HABITAT USE AND SEPARATION BETWEEN THE GIANT PANDA AND THE RED PANDA
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Journal of Mammalogy, 81(2):448–455, 2000
HABITAT USE AND SEPARATION BETWEEN THE GIANT PANDA
AND THE RED PANDA
FUWEN WEI,* ZUOJIAN FENG, ZUWANG WANG, AND JINCHU HU
Institute of Zoology, The Chinese Academy of Sciences, Beijing, 100080, China (FW, ZF, ZW)
Institute of Rare Animals & Plants, Sichuan Normal College, Nanchong,
Sichuan, 637002, China (JH)
Habitat use and separation between 2 sympatric species, the giant panda (Ailuropoda me-
lanoleuca) and the red panda (Ailurus fulgens), were studied in Yele Natural Reserve,
Mianning County of Sichuan Province, China, to elucidate the coexistence of these spe-
cialized carnivores. Nineteen variables describing proximate habitat structure were mea-
sured at each fecal-group site. We tested if habitat structure differed between pandas and
examined habitat separation between the species. Habitats used by each species differed
significantly. The 2 pandas exhibited different patterns in microhabitat use, although their
habitats overlapped in the study area. The giant panda occurred at sites on gentle slopes
with lower density of fallen logs, shrubs, and bamboo culms. Sites also were close to trees
and far from fallen logs, shrubs, and tree stumps. The red panda occurred at sites on steeper
slopes with higher density of fallen logs, shrubs, and bamboo culms. Sites also were close
to fallen logs, shrubs, and tree stumps. We conclude that microhabitat separation contributes
to coexistence of giant and red pandas in areas of sympatry.
Key words: Ailuropoda, Ailurus, coexistence, giant panda, microhabitat, red panda, resource par-
titioning
The giant panda (Ailuropoda melanoleu- Category II species, respectively, in the Na-
ca) and red panda (Ailurus fulgens) are en- tional Protected Animal List in China. Both
demic to the Himalayan–Hengduan Moun- also are listed by Convention on Interna-
tains. The giant panda now is found only in tional Trade in Endangered Species
Sichuan, Shaanxi, and Gansu provinces of (CITES) as Appendix I species. These 2
China (Hu et al. 1990; Schaller et al. 1985). species, in the order Carnivora, share a
The red panda, in contrast, has a larger number of anatomical and ecological char-
range than the giant panda, extending from acteristics. Both pandas represent not only
central Nepal eastward along the Himalayas monotypic genera, but also are sole repre-
through Bhutan, India, and Myanmar into sentatives of the subfamilies Ailuropodinae
China (Glatston 1994; Roberts and Gittle- and Ailurinae (Glatston 1989). Secondly,
man 1984; Wei et al. 1999a). In China, the they have specialized on bamboo diets and
2 panda species are sympatric in the Qion- share the same bamboo species in regions
glai, Minshan, Xiangling, and Liangshan of sympatry (Johnson et al. 1988; Reid et
mountains of Sichuan Province (Hu et al. al. 1991; Schaller et al. 1985; Wei et al.
1990; Schaller et al. 1985; Wei et al. 1995, 1996a). However, they retain the
1999a). short, relatively simple digestive tracts typ-
Giant and red pandas are treated as rare ical of other carnivores and cannot digest
animals and are listed as Category I and cellulose (Dierenfeld et al. 1982; Schaller
et al. 1985; Warnell et al. 1989; Wei et al.
* Correspondent: weifw@panda.ioz.ac.cn 1999b, 1999c). Finally, both pandas are
448May 2000 WEI ET AL.—GIANT AND RED PANDAS 449
confronted by the same environmental pres- dodendron, Lonicera, Sorbus, and Rosa. This for-
sures, such as habitat loss, population iso- est was the main habitat of both pandas. Alpine
lation, and human interference (Glatston shrub and meadow extended from 3,700 to 4,400
1994; Hu et al. 1990; Pan et al. 1988; m.
Schaller et al. 1985; Wei et al. 1999a). Re- Five species of bamboo (Bashania spanos-
tachya, Fargesia dulcicula, F. exposita, F. ferax,
cent studies have shown that both pandas
Yushania tineloata) occurred in the reserve, but
suffer high mortality in the wild: about 57%
B. spanostachya was dominant. This species,
for cubs of giant pandas in the Qionglai
which covered whole hillsides of our study ar-
Mountains (Wei and Hu 1994; Wei et al. eas, was the main food resource of both pandas.
1997a) and 86% for cubs of red pandas in Because the other 4 species occurred at low el-
Nepal (Yonzon 1989). evations and were disturbed extensively by hu-
Given their similar diets and similar hab- man activities, giant and red pandas seldom feed
itat, we evaluated how the 2 panda species on them.
coexist without competing for resources. Only 3 seasons could be distinguished in the
Our 1st objective was to determine what reserve. Winter lasted from October to March;
habitat characteristics are used by giant and spring lasted from April to June. Summer–au-
red pandas, and whether differential habitat tumn was about 3 months from July to Septem-
use provided significant separation between ber. At an elevation of 2,600 m, the average an-
the species. The 2nd objective was to un- nual rainfall, humidity, and temperature were
derstand how both pandas coexist in the 2,076.6 mm, 87.9%, and 7.18C, respectively.
same habitat while feeding on the same Annual temperatures ranged from 217.08C to
diet. 24.78C. Mean daily temperatures were highest
(15.98C) in July, and lowest (24.98C) in Janu-
MATERIALS AND METHODS ary.
Sampling.—Feces of the 2 pandas are similar
Study areas.—Field work was conducted in the in shape, but those of the giant panda are mas-
Yele Natural Reserve, Mianning County, south- sive (average length by width, 14.5 cm by 5.0
western Sichuan Province, Peoples Republic of cm). Feces of the red panda are small (4.4 cm
China (288509–298029N, 1018589–1028159E). The by 2.2 cm). Feces of an infant giant panda (7.5
reserve was 1 of 14 newly established reserves cm by 2.5 cm) are larger than those of an adult
of the Giant Panda Project in China and was in red panda, making it very easy to distinguish
the western Lesser Xiangling Mountains and
feces in the field.
southeastern Daxueshan Mountains. The reserve
Both giant and red pandas usually leave a
included about 242 km2 of rugged ridges and nar-
group of feces at feeding sites. Numbers in each
row valleys at elevations of 2,600–5,000 m. Our
group vary significantly. When feeding and rest-
research base was set at 3,100 m, and a concen-
ing for a short time, the giant panda often leaves
trated study area of about 25 km2 was in the up-
1–4 feces in a group, infrequently 5–10. During
per Shihuiyao Valley.
long rests, it leaves $10 feces (Reid and Hu
The vegetation showed characteristic vertical
zonation. Mixed coniferous and deciduous broad- 1991; Schaller et al. 1985). Numbers of feces in
leaf forest occurred below 2,800 m. The original a single red panda defecation are usually 8–15,
vegetation was dominated by Tsuga chinensis, and 15–30 or sometimes .100 are found in re-
Betula platyphylla, B. utilisi, and Acer. However, peatedly used sites, called latrines (Reid et al.
most of the forest has been degraded because of 1991; Wei et al. 1995; Yonzon 1989). Field ob-
cultivation, firewood cutting, and logging to servation indicated that the longer the animals
shrubland and meadow; these habitats are unsuit- spent at the feeding sites, the more fecal groups
able for pandas. Between 2,800 and 3,700 m, a were left. Therefore, a positive linear relation-
subalpine coniferous forest dominated in the re- ship existed between total time spent in feeding
serve. Dominant conifers were Abies fabri and sites and number of feces deposited (Reid and
Sabina pingii. Betula, Acer, and Prunus were the Hu 1991; Wei et al. 1996b). Because of the dif-
most common deciduous broadleafed trees. Dom- ficulty of observing activity of either panda in
inant shrubs were Bashania spanostachya, Rho- the field, fecal groups were selected as an indi-450 JOURNAL OF MAMMALOGY Vol. 81, No. 2
the 400-m2 square plot provided information
about canopy, slope, aspect, numbers of trees
and shrubs, diameter at breast height, distance
from fecal group to the nearest trees and shrubs,
numbers of fallen logs and tree stumps, and di-
ameter of the nearest fallen logs and tree stumps.
Nineteen habitat variables of giant and red pan-
das were measured (Appendix I). Because .1
fecal groups may have occurred in some of 20
m by 20 m plots, only 1 fecal group was used
as the plot center to measure habitat.
Statistical analysis.—A Bartlett-Box test was
used to evaluate the homogeneity of variance for
each variable between giant and red pandas.
One-way analysis of variance and Mann-Whit-
ney U-tests were used to test whether habitat
variables of both pandas differed. Discriminant
function analysis was used to examine habitat
FIG. 1.—Plot arrangement for habitat sam- separation between species. This method gen-
pling of sites used by giant and red pandas. erally has been applied to systematic studies in
the past, but has been used widely to measure
differences in habitat utilization patterns of dif-
rect index for quantifying habitat use of both ferent species in ecological studies (Beebee
pandas. 1985; Dueser and Shugart 1978; Marnell 1998;
Dueser and Shugart (1978) created a detailed Reinert 1984; Van Horne 1982). The stepwise
sampling technique that combined plots of var- method of discriminant function analysis was
ious sizes and shapes, as well as small transects. applied because it can be used as an exploratory
Van Horne (1982) and Reinert (1984) applied tool to identify predictor variables from poten-
similar methods to measure habitat use of deer tially useful parameters (Marnell 1998). That ap-
mice (Peromyscus) and snakes, respectively. proach entered variables into discriminant func-
Morrison et al. (1992) remarked that although tion analysis individually, and the variable that
designed for analysis of small-mammal habitat, minimized the overall Wilks’ lambda for the
these methods could be adapted easily for anal- function was selected for entry at each step. Var-
ysis of most terrestrial vertebrates. iable selection ended when no additional in-
We applied a similarly modified sampling crease in the accuracy of the discriminant func-
method to measure habitat of both pandas. From tion was achieved (Norusis 1994). Thus, that ap-
1995 to 1997, we walked through habitats of proach selected significant variables, which can
giant and red pandas in our study areas to search be best used to discriminate between sites of
for signs and fresh fecal groups left by both spe- both species. We selected the significantly dif-
cies. Our searches covered all habitats that could ferent variables between the species after pro-
be used by the 2 species. Whenever a fresh fecal cessing the parameter and nonparameter tests to
group was encountered, 3 independent sampling enter the stepwise approach. All tests above are
units were built and centered on the fecal loca- performed in SPSS for Windows (Norusis
tion. The sampling units included one 1.0-m2 1994).
bamboo plot at the center, two 20-m2 rectangular
transects (each 2 m by 10 m), and one 400-m2 RESULTS
square plot. At the center of each 100-m2 quad-
The mean and SD of the 19 habitat var-
rant within the 400-m2 square plot, an additional
1.0-m2 bamboo plot also was sampled (Fig. 1). iables demonstrated some differences be-
These five 1.0-m2 bamboo plots supplied a de- tween the 2 species (Table 1). Bartlett-Box
tailed measure of bamboo parameters such as univariate homogeneity of variance tests in-
density, height, basal diameter, and proportion of dicated that variances of 11 of 19 variables
old shoots. Two 20-m2 rectangular transects and were equal or homogeneous, whereas 8May 2000 WEI ET AL.—GIANT AND RED PANDAS 451
TABLE 1.—Mean, SD, and Bartlett-Box univariate homogeneity of variance tests of 19 habitat
variables of giant and red pandas.
Giant panda Red panda
(n 5 81) (n 5 92) Bartlett-Box test
Habitat variables X̄ SD X̄ SD F P
Canopy 3.11 0.69 3.27 0.63 0.6660 0.415
Slope 1.79 0.75 2.73 0.68 0.8699 0.351
Aspect 1.91 0.73 2.27 0.74 0.0343 0.853
Bamboo culm density 27.40 3.59 33.45 6.41 26.2771 ,0.000
Bamboo basal diameter 11.11 0.79 10.23 1.01 4.9203 0.027
Bamboo culm height 233.83 33.79 210.73 35.37 0.1757 0.675
Old shoot proportion 0.14 0.10 0.11 0.06 22.5347 ,0.000
Tree density 1.26 0.50 1.14 0.50 0.0009 0.976
Tree size 42.91 7.55 43.12 8.20 0.5768 0.448
Tree dispersion 2.44 0.55 4.02 1.40 64.0764 ,0.000
Shrub density 1.11 0.87 2.79 1.38 17.0255 ,0.000
Shrub size 21.61 4.77 22.08 4.39 0.6044 0.437
Shrub dispersion 4.03 1.81 1.95 0.96 33.3949 ,0.000
Fallen-log density 0.90 0.53 1.76 0.69 5.1923 0.023
Fallen-log size 30.61 6.69 31.31 6.78 0.0157 0.900
Fallen-log dispersion 4.18 1.44 1.98 0.97 12.8898 ,0.000
Tree-stump density 0.66 0.57 0.88 0.59 0.0923 0.761
Tree-stump size 29.81 5.71 30.73 6.82 2.6032 0.107
Tree-stump dispersion 4.30 1.20 2.49 1.11 0.5616 0.454
were unequal (Table 1). Although homo- habitat use. The discriminant function anal-
geneity of variance is an underlying as- ysis correctly classified 96.5% (167 of 173
sumption for analysis of variance, violation samples) of the habitat samples according
of this assumption is typical for ecological to species, 97.5% (79 of 81 samples) for
data and does not necessarily negate the giant pandas and 95.7% (88 of 92 samples)
derivation of biologically meaningful re- for red pandas. Although parameter and
sults from such analyses (Reinert 1984). nonparameter tests detected 13 and 12 po-
Because data obtained had nonnormal tential variables in identifying sites of giant
distributions, parametric and nonparametric and red pandas, the stepwise approach only
tests were applied to compare results. One- identified 8 predictor variables that ap-
way analysis of variance detected that 13 of peared to be most significant in discrimi-
19 variables differed significantly between nating sites of both species (Table 3).
giant and red pandas (P , 0.05; Table 2). Standardized canonical discriminant-
The Mann-Whitney U-test detected that 12 function coefficients and correlations be-
of 19 variables differed significantly be- tween discriminating variables and canoni-
tween species (P , 0.05). Results of para- cal discriminant functions can be used to
metric and nonparametric tests were almost judge the relative contribution to the power
the same relative to variables and level of of discriminant function. Larger absolute
probability, revealing that both species used values of correlations or coefficients indi-
different microhabitats. cate stronger contribution to the power of
The discriminant function analysis of the the function (Cooley and Lohnes 1971).
2 species was significant (eigenvalue 5 Correlations of the 8 indicator variables
3.577, Wilks’ l 5 0.219, x2 5 253.999, d.f. with the discriminant function fell within a
5 8, P , 0.001), which suggested that the narrow range of absolute values (0.304 and
2 species exhibited different patterns in 0.480; Table 3). Fallen-log dispersion con-452 JOURNAL OF MAMMALOGY Vol. 81, No. 2
TABLE 2.—One-way analysis of variance (ANOVA) and Mann-Whitney U-test for 19 habitat var-
iables of giant and red pandas.
ANOVA (d.f. 5 1, 172) Mann-Whitney U-test
Habitat
variables F P U P
Canopy 2.561 0.111 3,274.5 0.128
Slope 73.971 ,0.00 528.0 ,0.00
Aspect 10.204 0.002 2,760.0 0.002
Bamboo culm density 56.648 ,0.00 1,490.0 ,0.00
Bamboo basal diameter 39.984 ,0.00 1,842.0 ,0.00
Bamboo culm height 19.160 ,0.00 2,432.5 ,0.00
Old shoot proportion 5.174 0.024 3,237.0 0.137
Tree density 2.399 0.123 3,256.5 0.136
Tree size 0.030 0.863 3,690.5 0.914
Tree dispersion 90.484 ,0.00 1,049.0 ,0.00
Shrub density 89.276 ,0.00 1,084.0 ,0.00
Shrub size 0.449 0.504 3,369.5 0.278
Shrub dispersion 92.156 ,0.00 1,329.5 ,0.00
Fallen-log density 83.082 ,0.00 996.5 ,0.00
Fallen-log size 0.468 0.495 3,507.5 0.506
Fallen-log dispersion 140.659 ,0.00 537.0 ,0.00
Tree-stump density 6.114 0.014 2,680.5 0.001
Tree-stump size 0.911 0.341 3,530.0 0.551
Tree-stump dispersion 106.717 ,0.00 931.0 ,0.00
tributed most to the power of the discrimi- tree dispersion contributed least. Although
nant function, and bamboo density contrib- results of 2 analyses differed in some var-
uted least. Standardized coefficients of the iables, some were ranked the same. Because
8 selected variables also fell between 0.218 correlations and coefficients were similar,
and 0.407. Fallen-log dispersion contribut- those 8 variables seemed to contribute al-
ed most to the power of the function, but most equally to the power of the discrimi-
nant function and could be treated as indi-
cators in identifying sites of giant and red
TABLE 3.—Stepwise approach of discriminant
functional analysis for 13 significantly different
pandas.
habitat variables of giant and red pandas (max- The giant panda occurred at sites on gen-
imum significance of F to enter 0.05, minimum tle slopes with lower density of fallen logs,
significant of F to remove 0.1). shrubs, and bamboo culms. The sites also
were close to trees and far from fallen logs,
Correlation shrubs, and tree stumps. Conversely, the red
between
Standardized discriminating panda occurred at sites on steeper slopes
canonical variables and with higher density of fallen logs, shrubs,
discriminant canonical and bamboo culms. The sites were close to
function discriminant fallen logs, shrubs, and tree stumps.
Habitat variables coefficients functions
Fall-log dispersion 20.407 20.480 DISCUSSION
Shrub density 0.366 0.382
Microhabitat separation has been dem-
Slope 0.353 0.348
Fallen-log density 0.350 0.369 onstrated by a number of studies to be the
Shrub dispersion 20.336 20.388 most common form of niche partitioning in
Bamboo culm density 0.322 0.304 sympatric species of mammals (Brown and
Tree-stump dispersion 20.243 20.418 Lieberman 1973; Dueser and Shugart 1978;
Tree dispersion 0.218 0.385
Van Horne 1982; Wang 1995), birds (CodyMay 2000 WEI ET AL.—GIANT AND RED PANDAS 453
1978), reptiles (Reinert 1984), amphibians which happen to be associated with steeper
(Marnell 1998), and fishes (Werner and slopes.
Hall 1979). Habitat separation often is con-
sidered to be responsible for multispecies ACKNOWLEDGMENTS
coexistence (Schoener 1974). Our results This project was supported by the Zoo Berlin,
demonstrated that each panda used a dif- National Natural Science Foundation of China
ferent microhabitat and microhabitat sepa- (39870102 and 39730110), and the Young Sci-
ration contributed to coexistence of giant entist Funds of the Chinese Academy of Scienc-
es. P. Tang, E. Gutie, and W. Lu participated in
and red pandas in areas of sympatry.
parts of the fieldwork. The Sichuan Forest Bu-
Giant pandas, having larger body size, reau and Mianning Forest Bureau gave us assis-
used sites with lower densities of shrubs, tance during the field research. Previous ver-
fallen logs, and bamboo culms. Feeding and sions of the manuscript benefited from com-
moving in this more open microhabitat ments by G. B. Schaller, A. R. Glatston, T. Ful-
could reduce energy expenditures. Saving ler, R. Hoffmann, and D. G. Reid.
energy is important for the giant panda be-
cause its daily energy intake exceeds expen- LITERATURE CITED
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WEI, F., Z. FENG, Z. WANG, AND J. HU. 1999a. Current were randomly measured at each plot); OLD
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