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 collared 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 selected 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):16231632; 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 requirements of species initially were based on qualitative descriptions relating the presence or absence 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 often involves comparison of the use of habitat types or food items with the availability of those resources to the animal (Johnson 1980). Schamberger and ONeil (1986) emphasized that habitat-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 higherquality habitat. Habitat use and habitat selection are described and used differently. According to White and Gar1 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 actually chooses the habitats, and use of habitats is said to be selective if habitats are used disproportionately to their availability. We followed Johnsons terminology and consider that habitat selection mainly emphasizes the animals action of choosing the habitats and that habitat selection can be reflected by analyzing habitat use. Svardson (1949) and Hiden (1965) suggested that habitat selection involves 2 processes: primary selection of general environmental features under the different habitats and then further selection of specific habitat based on detailed features. According to Johnson (1980), animals follow an order in habitat selection: firstly, selection of geographical region, secondly, selection of a home range in the geographical region, and lastly, se-

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Fig. 1 Foping Nature Reserve in China and distributions of radiotelemetry locations (19911995) 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 areas with abundant bamboo, and feed almost exclusively on bamboo species. However, there is a substantial variation in the growth pattern (e.g., culm density, annual shoot production) and morphology (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 selection 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 selection 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 species, 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 environment 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 little 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 observations, and no statistical analysis was performed to test for habitat selection in Foping. Research using advanced methods to analyze giant panda habitat and habitat use during summer and winter has recently started in Foping NR. Yang et al. (1997, 1998a, 1998b) conducted statistical analyses on summer and winter habitats of giant pandas in Foping NR with focus on biophysical 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 selection, 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 landcover type, elevation, slope gradient, and slope aspect 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 southern slopes of the Qinling Mountains (3332 3345N, 1074010755E). The reserve covered approximately 290 km2, with elevations ranging from approximately 980 to 2,900 m. Four main watersheds cover the nature reserve, draining into the XiHe, DongHe, JinShuiHe and LongTanZi Rivers from the west to east. Average annual rainfall is about 920 mm. The lowest average temperature, about 3C, occurs in January, and the highest temperature, about 28C, 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 giant 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 conducted in 1990 (Yong et al. 1993), the giant panda population was about 64 pandas with an average density of 1 individual per 5 km2. About 300 people reside inside the nature reserve (Liu 2001).

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 westeast direction along the north boundary, and the towers for tracking during winter and spring were distributed in the DongHe River Valley in a southnorth 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 JulyAugust 1999. We used Global Positioning System (GPS) to record the locations of all survey plots. We adopted the line transect sampling method to get as many habitat types within the shortest route as possible. We surveyed 110 plots 10 m 10 m; Fig. 1), each of them containing 4 bamboo plots (1 m 1 m), to calculate bamboo parameters. Four bamboo plots were arranged in the centers of 4 small rectangles within the 10 m 10 m plot. We collected the following detailed information in each plot: (1) land cover types (i.e., conifer forest, mixed coniferbroadleaf forest, deciduous broadleaf forest, bamboo, shrub-grass-herb area, rock or bareland, farmland and settlements, and water); (2) tree layer (5 m) (i.e., species, number of stems, diameter at breast height [DBH], height, and canopy coverage per species, total canopy coverage 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, droppings, nesting signs).

Panda Habitat Map

We defined giant panda habitats based on land cover types, and we used these habitats to determine giant panda habitat selection. The giant panda habitat map created from using an integrated 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

METHODS Data
We assumed radiotracking data was able to reflect the principles of giant panda habitat selection. The 6 giant pandas (3 F and 3 M) tracked

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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, 19911995. Tracking period June 92May 93 June 92May 93 June 92May 93 Jan. 95Dec. 95 Jan. 95Dec. 95 Jan. 95Dec. 95 Tracking locations in different habitats cf a 15 23 45 19 15 17 134 12.6 b dbfcf 57 62 61 91 102 57 430 40.3 dbf 77 78 76 94 67 82 474 44.5 bam 0 1 12 3 1 2 19 1.8 shgr 1 0 0 0 0 5 6 0.6 fas 0 0 0 0 0 0 0 0.0 rab 0 0 0 0 0 3 3 0.3 wat 0 0 0 0 0 0 0 0.0 Total locations 150 164 194 207 185 166 1066 100

Panda 045 065 127 045 065 005 Total locations % of time in a year

Sex F M M F M M

a cf: conifer forest, dbfcf: mixed coniferbroadleaf forest, dbf: deciduous broadleaf forest, bam: bamboo (or mixed with meadow), shgr: shrubgrassherb 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) images (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 remote sensing data (TM band 1 to 5 and 7) and terrain data (elevation, slope, and aspect) were used in mapping. The habitat map with an overall mapping accuracy of 84%, together with terrain data, was used to extract the habitat information for all telemetry locations.

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 separately (Equation 2): 2 = [(observed frequency expected frequency)2 / expected frequency] (1) poi Za/2k [ poi (1 poi ) /n] 1/2 Pi poi + Za/2k [poi (1 poi ) /n ] 1/2 (2) where Pi was the calculated confidence interval for habitat type i, poi was the proportion of giant panda observations in habitat type i, n was the total number of observations, a was a level of significance, Za/2k was the upper standard normal table value corresponding to a probability of a/2k and represented the Bonferroni adjustment to maintain an experimentwise error rate of , 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 habitat 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 rejected. However, a lower interval boundary that exceeded the availability proportion would indicate selection of this habitat type. In contrast, if the upper interval boundary was less than the availability proportion, use of the habitats by giant

DATA ANALYSIS
We used radiotelemetry data from 4 giant pandas monitored for at least 1 year (total 6 years; 1,066 locations; Table 1) to estimate the proportional use of each habitat type, by overlaying the locations onto the digital habitat map (Liu 2001). We assumed 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 frequency 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 2 test to test for the goodnessof-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-

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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 components to giant pandas (trees and bamboos) was determined by comparing all locations with or without signs of giant panda presence. Based on Tians (1990) field observations and our own field surveys, we tested whether differences existed between giant panda presence and absence locations to the following parameters: total tree canopy coverage, 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.

ability. However, the remaining 4 habitats (i.e., mixed coniferbroadleaf forest, shrubgrassherb 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. qinlingensis) occurred under a tree canopy, we observed no differences among the tree structure parameters between areas where giant panda signs were present 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). However, 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).

RESULTS Habitat Use and Selection

We analysed giant panda habitat use data of 1year 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 locations in each habitat type was 13% for conifer forests, 40% for mixed coniferbroadleaf forests, 45% for deciduous broadleaf forests, 2% for bamboo groves, and <1% for shrubgrassherb areas and rock and bareland. Giant pandas were not located in farmland and settlements or water. Seasonal use of land-cover-based habitats and 3 topographic factors varied (Fig. 2). In winter, giant pandas stayed mostly in deciduous broadleaf forests and mixed coniferbroadleaf forests within an elevation of 1,600 to 1,800 m and a south-facing 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 2 goodness-of-fit analyses (Table 2A) showed significant differences between overall habitat availability and habitat use (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-

DISCUSSIONS
Panda habitat selection in Foping NR seems to vary seasonally, which likely corresponds to food availability. The overstory vegetation types and understory bamboo species have a vertical distribution 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 coniferbroadleaf forests occupy 70% of total NR area with a well-developed understory of B. fargesii, which provides giant pandas with an abundant 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 deciduous broadleaf forest, mixed conifer-broadleaf forest, 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 between 2,400 and 2,600 m in summer. Elevation between 1,950 and 2,160 m only have scattered bamboo 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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Fig. 2. Analysis of giant panda habitat use in 2 seasonal activity ranges in Foping Nature Reserve, China, 19911995.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 (16135 degrees), south (136225 degrees), west (226315 degrees), north (316360 and 045 degrees), and no aspect.

conducted during summer 1999 and by an analysis 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 areas 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, giant 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 2 analysis showed that giant pandas do not frequently select shrubgrassherb areas. This land-cover type has no tree and bamboo cover, and therefore it provides no value to giant pandas. However, mixed coniferbroadleaf forest was se-

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Table 2. Chi-square (2 ) analysis of giant panda habitat selection in Foping Nature Reserve, China, 19911995. A: Chi-square analysis (2 ) of all habitat types together. Habitat availability Habitat type 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 Area (km 2) Proportion 0.056 0.594 0.314 0.006 0.020 0.001 0.005 0.003 1 :paa Location of radiotracking Observed 134 430 474 19 6 0 3 0 1,066 Proportion 0.126 0.403 0.445 0.018 0.006 0.000 0.003 0.000 1 :poib Expected observationsc 60 633 334 6 22 1 6 4 1,066 2 test 91.47 65.06 58.42 26.62 11.45 1.45 1.36 3.63 259.47

16.5 174.2 92.0 1.7 6.0 0.4 1.6 1.0 293.4

B: Proportion availability and observed use of each habitat type and 95% C.I. C.I. on proportion of occurrence: Pi (at 95% experiment wise confidence coefficient) 0.098 0.362 0.403 0.007 0.001 0.002 Pcf 0.155 Pdbfcf 0.446 Pdbf 0.488 Pbam 0.029 Pshgr 0.012 0 Prab 0.007 0

Habitat type Conifer forest Mixed conifer-broadleaf forest Deciduous broadleaf forest Bamboo (or mixed with meadow) Shrub-grass-herb area Farmland and settlements Rock and bare land Water

Proportion of availability :pa 0.056 0.594 0.314 0.006 0.020 0.001 0.005 0.003

Proportion of observations :poi 0.126 0.403 0.445 0.018 0.006 0.000 0.003 0.000

Habitat selection Frequent Less frequent Frequent Frequent Less frequent Not selected In proportion Not selected

a The p is a proportion of the area of each habitat type to the total area (e.g., 16.5/293.4 = 0.056). a b The p is a proportion of observed locations in each habitat type to the total observed locations (e.g., 134/1066 = 0.126). oi c Expected locations of animals were calculated by multiplying p and the total observed locations (e.g., 0.056*1066 = 60). a

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 coniferbroadleaf forest may be important for giant pandas. Johnson (1980) suggested that lack of selection of a common 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 farmland 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 giant 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 conservation purposes. High and mature B. fargesii habitat is often regarded as less important to giant pandas. 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 mature B. fargesii habitats during spring season for foraging on thick bamboo shoots. The shoots consumed by giant pandas subsequently disappear from the groves, and giant panda droppings resulting from these shoots do not exist for >2 months because of easy decomposition. Thus, little sign was evident in the high and mature B. fargesii 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. qinlingensis 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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Fig. 3. Comparison of tree parameters with bamboo understory between giant panda-presence (white boxplot) and giant pandaabsence (grey boxplot) habitats (N = number of plots, NS = no significant difference, P = significance level, DBH = tree diameter at breast height), Foping Nature Reserve, China, JulAug 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.

the giant panda habitat, and consequently, allows managers to determine where the best habitat areas are for giant pandas in Foping NR, where potentially 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, managers 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 survey. We thank the State Forest Administration (SFA) of China for allowing us to carry out the field surveys in Foping Nature Reserve. We sin-

MANAGEMENT IMPLICATIONS
The characteristics of tree and bamboo layers influence giant panda habitat selection. Therefore, human activities on habitat, such as new cultivation, mushroom production, road construction, and development could affect giant pandas. Our analysis can provide strong support for researchers to map and assess the spatial pattern of

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Fig. 4. Comparison of bamboo parameters between giant panda-presence (white boxplot) and giant panda-absence (grey boxplot) habitats, Foping Nature Reserve, China, July-August 1999. N = number of plots, NS = no significant, S = significant difference 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.

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.

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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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