Ecological Indicators 36 (2014) 160–168

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Ecological Indicators
journal homepage: www.elsevier.com/locate/ecolind

Forest fragmentation and landscape connectivity change associated

with road network extension and city expansion: A case study in the
Lancang River Valley
Shiliang Liu a , Yuhong Dong b,∗ , Li Deng a , Qi Liu a , Haidi Zhao a , Shikui Dong a
a
School of Environment, State Key Laboratory of Water Environment Simulation, Beijing Normal University, No. 19, Xinjiekowai Street, Beijing 100875,
China
b
Research Institute of Forestry, Chinese Academy of Forestry, Beijing 100091, China

a r t i c l e i n f o a b s t r a c t

Article history: Human disturbances contribute significantly to the decrease in the quantity and connectivity of the
Received 5 January 2013 natural habitats. Taking Lincang City in Lancang River Valley as a case study, we focused on the forest
Received in revised form 11 July 2013 fragmentation and habitat connectivity loss associated with its road network and expansion in the past 15
Accepted 15 July 2013
years. Our findings show that, from 1991 to 2006, its forest habitat (>25 ha) decreased to 18.3% of the total
area due to fragmentation. Landscape fragmentation indices (NP, PLAND, and LPI) changed more than
Keywords:
the landscape shapes and the aggregation indices (SHAPE, FRAC, and DIVISION). Habitat loss occurred
Road effect zone
more often in the lower elevations and in areas near the city and road network. Landscape connectivity
Fragmentation
Habitat change
loss was 91.3%, which showed a more significant decrease than did the landscape pattern changes. With
Landscape connectivity sensitivity analysis of different animals’ dispersal abilities and landscape resistance consideration, our
Dispersal distance study reveals that landscape connectivity could be more effective in showing the potential ecological
effects caused by city and road network extension.
© 2013 Elsevier Ltd. All rights reserved.

1. Introduction
Landscape fragmentation as a breakup of a continuous habitat
into smaller, less connected patches can impose devastat-
ing and irreversible consequences on the regional biodiversity
(Trakhtenbrot et al., 2005; Brooks et al., 2006). Transportation
infrastructures built in line with a city’s expansion, particularly
road networks, cause both direct and indirect losses of habitat,
which, in turn, influence the abundance and distribution of plant
and animal species (Geneletti, 2003; Eigenbrod et al., 2007).
Most studies of the effects of roads on biodiversity are carried
out separately on different scales, with little effort to integrate
the results of these multiple scales (Laurance et al., 2004). On a
local scale, biodiversity characteristics such as mortality, move-
ment, distribution, and species richness are more often investigated
(Gibbs and Steen, 2005). At the landscape level, fragmentation
and land cover changes can be quantified using GIS methods for
habitat changes. However, biodiversity evaluations on a small
scale due to road construction are far from comprehensive due
to the lack of related knowledge, especially the species’ behav-
iors, while landscape-level analyses such as pattern and spatial
analysis often have limited ecological meanings for biodiversity
∗ Corresponding author. Tel.: +86 10 13681018528.
E-mail address: yhongdong@163.com (Y. Dong).
conservation given the absence of focal species (Fu et al., 2010;
Li et al., 2010).
Landscape connectivity is a major concern for the maintenance
of wildlife populations and is considered important for ecological
processes such as the movements of genes, individuals, species,
and populations over multiple scales, especially in fragmented
landscapes (Saura et al., 2011). Currently, landscape connectivity
is viewed both structurally (spatial arrangement of the habitat)
and functionally (focal species to the physical structure of the
landscape) (Saura and Torné, 2009). Many indices have been
developed and those based on graph theory (e.g. Probability of
Connectivity) perform well in practical landscape analysis (Bodin
and Saura, 2010; Pascual-Hortal and Saura, 2006). These new
indices with underlying ecological processes overcome the limi-
tations of small-scale empirical indices and large-scale landscape
pattern indices (Fu et al., 2010). However, studies on the ecological
effects caused by road network expansion mainly focus on pattern
analysis (Liu et al., 2011).
China’s road network has been growing rapidly during the past
three decades, in accordance with the government’s reform policy.
Landscape fragmentation and the subsequent effects of roads are
widely studied in many countries, but similar studies have just been
initiated in China (Forman et al., 2002; Li et al., 2010). As China con-
tinues to develop its western provinces, many high-level roads are
being constructed, especially in Yunnan Province where highways
account for 93% of the total transportation routes. In Yunnan, there

1470-160X/$ – see front matter © 2013 Elsevier Ltd. All rights reserved.
http://dx.doi.org/10.1016/j.ecolind.2013.07.018
 S. Liu et al. / Ecological Indicators 36 (2014) 160–168
Fig. 1. Study area of Yunnan Province, China.
are more than 18,000 higher plant species (51.6% of China’s total),
1836 vertebrate species (54.8%), and 15,000 seed plant species
(50%) (Yang et al., 2004). The richness value, endemic value, and
endemic rate in biodiversity all rank first in China, although this
province accounts for only 4.1% of the country’s land area, or eighth
largest, with a total area of 394,000 square kilometers (WWF, 1996).
Research on the ecological effects of road networks will therefore
have significant practical applications in this region.
Although biodiversity includes diversity at the genetic, species,
ecosystem, and landscape levels (Lindenmayer et al., 2000), species
could benefit from increasing connectivity from a landscape-level
perspective. Within this more restricted arena, the indicators of
biodiversity should consist of habitat pattern, habitat quality, and
connectivity. As landscapes are viewed as at least partially consist-
ing of highly interconnected linear features known as networks,
the network properties of connectivity are useful predictors of bio-
diversity (Forman, 1995). In conclusion, the analyses of habitat
fragmentation, heterogeneity, and connectivity at the landscape
level should supply additional important information within the
context of environmental management for biodiversity conserva-
tion.
In this paper, our study was conducted to investigate the degree
of habitat change due to road networks and cities’ expansion in
Yunnan Province. The objectives are as follows: (1) find the rela-
tionship between road network and habitat fragmentation and
(2) assess the landscape connectivity change by sensitivity analy-
sis of ecological processes. Here, the different dispersal/movement
distances are considered broadly representative of ecological pro-
cesses. The effectiveness of different methods from pattern and
process perspectives is also discussed.
2. Materials and methods
2.1. Study area
A 40 km × 40 km square zone (23.7◦ –24.1◦ N, 99.8◦ –100.2◦ E) in
Lincang City in southwest Yunnan Province, located in the middle
161
reach of the Lancang River, was selected as the study site (Fig. 1).
Two features of this zone make it suitable for this study: (1) inten-
sive road networks of different levels total 0.54 km km−2 with city
expansion and (2) biodiversity in this zone has been subjected to
severe disturbances by these roads.
2.2. Data acquisition
The road data was digitized based on the transportation maps
of Yunnan Province in 1990 and 2010 and rectified by 1:250,000
road databases from the National Fundamental Geographical Infor-
mation Center in 2002. According to Liu et al. (2008a), the roads
are divided into four categories: (1) first level (national roads),
(2) second level (mainly county, city-county, city roads), (3)
third level (mainly county, city-town roads), and (4) fourth level
(mainly town-village and village-village roads). Landsat TM images
(#131/043) in 1991 (February 17, 1991) and 2006 (December 11,
2006) were used for landscape mapping after interpretation and
supervised classification. We compared our classification of the
1991 images with land-use status maps (Yunnan land use status
map in 1990), and the classification of the 2006 images were val-
idated by using the ground truth data (53 points, 2009), which
indicated the accuracy level of the land cover map was 83.5%. The
images were classified into seven categories: grassland, farmland,
shrub land, forestland, water, urban land, and rural residential land.
2.3. Analysis methods
2.3.1. Habitat selection
According to local conditions and animals’ behavior, for connec-
tivity analysis, we selected forest land-cover types with a greater
than 30% closed canopy dominated by broadleaf and coniferous
forests as the habitat. The main forest types are Lithocarpus glaber
and Pinus kesiya Royle ex Gordon var. langbianensis (A.Chev) Gaussen.
According to the characteristics of the local environment and refer-
ring to previous studies, the minimum ecological threshold patch
162 S. Liu et al. / Ecological Indicators 36 (2014) 160–168

size was set at 25 ha (Pascual-Hortal and Saura, 2006; Saura et al., Table 1
Resistance values of different layers.
2011).
Layer Classification Buffer Resistance
2.3.2. Habitat change analysis Road First level 500 m 7
For habitat change analysis, the following commonly used pat- Second level 250 m 7
tern indices were examined: number of patches (NP), percentage Third level 100 m 7
Fourth level 25 m 7
of landscape (PLAND), largest patch index (LPI), mean shape index
(SHAPE), mean fractal dimension (FRAC), division index (DIVISION), Land cover Forest (except the source patches) – 2
and total change in habitat area from 1991 to 2006. These indices, Shrub – 2
Grassland – 3
indicating landscape fragmentation, shape, and aggregation infor-
Farmland – 4
mation, were calculated using the Fragstats 3.3 program to quantify Water – 5
the landscape changes (Mcgarigal et al., 2002). Furthermore, to test Urban land – 6
the sensitivity of pattern metrics for habitat loss to altered surface Rural residential land – 6
area and to perimeter, we calculated the selected pattern metrics Slope 0◦ –19.28◦ – 1
after serial removal of each habitat patch from a small to a large 19.29◦ –38.57◦ – 2
size in 1991 and correlated them with habitat area changes. 38.58◦ –57.86◦ – 3
57.87◦ –77.15◦ – 4

2.3.3. Landscape connectivity analysis

In this study, several local species were considered to determine
the dispersal distances for the analysis of landscape connectivity
in 1991 and 2006. The species that mainly inhabit the broadleaf
and coniferous forests include Prebytis phayrei, Lophura nythemera,
Phasianidae, and Phasianus colchicus. According to studies by Yun-
nan University on the dispersal distances of terrestrial mammals
and the investigative reports about animals in Lancang River val-
ley, the four comparative dispersal distances of 4 km, 8 km, 12 km,
and 16 km were defined for connectivity sensitivity analysis (van
Vuren, 1998).
The integral index of connectivity (IIC) selected for landscape
connectivity analysis was developed by Pascual-Hortal and Saura
(2006). It ranges from 0 to 1 and increases with improved connec-
tivity. It is given by:

n 
n
ai aj /(1 + nlij )
IIC =
A2L
i=1 j=1
where ai is the area of habitat patch i, aj is the area of habitat j, AL
is the total landscape area, and nlij is the number of linkages in the
shortest path between patches i and j with the threshold dispersal
distance.
The relative ranking of habitat patches by their contribution to
overall landscape connectivity based on the percentage of impor-
tance (dI) of each individual element is given by the following
expression (Pascual-Hortal and Saura, 2006):
I − I
dI(%) = 100 ×
I neous forest land cover were separated into several small parts.
where I is the IIC index value when the patch is present in the land-
scape and I is the IIC index value after removal of that patch. The
ranking of the dI value is used to indicate the influence of road
networks on the importance of patches.
Improved connectivity analysis with path determination for
animal movement uses effective distances and not just Euclidean
distances. The least-cost modeling method is increasingly used to
measure the effective distance between habitat patches.
In our calculation, we define the habitat patches as the source
layer. We define land cover, road network buffer zone, and slope
gradient as resistance layers. The first-, second-, third-, and
fourth-level roads are buffered at 500 m, 250 m, 100 m, and 25 m,
respectively, depending on the impact degree on the ecosystems
(Li et al., 2004). The slope gradient layer is classified into four levels
with an equal interval of 19.28% according to local conditions (Liu
et al., 2008a,b). The resistance values for classifying each layer are
shown in Table 1 (Marulli and Mallarach, 2005). The overlay of dif-
ferent resistance layers was converted to a raster format with a grid
size of 25 m × 25 m for distance calculations in ArcGIS. Effective
distance in the least-cost modeling method requires two raster-
based data inputs, including the resistance layer, indicating the
travel cost through each cell and the source patch layer to which
the cumulative travel cost for each cell is measured. The output is
a raster cost surface indicating the effective distance from every
cell in the landscape to a source patch (Adriaensen et al., 2003;
Levin et al., 2007).
3. Results
3.1. Habitat changes from 1991 to 2006
3.1.1. Landscape fragmentation
Fig. 2 shows the landscape dynamics from 1991 to 2006. Forest-
land as habitat patches occupied 35.2% and 18.9% of the area in
(1)
1991 and 2006, and shrub land occupied 18.1% and 36.8%, respec-
tively. So 18.3% of the study area was converted from natural habitat
into other land uses as a whole, thus decreasing the area from
525.27 km2 to 226.81 km2 and becoming more fragmented during
this time period. The average habitat patch area decreased from
43.77 km2 to 14.18 km2 . The largest habitat patch reduced in size
from 497.78 km2 to 123.21 km2 .
As shown in Fig. 2, the characteristics of the habitat changed in
line with the land cover dynamics. The forests in the central and
south-east part of the area almost vanished. Approximately 98% of
the habitats lost from 1991 to 2006 were converted into shrub land
that cannot be considered as a habitat. Large patches of homoge-
(2)
However, the bulk of patches maintained the natural and semi-
natural attributes as total area percentage of forest land and shrub
land accounted for more than 53% and changed little from 1991 to
2006.
The changes of different selected landscape metrics are shown
in Fig. 3. In the study area, NP increased from 25 to 29. PLAND and
LPI decreased greatly from 32.8 to 14.2 and from 30.9 to 7.7, respec-
tively, indicating the habitat patches became more fragmented. The
slight decline of SHAPE and FRAC that reflected shape complexity
also indicated that the landscape was composed of less complex
patches. DIVISION is based on the cumulative patch area distribu-
tion and is interpreted as the probability that two randomly chosen
pixels in the landscape are not situated in the same patch of the cor-
responding patch type. The results showed that the value increased
from 0.89 to 0.99, indicating the proportion of the habitat patch type
decreased and the patches decreased in size.
We also tested the sensitivity of pattern metrics for habitat loss
to altered surface area and to perimeters based on the landscape
in 1991. Fig. 4 shows the relationship between pattern metrics
 S. Liu et al. / Ecological Indicators 36 (2014) 160–168
Fig. 2. Forest changes from 1991 to 2006.
Fig. 3. Landscape pattern index values for habitats in 1991 and 2006.
changes with area changes under the scenarios of one habitat
patch removal each time. The results show that NP, PLAND, and LPI
are positively related with area changes, while SHAPE, FRAC, and
DIVISION have large fluctuations. There also exists a high linear cor-
relation coefficient between area and perimeter (R2 = 0.989). These
results indicate that the fragmentation indices performed better for
habitat loss than the shape indices in the study area.
3.1.2. Relationship between habitat loss and influencing factors
With GIS spatial analysis, we identified the habitat loss
distribution. Using GIS zonal statistics analysis, we calculated
163
the average distances from the habitats to the nearest road
and city center and the average elevation of the habitats.
The distances were reclassified into five levels with equal
intervals (Fig. 5).
We calculated that the total area of habitat patches with the
average distances from 361 m to 720 m to the road network (light-
blue area) was about 262 km2 . As shown in Fig. 5, habitat changes
(the red boundary line) mainly occurred in those patches with a
light-blue color, which indicated there is a concentration of the
habitat loss within a 720-m distance zone. Also, the habitats in
the elevation ranging from 2025 m to 2301 m have been reduced
dramatically.
Fig. 6 shows the average values for elevation and distances from
the city and roads of the habitats in 1991 and 2006. The average
elevation of the habitats was 2287 m in 2006, which was a little
higher than in 1991 (2167 m). The average distance from the habitat
to the city in 1991 was 15,336 m, which increased to 18,518 m in
2006. The average distances to roads also increased from 814 m in
1991 to 1077 in 2006.
3.2. Landscape connectivity change
Based on Eq. (1), we calculated the IIC values from 1991 to
2006. The results show that landscape connectivity increased
with the dispersal distances. In 1991, the IIC indices were 0.054,
164 S. Liu et al. / Ecological Indicators 36 (2014) 160–168
Fig. 4. The landscape pattern metrics changed with the habitat area loss.
0.064, 0.0945, and 0.0965 under 4 km, 8 km, 12 km, and 16 km dis-
persal distances, respectively, and decreased in 2006 to 0.0060,
0.0062, 0.0065, and 0.0071, respectively. The landscape con-
nectivity decreased 88.5%, 90.3%, 93.8%, and 92.6% for 4 km,
8 km, 12 km, and 16 km dispersal distances, respectively, for an
average of 91.32%. Obviously, the fragmentation and isolation
of the habitats, especially the habitat loss in the southeastern
part, caused this significant reduction of landscape connectiv-
ity. The sensitivity analysis of serial dispersal distances shows
that the connectivity loss reached a maximum at the 12-km
threshold.
Fig. 7 shows the habitat importance distribution based on the
DI index in Eq. (2) for the two periods under different sensitivity
levels. The importance is classified into five equal interval levels:
low importance, medium low importance, medium importance,
medium-to-high importance and high importance. The distribu-
tion of habitat importance varied greatly in 1991 and 2006 and it
became more fragmented and isolated. The habitat patches, which
were presented as a well-connected network in 1991, were divided
into three components in the northern, western, and eastern parts
in 2006. The eastern component was degraded due to its complete
isolation from the other patches. The patches of high importance
were not well protected from road network disturbance and expe-
rienced a large extent loss of 96.2%.
Fig. 7 shows the potential dispersal paths between different
patches. Habitat patches in 1991 had 20, 18, 10, and 8 least-cost
dispersal paths between habitat patches at respective threshold
distances of 16 km, 12 km, 8 km, and 4 km. In 2006, the number of
dispersal paths decreased to 16 at the 16-km threshold and 12 at
the 12-km threshold. However, they remained the same at the 8-
km and 4-km thresholds. Though the main path loss occurred in the
north, which contributed to patch isolation in that region, there was
no real path between the three main components in 2006. From this
aspect, corridor design for improving connectivity is needed more
between these patches.
4. Discussion
This case study combined landscape pattern and landscape con-
nectivity methods to assess the ecological effects of road network
extension and city expansion. An improved connectivity index (IIC)
with its depiction of important habitats was used and different
dispersal/movement distances broadly representative of selected
ecological processes were compared for sensitivity analysis.
4.1.1. Landscape pattern analysis with GIS methods
Landscape patterns can be directly estimated by various land-
scape pattern metrics. These metrics are measures for quantifying
the composition and configuration of ecosystems across a region
(Mcgarigal et al., 2002). For assessing the potential effects of road
construction or city expansion, a comparison of landscape pattern
metrics is a basic way for describing the landscape changes due to
all disturbances. However, pattern analysis alone may not be suffi-
cient as landscape changes due to disturbances are limited within
a specific zone known as the disturbance zone (Liu et al., 2008a,b).
Moreover, it is difficult to discern the exact disturbance distances of
every pattern index or ecological factor. Usually, GIS methods such
as buffer analysis and spatial statistics are used for the analyses.
In our case, we compared the changes in the landscape fragmen-
tation indices, landscape shape, and aggregation indices. We then
used spatial analysis to identify the relationship between habitat
loss and the influencing factors. Our results show that most of the
selected indices changed greatly. More specifically, zonal statis-
tics show that the elevation of habitat increased about 5.5% from
1991 to 2006, while the distances of habitats to the city and roads
 S. Liu et al. / Ecological Indicators 36 (2014) 160–168
Fig. 5. Habitats in areas with elevation and different distances to roads. (For interpretation of the references to color in text, the reader is referred to the web version of the
article.)
increased 20.7% and 32.3%, respectively (Fig. 6). Obviously, those
patches near the road network and city were more affected by the
landscape fragmentation. Previous studies show the roads as the
driving forces in defining landscape patterns and the developments
and changes in land use near road corridors, often causing further
fragmentation (Hawbaker et al., 2005). These results clearly eluci-
date the effects of road construction and city expansion on habitat
losses.
4.2. Advantage of landscape connectivity method
A substantial shortcoming is that pattern metrics have not been
explicitly linked with ecological processes as indirect ecological
effects. Previously, the road effect on landscape pattern and the
regional ecological risk in Yunnan Province were studied, but with-
out the advent of ecological processes (Liu et al., 2008a,b; Wen et al.,
2007).
165
Roads could be considered as the sources of fragmentation,
barriers for most of the species related, and causes of landscape
function degradation (Fu et al., 2010). For the ecological process,
especially species movement, landscape connectivity has attracted
much attention with the natural habitats being lost and fragmented
due to urban expansion, transport development, and intensive agri-
culture (Foltête et al., 2012). In a review of landscape connectivity
metrics, the more popular CS22 program can quantify the impor-
tance of habitat patches in terms of connectivity (Saura et al., 2011).
Owing to the combination with the preferred IIC index of func-
tional connectivity, the habitat of high conservation importance
could provide references for the improvement of management
practices. Though scale variation will affect the precise location
and orientation of linear elements of effective distances, sensitiv-
ity analysis can be compared in a quantitative way at the landscape
level as well as for individual grid cells (Foltête and Giraudoux,
2012).
166 S. Liu et al. / Ecological Indicators 36 (2014) 160–168
Fig. 6. Average values of the habitat distribution (elevation, distances from city and
roads) in 1991 and 2006.
4.3. Comparison of landscape pattern and landscape connectivity
analysis
In our study, it is obvious that the landscape metrics vary greatly
for different index groups. For the change ratios, landscape frag-
mentation indices (NP, PLAND, and LPI) have larger values than
landscape shape indices (SHAPE, FRAC) and landscape aggrega-
tion index (DIVISION). The rates of change of NP, PLAND, and LPI
indices were 16.0%, −46.8% and −72.8%, while those for the SHAPE,
FRAC, and DIVISION indices were −11.1%, −0.67%, and 10.6%. But for
the IIC index, the values for different dispersal distances decreased
more significantly than pattern indices from 88.5% to 93.8%, which
means the potential effects on species movement may be more
severe than the direct effects on the pattern. Furthermore, the
dI values of each habitat under different movement abilities can
be shown explicitly on a map and quantitatively classified into
Fig. 7. Landscape connectivity in 1991 and 2006 for different dispersal sets.
different importance levels, which is helpful for biodiversity con-
servation. Until now, few studies have incorporated a series of
species’ movements in the quantification of road effects. The effec-
tiveness of pattern and connectivity analysis implies the direct and
the potential effects of road construction and city expansion.
4.4. Landscape connectivity and species consideration
Empirical proofs frequently find that habitat loss and fragmen-
tation result, and will continue to result, in the loss of biodiversity
both in terms of species and genetic variation (Bailey, 2007). During
the habitat change, landscape connectivity is a significant consid-
eration for species’ population survival, which can be interpreted
as the degree to which the landscape impedes the dispersal move-
ment of populations across habitat patches (Taylor et al., 1993).
In particular, integrating landscape connectivity into biodiversity
conservation can identify the patches that should be given extra
protection (Wu et al., 2010).
Estimations of the dispersal behavior and movement abilities
of the focal species are necessary to measure functional connectiv-
ity. As field observations of animal species may take a long time
and often provide a meaningless and insufficient overview of these
connections, a collective library of empirical studies in which eco-
logical responses are related to particular landscape patterns was
difficult to build. Therefore, sensitivity analysis used for compari-
son could complement pattern analysis and provide a quantitative
description for the impact on species’ movements.
Until now, the studies of behaviors of plants and animals in Yun-
nan Province are still limited. In the published literature (Table 2),
the dispersal abilities below or near 1 km are reported more often,
and so the long-dispersal abilities need to be investigated more
thoroughly, not just qualitatively described (Gan and Hu, 2008).
However, quantitatively predicting and evaluating the effects of
human disturbance on the flow of species among habitats is still a
challenge for biodiversity protection. In our studies, although fea-
tures of the species can be identified, species type and dynamic
changes were more complicated to predict in more detail.
 S. Liu et al. / Ecological Indicators 36 (2014) 160–168
Table 2
Dispersal abilities of different species in Yunnan Province as reported in research studies.
Type Name
Plant Himalayan Mulberry (Morus macroura)
Plant Wild banana (Musa acuminata)
Plant Chinese tulip tree (Liriodendron chinense)
Animal Tiger (Pantheratigris)
Animal Black-crested Gibbon (Hylobates concolor furvogaster)
5. Conclusions
In conclusion, this study analyzes the landscape changes of habi-
tat due to fragmentation and connectivity loss under the influence
of road networks and city expansion using two types of indicators.
We found a positive relationship between city and road exten-
sion, habitat fragmentation, and connectivity loss. Spatial analysis
showed that distances from habitat to the city and roads increased
greatly from 1991 to 2006, which indicated that the city and road
extensions would be the driving forces for habitat loss. The results
also verified that connectivity loss would be severer due to road
construction and city expansion than landscape pattern change,
which means the potential effects of road networks on species may
be severer than the direct effect at the regional scale. Landscape
connectivity as large-scale assessment relevant to key species char-
acteristics can be further integrated with other detailed small-scale
investigations. Also, the habitats of high importance derived by IIC
analysis should be given more conservation priority.
In the case study area, it is necessary to establish networks
of habitats connected by protected corridors and buffer zones to
maintain the critical elements of the landscapes. Remnant forest
patches in the southern and western parts should be under protec-
tion and connected with each other to sustain their connectivity.
As the “Grain for Green Project” in China provided opportunities
to conserve and restore the vegetation, natural forests are urgently
needed to restore this region.
Acknowledgments
The paper was financially supported by Nonprofit Environment
Protection Specific Project of China (No. 201209029-4) and the
National Natural Sciences Foundation of China (No. 50939001).
References
Adriaensen, F., Chardon, J.P., DeBlust, G., Swinnen, E., Villalba, S., Gulinck, H., Matthy-
sen, E., 2003. The application of ‘least-cost’ modelling as a functional landscape
model. Landsc. Urban Plan. 64, 233–247.
Bailey, S., 2007. Increasing connectivity in fragmented landscapes: an investiga-
tion of evidence for biodiversity gain in woodlands. Forest Ecol. Manage. 238,
7–23.
Bodin, Ö., Saura, S., 2010. Ranking individual habitat patches as connectivity
providers: integrating network analysis and patch removal experiments. Ecol.
Model. 221, 2393–2405.
Brooks, T.M., Mittermeier, R.A., da Fonseca, G.A.B., Gerlach, J., Hoffmann, M.,
Lamoreux, J.F., Mittermeier, C.G., Pilgrim, J.D., Rodrigues, S.L., 2006. Global bio-
diversity conservation priorities. Science 313, 58–61.
Eigenbrod, F., Hecnar, S.J., Fahrig, L., 2007. Accessible habitat: an improved measure
of the effects of habitat loss and roads on wildlife populations. Landsc. Ecol. 23,
159–168.
Foltête, J.C., Clauzel, C., Vuidel, G., 2012. A software tool dedicated to the modelling
of landscape networks. Environ. Modell. Softw. 38, 316–327.
Foltête, J.-C., Giraudoux, P., 2012. A graph-based approach to investigating the
influence of the landscape on population spread processes. Ecol. Indic. 18,
684–692.
Forman, R.T.T., 1995. Land Mosaics: The Ecology of Landscape and Regions.
Cambridge University Press, Cambridge.
Forman, R.T., Reineking, B., Hersperger, A.M., 2002. Road traffic and nearby grassland
bird patterns in a suburbanizing landscape. Environ. Manage. 29, 782–800.
167
Dispersal distance (km) Source
0.5 Tang et al. (2008)
0.2 Tang et al. (2007)
0.1 Huang et al. (1998)
10 (female) 33 (male) Seidensticker and Lumpkin (1991)
1.3 Sheeran et al. (1998)
Fu, W., Liu, S., Degloria, S.D., Dong, S., Beazley, R., 2010. Characterizing the
“fragmentation–barrier” effect of road networks on landscape connectivity:
a case study in Xishuangbanna, Southwest China. Landsc. Urban Plan. 95,
122–129.
Gan, H.X., Hu, H.B., 2008. Biodiversity conservation corridor design based on habitat
selection of gaur (Bos Gaurus): a case study from Xishuangbanna, China. Chin. J.
Ecol. 27, 2153–2158 (Chinese article with English abstract).
Geneletti, D., 2003. Biodiversity impact assessment of roads: an approach based on
ecosystem rarity. Environ. Impact Assess. Rev. 23, 343–365.
Gibbs, J.P., Steen, D.A., 2005. Trends in sex ratios of turtles in the United States:
Implications of road mortality. Conserv. Biol. 19, 552–556.
Hawbaker, T.J., Radeloff, V.C., Hammer, R.B., Clayton, M.K., 2005. Road density and
landscape pattern in relation to housing density, and ownership, Land Cover,
and Soils. Landsc. Ecol. 20, 609–625.
Huang, S.Q., Guo, Y.H., Chen, J.K., Wang, Q.F., 1998. Primary measurement of
pollen flow in Liriodendron chinense using RAPD markers. Chin. Sci. Bull. 43,
1819–1822.
Laurance, S.G.W., Stouffer, P.C., Laurance, W.F., 2004. Effects of road clearing on
movement patterns of understory rainforest birds in central Amazonia. Conserv.
Biol. 18, 1099–1109.
Levin, N., Lahav, H., Ramon, U., Heller, A., Nizry, G., Tsoar, A., Sagi, Y., 2007. Landscape
continuity analysis: a new approach to conservation planning in Israel. Landsc.
Urban Plan. 79, 53–64.
Li, S.C., Xu, Y.Q., Zhou, Q.F., Wang, L., 2004. Statistical analysis on the relationship
between road network and ecosystem fragmentation in China. Prog. Geogr. 23,
78–85 (in Chinese with English abstract).
Li, T., Shilling, F., Thorne, J., Li, F., Schott, H., Boynton, R., Berry, A.M., 2010. Frag-
mentation of China’s landscape by roads and urban areas. Landsc. Ecol. 25,
839–853.
Lindenmayer, D.B., Margules, C.R., Botkin, D.B., 2000. Indicators of biodiver-
sity for ecological sustainable forest management. Conserv. Biol. 14, 941–
950.
Liu, S.L., Cui, B.S., Dong, S.K., Yang, Z.F., Yang, M., Holt, K., 2008a. Evaluating
the influence of road networks on landscape and regional ecological risk
– a case study in Lancang River Valley of Southwest China. Ecol. Eng. 34,
91–99.
Liu, S., Deng, L., Zhao, Q., DeGloria, S.D., Dong, S., 2011. Effects of road network
on vegetation pattern in Xishuangbanna, Yunnan Province, Southwest China.
Transport. Res. Part D: Transport Environ. 16, 591–594.
Liu, S., Wen, M., Cui, B., Fu, W., Yang, M., 2008b. Definition and spatial differentia-
tion of road-effect zone: a case study in Longitudinal Range-Gorge Region. Prog.
Geogr. 27, 122–128.
Marulli, J., Mallarach, J.M., 2005. A GIS methodology for assessing ecological connec-
tivity: application to the Barcelona Metropolitan Area. Landsc. Urban Plan. 71,
243–262.
Mcgarigal, K., Cushman, S.A., Neel, M.C., Ene, E., 2002. Fragstats: Spatial Pattern
Analysis Software for Categorical Maps. University of Massachusetts, Amherst.
Pascual-Hortal, L., Saura, S., 2006. Comparison and development of new graph-based
landscape connectivity indices: towards the priorization of habitat patches and
corridors for conservation. Landsc. Ecol. 21, 959–967.
Saura, S., Estreguil, C., Mouton, C., Rodríguez-Freire, M., 2011. Network analy-
sis to assess landscape connectivity trends: application to European forests
(1990–2000). Ecol. Indic. 11, 407–416.
Saura, S., Torné, J., 2009. Conefor Sensinode 2.2: a software package for quantifying
the importance of habitat patches for landscape connectivity. Environ. Model.
Softw. 24, 135–139.
Seidensticker, J., Lumpkin, S., 1991. The Great Cats. Rodale Press, Emmaus.
Sheeran, L.K., Zhang, Y., Poirier, F.E., Yang, D., 1998. Preliminary report on the behav-
ior of the Jingdong black gibbon (Hylobates concolor furvogaster). Trop. Biodiv. 5,
113–125.
Tang, Z.H., Cao, M., Sheng, L.X., Ma, X.F., Walsh, A., Zhang, S.Y., 2008. Seed dispersal
of Morus macroura (Moraceae) by two frugivorous Bats in Xishuangbanna, SW
China. Biotropica 40, 127–131.
Tang, Z.H., Sheng, L.X., Ma, X.F., Cao, M., Parsons, S., Ma, J., Zhang, S.Y., 2007. Temporal
and spatial patterns of seed dispersal of Musa acuminata by Cynopterus sphinx.
Acta Chiropterol. 9, 229–235.
Taylor, P.D., Fahrig, L., Henein, K., Merriam, G., 1993. Connectivity is a vital element
of landscape structure. Oikos 68, 571–573.
Trakhtenbrot, A., Nathan, R., Perry, G., Richardson, D.M., 2005. The importance
of long-distance dispersal in biodiversity conservation. Divers. Distrib. 11,
173–181.
168 S. Liu et al. / Ecological Indicators 36 (2014) 160–168

van Vuren, D., 1998. Mammalian dispersal and reserve design. In: Caro, T (Ed.),
Behavioral Ecology and Conservation Biology. Oxford University Press, New
York.
Wen, M.X., Liu, S.L., Cui, B.S., 2007. Research on spatio-temporal change of ecological
capacity and driving forces in LRGR. Chin. Sci. Bull. 52, 1–8.
World Wildlife Fund, 1996. China Biodiversity Conservation Summaries. Chinese
Forestry Press, Beijing.
Wu, C.G., Zhou, Z.X., Wang, P.C., Xiao, W.F., Teng, M.J., 2010. The concept and mea-
surement of landscape connectivity and its applications. Acta Ecol. Sin. 30,
1903–1910.
Yang, Y.M., Tian, K., Hao, J.M., Pei, S.J., Yang, Y.X., 2004. Biodiversity and biodiversity
conservation in Yunnan, China. Biodiv. Conserv. 13, 813–826.