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Landscape and Urban Planning 95 (2010) 122129

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Characterizing the fragmentationbarrier effect of road networks on landscape connectivity: A case study in Xishuangbanna, Southwest China
Wei Fu a , Shiliang Liu a, , Stephen D. Degloria b , Shikui Dong a , Robert Beazley c
a b c

School of Environment, State Key Laboratory of Water Environment Simulation, Beijing Normal University, No. 19, Xinjiekowai Street, Beijing 100875, China Department of Crop and Soil Sciences, Cornell University, Ithaca, NY 14853, USA Graduate Student Department of Natural Resources, College of Agriculture and Life Sciences, Fernow Hall 302, Cornell University, Ithaca, NY 14853, USA

a r t i c l e

i n f o

a b s t r a c t
The fragmentationbarrier effect of road network expansion on landscape function draws attention throughout the world. Presently, there is a lack of quantitative analysis to integrate different ecological effects of road networks at landscape scale to depict, compare and evaluate landscape changes. In this study, the Probability of Connectivity (PC) index was used to evaluate the effects of road networks on landscape connectivity for an area in Xishuangbanna, Southwest China. The results indicate that the fragmentation effect of road networks decreased in PC value by 15.81%, the barrier effect of road networks decreased in PC value by 11.73% and the combination of the two effects decreased PC value by 32.78%. In conclusion, the combined fragmentation and barrier effects of road networks considerably degraded landscape connectivity more than either individual effect. In addition, the fragmentation effect inuenced connectivity to a greater degree for ecological processes having low movement abilities. The barrier effect inuenced connectivity to a greater degree for medium to high movement abilities. The combined effects inuenced connectivity for those ecological processes of low movement ability within the study area. 2009 Elsevier B.V. All rights reserved.

Article history: Received 11 February 2009 Received in revised form 16 November 2009 Accepted 4 December 2009 Available online 12 January 2010 Keywords: Landscape ecology Effective distance Fragmentation effect Barrier effect Connectivity index

1. Introduction As early as 1970, wildlife biologists began publishing research on the effect of roads on wildlife populations as barriers to movement (Bhattacharya et al., 2003; Mech et al., 1988; Oxley et al., 1974), sources of mortality (Bellis and Graves, 1971; Dodd et al., 2004) and causes of behavior modication (Rost and Bailey, 1979; Tigas et al., 2002) at less than ecosystem scale. Coincident with the development of landscape ecology and landscape scale analysis, attention to road ecology has turned to the broader scale effects of landscape and habitat fragmentation and, specically, the qualitative effect of roads on fragmentation and interactions with the landscape to sustain ecological processes (Cofn, 2007). Quantitative methods to assess the impacts of road networks on landscape connectivity at broad spatial scales, however, are lacking and need to be developed. The concept and measurement of landscape connectivity are needed to overcome the limitations of empirical indices at small scale and landscape pattern indices at large scale for characterizing landscape change. Landscape connectivity has been dened as the

Corresponding author. Tel.: +86 13522671206; fax: +86 1058800397. E-mail addresses: (W. Fu), (S. Liu), (S.D. Degloria), (S. Dong), (R. Beazley). 0169-2046/$ see front matter 2009 Elsevier B.V. All rights reserved. doi:10.1016/j.landurbplan.2009.12.009

degree to which the landscape facilitates or impedes movement among resource patches (Taylor et al., 1993). Such connectivity is considered important for ecological processes such as movement of genes, individuals, species, and populations over multiple spatiotemporal scales, especially in fragmented landscapes (Minor and Urban, 2008). Landscape connectivity is currently viewed either structurally, where connectivity is entirely based on landscape structure (usually habitat contiguity), or functionally, where behavioral responses to landscape elements (patches and edges) are considered with the spatial structure of the landscape (Tischendorf and Fahrig, 2000). Being an essential issue of landscape analysis, approaches and indices using different kinds of measurements have been suggested (Marulli and Mallarach, 2005; Pascual-Hortal and Saura, 2006; Minor and Urban, 2008). Among many indices developed, a newly proposed index Probability of Connectivity (PC) which based on graph theory performs well in practical landscape analysis for many good properties in expressing landscape dynamics (Saura and Pascual-Hortal, 2007). Recent applications of this index, however, have not used effective (minimum-cost) distances to combine features of the landscape matrix with features of source patches for connectivity analysis yet. These relevant evaluations of connectivity broadly applied in biological conservation and open area planning at large scale need to consider the impacts of road networks. Currently, road construction, especially high-level road construction is occurring very rapidly in China. In 2007, the total length

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Fig. 1. Location of study area in Xishuangbanna. Legend: China, Yunnan Province, Xishuangbanna, study area, expressway, rst level road, second level road, third level road, fourth level road.

of roads in China reached 357 104 km including 5.36 104 km of expressways. To realize the modernization of its transportation network, Chinas national road construction plan calls for 200 104 km of highways, including 6.5 104 km of expressways by 2010; by 2050, the plan calls for 400 104 km of highways. The Great Development in West China project has resulted in many high-level roads being constructed in Yunnan Province to connect China with Southeast Asia (Liu et al., 2008). As highways account for more than 93% of the total transportation routes in Yunnan, Xishuangbanna, as a border administrative region under the development of tourism, inevitably experiences the drastic pressure of road network expansion on local ecosystems. In addition, Xishuangbanna is a key biogeographic area and a hotspot for biodiversity (Myers, 1998). Therefore, research on the effects of road networks on landscape connectivity in this region of China will have signicant practical importance for biodiversity conservation and road network planning. The objectives of this study are to: (1) assess landscape changes due to road networks in a typical landscape in southwest China by applying a landscape connectivity index combined with effective distances; and (2) characterize the barrier-fragmentation effect of road networks on specied landscapes under different ecological process scenarios. The focused ecological processes are those that occur in suitable patches and could benet from increasing connectivity from a landscape-level perspective. The different dispersal/movement distances are considered broadly representative of different ecological processes in the study. 2. Materials and methods 2.1. Study area Xishuangbanna is situated in a transitional zone from tropical Southeast Asia to subtropical East Asia, and is at the junction of the

Indian and Burmese plates of Gondwana and the Eurasian plate of Laurasia which has great biological diversity and typical ecosystems (Zhu et al., 2006). The elevation varies from 475 m to 2430 m above sea level. Annual mean temperature is approximately 21 C and the annual precipitation is over l500 mm. Primary vegetation can be organized into four main types: tropical rain forest, tropical seasonal moist forest, tropical montane evergreen broad-leaved forest and tropical monsoon forest (Zhu, 2006). Human population increased from 220,000 in 1953 to 990,000 in 2000 in Xishuangbanna, most of which is now distributed in the city of Jinghong. Due to increasing population pressures, large areas of tropical rain forest and shifting cultivation lands at lower elevations have been converted to rubber plantations during the past 50 years, thereby inducing the harvest of forests at high elevations or steep slopes to meet the requirement for new arable land (Li et al., 2007). Roads motivated by human encroachment in the study area promote landscape modication signicantly (Li et al., 2009). The location of the study area (21.721.8 N, 100.4100.8 E) is situated near the boundary of Jinghong city and Mengla county for our landscape connectivity analysis independent of impacts of the Lancang River which was excluded from our study (Fig. 1). Within the entire region of Xishuangbannan, the study area possesses high biodiversity. Forest land occupies 40.08% of the land area (1600 km2 ) including nearly all four types of vegetation in the study area. Therefore, there are sufcient and diverse forest patches to accommodate ecological processes in the study region. However, human exploitative activities that are supported by the high density of road networks (0.006 km/km2 ) lead to severe disturbance of the regional landscape. 2.2. Data acquisition The road data in vector format for this study were digitized using the present transportation map of Yunnan province and also


W. Fu et al. / Landscape and Urban Planning 95 (2010) 122129

conrmed using the 1:250,000 scale road database produced by the National Fundamental Geographical Information Centre in 2005. Roads in this area can be divided into ve levels: expressway, rst level (national road), second level (mainly county citycounty city road), third level (mainly county citytown road) and fourth level (mainly townvillage and villagevillage road). Landsat TM images in 2007 were classied as forest, grassland, farmland, saline land and marshland, urban land, rural residential land, and construction land for land cover mapping. Then the map was validated using ground truth data which indicated the overall accuracy of at least 90% and Kappa coefcient of 0.88. 2.3. Connectivity analyses The methodology in this study is to analyze the fragmentation and barrier effects of road networks using the landscape connectivity index, PC. The fragmentation effect is related to the dissection processes of increasing the number of landscape patches, decreasing interior habitat area, or increasing the extent of opening edges (Li and Reynolds, 1993). The barrier effect is with regard to patch isolation effects which inhibit the movement of organisms for focal ecological processes occurring between patches (Richardson et al., 1997). The parameters of resource patch size reduction and the resistance value of landscape elements in PC calculations represent the two effects of road networks. The calculation of the index proceeds under different scenarios of ecological processes such as plants or animals dispersal movement dened by movement abilities. In each scenario, the landscape without road network was set as the control test of the landscape to distinguish the effects of roads from other landscape elements on ecological processes. Thus, the analysis consists of several steps as described below. 2.3.1. Resource patch selection and fragmentation effect quantication According to Liu and Li (2008), the rst processing step of resource patch selection should be carried out following these three principles: (i) the patch should have high landscape suitability, (ii) the patch area should be large enough to sustain ecological processes, and (iii) accessibility from any resource patch to others should be considered. Landscape suitability here means the tness of the landscape type to support a particular ecological process such as decomposition, nitrogen cycling, pollination, seed dispersal, animal mitigation, bioturbation and so forth (Forman and Deblinger, 2000; Nichols et al., 2008). A series of dispersal movements of plants or animals that could benet from improved connectivity in specied patches was taken into account in this study. However, available seed dispersal distance information seems more than any other ecological process in Xishuangbanna. Most of the seeds are reported to travel a short distance to other places within approximately 1 km (Tang et al., 2008; Zhou et al., 2007), while others have long dispersal distances to as much as 27 km, though this rarely occurs (Xiao and Gong, 2006). Without additional research related to seed dispersal by large mammals, the long-distance seed dispersals in Xishuangbanna may exist but are rarely studied as compared with the discovery of 512 km seed dispersal distances by elephants in other parts of world (Theuerkauf et al., 2000). High landscape suitability is indicated by the forest land cover type as this type accommodates more ecological processes of interest in this study (Liu et al., 2002). The size of patches is both processand region-specic. The lack of references regarding patch size for selected ecological processes in Xishuangbanna made the selection of patch size difcult. For identifying and comparing the degree of impact, a set of minimum-sized patches needs to include as many ecological processes as possible based on the methods of PascualHortal and Saura (2007) and Platt and Lowe (2002). Accessibility

is considered in determining size of the study area and in modeling dispersal movement distance for our patch ecological processes within the study area. Finally, we dened resource patches in the study area as forest land cover with canopy density higher than 30% and size larger than 25 ha based on the characteristics of the environment and related researches mentioned previously (Liu et al., 2002; Platt and Lowe, 2002; Pascual-Hortal and Saura, 2007). The fragmentation effect was quantied by the reduction in area of resource patches due to the occupancy and incision by road networks. Patches were intersected by road networks with the true width as follows: expressway, 30 m; rst level road, 15 m; second level road, 10 m; third level road, 7 m; fourth level road, 3.5 m. 2.3.2. Scenario analysis According to Kahn and Wiener (1967), scenarios were dened as hypothetical sequences of events constructed with the purpose of focusing attention on causal processes and decision points. The quantitative (modeling) and qualitative (narrative) traditions of scenario analysis have been applied in different elds of study in addition to addressing environmental issues of concern here. To test road network effects on landscape connectivity, this study modeled a series of ecological processes (dispersal movements of species) that could benet from improved connectivity in suitable patches. Ecological processes are represented as different dispersal/movement distances (scenarios) broadly taking place in the study landscape. According to studies on dispersal distances for different assemblages of species (Sutherland et al., 2000; Van Vuren, 1998), four scenarios are dened in the model of PC with the different distances and the same direct dispersal probability (pij ) set at 0.5. For each scenario, the same value of 0.5 will indicate the probability of the occurrence of a random event. So, the direct probability of the occurrence of dispersal movement between any two patches equals 0.5 based on probability theory at cost-distances of 12,000 m, 8000 m, 4000 m and 2000 m, respectively. 2.3.3. Effective distance calculation Connectivity analysis may consider effective distances and not just Euclidean distances. Two layers, a source layer and a friction/resistance layer, form the input of the effective distance computation. The source layer indicates the resource patches from which effective distance is calculated. The resistance layer indicates both the resistance value and the geographical position and orientation of all relevant landscape elements (Adriaensen et al., 2003). Herein, the road data are treated as an individual layer other than the mixture with other landscape elements in the land cover map (Marulli and Mallarach, 2005). The road networks buffer zone layer generated by the specic ecological zone constitutes the resistance layer together with the land cover and slope gradient layer (Li et al., 2004). The land cover types are in accordance with the classication of the TM image. The slope gradient layer is classied into ve types with equal interval of 19%. The resistance values for classifying each layer are dened with reference to the literature (Marulli and Mallarach, 2005) and consultation with local experts (Table 1). Then, the resistance layer was converted to a raster with grid size of 25 m 25 m for distance calculations following the calculating equation of effective distances (Yu, 1999). 2.3.4. Landscape connectivity calculation The probability of connectivity index (PC) is dened as the probability that two individuals in dispersal movements randomly placed within the landscape fall into habitat areas that are accessible from each other (interconnected) given a set of n habitat patches and the connections (pij ) among them. In this study, the PC index represents the probability of occurrence of focal ecological processes dened before. The PC index is calculated by the following

W. Fu et al. / Landscape and Urban Planning 95 (2010) 122129 Table 1 Resistance values of different layers. Layer Road Classication Expressway First level Second level Third level Fourth level Forest (except the source patches) (28.24%) Grassland (22.34%) Farmland (15.54%) Saline land and marshland (0.05%) Urban land (11.23%) Rural residential land (8.25%) Construction land (2.51%) 019% 2039% 4059% 6079% Buffer 1000 m 500 m 250 m 100 m 25 m Daily trafc volume 25,000 10,00025,000 200010,000 2002000 100200


Resistance 8 8 8 8 8 2 2 3 4 6 6 6 1 2 3 4

Land cover


expression (Saura and Pascual-Hortal, 2007): PC =

n i=1 n a a p j=1 i j ij A2 L

3.1. Fragmentation effect on landscape connectivity (1) As the result of the intersection of the resource patches by road networks, the resource patches in the study area increased in number to 124, but decreased in area to 10.61% of the study area. Using Eq. (1), road networks decreased PC in different scenarios by 15.81% on average. The effect of fragmentation on resource patches and paths of landscape connectivity are represented through the p ij selection above 0.5 and the classication of dPC of the patches for each scenario (Fig. 2). According to Fig. 2, compared with the landscape without road networks, the paths increased mainly among the fragments of the former resource patches (without road network). However, the length and location of the increased paths varied with different ecological process scenarios. In the entire landscape of the study area, the total PC value decreased by 3.94%, 6.52%, 16.32% and 36.47% in different scenarios with pij of 0.5 at distance of 12,000 m, 8000 m, 4000 m, and 2000 m, respectively. At the patch scale, the importance of each patch changed only slightly as the movement ability decreased (from top to bottom in Fig. 2). In each scenario, the large patches of high importance (dPC) reduced in area as the surrounding patches increased to the average importance. The fragmentation induced by road networks generally had negative inuence on the landscape in sustaining connectivity in the study area, especially on those patches directly crossed by road networks. The fragmentation affected to a greater degree landscapes that sustain ecological processes of low movement ability (dispersal distance at approximate 2000 m). In other words, the movements of those ecological processes which were often constrained in a limited number of patches within small extent were more vulnerable to fragmentation of the patches. 3.2. Barrier effect on landscape connectivity The quantication of the barrier effect using the resistance value increased the effective distances between resource patches among 2415 paths associated with the scenarios. Based on Eq. (2), the decreased rate of the PC index value was 11.73% on average in the study area. The barrier effect of road networks generally decreased the connection paths (p 0.5) between the patches (Fig. 3). In addiij tion, the number of paths removed by the overlay of the road networks was 56, 20, 7 and 3, respectively with movement ability decreased. Due to the removal of paths, PC value of the entire study area decreased by 15.30%, 15.57%, 11.33% and 4.72%, in different scenarios with pij of 0.5 at distance of 12,000 m, 8000 m,

where ai and aj are the areas of the resource patches i and j, and AL is the total landscape area (area of the study region); p is dened ij as the maximum product probability of all possible paths between patches i and j (including single-step paths); and p is determined ij by the pij which is following a negative exponential as a function of effective distances set in the study. It is given by the following expression: pij = edij (2)

where pij is the direct dispersal probability, dij is the effective distances between each two patches, is a constant which is determined by a pair of pij and dij values. Conefor Sensinode 2.2 (Saura and Torn, 2009) was used for all index calculations. Landscape connectivity of the study area is calculated under different movement ability scenarios with pij set at 0.5 when dij equals 2000 m, 4000 m, 8000 m, and 12,000 m, respectively. Also, the paths between any two patches with p over 0.5 are selected to depict the ij degree of connection. 2.3.5. Percentage of importance calculation The relative ranking of landscape elements by their contribution to overall landscape connectivity according to a certain index (I) can be obtained by calculating the percentage of importance (dI) of each resource patch (Pascual-Hortal and Saura, 2006). It is given by: dI (%) = II 100 I (3)

where I is the PC index value when the resource patch is present in the landscape and I is the PC index value after removal of that patch (e.g. after loss of a certain habitat patch). The ranking of value of dPC over the landscape indicates the importance of patches in sustaining landscape connectivity under the inuence of road networks. The importance of the patches is classied into three levels based on PC as high importance, medium importance, and low importance. 3. Results and discussions After the selection of resource patches, there were 70 patches occupying 10.91% of the 40 km 40 km extent of the study area.


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Fig. 2. Fragmentation effect on landscape connectivity under different ecological processes. Legend: patch, path, study area, low importance, medium importance, high importance. (AD) Four scenarios with pij set at 0.5 at dij of 12,000 m, 8000 m, 4000 m and 2000 m, respectively. (i) With road network overlay; (ii) without road network overlay. Path: cost-distance path with p over 0.5. Importance: classied as ij high, medium, and low based on dPC with equal interval.

Fig. 3. Barrier effect on landscape connectivity under different ecological processes. Legend: patch, path, study area, low importance, medium importance, high importance. (AD) Four scenarios with pij set at 0.5 at dij of 12,000 m, 8000 m, 4000 m and 2000 m, respectively. (i) With road network overlay; (ii) without road network overlay. Path: cost-distance path with p over 0.5. Importance: classied as high, ij medium, and low based on dPC with equal interval.

4000 m, 2000 m, respectively. The importance of each patch, however, changed under different ecological process scenarios except for a patch of which the importance changed from medium to low with increase in movement ability. Road networks had a negative effect on landscape connectivity as a typical barrier and changed the importance of certain patches in accommodating the ecological processes in general by increasing the effective distance and then reducing the maximum probability of the movement between any two patches. For different ecological processes (dened by the pij = 0.5 at different distances), the road

network exhibited different degrees of barrier effect on the landscape, generally, with the movement ability decreased, the negative effect degree increased and then decreased, reaching the maximum at 8000 m (pij = 0.5). Results indicated that certain ecological processes of higher movement ability (dispersal movement distance at approximate 8000 m) within the large extent of the landscape were more inuenced than those within the small extent by the networks of roads as a type of barrier. However, the barrier effect merely changed the importance of patches for landscape connectivity under different ecological process scenarios

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3.3. Combined effects on landscape connectivity Up to this point, we have evaluated the fragmentation and barrier effects individually on landscapes to sustain the connectivity for different ecological processes. The combined effect of the two will be analyzed here and compared with the two individual effects. Both the reduction in source patches and the increases in resistance value were calculated. The combined effects of fragmentation and barriers produced an average decreased rate of PC value in comparison to the control test of 32.78% under different ecological process scenarios. This result was higher than the

Fig. 5. Comparison of different effects under different ecological processes. Legend: AD. Label: PC num (1015 m2 ), effects. Control: without road network overlay; f: fragmentation effects; b: barrier effects; c: combined effects. (AD) Four scenarios with pij set at 0.5 at dij of 12,000 m, 8000 m, 4000 m and 2000 m, respectively.

fragmentation effect of 15.81% and barrier effect of 11.73%, individually. The combined effects made the importance changes of patches for landscape connectivity a little different from those under individual effects (Fig. 4). Although it is the same as the individual effect that most of the patches importance decreased except that few patches decreased, the changes of importance took place in different patches at different degree. The total decreased PC value rate to control test was 26.34%, 27.25%, 31.62% and 45.92%, with pij of 0.5 at distance of 12,000 m, 8000 m, 4000 m, and 2000 m, respectively which are higher than the individual effects. Comparing the different effects, the combined effects of road networks in the study area resulted in a signicantly higher decrease in PC value than the two effects considered independently as shown in Fig. 5. Although each effect decreased landscape connectivity, the degree of decrease differed with movement abilities of ecological process dened before. The fragmentation affected the ecological processes of low movement abilities (dispersal movement distance at approximate 2000 m) to a greater extent. The barrier effect affected the medium high movement abilities (dispersal movement distance at approximate 8000 m) to a greater extent. The combined effects affected the ecological processes of low movement abilities (dispersal movement distance at approximate 2000 m) to a greater extent. To summarize, the combined effects, rather than the individual effects, of road networks may considerably decrease landscape connectivity and further degrade the landscape function for facilitating movement of organisms. 4. Summary and conclusions This case study utilized landscape connectivity as a new thrust for the study of ecological effects of road networks. The study took different dispersal/movement distances broadly representative of selected ecological processes to analyze landscape pattern and associated function changes related to road networks and the surrounding landscape. Our results indicate that (i) the PC index is a good measure of landscape dynamics, (ii) landscape connectivity is a simple approach for scenario analysis, and (iii) the analysis of landscape connectivity is useful for advancing our understanding of the ecological effect of road networks. Previously, many studies have concentrated on the effects of road networks at large spatial scales related to land cover changes (Liu et al., 2006), habitat fragmentation (Theobald et al., 1997), barrier effects (Forman and Deblinger, 2000), and function change (Wen et al., 2007). Roads are considered as sources of fragmen-

Fig. 4. Combined effects on landscape connectivity under different ecological processes. Legend: patch, path, study area, low importance, medium importance, high importance. (AD) Four scenarios with pij set at 0.5 at dij of 12,000 m, 8000 m, 4000 m and 2000 m, respectively. (i) With road network overlay; (ii) without road network overlay. Path: cost-distance path with p over 0.5. Importance: classied as high, ij medium, and low based on dPC with equal interval.


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Fig. 6. The impact degree of the different roads in quantication of barrier effect. Legend: cross-road movement. Accumulated resistance = 8 40 = 320. dij = 8(40d0 + 39 25). Expressway accumulated resistance = 8 1 = 8. dij = 8 d0 . Fourth level road. dij : effective distance of the expressway; d distance of the rst grid the movement arrived (equal set for the comparison of the two levels of roads).
ij :

effective distance of the fourth level road; d0 : the

tation, barriers for most of the species considered, and causes of landscape function degradation. Road networks appear to have a disadvantage over other anthropogenic causes as elicitors of landscape fragmentation and barrier to the ecological processes within the landscape (Forman et al., 2003). With the extension and connection of road networks, the landscape becomes more fragmented and less well connected. Few studies, however, have incorporated a series of ecological processes in the quantication of road effects due to the large requirements of data to characterize the behavior of target species and logistical difculties of investigations at such scales. Estimation of the road effect zone and the selection of target species may be very difcult at landscape scale for the reason that the extent of road effect zones may be different for different regions (Forman and Deblinger, 2000; Li et al., 2004). In addition, target species cannot represent all of the important species found in a given study area. Therefore, the road effect zone and the dispersal distances of species as reported in the literatures are utilized to quantify the road effect and to integrate ecological processes in the study. The fragmentation effect was quantied by the direct dissection of resource patches while the barrier effect was quantied by the allocation of the cost in effective distance calculations. Using the same cost value of the road networks may seem counterintuitive, but the large extent of the ecological effect zones for high-level roads could reveal the higher barrier effect on the ecological processes than the lower level roads. The large extent of the ecological effect zone has already been considered as having a higher degree of impact by higher level roads (Li et al., 2004). This impact, through crossroad movement of ecological processes, is the focus movement for

quantifying the barrier effect (Dyer et al., 2002). The higher level roads (such has expressway) represent higher accumulated resistance values and effective distances than the lower level roads (such as fourth level roads) (see Fig. 6). This study evaluated landscape changes by combining quantitative methods with the PC index to characterize the effect of road networks on landscape connectivity. Herein, the separate and combined fragmentationbarrier effects were calculated and compared on the landscape by PC in scenarios of different ecological processes. In conclusion, the fragmentationbarrier effect had a negative effect on landscape connectivity. The combined inuence of the two effects had much higher degree of impact than the individual effects. The different ecological processes may endure different degrees of threats from different types of road network effects: the ecological processes of low movement abilities (dispersal movement distance at approximate 2000 m) are impacted more from the fragmentation effect the combined effects of fragmentation and barriers; those of medium high movement abilities (dispersal movement distance at approximate 8000 m) are impacted more by the barrier effect. While this study provided an example of the usefulness of land connectivity analysis in road network assessment, additional studies are still required. For example, more detailed investigations related to dispersal movement behavior of target species need to be conducted at the same spatial scale for improving the application of the PC index for biodiversity conservation. Furthermore, the study only considered two effects of road networks for the specied landscape in one typical study area. The applicability of our results is only constrained for road network planning and assessment in this

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area, and should not be used to develop policies to prevent landscape connectivity from other negative effects of road networks. Also, additional analysis should be expanded to other landscapes of high heterogeneity with integration of detailed dispersal information of species to identify the ecological effects of road networks at suitable spatial scales using the PC index as the metric for reference. Acknowledgments The paper was nancially supported by the National Natural Science Foundation of China (40871237), National Basic Research Program of China (No. 2003CB415104) and was inspired by Santiago Saura and Luca Pascual-Hortals research on landscape connectivity measurements (PC) and software (Conefor Sensinode 2.2) development. References
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