the management of urban ecosystems and generate practical implications for biodiversity conservation in Taipei (Jim & Chen, 2008). Study area and methods Study area Taipei city is situated in the lowest portion of the Taipei Basin, occupying 27,127 ha, of which 66% has been urbanized. It is surrounded by hills on the north, east and south sides, with Danshui River and its floodplain opening to the west. The wellvegetated surrounding hills occupy about half of the city’s territory. The main landform is alluvial plain deposited by the Danshui River in the last few millennia. Others include riverine marsh, river terraces, lower hillslopes and water surfaces. The subtropical monsoon climate has an annual average temperature of 23.5 C and rainfall of 2153 mm mainly deposited in the warm months. The climate supports continuous plant growth for most species, except some deciduous elements that avoid the dry season (Huang, 1993). Data This study focused on landscape trees in the built-up portion of Taipei city. They are largely cultivated and managed by the Parks and Street Lights Office of the Public Works Department (Parks and Street Lights Office, 2000, 2005). The analysis was based on a tree census in the study area (including species identity and site location) conducted in 2005 with the help of this Office. All landscape trees were recorded, except those situated in small (<1 ha) neighbourhood parks. To investigate the relationship between habitat conditions and human disturbance, 12 street sections (Table 2), 10 urban parks (Table 4), 10 riverside parks (Table 6) were evaluated as representative samples that accommodate most landscape trees and main site variations. Species diversity indices and statistical computation Species richness (S) denotes the total number of tree species in a certain sample area. Amongst different species diversity indices (Greig-Smith, 1983), Simpson (Simpson, 1949), Shannon–Wiener (Lloyd, Zar, & Karr, 1968) and Evenness were chosen. The Simpson index describes the probability that a second individual drawn from a population (vegetation community) should be of the same species as the first. The Shannon–Wiener index indicates the species heterogeneity of a vegetation community. The Evenness index refers to the distribution pattern of the individuals between the species. The computations were preformed by the Species Diversity and Richness (III) software package (Seaby & Henderson, 2004). Pearson correlations C.Y. Jim, W.Y. Chen / Applied Geography 29 (2009) 577–587 579 were computed to explore the relationship between species diversity and site area, development history (denoting the intensity of human disturbance) with the help of SPSSPC 12 software. A biomass index is proposed to indicate the contribution of individual species to the treescape and the potential provision of ecosystem services. It is computed by B ¼ (Tree count Final height score)/100, where species with small (5–9 m), medium (9–18 m) and large (>18 m) final height are assigned respectively scores of 1, 2 and 3. Urban forests could bring extensive benefits: microclimate amelioration, energy saving, improved water, air and soil quality, mitigation of storm water runoff, reduced carbon dioxide emission, increased property value, and community vitality (Macro & McPherson, 2002; McPherson et al., 1997; Xiao, McPherson, Simpson, & Ustin, 1998). The average life span of urban trees in stressful sites is about 10–25 years (Urban, 1989). Trees that can remain could reach their biological potential size, to realize their potential benefits. Without data on age structure and size distribution, an analysis based on actual tree dimensions could not be conducted at this stage. Results and discussion Street trees Taipei’s street trees are composed of 33,656 trees from 49 species. Species composition is dominated by a small cohort of popular species (Table 1). The top five contributing 61% are Cinnamomum camphora, Ficus microcarpa, Koelreuteria elegans, Melaleuca quinquenervia and Bischofia javanica. The street tree population showed distinct species composition. Six species are common to all three habitats, five shared between streets and urban parks, only one between streets and riverside parks, and eight are exclusively planted in streets. The conifers and palms are minor components. The tropical broadleaf growth form and species with large final size (>18 m tall) are dominant (Tables 1 and 7). Some 48% of the trees are evergreen broadleaves, and 42% deciduous broadleaves. The 12 deciduous broadleaved species prevail over six evergreen broadleaved ones. Nine species could attain large final size, versus eight for medium (9–18 m tall) and only three small (5–9 m tall). Table 1 The composition, landscape features, provenance and quantities of the top 20 landscape tree species in streets. Rank Species Family Growth forma Final heightb 1 2 3 4 5 6 7 8 9 10 11 12 13 Cinnamomum camphora Ficus microcarpa Koelreuteria elegans Melaleuca quinquenervia Bischofia javanica Liquidambar formosana Roystonea regia Ulmus parvifolia Bombax ceiba Ficus religiosa Peltophorum pterocarpum Lagerstroemia speciosa Alstonia schloaris Lauraceae Moraceae Sapindaceae Myrtaceae Euphorbiaceae Hamamelidaceae Arecaceae Ulmaceae Bombacaceae Moraceae Caesalpiniaceae Lythraceae Apocynaceae BLE BLE BLD BLE BLD BLD Palm BLD BLD BLD BLD BLD BLE Large Large Small Large Large Large Large Small Large Large Medium Medium Medium 14 15 16 Terminalia mantalyi Ficus elastica Erythrina indica Combretaceae Moraceae Fabaceae BLD BLE BLD Medium Large Medium 17 Pongamia pinnata Fabaceae BLD Medium 18 Mangifera indica Anacardiaceae BLE Medium 19 20 Phoenix roebelenii Pistacia chinensis Arecaceae Anacardiaceae Palm BLD Small Medium Total Average a Provenance Native Native Native India, Malaysia, Australia Native Native Central America Native India India, Sri Lanka, Myanmar Tropical Asia, Australia India, Australia India, Malaysia, Java, Philippines Tropical Africa India, Malaysia, Java India, Malaysia, Pacific islands India, Malaysia, S. China, Ryukyu, Australia India, Myanmar, Malaysia India, Indochina Native Tree quantity Species biomass Tree count (no.) Tree frequency (%)c Biomass indexd 6792 5873 3675 2122 2042 1907 1767 1208 1110 1065 764 751 739 20.18 17.45 10.92 6.30 6.07 5.67 5.25 3.59 3.30 3.16 2.27 2.23 2.20 203.76 176.19 36.75 63.66 61.26 57.21 53.01 12.08 33.30 31.95 15.28 15.02 14.78 24.70 21.36 4.45 7.72 7.43 6.93 6.43 1.46 4.04 3.87 1.85 1.82 1.79 562 455 418 1.67 1.35 1.24 11.24 13.65 8.36 1.36 1.65 1.01 309 0.92 6.18 0.75 259 0.77 5.18 0.63 235 188 0.70 0.56 2.35 3.76 0.28 0.46 32,241 1612 95.80 4.79 824.97 41.25 100.00 5.00 Biomass index (%)e For tree growth form, BLE denotes broadleaf evergreen, and BLD broadleaf deciduous. This is the final tree height that could be attained in Taiwan: small trees are 5–9 m tall, medium 9–18 m and large >18 m. Tree frequency (%) ¼ (Tree count/Total tree number in street habitat) 100%. d Species biomass index ¼ (Tree count Final height score)/100, where large, medium and small final heights are assigned values of 3, 2 and 1. It indicates the biomass and hence landscape impact of a species upon reaching its final dimension. e Biomass index (%) ¼ (Species biomass index/Total biomass index) 100%. b c 580 C.Y. Jim, W.Y. Chen / Applied Geography 29 (2009) 577–587 Of the 20 most frequent species in street habitats, 13 are exotic, with 11 originated from Asia and Australia (Table 1). South and Southeast Asia are the principal sources of introduced species. Exotic trees contribute 31% of the total tree count and 33% of attainable biomass. The seven native species lead by sheer numbers at 64% by trees and 67% by biomass. The large proportion of natives could offer suitable habitat, shelter and forage for native wildlife to enrich urban ecology. The limited colourful blooms in Taipei’s streets are mainly offered by exotics. Streets contain a large number of species with large final dimensions, indicating their potential ability to offer a high leaf area index to bestow environmental and ecological benefits (Gatrell & Jensen, 2002). Daan has the most street trees (8470 trees, 25.17%), and Nangang has the least (627 trees, 1.86%) (Table 2). Street species composition demonstrates significant district variations. The city’s 49 species are unevenly distributed, ranging from 23 species in the dense core districts of Wanhua and Zhongshan to only four in Nangang. Individual districts showed a unique species assemblage. Wanhua has the highest species density at 4.0 species/km2 by urbanized area and 21 species/km2 by road area. Wanhua is the oldest centre, with a long and varied development history. Different street tree species have been introduced at different periods tied to changing landscape preference. The initiation of urban greening under Japanese influence (1895–1945) shifted to indigenous persuasion after the Second World War, and then to recent systematic green-city planning after 1995 (Public Works Department, 2004). Nangang is an outlier with only four species, and the lowest species density. It is a newly built industrial centre receiving less landscaping attention. For the whole city, species diversity in streets is moderate (Simpson index D ¼ 9.75, Shannon–Weiner index H ¼ 2.71; Table 2), which is slightly lower than the average tree diversity in natural forests in the Taipei region (at H ¼ 2.90; Su, 1994). The stressful street environment has not limited the growth of many species. Changes in species selection over time have accumulated more species, as exemplified by the highest diversity of the oldest Wanhua district (D ¼ 8.11, H ¼ 2.41), followed by Daan (D ¼ 7.89, H ¼ 2.34). The relatively new and peripheral Nangang has the least (D ¼ 1.83, H ¼ 0.81). For Evenness index, the old and compact Wanhua (E ¼ 0.62) and Daan (E ¼ 0.60) have the highest scores, denoting a relatively equitable representation of different species. Nangang scores the lowest (E ¼ 0.21), reflecting that its small tree population is heavily represented by common species. The large street tree stocks in Shilin, Songshan, and Zhongshan have lifted the D and H values, but their E values have been suppressed by the occurrence of uncommon to rare species. The results indicate the limited species palette of street trees. Growth environment in different streets tend to have similar soil and atmospheric conditions. Street trees serve similar environmental benefits such as air pollutant removal (McPherson et al., 1997), wildlife habitats (Clark, Matheny, Cross, & Wake, 1997) and ornamental functions (McPherson, Simpson, Peper, & Xiao, 1999). They share similar management concerns and challenges (Parks and Street Lights Office, 2005). The common physical and physiological constraints also restrict species selection. Usually, the relatively narrow roadside corridor and underground utilities severely confined tree growth in compact city environment (Jim, 1992). The heavy shading, air pollution, poor soil quality, restricted soil volume and soil compaction would exclude many species from roadside use (Bassuk & Whitlow, 1987; Jim, 1999). The need for headroom and lateral clearance for vehicular and pedestrian traffic, and avoidance Table 2 The abundance, distribution and species diversity of landscape trees in streets in the 12 districts. Code District Land area Tree Stock Beitou Daan Datong Nangang Neihu Shilin Songshan Wanhua Wenshan Xinyi Zhongshan Zhongzheng Total Average or aggregate Range a b c d e 1691 1093 478 972 1446 1897 825 575 1635 1038 998 704 13,352 1113 1419 Road area Species diversity index Species Tree richness density (trees/ Sa ha)b Species density (species/ km2)c Tree density (trees/ ha)b Species Tree-species Person- Simpson Shannon– Evenness density ratiod tree diversity Wiener index, E (species/ ratioe index, D diversity 2 c km ) index, H 11 19 7 4 20 16 19 23 9 12 23 13 0.93 7.75 2.03 0.65 1.00 2.08 7.16 4.71 0.58 0.87 2.95 4.56 0.65 1.74 1.46 0.41 1.38 0.84 2.30 4.00 0.55 1.16 2.30 1.85 9.33 34.22 8.86 7.64 5.98 20.23 43.43 25.26 5.52 6.56 11.63 21.46 6.52 7.68 6.40 4.87 8.28 8.19 13.97 21.45 5.26 8.68 9.09 8.70 143.09 445.79 138.43 156.75 72.25 246.94 310.95 117.74 105.00 75.58 128.04 246.69 157.89 36.93 132.41 180.02 180.79 73.13 34.94 72.79 272.99 255.95 73.63 49.36 6.36 7.89 3.21 1.83 4.38 4.29 4.47 8.11 2.74 7.11 6.43 4.95 2.03 2.34 1.39 0.81 1.85 1.72 2.01 2.41 1.33 2.11 2.15 1.82 0.52 0.60 0.36 0.21 0.48 0.44 0.52 0.62 0.34 0.54 0.55 0.47 2000 33,656 49 167 2805 2.52 0.37 16.83 2.45 686.86 77.89 9.75 2.71 0.70 7.17 3.59 37.91 16.58 373.54 238.05 6.07 1.60 0.41 Urbanized Road Tree (ha) (ha) count (no.) D1 D2 D3 D4 D5 D6 D7 D8 D9 D10 D11 D12 Urbanized area 169 248 109 82 241 195 136 107 171 138 253 149 171 1574 8470 969 627 1445 3951 5908 2708 945 907 2945 3207 7843 19 S ¼ Number of tree species. Tree density ¼ Tree count/Land area in ha. Species density ¼ Species richness/Land area in km2. Tree-species ratio ¼ Tree count/Species richness. Person-tree ratio ¼ Population/Tree count. C.Y. Jim, W.Y. Chen / Applied Geography 29 (2009) 577–587 581 of nuisance to road users and adjacent buildings would preclude more species (Galvin, 1999). The high mortality rate of street trees (McPherson, 1994) implies that the species that could not adapt to the harsh streetside milieu would be sieved and eliminated. On the other hand, heavy long tenure of human influence and continual replacement of lost and weak trees with better species (Galvin, 1999) could enrich species diversity. Different species in different road sections reflects conscious decisions to bring inter-site diversity despite the relatively homogeneous and stressful conditions. Wanhua is the oldest centre of Taipei that has experienced different development periods. The development and redevelopment processes have permitted the adoption of different species in tandem with changing landscape styles and preferences. Nangang is a newly built industrial centre, where the relatively uniform urban design and low intensities of human activities have brought monotonous landscapes. A snapshot of the street tree-species composition displays the resultant of continual changes and frequent tree replacements. Urban park trees Urban parks in Taipei have a rather short development history. Except 228 Peace Park opened in 1899, others were installed since the 1970s. The ten main parks contain 139 tree species and 33,342 trees (Table 3). The top 34 park species account for 82% of the population. The top three species, all natives, contribute 33% of tree counts and 47% of potential biomass. Domination by popular species is somewhat subdued in comparison with street trees. Besides F. microcarpa, which is widely planted in all three habitats, only two species with frequency >2% (Liquidambar formosana and C. camphora) are shared between streets and parks. The city has 51 species with tree count >50, but only 11 species are shared between parks and streets, and 18 species are found exclusively in parks. The divergence in species composition between parks and streets is conspicuous. Differences in site conditions and landscape intentions have generated marked species differentiation. Urban parks are mainly semi-natural sites with less human modifications than street habitats. The notable woodland-relic components in parks indicated inheritance from the natural or semi-natural sites (Parks and Street Lights Office, 2000). They serve multiple functions with multiple habitat conditions. The domination by native species attains 49% by tree count which falls below street trees, and 65% by potential biomass which is comparable to street trees (Table 7). The preponderance of broadleaved species, about two-third of which are evergreen, is somewhat diluted by the notable presence of palms. Conifers are minor elements, with only two species (Juniperus chinensis and Cupressus funebris) contributing 3.92% of the tree population amongst the top 34 urban parks species (Table 3). Final height of park trees is mainly large. With a potential biomass of 63%, species with large final dimensions will in time impose increasingly conspicuous landscape impacts on the city. The familiar species such as native B. javanica, C. camphora, F. microcarpa, L. formosana and Machilus kusanoi, and exotic Ficus benjamina, M. quinquenervia, Bombax ceiba and Pterocarpus indicus, will in due course notably raise their landscape and environmental benefits. The large and popular trees are known for their sizeable stature, wide crown, dense foliage and excellent shading effect. However, they do not present showy flowers to enliven the parks in the visual and olfactory senses. Amongst the top ten, only the exotic Prunus serrulata (rank 4) displays ornamental blossoms. Two flowering species in line are exotic Cassia fistula (rank 14) and native K. elegans (rank 15). Despite the presence of 139 park tree species and the general expectation for decorative vegetation in parks, few species with attractive blooms have been planted. Only seven of the top 34 species display ornamental flowers (Table 3), indicating a strong preference for foliage trees with key impacts on the parkscape. By species count, exotics are dominant with 20 common species versus 14 of native provenance (Table 3). Six of the seven flowering species are exotics and only one species (K. elegans) is native (Table 3). Exotic trees thus impart a more prominent presence due to their attractive blossoms. Overall, exotics contribute significantly to species diversity in parks (Table 7). Of the ten major parks, Yangming Park is the biggest with the most trees (10,950 trees or one-third of total park trees) and the maximum tree-species ratio of 304 (Table 4). However, with only 36 species it has the lowest diversity indices (D ¼ 4.898, H ¼ 2.063). Yangming Park is located at edge of the Yangming Mountain Nature Park in a countryside setting. The vegetation is mainly secondary forest planted after destruction of the primary forest in the twentieth century. The massive deforestation has drastically impoverished native species diversity. The relatively young secondary forest has not recuperated from biotic pauperization, although some native species have invaded spontaneously (Wang, Hang, & Lin, 2004). Daan Forest Park has the maximum species richness at 92. It also has the highest diversity indices (D ¼ 21.370, H ¼ 3.493). Overall, there was no significant statistical association between species diversity (H) and park area, opening year and terrain type (all P > 0.10). The variation of E shows the same pattern as H across urban parks, with Yangming registering the least equitability amongst the constituent species, and Daan the highest (Table 4). The small differences between the H and D rankings of urban parks are related to the abundance of common species in Bihu, Qingnian and Jiancheng Parks. The average Shannon–Wiener index of urban parks in Taipei (H ¼ 3.76) is higher than the average tree diversity in the floristically impoverished secondary forest of the region (H ¼ 2.90; Su, 1994). The findings imply that human-oriented functions could accumulate species diversity in urban parks to a high level. For comparison, the study of Kühn, Brandl, & Klotz (2004) found that plant diversity in urban area was higher in German cities than surrounding areas. Taipei has witnessed some recent changes in the basic tenet of park design and use, triggered by the vision of sustainable and ecological city (Lo & Lin, 2007). It has moved notably away from formal-manicured to ecological-naturalistic, and from passive open-space amenity to user-nature interaction (Tu, 2002). Vegetation species diversity has been emphasized in the recently established parks (such as Daan Forest Park added recently to the stock). Comparing with streets, urban parks in general are blessed with a genial environment with subdued negative growth factors. Both site limitations and deleterious 582 C.Y. Jim, W.Y. Chen / Applied Geography 29 (2009) 577–587 Table 3 The composition, landscape features, provenance and quantities of the top 34 landscape tree species in urban parks. Rank Species Family Growth forma Final heightb Provenance Tree count (no.) 1 2 3 4 Ficus microcarpa Machilus kusanoi Sphaeropteris lepifera Prunus serrulata Moraceae Lauraceae Cyatheaceae Rosaceae BLE Large BLE Large Tree fernf Medium BLD Small 5 6 Ficus benjamina Juniperus chinensis Moraceae Cupressaceae BLE Conifer Large Medium 7 Alstonia scholaris Apocynaceae BLE Medium 8 9 10 11 12 13 14 15 Arecaceae Hamamelidaceae Lauraceae Salicaceae Euphorbiaceae Euphorbiaceae Caesalpiniaceae Sapindaceae Palm BLD BLE BLD BLD BLE BLD BLD Medium Large Large Small Large Small Medium Medium Moraceae BLE Small Native 17 18 Chrysalidocarpus lutescens Liquidambar formosana Cinnamomum camphora Salix babylonica Bischofia javanica Mallotus paniculatus Cassia fistula Koelreuteria elegans ssp. formosana Ficus microcarpa cv. golden leaves Trema orientalis Pongamia pinnata Native Native Native Mainland China, Japan, Korea India, Malaysia Mainland China, Japan India, Malaysia, Java, Philippines Madagascar Native Native Mainland China Native Native India Native Ulmaceae Fabaceae BLE BLD 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 Ternstroemia gymnanthera Melaleuca quinquenervia Garcinia spicata Ulmus parvifolia Roystonea regia Archontophoenix alexandrae Bauhinia ’Blakeana’ Bombax ceiba Pterocarpus indicus Cupressus funebris Prunus persica f. duplex Syzygium grijsii Acacia confusa Prunus mume Eucalyptus robusta Sterculia foetida Theaceae Myrtaceae Clusiaceae Ulmaceae Arecaceae Arecaceae Caesalpiniaceae Bombacaceae Fabaceae Cupressaceae Rosaceae Myrtaceae Mimosaceae Rosaceae Myrtaceae Sterculiaceae BLE BLE BLE BLD Palm Palm BLE BLD BLD Conifer BLD BLE BLE BLD BLE BLD Medium Native Medium India, Malaysia, S. China, Ryukyu, Australia Small Native Large India, Malaysia, Australia Medium India, Sri Lanka Small Native Large Central America Medium Queensland Small Hong Kong Large India Large Tropical Asia Large Mainland China Small Mainland China Small Native Small Native Small Mainland China Large Australia Large Tropical Asia 16 Total Average Tree quantity Species biomass Biomass Tree frequency indexd (%)c Biomass index (%)e 4298 3313 3305 1818 12.89 9.94 9.91 5.45 128.94 99.39 66.10 18.18 20.74 15.99 10.63 2.92 1215 1012 3.64 3.04 36.45 20.24 5.86 3.26 839 2.52 16.78 2.70 827 771 765 696 664 625 523 507 2.48 2.31 2.29 2.09 1.99 1.87 1.57 1.52 16.54 23.13 22.95 6.96 19.92 6.25 10.46 10.14 2.66 3.72 3.69 1.12 3.20 1.01 1.68 1.63 414 1.24 4.14 0.67 406 395 1.22 1.18 8.12 7.90 1.31 1.27 378 376 357 348 335 334 321 306 300 295 292 285 267 265 265 246 1.13 1.13 1.07 1.04 1.00 1.00 0.96 0.92 0.90 0.88 0.88 0.85 0.80 0.79 0.79 0.74 3.78 11.28 7.14 3.48 10.05 6.68 3.21 9.18 9.00 8.85 2.92 2.85 2.67 2.65 7.95 7.38 0.61 1.81 1.15 0.56 1.62 1.07 0.52 1.48 1.45 1.42 0.47 0.46 0.43 0.43 1.28 1.19 27,363 805 82.05 2.41 621.66 18.28 100.00 2.94 a For tree growth form, BLE denotes broadleaf evergreen, and BLD broadleaf deciduous. The final tree height that could be attained in Taiwan: small trees are 5–9 m tall, medium 9–18 m and large >18 m. Tree frequency ¼ (Tree count/Total tree number in urban park habitat) 100%. d Biomass dominance index ¼ (Tree count Final height score)/100, where large, medium and small final heights are assigned values of 3, 2 and 1. It indicates the biomass and hence landscape impact of a species upon reaching its final dimension. e Biomass index (%) ¼ (Species biomass index/Total biomass index) 100%. f The tree fern has a palm-like structure and appearance in landscape applications; it is grouped under the palm growth form in this study. b c human impacts are less arresting, and more management inputs and cares are regularly introduced to foster tree performance. This tree growth regime is conducive to longevity, stability in the tree population and a general resistance to changes. Unlike the more changeable street trees, in urban parks the initial tree flora will tend to linger for decades. Riverside park trees Taipei originated as a river port. The Danshui, Jilong and Xindianxi Rivers and associated tributaries form an elaborate watercourse network in the city. The fluvial marsh and banks in Taipei are part of the drainage and flood management system of the drainage basin (Lin & Jiao, 2005). The edge strips along the urban stretch of the river serve as reserved river channels in times of unusual peak discharge to prevent flooding. Such lands also serve as extensive recreational grounds. The riverside C.Y. Jim, W.Y. Chen / Applied Geography 29 (2009) 577–587 583 Table 4 The abundance, distribution and species diversity of landscape trees in ten main urban parks. Code Park District Terrain U1 U2 U3 U4 U5 U6 U7 U8 U9 U10 Zhongzheng Neihu Daan Datong Wanhua Songshan Shilin Zhongshan Beitou Shilin River plain 7.15 1899 Lake & environs 16.39 1987 River plain 25.93 1992 Lower hillslope 1.06 1980 Lower hillslope 24.44 1975 River terrace 2.68 1973 River terrace 2.63 1996e River terrace 19.50 1978 Lower hillslope 125.95 1977 Lower hillslope 10.01 1986 228 Peace Park Bihu Park Daan Forest Park Jiancheng Park Qingnian Park Sanmin Park Shilin Park Xinsheng Park Yangming Park Zhishan Park Total or aggregate Average Range a b c d e Area (ha) 235.74 23.57 124.88 Opening Tree year count (no.) 1339 1195 7675 306 4626 428 2531 3128 10,950 1164 Species TreeSimpson Shannon– Evenness Species Tree density species diversity Wiener richness density index, E a b d (trees/ha) (species/ ratio index D diversity S km2)c index, H 62 28 92 18 63 23 69 31 36 34 187.22 72.92 296.00 287.59 189.28 159.58 961.77 160.41 86.94 116.33 8.67 1.71 3.55 16.92 2.58 8.58 26.22 1.59 0.29 3.40 21.60 13.11 42.68 12.86 83.42 21.37 17.00 5.65 73.43 8.97 18.61 5.30 36.68 9.64 100.90 5.20 304.17 4.90 34.24 9.09 3.16 2.83 3.49 2.09 3.00 2.31 2.90 2.40 2.06 2.64 0.62 0.56 0.69 0.41 0.59 0.46 0.57 0.47 0.41 0.52 33,342 139 3334 14 10,644 74 141.43 0.59 239.87 21.00 3.76 0.74 888.85 25.93 287.17 16.47 1.43 0.28 S ¼ number of tree species. Tree density ¼ Tree count/Land area in ha. Species density ¼ Species richness/Land area in km2. Tree-species ratio ¼ Tree count/Species richness. Previously the official residence of the President; opened to the public as a park since 1996. parks increase the stock of green spaces in Taipei, and offer alternative outdoor recreational venues with reduced facility and management. Despite the rather simple and uniform site condition, 62 tree species have been found in riverside areas (Table 6), a level that lies between street and park habitats. Species richness of individual venues is notably lower than urban parks (Table 6). The top ten species account for 79% of the trees, and top 20 for 91%, indicating a high degree of dominance by popular species (Table 5) that is comparable to street trees but higher than urban parks. Besides the almost ubiquitous native F. microcarpa (rank 1), species with natural affinity for water have been preferentially planted, such as exotic Salix babylonica (rank 2) and Pongamia pinnata (rank 5), Garcinia spicata (rank 7), Pithecellobium dulce (rank 8), and native Palaquium formosanum (rank 9). Only five species are shared with urban parks, merely one with streets, and eight species are exclusive to riverside parks. The pronounced divergence in arboreal components signifies the unique landscape and ecological exclusivity of riverside parks. Like street and urban park sites, riverside parks have more exotics than natives (eight of the top ten species are exotic). The proportion of native trees is the lowest of the three habitats. The biomass index of natives exceeds exotics, implying that the exotics have smaller final dimensions. Riverside parks have the highest proportion of broadleaved trees, especially evergreens, reaching 53% by tree count and 70% by biomass (Table 7). Palm presence lies between streets and urban parks; trees with large final size also lie between them. A total of 5449 trees are enumerated in ten sampled riverside parks, 51% of which dwell in Huazhong Riverbank Park (Table 6). As the largest riverside park, it also has the highest species richness (26 species) and diversity index (H ¼ 2.31). The ranking of E is similar to H. There is slight difference between H and D, which is related to the occurrence of rare species. For example, Caihong Riverside Park has the highest D, which has species with a solitary member, including Ficus elastica, Erythrina corallodendron and F. benjamina. There is no significant statistical relationship between species richness, diversity, site area and opening year (all P > 0.10). Although both park habitats have generous provision of growth spaces, riverside parks are subject to stringent inherent site limitations and they are established for a different purpose. The high water table and wet soil conditions have imposed constraints on species choice. The preference for water-edge and stress-tolerant species reflects this site reality. The amount of flowering or decorative species is the lowest amongst the three habitats. Some trees were inherited from spontaneous growths on river sediments to augment species diversity. Most species do not require horticultural care. The sites could be periodically flooded and impacted by fast-running water during periods of high discharge, and trees intolerant of such stresses could be harmed or eliminated. Trees are deliberately planted at a low density to avoid obstructing water flow. Some sites serve as protected areas or sanctuary for birds and riverine wetland vegetation, as exemplified by Guandu and Huazhong. Planting too many landscape trees is avoided, lest the natural processes of wetland ecosystem are disturbed and its ecological function as wildlife habitat is compromised (Henry & Amoros, 1995). Overall, the pragmatic species choice and planting pattern match the challenging site conditions, and shared use of the land amongst green space, river discharge and nature conservation. Riverside lands are co-used on a time-sharing basis between amenity space and river discharge. As they are designed and managed to perform dualistic functions, the choice of suitable species would be more restricted. With less latitude for species selection and the need to maintain a sparse tree cover to facilitate water discharge, the tree flora is notably less varied than urban parks. With low tree cover and limited recreational facilities, riverside parks attract less patronage and visitor impacts. 584 C.Y. Jim, W.Y. Chen / Applied Geography 29 (2009) 577–587 Table 5 The composition, landscape features, provenance and quantities of the top 20 landscape tree species in riverside parks. Rank 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 Species Growth forma Final heightb Provenance Moraceae Salicaceae Arecaceae BLE BLD Palm Large Small Medium Native Mainland China Madagascar Moraceae Fabaceae Ulmaceae Clusiaceae Mimosaceae Sapotaceae Arecaceae Moraceae Lythraceae Mimosaceae Combretaceae Moraceae Apocynaceae Mimosaceae Hamamelidaceae Euphorbiaceae Lauraceae BLE BLD BLD BLE BLE BLE Palm BLE BLD BLE BLD BLE BLE BLD BLD BLD BLE Large Medium Large Medium Medium Large Small Large Small Medium Medium Small Medium Small Large Large Large India, Malaysia India, Malaysia, S. China, Australia Native India, Sri Lanka Tropical America Native Mainland China, Ryukyu India, Malaysia, Java India, Mainland China Native Native Native India, Malaysia, Java, Philippines Tropical America Native Native Native Family Ficus microcarpa Salix babylonica Chrysalidocarpus lutescens Ficus benjamina Pongamia pinnata Celtis sinensis Garcinia spicata Pithecellobium dulce Palaquium formosanum Livistona chinensis Ficus elastica Lagerstroemia indica Acacia confuse Terminalia catappa Ficus septica Alstonia scholaris Leucanea leucocephala Liquidambar formosana Bischofia javanica Cinnamomum camphora Total Average Tree quantity Species biomass Tree count (no.) Tree frequency (%)c Biomass indexd Biomass index (%)e 1710 730 394 31.38 13.40 7.23 51.3 7.3 7.88 44.65 6.35 6.86 296 291 198 189 179 164 129 110 84 84 71 65 62 57 55 49 46 5.43 5.34 3.63 3.47 3.29 3.01 2.37 2.02 1.54 1.54 1.30 1.19 1.14 1.05 1.01 0.90 0.84 8.88 5.82 5.94 3.78 3.58 4.92 1.29 3.3 0.84 1.68 1.42 0.65 1.24 0.57 1.65 1.47 1.38 7.73 5.07 5.17 3.29 3.12 4.28 1.12 2.87 0.73 1.46 1.24 0.57 1.08 0.50 1.44 1.28 1.20 4963 248 91.08 4.55 114.89 5.74 100.00 5.00 a For tree growth form, BLE denotes broadleaf evergreen, and BLD broadleaf deciduous. The final tree height that could be attained in Taiwan: small trees are 5–9 m tall, medium 9–18 m and large >18 m. Tree frequency (%) ¼ (Tree count/Total tree number in riverside habitat) 100%. d Biomass dominance index ¼ (Tree count Final height score)/100, where large, medium and small final heights are assigned values of 3, 2 and 1. It indicates the biomass and hence landscape impact of a species upon reaching its final dimension. e Biomass index (%) ¼ (Species biomass index/Total biomass index) 100%. b c Table 6 The abundance, distribution and species diversity of landscape trees in ten main riverside parks. Code Park District R1 R2 Shilin Neihu Bailing Left Bank Caihong Riverbank Park R3 Chengmei Left Bank R4 Dajia Riverbank Park R5 Fuhe Riverbank Park R6 Guandu Riverbank Park R7 Guting Riverbank Park R8 Huazhong Riverbank Park R9 Yanping Riverbank Park R10 Yingfeng Riverbank Park Total Average Range a b c d Area (ha) Opening Tree count Species year (no.) richness, Sa Tree density (trees/ ha)b Species density (species/ km2)c Tree-species Simpson ratiod diversity index, D Shannon– Wiener diversity index, H Evenness index, E 25.48 1992 29.00 1998 392 204 13 13 15.39 7.03 0.51 0.45 30.15 15.69 1.72 7.83 1.06 2.18 0.26 0.53 2.20 1990 63 6 28.64 2.73 10.50 3.15 1.29 0.31 36.00 1997 355 17 9.86 0.47 20.88 6.90 2.20 0.53 Wenshan 2.20 1984 510 9 231.82 4.09 56.67 2.46 1.16 0.28 Beitou 2.34 1994 163 8 69.80 3.43 20.38 5.60 1.85 0.45 Zhongzheng 23.35 1988 215 16 9.21 0.69 13.44 3.92 1.88 0.46 Wanhua 54.35 1988 2782 26 51.19 0.48 107.00 6.68 2.31 0.56 Datong 8.78 1967 597 13 68.03 1.48 45.92 2.06 1.15 0.28 53.00 1998 168 13 3.17 0.25 12.92 5.63 2.02 0.49 5449 545 2719 62 23.02 0.26 87.89 7.44 2.75 0.67 228.65 3.85 96.50 6.12 1.25 0.30 Nangang Zhongshan Songshan 236.69 23.67 52.15 S ¼ number of tree species. Tree density ¼ Tree count/Land area in ha. Species density ¼ Species richness/Land area in km2. Tree-species ratio ¼ Tree count/Species richness. 20.00 C.Y. Jim, W.Y. Chen / Applied Geography 29 (2009) 577–587 585 Table 7 Variations in tree growth form, final height and provenance of the top ranking 51 tree species that occur in three main urban forest habitats.a Final heightb Growth form c Species provenance Broadleaf Broadleaf Conifer Palm evergreen deciduous Total Large Medium Small Total Native Exotic-Asiad Exotic-others Total Street Species count (no.) Tree count (no.) Tree frequency (%) Species biomass index Species biomass (%) 6 16,240 48.25 477.22 57.85 12 13,999 41.59 292.39 35.44 0 0 0.00 0.00 0.00 2 2002 5.95 55.36 6.71 20 32,241 95.80 824.97 100.00 9 23,133 68.73 693.99 84.12 8 3990 11.86 79.80 9.67 3 5118 15.21 51.18 6.20 20 32,241 95.80 824.97 100.00 7 21,685 64.43 551.01 66.79 11 8227 24.44 209.71 25.42 2 2329 6.92 64.25 7.79 20 32,241 95.80 824.97 100.00 Urban park Species count (no.) Tree count (no.) Tree frequency (%) Species biomass index Species biomass (%) 15 14,124 42.35 361.90 58.22 13 7131 21.38 131.30 21.12 2 1496 4.49 33.27 5.35 3 4612 13.83 95.19 15.31 33 27,363 82.05 621.66 100.00 13 13,149 39.43 394.47 63.45 10 8505 25.51 170.10 27.36 11 5709 17.11 57.09 9.18 34 27,363 82.05 621.66 100.00 14 16,346 49.01 401.86 64.64 18 9855 29.56 193.21 31.08 2 1162 3.48 26.59 4.28 34 27,363 82.05 621.66 100.00 Riverside park Species count (no.) Tree count (no.) Tree frequency (%) Species biomass index Species biomass (%) 10 2905 53.31 80.71 70.25 8 1535 28.17 25.01 21.77 0 0 0.00 0.00 0.00 2 523 9.60 9.17 7.98 20 4963 91.08 114.89 100.00 8 2628 48.23 78.84 68.62 7 1270 23.31 25.40 22.11 5 1065 19.54 10.65 9.27 20 4963 91.08 114.89 100.00 9 2442 44.82 70.41 61.28 8 1891 34.70 32.45 28.24 3 630 11.56 12.03 10.47 20 4963 91.08 114.89 100.00 All habitats Species count (no.) Tree count (no.) Tree frequency (%) Species biomass index Species biomass (%) 21 33,269 51.53 919.83 58.91 23 22,665 35.10 448.70 28.73 2 1496 2.32 33.27 2.13 5 7137 11.05 159.72 10.23 51 64,567 100.00 1561.52 100.00 17 38,910 60.26 1167.30 74.75 18 13,765 21.32 275.30 17.63 16 11,892 18.42 118.92 7.62 51 64,567 100.00 1561.52 100.00 19 40,473 62.68 1023.28 65.53 27 19,973 30.93 435.37 27.88 5 4121 6.38 102.87 6.59 51 64,567 100.00 1561.52 100.00 a b c d Only the common species included in Tables 1, 4 and 6 have been included in the computations. The final tree height that could be attained in Taiwan: small trees are 5–9 m tall, medium 9–18 m and large >18 m. The tree fern Sphaeropteris lepifera has been grouped under the palm growth form in view of its palm-like appearance and landscape applications. Including Australia as the Australasian biogeographical realm. Trees in riverside parks are expected to persist, and many could realize their biological life span and dimensions to sustain their ecosystem functions. Conclusion and implications The high-density and compact urban form of Taipei has not stifled flora diversity and spatial variability of its urban forests. Our findings suggest that the tree stock in three main habitats of Taipei tend to diverge and is envisaged to maintain their divergence. Natural and human factors have differential effects on tree diversity in different habitats. The interplay of inherent site characteristics and management intervention has jointly determined the tree-species pattern. Similar to landscape ecological changes in countryside areas, dominating human factors interact with biophysical site conditions to influence the resulting vegetation composition (Serra, Pons, & Sauri, 2008). Street trees are beset by a rather uniformly stressful growth regime and unvarying site preparation and design, and they share common but narrow planting objectives. The acute disturbances of street environment might bring extensive premature tree decline, high turnover rate and low floristic stability. Species composition in urban parks is influenced by the varied functions assigned to different venues and trees inherited from pre-development vegetation, moderated by diverse inherent site conditions, varying site preparation and design, and evolving landscape planting fads. Riverside parks take an intermediate position between streets and urban parks, the species diversity of which is mainly determined by their special environmental constraints and function of facilitating water discharge. Understanding the status and dynamics of species diversity marks the first step in nature and biodiversity conservation in urban areas (Alvey, 2006). Based on our findings, we have deduced some implications to promote and preserve urban biodiversity in urban areas. On the one hand, introducing more indigenous species could create biologically rich urban forests to benefit not only humans but also the sustainability of urban ecosystems, especially for street habitats with a small species cohort. Such simple composition could accentuate the biotic hazard of tree diseases and pests, and reduce the stability of its vegetation communities. The capability to attract wildlife into the city’s linear street tree belts is also limited by a simplistic and overly exotic species assemblage. On the other hand, biodiversity in urban ecosystems could be enhanced by creating or enhancing a wide range of habitats (Goode, 1993; Wheater, 1999), and varied habitats could be inserted in tandem with the city’s development and redevelopment (Harrison & Davies, 2002). In Taipei, beautifying the urban landscape and providing recreational grounds remain the principal aims of the open-space programme (Research, Development and Evaluation Commission, 1997). Most parks were or will be built to meet these overriding objectives (Public Works Department, 2004). 586 C.Y. Jim, W.Y. Chen / Applied Geography 29 (2009) 577–587 However, biodiversity conservation has not been firmly incorporated in the long-term planning vision (Yang, 1989). Two recently built natural parks have been designed mainly for recreation and ecological education purposes, and nature conservation is considered as a pleasant by-product. In the continual expansion of the city into its peri-urban countryside, areas with high ecological value could be identified for protection as future urban natural enclaves. Measures could be taken to forestall the impacts on the high quality forests due to continued urban sprawl at the city’s hilly fringe (MacDonald & Rudel, 2005). A move away from the deeply ingrained conventional thinking is needed to embrace new ideas and practices in urban green space design. Recent studies in urban ecology and biogeography have provided ample evidence of high biodiversity in urban areas. Empirical findings suggested species enrichment in urbanized areas in comparison with the surrounding countryside (Kühn, Brandl, & Klotz, 2004). A similar effect was found in this case study. Social forces could impose notable increase in green spaces and species composition also in rural areas by social recomposition leading to re-landscaping of gentrified villages (Phillips, Page, Saratsi, Tansey, & Moore, 2008). Specifically, such species enrichment seems to be confined to targeted functions while the natural environment still plays a certain role in the diversification process. However, due to the shortage of historical record of tree plantation and introduction, quantitative analysis of the relationship between tree biodiversity and development history so far is impossible. Regular long-term observations could explore tree-species dynamics in Taipei and other subtropical cities. Further study is needed to ascertain whether such a trend could result in homogenization due to increase in introduced species at the expense of native, or diversification due to co-existence of native and exotic species. 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