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GIANT PANDA HABITAT SELECTION IN FOPING NATURE RESERVE, CHINA

XUEHUA LIU, 1 Department of Environmental Science and Engineering, Tsinghua University, Beijing 100084, China ALBERTUS G. TOXOPEUS, ACE Division, ITC, P.O. Box 6, 7500 AA Enschede, the Netherlands ANDREW K. SKIDMORE, ACE Division, ITC, P.O. Box 6, 7500 AA Enschede, the Netherlands XIAOMING SHAO, College of Biological Sciences, Chinese Agricultural University, Beijing 100094, China GAODI DANG, Huangjiawan Road 12, Foping Nature Reserve, Shaanxi, P. R. China TIEJUN WANG, Huangjiawan Road 12, Foping Nature Reserve, Shaanxi, P. R. China HERBERT H. T. PRINS, Tropical Nature Conservation and Vertebrate Ecology Group, Wageningen University, Bornesteeg 69, 6708 PD Wageningen, the Netherlands

Abstract: Little is known about habitat selection of the giant panda (Ailuropoda melanoleuca), especially about the relationship between giant panda presence and bamboo and tree structures. We presented data on giant panda habitat use and selection in Foping Nature Reserve (NR), China. We used 1,066 radiotracking locations for 6 col- lared individuals to analyze giant panda habitat selection, and we used 110 plots to reveal the structure of giant panda habitat and its relationship with giant panda presence. We found that (1) giant pandas in Foping NR select- ed mostly 3 habitats: conifer forest, deciduous broadleaf forest, and Fargesia bamboo groves. (2) In winter, giant pandas selected deciduous broadleaf forest within elevations of 1,600 to 1,800 m with a south-facing slope of 10 to 20 degrees. In summer, giant pandas selected conifer forest within elevations of 2,400 to 2,600 m and a slope of 20 to 30 degrees. (3) Giant pandas selected the Bashaina fargesii bamboo area with short and dense culms in winter, while they selected the Fargesia qinlingensis bamboo area with a high coverage of tall and thick culms in summer. We concluded that giant pandas in Foping NR do select their preferred habitats. These findings may be used to guide the human activities in the reserve with consideration of giant panda habitat conditions.

JOURNAL OF WILDLIFE MANAGEMENT 69(4):1623–1632; 2005

Key words: Ailuropoda melanoleuca, China, Foping Nature Reserve, giant panda, habitat selection, habitat use, giant panda presence, giant panda absence.

Habitat is any spatial unit that can be occupied by an individual animal, no matter how briefly (Baker 1978). Analysis of habitat selection has been a common and important aspect of wildlife science (Alldredge and Ratti 1986). Habitat re- quirements of species initially were based on qual- itative descriptions relating the presence or ab- sence of species to the general type or structure of vegetation. In recent years, however, there has been a growing interest in quantitative techniques to gain insights into habitat-selection patterns of animals (Capen 1981). The ecological research of- ten involves comparison of the use of habitat types or food items with the availability of those re- sources to the animal (Johnson 1980). Scham- berger and O’Neil (1986) emphasized that habi- tat-use data were useful to document the species’ use of particular areas within its range based on 2 assumptions: (1) a species will select and use areas that are best able to satisfy its life requirements; and (2) as a result, greater use will occur in higher- quality habitat. Habitat use and habitat selection are described and used differently. According to White and Gar-

1 E-mail: xuehua-hjx@tsinghua.edu.cn

rott (1990), habitat use means that locations taken for each animal are classified as to the habitats in which they occur, thus the percentage of time each animal spends in a particular habitat can be estimated. If 1 habitat is preferred, then more time will be spent in this habitat than expected by chance alone. The term habitat selection has been widely used (Alldredge and Ratti 1986; Reid and Hu 1991; Augustine et al. 1995; Wei et al. 1996, 1999; Babaasa 2000). Johnson (1980) defined habitat selection as process in which an animal ac- tually chooses the habitats, and use of habitats is said to be selective if habitats are used dispropor- tionately to their availability. We followed John- son’s terminology and consider that habitat selec- tion mainly emphasizes the animals’ action of choosing the habitats and that habitat selection can be reflected by analyzing habitat use. Svard- son (1949) and Hiden (1965) suggested that habi- tat selection involves 2 processes: primary selec- tion of general environmental features under the different habitats and then further selection of specific habitat based on detailed features. Ac- cording to Johnson (1980), animals follow an or- der in habitat selection: firstly, selection of geo- graphical region, secondly, selection of a home range in the geographical region, and lastly, se-

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IN FOPING • Liu et al. J. Wildl. Manage. 69(4):2005 Fig. 1 Foping Nature Reserve in

Fig. 1 Foping Nature Reserve in China and distributions of radiotelemetry locations (1991–1995) and 110 field survey plots.

lection of habitats within their home range. Wiens (1981) indicated that habitat selection may occur at different spatial scales and need not be based on the same criteria at each scale. Schaller et al. (1985) indicated that little is known about habitat selection of giant pandas, except that they seem to occur in mountainous areas, live in ar- eas with abundant bamboo, and feed almost exclu- sively on bamboo species. However, there is a sub- stantial variation in the growth pattern (e.g., culm density, annual shoot production) and morphol- ogy (e.g., culm height, basal diameter) of bamboo culms under different environmental conditions, and that may affect giant panda habitat selection (Reid and Hu 1991). Wei et al. (1996) reported that some research was done on giant panda habitat se- lection by Reid and Hu (1991) in Wolong NR. Based on that research, Wei et al. (1996, 1999) applied the same methods to analyze giant panda habitat selec- tion in Mabian Dafengding NR and compared the habitat selection between giant pandas and red pandas (Ailurus fulgens styani) in Yele NR. Two main habitats are used by giant pandas in Foping NR: winter habitat with bamboo species Bashaina fargesii and summer habitat with bamboo species Fargesia qinlingensis (Pan et al. 1988, Yong et al. 1994, Liu 2001). Tian (1986, 1989, 1990,

1991) researched characteristics of bamboo spe- cies, including flowering, within the entire range of the Qinling Mountains. However, nothing has been published about the relationship between the bamboo, the tree canopy, and giant panda presence. Tian (1990) reported that giant pandas do not feed in areas where bamboo stems are very dense or very sparse, where bamboo closure is very high or very low, where the understory environ- ment is dark, and where the slopes are steep. The field survey conducted in Foping NR in 1984 showed that 20% of the bamboo area received lit- tle or no use by giant pandas (Tian 1990). Tian (1990) also observed that giant pandas selected only 1- to 2-year-old bamboo stems for forage. These findings were based only on field observa- tions, and no statistical analysis was performed to test for habitat selection in Foping. Research using advanced methods to analyze gi- ant panda habitat and habitat use during summer and winter has recently started in Foping NR. Yang et al. (1997, 1998a, 1998b) conducted statis- tical analyses on summer and winter habitats of gi- ant pandas in Foping NR with focus on biophysi- cal and abiotic factors affecting habitat use. Ren et al. (1998) studied the relationship between plant species richness and elevation. However, a

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quantitative analysis of giant panda habitat selec- tion, particularly regarding use of bamboo and vegetative structure, has not been conducted in Foping NR. Moreover, no radiotelemetry data have been used to determine giant panda habitat selection. We examined the relationship between giant panda presence and habitat factors such as land- cover type, elevation, slope gradient, and slope as- pect by using radiotelemetry data (Liu et al. 2002), survey plot data, and mapping results from Liu (2001). Our goals were to gain insights into habitat selection of giant pandas, to examine differences between giant panda habitats, and to determine the specific characteristics of giant panda habitat. Our objective was to provide managers, researchers, and local people with detailed information about giant panda habitat selection so that conflicts between conservation and development can be avoided.

STUDY AREA

Foping NR was located in the southern portion of Shaanxi province (Fig. 1), on the central south- ern slopes of the Qinling Mountains (33°32′33°45′N, 107°40′107°55′E). The reserve covered approximately 290 km 2 , with elevations ranging from approximately 980 to 2,900 m. Four main wa- tersheds cover the nature reserve, draining into the XiHe, DongHe, JinShuiHe and LongTanZi Rivers from the west to east. Average annual rain- fall is about 920 mm. The lowest average tempera- ture, about –C, occurs in January, and the high- est temperature, about 28°C, occurs in July. The main vegetation types were conifer forests, mixed conifer-broadleaf forests, deciduous broadleaf forests, shrub, and meadow (Ren et al. 1998, CVCC 1980). There were 2 main bamboo species for gi- ant panda forage: B. fargesii and F. qinlingensis (Pan et al. 1988; Tian 1989, 1990; Yong et al. 1994; Ren et al. 1998). B. fargesii was generally distributed in areas below 1,900 m, whereas F. qinlingensis occured mainly above 1,900 m. According to surveys con- ducted in 1990 (Yong et al. 1993), the giant panda population was about 64 pandas with an average density of 1 individual per 5 km 2 . About 300 peo- ple reside inside the nature reserve (Liu 2001).

METHODS

Data

We assumed radiotracking data was able to re- flect the principles of giant panda habitat selec- tion. The 6 giant pandas (3 F and 3 M) tracked

with telemetry collars in different periods from May 1991 through December 1995 (Liu et al. 2002) and the 59 telemetry towers were located for telemetry monitoring. The towers for tracking during summer and fall were distributed along the top of GuangTouShan Mountain in a west–east di- rection along the north boundary, and the towers for tracking during winter and spring were dis- tributed in the DongHe River Valley in a south–north direction. We checked all 1,760 telemetry locations for accuracy and kept 1,639 locations (Liu et al. 2002; Fig. 1). We used 1 telemetry location per day to calculate giant panda habitat use and selection. We collected habitat data based on a field survey conducted during July–August 1999. We used Global Positioning System (GPS) to record the lo- cations of all survey plots. We adopted the line transect sampling method to get as many habitat types within the shortest route as possible. We sur- veyed 110 plots 10 m × 10 m; Fig. 1), each of them containing 4 bamboo plots (1 m × 1 m), to calcu- late bamboo parameters. Four bamboo plots were arranged in the centers of 4 small rectangles within the 10 m × 10 m plot. We collected the fol- lowing detailed information in each plot: (1) land cover types (i.e., conifer forest, mixed conifer- broadleaf forest, deciduous broadleaf forest, bam- boo, shrub-grass-herb area, rock or bareland, farmland and settlements, and water); (2) tree layer (≥5 m) (i.e., species, number of stems, di- ameter at breast height [DBH], height, and canopy coverage per species, total canopy cover- age in 10 m × 10 m plot); (3) bamboo layer (i.e., species, number of culms, basal diameter [BD] per culm, average bamboo height in bamboo plot, and total bamboo coverage in bamboo plot); (4) terrain (i.e., elevation, slope gradients, aspects); and (5) panda signs (i.e., feeding signs, drop- pings, nesting signs).

Panda Habitat Map

We defined giant panda habitats based on land cover types, and we used these habitats to deter- mine giant panda habitat selection. The giant panda habitat map created from using an inte- grated neural network and expert system (Liu 2001) was used as the base in this study. The ground-truthing data consisted of 160 points with records of land cover types from field surveys in 1999, including 110 plots mentioned previously with detailed habitat information and 50 survey points with only information of land cover types (such as water, farmland and settlements, rock

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Table 1. Frequency of use (locations, 1 location per day) of habitat types by giant pandas in Foping Nature Reserve, China,

1991–1995.

 

Tracking

Tracking locations in different habitats

 

Total

Panda

Sex

period

cf a

dbfcf

dbf

bam

shgr

fas

rab

wat

locations

045

F

June 92–May 93

15

57

77

0

1

0

0

0

150

065

M

June 92–May 93

23

62

78

1

0

0

0

0

164

127

M

June 92–May 93

45

61

76

12

0

0

0

0

194

045

F

Jan. 95–Dec. 95

19

91

94

3

0

0

0

0

207

065

M

Jan. 95–Dec. 95

15

102

67

1

0

0

0

0

185

005

M

Jan. 95–Dec. 95

17

57

82

2

5

0

3

0

166

Total locations % of time in

134

430

474

19

6030

1066

a year

12.6 b

40.3

44.5

1.8

0.6

0.0

0.3

0.0

100

a cf: conifer forest, dbfcf: mixed conifer–broadleaf forest, dbf: deciduous broadleaf forest, bam: bamboo (or mixed with meadow), shgr: shrub–grass–herb area, fas: farmland and settlements, rab: rock and bareland, and wat: water area.

b 12.6 = 134/1066*100.

and bareland). Stratified random sampling method was used to map the giant panda habitats, through which 2 independent sample sets were created from 160 points. One sample set was used to classify the Landsat Thematic Mapper (TM) im- ages (acquired in July 1997) and another set to test the created habitat map. To overcome the impacts on mapping from complex terrain and various ground objects, 9 GIS data layers, containing re- mote sensing data (TM band 1 to 5 and 7) and ter- rain data (elevation, slope, and aspect) were used in mapping. The habitat map with an overall map- ping accuracy of 84%, together with terrain data, was used to extract the habitat information for all telemetry locations.

DATA ANALYSIS

We used radiotelemetry data from 4 giant pandas monitored for at least 1 year (total 6 years; 1,066 lo- cations; Table 1) to estimate the proportional use of each habitat type, by overlaying the locations onto the digital habitat map (Liu 2001). We as- sumed that all giant pandas had equal access to the different habitats and that all habitats in the nature reserve were available to the tracked giant pandas. We partitioned our data to determine the fre- quency of giant panda occurrence by habitats in 2 seasonal activity ranges to understand how giant pandas utilised habitat in winter and summer. We also used other physical environmental factors (i.e., elevation, slope gradient, slope aspect) to evaluate the effect of terrain factors on giant panda habitat use. We performed a χ 2 test to test for the goodness- of-fit of utilized habitats to available habitats to gain an insight into giant panda habitat selection (Neu et al. 1974, Byers et al. 1984, White and Gar-

rot 1990). We tested 2 null hypotheses: (1) habitat use occured in proportion to habitat availability, considering all habitats simultaneously (Equation 1); and (2) habitat use occured in proportion to habitat availability considering each habitat sepa- rately (Equation 2):

χ 2 = [(observed frequency – expected frequency) 2 /

expected frequency]

(1)

p oi Z a/2k [ p oi (1 p oi ) /n] 1/2

[p oi (1

p oi ) /n] 1/2

P i p oi + Z a/2k

(2)

where P i was the calculated confidence interval for habitat type i, p oi was the proportion of giant panda observations in habitat type i, n was the to- tal number of observations, a was a level of signif- icance, Z a/2k was the upper standard normal table value corresponding to a probability of a/2k and represented the Bonferroni adjustment to main- tain an experimentwise error rate of α, and k was the number of habitats. We used Equation 1 to test whether a significant difference existed between observed and expected frequencies of giant panda location within the habitats. If there was a significant difference, the first null hypothesis was rejected indicating habi- tat selection. We subsequently used Equation 2 to calculate confidence intervals and to test which habitats were selected. If the confidence interval included the available proportion for a habitat type, the second null hypothesis cannot be re- jected. However, a lower interval boundary that exceeded the availability proportion would indi- cate selection of this habitat type. In contrast, if the upper interval boundary was less than the availability proportion, use of the habitats by giant

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pandas would be less than expected, indicating avoidance. The availability (i.e., area) of each habitat type was based on the digital habitat map (Liu 2001). The importance of habitat structure compo- nents to giant pandas (trees and bamboos) was de- termined by comparing all locations with or with- out signs of giant panda presence. Based on Tian’s (1990) field observations and our own field sur- veys, we tested whether differences existed be- tween giant panda presence and absence locations to the following parameters: total tree canopy cov- erage, total bamboo coverage, number of tree stems, number of bamboo culms, height of tree and bamboo species, DBH of tree stems, and BD of bamboo culms. We used the Mann-Whitney U test for all comparisons.

RESULTS Habitat Use and Selection

We analysed giant panda habitat use data of 1- year periods for 4 giant pandas; 2 giant pandas were tracked for 2 years, resulting in a sample size of 6 years (Table 1). The average percentage of lo- cations in each habitat type was 13% for conifer forests, 40% for mixed conifer–broadleaf forests, 45% for deciduous broadleaf forests, 2% for bam- boo groves, and <1% for shrub–grass–herb areas and rock and bareland. Giant pandas were not lo- cated in farmland and settlements or water. Seasonal use of land-cover-based habitats and 3 topographic factors varied (Fig. 2). In winter, gi- ant pandas stayed mostly in deciduous broadleaf forests and mixed conifer–broadleaf forests within an elevation of 1,600 to 1,800 m and a south-fac- ing slope of 11 to 20 degrees. In summer, giant pandas often used conifer forests and mixed conifer-broadleaf forests within elevations from 2,400 to 2,600 m and northern and western slopes from 21 to 30 degrees. About 9% of the tracking locations collected during winter were in the no aspect class, compared to only 1% during summer; this suggests that giant pandas used flat areas more frequently during winter. The χ 2 goodness-of-fit analyses (Table 2A) showed significant differences between overall habitat availability and habitat use (χ 2 = 259, df = 7, P < 0.001). Thus, giant pandas showed habitat selection when considering all habitats. Three habitats were frequently selected by giant pandas:

conifer forest, deciduous broadleaf forest, and bamboo groves (Table 2B). Giant pandas used rock and bareland areas in proportion to its avail-

ability. However, the remaining 4 habitats (i.e., mixed conifer–broadleaf forest, shrub–grass–herb area, farmland and settlements, and water) were used less than expected based on availability.

Panda Presence and Habitat Structure

When bamboo species (B. fargesii or F. qinlingen- sis) occurred under a tree canopy, we observed no differences among the tree structure parameters between areas where giant panda signs were pre- sent or absent (all P > 0.05; Fig. 3). In B. fargesii groves, we found no differences for total bamboo coverage and average BD of bamboo culms between areas with and without giant panda sign (P = 0.798 and 0.186, respectively; Fig. 4). How- ever, giant pandas selected short and dense bamboo groves (P = 0.004 and 0.001, respectively; Fig. 4). In F. qinlingensis groves, only the density of bamboo culms was similar between areas where giant panda sign was present or absent (P = 0.221; Fig. 4), but areas with giant panda sign had greater bamboo coverage and taller and thicker bamboo culms (P = 0.037, 0.004, and < 0.001, respectively; Fig. 4).

DISCUSSIONS

Panda habitat selection in Foping NR seems to vary seasonally, which likely corresponds to food availability. The overstory vegetation types and un- derstory bamboo species have a vertical distribu- tion along the elevation gradient (Liu 2001). Analysis of radiotracking data showed that areas below 1,950 m were winter habitat, and areas above 2,160 m were summer habitat (Liu et al. 2002). In the winter range, deciduous broadleaf forests and mixed conifer–broadleaf forests occupy 70% of to- tal NR area with a well-developed understory of B. fargesii, which provides giant pandas with an abun- dant food supply. Summer habitat areas (i.e., 20% of total NR area) consist of conifer forest and mixed conifer-broadleaf forest with F. qinlingensis as the giant pandas’ primary food. Giant pandas spent 45%, 40%, and 13% of their time in decidu- ous broadleaf forest, mixed conifer-broadleaf for- est, and conifer forest, respectively; this indicates that giant panda habitat selection coincides with seasonal changes (Table 1). Pandas did not use the entire elevation range evenly in Foping NR; they mainly stayed in areas between 1,600 and 1,800 m in winter and areas be- tween 2,400 and 2,600 m in summer. Elevation be- tween 1,950 and 2,160 m only have scattered bam- boo groves, and few locations or signs of resident giant pandas have been recorded there (Fig. 2B). This observation is supported by our field surveys

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A B C D
A
B
C
D

Fig. 2. Analysis of giant panda habitat use in 2 seasonal activity ranges in Foping Nature Reserve, China, 1991–1995.A: Use of land-cover-types: conifer forest (cf), mixed conifer-broadleaf forest (dbfcf), deciduous broadleaf forest (dbf), bamboo (or mixed with meadow; bam), shrub-grass-herb area (shgr), rock and bareland (rab), farmland and settlements (fas), and water (wat). B:

Use of elevation. C: Use of slope gradients. D: Use of slope aspects: east (16–135 degrees), south (136–225 degrees), west (226–315 degrees), north (316–360 and 0–45 degrees), and no aspect.

conducted during summer 1999 and by an analy- sis of radiotelemetry data (Liu et al. 2002). This transitional habitat (Liu 2001) was used by giant pandas to move between the 2 seasonal activity ranges in June and September. It has been reported that giant pandas select ar- eas with a gentle slope gradient (Ouyang et al. 1996), and our results support this finding. The frequency of giant panda use of areas with slopes >30 degrees was greater in summer than in winter (27% in summer and 0% in winter), which agreed

with Yang et al. (1998a, 1998b). In Wolong NR, gi- ant pandas selected flat areas or gentle slopes (10 to 20 degrees) during the entire year (Ouyang et al. 1996). One reason for this difference could be that summer habitat in Foping NR is limited and mostly had slopes of 20 to 30 degrees. Our χ 2 analysis showed that giant pandas do not frequently select shrub–grass–herb areas. This land-cover type has no tree and bamboo cover, and therefore it provides no value to giant pandas. However, mixed conifer–broadleaf forest was se-

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Table 2. Chi-square (χ 2 ) analysis of giant panda habitat selection in Foping Nature Reserve, China, 1991–1995.

A: Chi-square analysis (χ 2 ) of all habitat types together.

 

Habitat availability

Location of radiotracking

Expected

χ 2

Habitat type

Area (km 2 )

Proportion :p a a

Observed

Proportion :p oi b

observations c

test

Conifer forest Mixed conifer-broadleaf forest Deciduous broadleaf forest Bamboo (or mixed with meadow) shrub-grass-herb area Farmland and settlements Rock and bareland Water Total

16.5

0.056

134

0.126

60

91.47

174.2

0.594

430

0.403

633

65.06

92.0

0.314

474

0.445

334

58.42

1.7

0.006

19

0.018

6

26.62

6.0

0.020

6

0.006

22

11.45

0.4

0.001

0

0.000

1

1.45

1.6

0.005

3

0.003

6

1.36

1.0

0.003

0

0.000

4

3.63

293.4

1

1,066

1

1,066

259.47

B: Proportion availability and observed use of each habitat type and 95% C.I.

 

Proportion of

Proportion of

C.I. on proportion of occurrence: P i (at 95% experiment wise

Habitat

Habitat type

availability :p a

observations :p oi

confidence coefficient)

selection

Conifer forest

0.056

0.126

0.098

Mixed conifer-broadleaf forest

0.594

0.403

0.362

Deciduous broadleaf forest

0.314

0.445

0.403

Bamboo (or mixed with meadow)

0.006

0.018

0.007

Shrub-grass-herb area

0.020

0.006

–0.001

Farmland and settlements

0.001

0.000

Rock and bare land

0.005

0.003

–0.002

Water

0.003

0.000

P cf

P dbfcf

0.155

0.446

P

P

P

0

P rab

0

dbf

bam

shgr

0.488

0.029

0.012

0.007

Frequent

Less frequent

Frequent

Frequent

Less frequent

Not selected

In proportion

Not selected

a The p

b The p oi is a proportion of observed locations in each habitat type to the total observed locations (e.g., 134/1066 = 0.126).

c Expected locations of animals were calculated by multiplying p a and the total observed locations (e.g., 0.056*1066 = 60).

a

is a proportion of the area of each habitat type to the total area (e.g., 16.5/293.4 = 0.056).

lected less than expected by giant pandas (Table 2B). We suggest that this finding was due because

this habitat type covered a large area of Foping NR

(60%). Giant pandas often occupied this habitat type in summer and winter. Therefore, despite the lack of selection, mixed conifer–broadleaf forest may be important for giant pandas. Johnson (1980) suggested that lack of selection of a com- mon habitat types at 1 scale does not necessarily imply that the habitat type is not important. The abundance of a particular habitat type may have been selected for at a broader scale. Giant pandas avoided areas influenced by humans, such as farm- land and settlements. During the winter season, giant pandas stay in

areas with B. fargesii and frequently select bamboo groves called “ZhuYangZi” by local people, which means that the bamboo culms are short (about 2

m high) and dense with many culms and

branches, caused by multiple feeding events by gi-

ant pandas (Fig. 5A). The structural parameters of

ZhuYangZi bamboo groves are different from those of the normal B. fargesii groves with tall

(about 4 m) and sparse culms without branches

(Fig. 4, Fig. 5B). It may be important to separate these 2 different B. fargesii habitats for conserva- tion purposes. High and mature B. fargesii habitat is often regarded as less important to giant pan- das. However, during our surveys in summer 1999, we found remains of giant panda droppings and piles of the remaining parts of bamboo shoots. Therefore, giant pandas used the high and ma- ture B. fargesii habitats during spring season for foraging on thick bamboo shoots. The shoots con- sumed by giant pandas subsequently disappear from the groves, and giant panda droppings re- sulting from these shoots do not exist for >2 months because of easy decomposition. Thus, lit- tle sign was evident in the high and mature B. far- gesii groves, which, in turn, may lead to the wrong conclusion that giant pandas do not use the high mature B. fargesii groves. F. qinlingensis bamboo in summer habitats has shorter and thinner culms, and they grows more densely. We found that giant pandas selected F. qin- lingensis bamboo groves with higher coverage and taller and thicker culms during summer. Because the biomass of individual F. qinlingensis culms is rel-

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IN FOPING • Liu et al. J. Wildl. Manage. 69(4):2005 Fig. 3. Comparison of tree parameters

Fig. 3. Comparison of tree parameters with bamboo understory between giant panda-presence (white boxplot) and giant panda- absence (grey boxplot) habitats (N = number of plots, NS = no significant difference, P = significance level, DBH = tree diameter at breast height), Foping Nature Reserve, China, Jul–Aug 1999. Statistical outliers (o) and extreme outliers (*) are noted.

atively low compared with B. fargesii, and summer habitat is steep, giant pandas likely select bamboo groves with higher bamboo biomass (i.e., taller culms, thicker basal diameter, and higher coverage due to more culms and leaves) while avoiding steep slopes. Bamboo densities in areas with and without giant panda sign in high-elevation areas were not different. This finding is similar to those in Wolong NR, where giant pandas select B. fangiana in high elevations (Reid and Hu 1991). Most bamboo groves in high-elevation areas grow densely.

MANAGEMENT IMPLICATIONS

The characteristics of tree and bamboo layers in- fluence giant panda habitat selection. Therefore, human activities on habitat, such as new cultiva- tion, mushroom production, road construction, and development could affect giant pandas. Our analysis can provide strong support for re- searchers to map and assess the spatial pattern of

the giant panda habitat, and consequently, allows managers to determine where the best habitat ar- eas are for giant pandas in Foping NR, where po- tentially important habitat areas for giant pandas could be out of the nature reserve, and where the local economical developing and infrastructure constructing should prohibited. As such, man- agers can use this information to reduce future impacts on giant panda habitat areas.

ACKNOWLEDGMENTS

We received support from the National Natural Science Foundation of China (30230080). We are grateful to J. J. Zhao who helped us with field data collection and devoted his entire life to giant panda conservation; unfortunately, he passed away in 2000 during the 3rd national giant panda sur- vey. We thank the State Forest Administration (SFA) of China for allowing us to carry out the field surveys in Foping Nature Reserve. We sin-

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PANDAS’ HABITAT SELECTION IN FOPING • Liu et al. 1631 Fig. 4. Comparison of bamboo parameters

Fig. 4. Comparison of bamboo parameters between giant panda-presence (white boxplot) and giant panda-absence (grey box- plot) habitats, Foping Nature Reserve, China, July-August 1999. N = number of plots, NS = no significant, S = significant differ- ence at 95% C.I., P = significance level, DBH = tree diameter at breast height, 1 = Bashania fargesii, 2 = Fargesia qinlingensis). Statistical outliers (o) and extreme outliers (*) are noted.

A B
A
B

Fig. 5. (A) ZhuYangZi B. fargesii habitat with short (about 2 m) and dense culms with many branches caused by multiple feeding events by giant pandas, and (B) high mature B. fargesii habitat with tall (about 4 m) and sparse culms with no branches, Foping Nature Reserve, China.

1632 GIANT PANDAS’ HABITAT SELECTION IN FOPING • Liu et al.

J. Wildl. Manage. 69(4):2005

cerely thank the directors of the nature reserve (J. Zhang, X. J. Wang, and D. H. Zhao) for their support. Without them, the fieldwork could not have been carried out. We thank L. Kumar and U. B. Nidumolu for reviewing and commenting on an earlier draft of our manuscript.

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Associate Editor: Hall.