Habitat use by giant pandas Ailuropoda melanoleuca in the Wanglang Nature Reserve, Sichuan, China
© Kang et al.; licensee Springer. 2013
Received: 28 September 2012
Accepted: 10 May 2013
Published: 30 September 2013
To better understand the ecological requirements of the giant panda Ailuropoda melanoleuca in the wild, field surveys were carried out at both the microhabitat scale and foraging site scale in Wanglang National Nature Reserve, Sichuan, China.
The results indicated that (1) at the microhabitat scale, giant pandas usually occupied habitats with a high fallen-log density, lower shrub density, and bamboo coverage of 50% to 75%; (2) at the foraging site scale, pandas usually used sites with higher bamboo densities and taller and larger-diameter bamboo; and (3) giant pandas may abandon plots when the proportion of young bamboo decreases below average in the environment.
The availability of young bamboo is an important driving force in habitat selection by giant pandas, which could provide important reference for the conservation of giant pandas and their habitats.
The giant panda Ailuropoda melanoleuca is a flagship species for efforts related to habitat conservation, is one of the most important rare and endangered species in the world, and is a category I protected species on the Red List of China (Wei et al. 2000). Despite efforts by the government, international organizations, and local people, loss and fragmentation of panda habitat threaten the species throughout its range (State Forestry Administration 2006; Shen et al. 2008). Thus, a deeper understanding of the habitat requirements of the giant panda is needed to develop specific conservation strategies for this endangered species.
A knowledge on the habitat requirements of animal species provides important information needed for conservation planning and policy making (Pan et al. 1998; Hu 2001; Ouyang et al. 2001; Wang et al. 2002; Liu et al. 2005 2006; Yang et al. 2006; Xu et al. 2006; Swaisgood et al. 2010; Wang et al. 2010; Wei et al. 2011), but species-habitat relationships can vary among different regions (Hu 2001; Wei et al. 2011; Panthi et al. 2012). For example, an abundance of research indicates that giant pandas usually use primary forests (Ran et al. 2003 2004; Wang et al. 2006), although some research suggests that they also use secondary forests (Zhang et al. 2002a). Others reported that pandas use both primary and secondary forests equally with no significant habitat preference (Zeng et al. 2002; Zeng et al. 2003), so land managers should use caution if they wish to apply one region’s conservation measures to another region (Wei et al. 2011). However, understanding the habitat requirements of an animal species in a specific area and then developing site-specific conservation strategies should be considered because this knowledge is important for managers who wish to implement effective conservation measures.
We report the results of field surveys on giant panda habitat in the Wanglang Nature Reserve. We identified variables related to giant panda habitat at the microhabitat scale and foraging site scale (Zhang et al. 2002b). The objective of our study was to discriminate habitat features which can be used to explain the habitat use of pandas in this area. By studying foraging sites used by giant pandas, we aimed to draw some general conclusions related to factors that drive habitat selection. These insights should deepen our understanding on the habitat requirements of giant pandas and facilitate identification of appropriate management strategies in this area.
Habitat use at the microhabitat scale
In January, March, May, and June 2011, we conducted field surveys along 40 transects (Figure 1) in Wanglang Nature Reserve and repeated the surveys in April, May, July, and August 2012. All transects were established along an elevational gradient to ensure that each sampled transect contained typical and representative panda habitat. Each transect was at least 2 km long.
Descriptions and definitions of variables used in this study
Slope aspect (deg)
Nine categories: no aspect (slope of <5°), north, northeast, east, southeast, south, southwest, west, and northwest
Five categories: <10°, 10° to 20°, 20° to 30°, 30° to 40°, and ≥40°
Canopy of overstory in sampling plot, four categories: <25%, 25% to 50%, 50% to 75%, and ≥75%
Shrub coverage (%)
Coverage of shrubs in the sampling plot, four categories: <25%, 25% to 50%, 50% to 75%, and ≥75%
Bamboo coverage (%)
Coverage of bamboo in the sampling plot, four categories: <25%, 25% to 50%, 50% to 75%, and ≥75%
Tree size (cm)
Average breast diameter (DBH) of trees in each 100-m2 square plot nearest to the center of a 400-m2 plot
Tree dispersion (m)
Average distance of the nearest tree to the center in each 100-m2 square plot
Average number of trees in two 20-m2 rectangular transects in a 400-m2 plot
Shrub size (cm)
Average DBH of shrubs in each 100-m2 square plot nearest to the center of a 400-m2 plot
Shrub dispersion (m)
Average distance of the nearest shrub to the center in each 100-m2 square plot
Average number of shrubs in two 20-m2 rectangular transects in a 400-m2 plot
Total number of fallen logs (>10 cm in diameter) in a 400-m2 plot
Tree stump density
Total number of tree stumps (> 10 cm in diameter) in a 400-m2 plot
Foraging site scale
Number of culms in a 1 × 1-m bamboo site
Old-bamboo size (mm)
Average basal diameter of culms in a 1 × 1-m bamboo site (five old bamboo culms were measured randomly at each site)
Old-bamboo height (m)
Average height of culms in a 1 × 1-m bamboo site (five old bamboo culms were measured randomly at each site)
Number of young bamboo culms in a 1 × 1-m bamboo site
Young-bamboo proportion (%)
Proportion of young bamboo culms in a 1 × 1-m bamboo site
Habitat use at the foraging site scale
Five 1 × 1-m bamboo sites were established to investigate bamboo characteristics, such as bamboo density, basal diameter of old bamboo, height of old bamboo, young-bamboo density, and the proportion of young bamboo at each site (Table 1), and they were at the center of 20 × 20-m plots and the center of each 10 × 10-m plot in 2012 (Wei et al. 2000; Zhang et al. 2006 2009). However, only four 1 × 1-m bamboo sites were randomly placed in each 10 × 10-m plot in 2011, but we also used these data as a reference.
The bamboo sites surveyed in the panda-use plots were divided into two categories: forage and non-forage sites. A forage site was defined as a small area where remnants of bamboo foraged by giant pandas were found (Zhang et al. 2009). Bamboo sites surveyed in the non-use plots were defined as control sites that reflected the surrounding environment. In total, 1,456 bamboo sites (151 forage sites, 413 non-forage sites, and 892 control sites) were sampled for analysis.
To describe habitat characteristics and compare habitat differences of panda-use and non-use plots, we used a χ 2 test to compare discrete variables; for continuous variables, we used independent-sample t tests when data were normally distributed and the Mann–Whitney U test when the distributional assumptions were not met. To identify factors that differentiated microhabitat characteristics of panda-use and non-use plots, variables showing a significant difference between these two types of plots were subsequently analyzed with a logistic regression analysis. Also, to ensure that variables were independent, only variables with clear biological meaning were considered during subsequent analysis for those with absolute correlation coefficients of >0.70 (Schweiger et al. 2012).
To describe habitat characteristics of giant pandas at a foraging site scale, we used a one-way analysis of variance (ANOVA) test to compare means of variables among different bamboo site groups when data were normally distributed and a Kruskal-Wallis test when the distributional assumptions were not met. For those variables showing a significant difference, we then used a least significant difference (LSD) multiple comparison to test whether or not significant differences existed between all members when data were normally distributed and a Games-Howell multiple-comparison test when the distributional assumptions were not met. Furthermore, when the data type of the proportion of young bamboo was measured using percent, we used the Kruskal-Wallis test and Games-Howell multiple-comparison test to analyze such variables.
Habitat use at the microhabitat scale
χ 2 test of discrete variables in panda-use and non-use plots
Use plots (n= 186)
Non-use plots (n= 329)
χ 2 = 29.33, d.f. = 8, p = 0.00
χ 2 = 7.15, d.f. = 4, p = 0.13
χ 2 = 3.74, d.f. = 3, p = 0.29
χ 2 = 19.45, d.f. = 3, p = 0.00
χ 2 = 140.24, d.f. = 3, p = 0.00
Mann–Whitney U test of continuous variables in panda-use and non-use plots
Mean (mean rank)
32.20 (250.30) (n = 180)
29.07 (233.80) (n = 299)
2.86 (218.27) (n = 180)
3.08 (253.08) (n = 299)
2.51 (252.60) (n = 186)
2.55 (261.05) (n = 329)
2.41 (280.19) (n = 184)
2.03 (243.21) (n = 328)
2.05 (265.97) (n = 182)
1.88 (247.30) (n = 325)
4.74 (212.09) (n = 186)
7.24 (283.96) (n = 329)
1.96 (283.17) (n = 186)
1.65 (243.77) (n = 329)
Tree stump density
0.77 (272.83) (n = 186)
0.65 (249.62) (n = 329)
Logistic regression analysis (backward stepwise (conditional) method) of seven microhabitat variables with significant differences
Habitat use at the foraging site scale
ANOVA and Kruskal-Wallis test for foraging site variables among different bamboo site groups
Mean (SD) or Mean (rank)
F or χ 2
Forage sites (n= 151)
Non-forage sites (n= 413)
Control sites (n= 892)
Multiple comparisons of four foraging site variables with significant differences
95% Confidence interval
Many factors affect habitat use by giant pandas, such as food distribution, concealment conditions, body size, and so on (Wei et al. 2000; Zhang et al. 2009). Of the 13 variables tested in this study, 7 significantly differed between panda-use and non-use plots (Table 2); however, only bamboo coverage, shrub density, and fallen-log density were entered into the prediction equation, indicating that they were the most important indicators of habitat use by giant pandas.
Giant pandas feed almost exclusively on bamboo (Hu 2001; Yang et al. 2006; Zhang et al. 2009), so the life of giant pandas is directly influenced by the availability of bamboo. Habitat with lower bamboo coverage is not a suitable giant panda habitat because of the additional energy required for pandas to forage for food in such areas (Hu 2001). However, areas with high bamboo coverage impede the travel of pandas requiring the expenditure of additional energy (Hu 2001). Thus, habitat with medium levels of bamboo coverage provides adequate food for giant pandas and also allows pandas to more easily move across the landscape.
Fallen logs usually serve as passageways connecting different habitat areas and allow pandas to travel more easily within their habitat. Additionally, an abundance of fallen wood is an important characteristic of old-growth forests; based on this study, giant pandas typically occupy habitats with high fallen-log densities, which is consistent with previous reports stating that giant pandas require old-growth forest habitats (Zhang et al. 2011). Furthermore, shrubs are not a food of giant panda, and habitats with lower shrub densities could reduce the energy expenditure of giant panda when moving about (Wei et al. 2000).
Foraging site use
A panda can eat 10~18 kg of fresh leaves or stems or about 40 kg of new shoots per day and will spend more than 50% of the day foraging (Hu 2001; Zhang et al. 2009), so pandas need to conserve energy and search for high-quality food. Bamboo sites with a high density of bamboo, tall bamboo, and large-diameter bamboo stems provide high levels of biomass and high-quality and adequate food resources for giant pandas; this allows pandas to maximize their energy and nutrient intake in this type of foraging site and helps them reduce energy expenditures when moving in search of food (Zhang et al. 2009).
Giant pandas primarily feed on young bamboo shoots (Hu 2001; Zhang et al. 2009), and young bamboo provides high-quality food for giant pandas. This research confirms that proportions of young bamboo at both forage and non-forage sites were all significantly lower than that at control sites. Furthermore, no significant difference was found between forage and non-forage sites, which indicates that giant pandas would need to leave their current use plots when the proportion of young bamboo falls below average in the environment and search for sites with higher-quality food.
In conclusion, the availability of young bamboo is an important driving force in habitat selection by giant pandas.
This study was supported by the National Key Technology Research and Development Program in the 11th Five-Year Plan of China (2008BADB0B04) and the National Natural Science Foundation of China (31170500). We would like to thank Xiaorong Wang, Zhiwei Yuan, Chunping Luo, Chunping Liang, Yong Zheng, Jie Ouyang, Jixu Zhao, Gaoshan Lan, Hualong Zhou, Xiaorong Xie, Yunxi Li, Xiong Gao, Wenlong Jiang, Yubo Zhang, and Lijuan Duan for participating in the fieldwork.
- Chen YP, Jiang SW, Zhao LJ, Huang JZ: Surveillance on giant panda habitats in Wanglang Nature Reserve, Sichuan. Sichuan J Zool 2003, 22: 49–50. (in Chinese with English summary)Google Scholar
- Hu JC: Research on the giant pandas. Shanghai: Shanghai Scientific and Technological Press; 2001. (in Chinese)Google Scholar
- Liu XH, Toxopeus AG, Skidmore AK, Shao XM, Dang GD, Wang TJ, Prins HHT: Giant panda habitat selection in Foping Nature Reserve, China. J Wildl Manage 2005, 69: 1623–1632. 10.2193/0022-541X(2005)69[1623:GPHSIF]2.0.CO;2View ArticleGoogle Scholar
- Liu XH, Skidmore AK, Bronsveld MC: Assessment of giant panda habitat based on integration of expert system and neural network. Chin J Appl Ecol 2006, 17: 438–443. (in Chinese with English summary)Google Scholar
- Ouyang ZY, Liu JG, Xiao H, Tan YC, Zhang HM: An assessment of giant panda habitat in Wolong Nature Reserve. Acta Ecol Sin 2001, 21: 1869–1874. (in Chinese with English summary)Google Scholar
- Pan WS, Gao ZS, Lü Z: The natural refuge for giant pandas in Qinling. Beijing: Peking University Press; 1998. (in Chinese)Google Scholar
- Panthi S, Aryal A, Raubenheimer D, Lord J, Adhikari B: Summer diet and distribution of the red panda ( Ailurus fulgens fulgens ) in Dhorpatan Hunting Reserve, Nepal. Zool Stud 2012, 51: 701–709.Google Scholar
- Ran JH, Liu SY, Wang HJ, Sun ZY, Zeng ZY, Liu SC: Effect of grazing on giant pandas’ habitat in Yele Nature Reserve. Acta Theriol Sin 2003, 23: 288–294. (in Chinese with English summary)Google Scholar
- Ran JH, Zeng ZY, Wang H, Liu SY, Wang HJ, Liu SC: A comparative study on habitat preference of giant pandas in primary and secondary forests. J Beijing For Univ 2004, 26: 8–14. (in Chinese with English summary)Google Scholar
- Schweiger AK, Ursula NM, Margit Z: Small-scale habitat use of black grouse ( Tetrao tetrix L.) and Rock Ptarmigan ( Lagopus muta helvetica Thienemann) in the Austrian Alps. Eur J Wildl Res 2012, 58: 35–45. 10.1007/s10344-011-0537-7View ArticleGoogle Scholar
- Shen GZ, Feng CY, Xie ZQ, Ouyang ZY, Li JQ, Pascal M: Proposed conservation landscape for giant pandas in the Minshan Mountains, China. Conserv Biol 2008, 22: 1144–1153. 10.1111/j.1523-1739.2008.01038.xView ArticlePubMedGoogle Scholar
- State Forestry Administration: The third national survey report on giant panda in China. Beijing: Science Press; 2006. (in Chinese)Google Scholar
- Swaisgood RR, Wei FW, Wildt DE, Kouba AJ, Zhang ZJ: Giant panda conservation science: how far we have come. Biol Lett 2010, 6: 143–145. 10.1098/rsbl.2009.0786PubMed CentralView ArticlePubMedGoogle Scholar
- Wang MJ, Li JQ: Research on habitat restoration of giant panda after a grave disturbance of earthquake in Wanglang Nature Reserve, Sichuan Province. Acta Ecol Sin 2008, 28: 5848–5855. (in Chinese with English summary)View ArticleGoogle Scholar
- Wang H, Li SG, Pan WS: Population viability analysis of giant panda ( Ailuropoda melanoleuca ) in Qinling Mountains. Acta Sci Nat Univ Peking 2002, 38: 756–761. (in Chinese with English summary)Google Scholar
- Wang JH, Huang HL, Teng JR, Yang WY, Ma T: Habitat selection by giant pandas in west of Baishuijiang Nature Reserve. Sichuan J Zool 2006, 25: 771–775. (in Chinese with English summary)Google Scholar
- Wang TJ, Ye XP, Skidmore AK, Toxopeus AG: Characterizing the spatial distribution of giant pandas ( Ailuropoda melanoleuca ) in fragmented forest landscapes. J Biogeogr 2010, 37: 865–878. 10.1111/j.1365-2699.2009.02259.xView ArticleGoogle Scholar
- Wei FW, Wang ZW, Hu JC: Habitat use and separation between the giant panda and the red panda. J Mammal 2000, 81: 448–455. 10.1644/1545-1542(2000)081<0448:HUASBT>2.0.CO;2View ArticleGoogle Scholar
- Wei FW, Zhang ZJ, Hu JC: Research advances and perspectives on the ecology of wild giant panda. Acta Theriol Sin 2011, 31: 412–421. (in Chinese with English summary)Google Scholar
- Xu WH, Ouyang ZY, Jiang ZY, Zheng H, Liu JG: Assessment of giant panda habitat in the Daxiangling Mountain Range, Sichuan, China. Biodivers Sci 2006, 14: 223–231. (in Chinese with English summary) 10.1360/biodiv.050191View ArticleGoogle Scholar
- Yang CH, Zhang HM, Zhou XP, Wang PY, Wang XM: Review of habitat selection in the giant panda ( Ailuropoda melanoleuca ). Acta Ecol Sin 2006, 26: 3442–3453. (in Chinese with English summary)Google Scholar
- Zeng ZY, Yue BS, Ran JH, Liu SY, Chen YP, Jiang SW: Panda’s exploitation of habitats at Wanglang Nature Reserve. J Sichuan Univ (Nat Sci) 2002, 39: 1140–1144. (in Chinese with English summary)Google Scholar
- Zeng T, Ran JH, Liu SY, Sun ZY, Liu SC, Zeng ZY: Habitat exploitation by panda in the Baihe Nature Reserve. Chin J Appl Environ Biol 2003, 9: 405–408. (in Chinese with English summary)Google Scholar
- Zhang ZJ, Hu JC, Wu H: Comparison of habitat selection of giant pandas and red pandas in the Qionglai Mountains. Acta Theriol Sin 2002, 22: 161–168. (in Chinese with English summary)Google Scholar
- Zhang ZJ, Wei FW, Hu JC: Habitat selection and separation: a case for the giant panda and the red panda. J China West Normal Univ (Nat Sci) 2002, 28: 111–116. (in Chinese with English summary)View ArticleGoogle Scholar
- Zhang ZJ, Wei FW, Li M, Hu JC: Winter microhabitat separation between giant and red pandas in Bashania faberi bamboo forest in Fengtongzhai Nature Reserve. J Wildl Manage 2006, 70: 231–235. 10.2193/0022-541X(2006)70[231:WMSBGA]2.0.CO;2View ArticleGoogle Scholar
- Zhang ZJ, Zhan XJ, Yan L, Li M, Hu JC, Wei FW: What determines selection and abandonment of a foraging patch by wild giant pandas ( Ailuropoda melanoleuca ) in winter? Environ Sci Pollut Res 2009, 16: 79–84. 10.1007/s11356-008-0066-4View ArticleGoogle Scholar
- Zhang ZJ, Swaisgood RR, Zhang SN, Nordstrom LA, Wang HJ, Gu XD, Hu JC, Wei FW: Old-growth forest is what giant pandas really need. Biol Lett 2011, 7: 403–406. 10.1098/rsbl.2010.1081PubMed CentralView ArticlePubMedGoogle Scholar
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