Plant Ecology

, Volume 192, Issue 1, pp 97–106 | Cite as

Clonal regeneration of an arrow bamboo, Fargesia qinlingensis, following giant panda herbivory

  • Wei WangEmail author
  • Scott B. Franklin
  • John R. Ouellette
Original Paper


Characteristics of giant panda herbivory sites and clonal regeneration of an arrow bamboo Fargesia qinlingensis following giant panda grazing were studied in the Qinling Mountains of China. Three types of plots were located in a pandas’ summer habitat in 2002: herbivory (naturally grazed by giant pandas), clipped (simulated panda herbivory), and control. Average area of herbivory sites was 2.92 m2 and average distance from herbivory sites to the closest tree (dbh > 10 cm) was 1.0 m. Pandas avoided thin bamboo culms with basal diameters <5 mm. Average height of stumps of culms grazed by panda was 0.67 m and average density of grazed culms was 9.0 culms m−2. Annual culm mortality rate was significantly greater in herbivory and clipped plots than in control plots while annual recruitment rate was not significantly different among the three plot types in 2003. Neither recruitment rate nor mortality rate were significantly different among the three plot types in 2004. Annual recruitment rate was significantly greater than annual mortality rate only in control plots in both 2003 and 2004, suggesting static ramet dynamics in disturbed plots (herbivory and clipped). Density of new shoots was not significantly different, but basal diameter of new shoots was significantly less in herbivory plots compared to control plots in 2002. Differences of annual mortality rate and growth of new shoots found between control plots and herbivory plots suggest that clonal regeneration of F. qinlingensis culms was negatively affected by giant panda grazing. Therefore, no evidence of a clonal integration compensatory response to panda herbivory was found in F. qinlingensis.


Basal diameter Clonal integration Culm Fertilizer effect Mortality New shoot Recruitment 



We are grateful to Dr. Arnold van der Valk and two anonymous reviewers whose comments greatly improved the manuscript. We would like to thank the Foping National Nature Reserve for logistic support of our fieldwork in the reserve. We also thank Gaodi Dang, Leigang Zhao, Xichun Du, and Haining Li for their assistance in the field. This study was supported by the Memphis Zoo.


  1. Alpert P (1999) Clonal integration in Fragaria chiloensis differes between populations: ramets from grassland are selfish. Oecologia 120:69–76CrossRefGoogle Scholar
  2. Baldwin IT (1990) Herbivory simulations in ecological research. Trends Ecol Evol 5:91–93CrossRefGoogle Scholar
  3. Bergelson J, Juenger T, Crawley MJ (1996) Regrowth following herbivory in Ipomopsis aggregate: compensation but not overcompensation. Am Nat 148:744–755CrossRefGoogle Scholar
  4. Bullock JM, Hill BC, Silvertown J (1994) Tiller dynamics of two grasses-responses to grazing, density and weather. J Ecol 82:331–340CrossRefGoogle Scholar
  5. Campbell J, Qin Z, Zhang X (1983) Flowering and population dynamics of bamboo stems in the range of giant pandas, China. World Wildlife Fund, Mimeo ReportGoogle Scholar
  6. Danell K, Niemelä P, Varvikko T, Vuorisalo T (1991) Moose browsing on Scots pine along a gradient of plant productivity. Ecology 72:1624–1633CrossRefGoogle Scholar
  7. Dyer MI, Turner CL, Seastedt TR (1991) Mowing and fertilization effect on productivity and spectral reflectance in Bromus inermis plots. Ecol Appl 1:443–452CrossRefGoogle Scholar
  8. Gadgil M, Prasad SN (1984) Ecological determinants of life history evolution of two Indian bamboo species. Biotropica 16:161–172CrossRefGoogle Scholar
  9. Gough L, Goldberg DE, Hershock C, Pauliukonis N, Petru M (2002) Investigating the community consequences of competition among clonal plants. Evol Ecol 15:547–563CrossRefGoogle Scholar
  10. Haukioja E (1991) The influence of grazing on the evolution, morphology and physiology of plants as modular organisms. Phil Trans Biol Sci 333:241–247CrossRefGoogle Scholar
  11. Herms DA, Mattson WJ (1992) The dilemma of plants: to grow or defend. Quart Rev Biol 67:283–335CrossRefGoogle Scholar
  12. Hu J, Reid DG, Dong S, Wang W, Huang Y (1990) Giant panda behavior and carring capacity following a bamboo die-off. In: Hu J (ed) Research and progress in biology of the giant panda. Sichuan Publishing House of Science of Technology, Chengdu, China, pp 246–262Google Scholar
  13. Janzen DH (1976) Why bamboos wait so long to flower. Annu Rev Ecol Syst 7:347–391CrossRefGoogle Scholar
  14. Jónsdóttir IS, Callaghan TV (1989) Localized defoliation stress and the movement of 14C-photoassimilates between tillers of Carex bigelowii. Oikos 54:211–219CrossRefGoogle Scholar
  15. Li Y (2002) Studies on the taxonomy and distribution of bamboo as the main food of giant panda and the biomass of Bashania fargesii in Mt. Qinling. MA Thesis, Northwest University, Xi’an, ChinaGoogle Scholar
  16. Li R, Werger MJA, de Kroon H, During HJ, Zhong ZC (2000) Interactions between shoot age structure, nutrient availability and physiological integration in the giant bamboo Phyllostachys pubescens. Plant Biol 2:437–446CrossRefGoogle Scholar
  17. Linderman M, Bearer S, An L, Tan Y, Ouyang Z, Liu J (2005) The effects of understory bamboo on broad-scale estimates of giant panda habitat. Biol Conserv 121:383–390CrossRefGoogle Scholar
  18. Liu X (2001) Mapping and modeling the habitat of giant pandas in Foping Nature Reserve, China. PhD Dissertation, Wageningen University, The NetherlandsGoogle Scholar
  19. Long Y, Lu Z, Wang D, Zhu X, Wang H, Zhang Y, Pan W (2004) Nutritional strategy of giant pandas in the Qinling Mountains of China. In: Lindburg D, Baragona K (eds) Giant pandas: biology and conservation. University of California Press, Berkeley/Los Angeles, CA, pp 90–100Google Scholar
  20. Mackinnon J, Sha M, Cheung C, Carey G, Zhu X, Melville D (1996) A biodiversity review of China. WWF International, Hong KongGoogle Scholar
  21. McClure F (1966) Bamboos—a fresh perspective. Harvard University Press, Cambridge, MA, USAGoogle Scholar
  22. McNaughton SJ (1984) Grazing lawns: animals in herds, plant form, and coevolution. Am Nat 124:863–886CrossRefGoogle Scholar
  23. Paige KN (1999) Regrowth following ungulate herbivory in Ipomopsis aggregata: geographic evidence for overcompensation. Oecologia 118:316–323CrossRefGoogle Scholar
  24. Pan W, Gao Z, Lu Z, Xia Z, Zhang M, Ma L, Meng G, Zhe X, Liu X, Cui H, Chen F (1988) The giant panda’s natural refuge in the Qinling Mountains. Beijing University Press, Beijing, ChinaGoogle Scholar
  25. Peltzer DA (2002) Does clonal integration improve competitive ability? A test using aspen (Populus tremuloides [Salicaceae]) invasion into prairie. Am J Bot 98:494–499Google Scholar
  26. Peterson CJ, Squires ER (1995) Competition and succession in an aspen-white-pine forest. J Ecol 83:449–457CrossRefGoogle Scholar
  27. Reid DG, Hu J (1991) Giant panda selection between Bashania fangiana bamboo habitats in Wolong Reserve, Sichuan, China. J Appl Ecol 28:228–243CrossRefGoogle Scholar
  28. Reid DG, Taylor AH, Hu J, Qin Z (1991) Environmental influences on bamboo Bashania fangiana growth and implications for giant panda conservation. J Appl Ecol 28:855–868CrossRefGoogle Scholar
  29. Ren Y (1998) Plants of giant panda’s habitat of Qinling Mountain. Shaanxi Science and Technology Press, Xi’an, ChinaGoogle Scholar
  30. Romme WH, Turner MG, Wallace LL, Walker JS (1995) Aspen, elk, and fire in northern Yellowstone National Park. Ecology 76:2097–2106CrossRefGoogle Scholar
  31. Rooke T (2003) Growth responses of a woody species to clipping and goat saliva. Afr J Ecol 41:324–328CrossRefGoogle Scholar
  32. Rooney TP (1997) Escaping herbivory: refuge effects on the morphology and shoot demography of the clonal forest herb Maianthemum canadense. J Torr Bot Club 124:280–285CrossRefGoogle Scholar
  33. Rounds R (1979) Height and species as factors determining browsing of shrubs by wapiti. J Appl Ecol 16:227–241CrossRefGoogle Scholar
  34. Sackville Hamilton NR, Schmid B, Harper JL (1987) Life-history concepts and the population biology of clonal organisms. Proc Roy Soc London B 232:35–57CrossRefGoogle Scholar
  35. Saitoh T, Seiwa K, Nishiwaki A (2002) Importance of physiological integration of dwarf bamboo to persistence in forest understorey: a field experiment. J Ecol 90:78–85CrossRefGoogle Scholar
  36. SAS (2001) SAS users guide: Statistics, 8.2 edition. SAS institute, Cary, NC, USAGoogle Scholar
  37. Schaller GB, Hu J, Pan W, Zhu J (1985) The giant pandas of Wolong. The University of Chicago Press, Chicago, USAGoogle Scholar
  38. Schmid B, Miao SL, Bazzaz FA (1990) Effects of simulated root herbivory and fertilizer application on growth and biomass allocation in the clonal perennial Solidago canadensis. Oecologia 84:9–15CrossRefGoogle Scholar
  39. Slade AJ, Hutchings MJ (1987) Clonal integration and plasticity in foraging behaviour in Glechoma hederacea. J Ecol 75:1023–1036CrossRefGoogle Scholar
  40. Solangaraachi SM, Harper JL (1987) The effect of canopy filtered light on the growth of white clover (Trifolium repens L). Oecologia 72:372–375CrossRefGoogle Scholar
  41. Stuefer JF, Erschamber B, Huber H, Suzuki J-I (2002) The ecology and evolutionary biology of clonal plants: an introduction to the proceedings of Clone-2000. Evol Ecol 15:223–230CrossRefGoogle Scholar
  42. Swihart RK, Picone PM (1998) Selection of mature growth stages of coniferous browse in temperate forests by white-tailed deer (Odocoileus virginianus). Am Midl Nat 139:269–274CrossRefGoogle Scholar
  43. Taylor AH, Qin Z (1987) Culm dynamics and dry matter production of bamboos in the Wolong and Tangjiahe giant panda reserves, Sichuan, China. J Appl Ecol 24:419–433CrossRefGoogle Scholar
  44. Taylor AH, Qin Z. (1988a) Regeneration from seed of Sinarundinaria fangiana, a bamboo, in the Wolong giant panda reserve, Sichuan, China. Am J Bot 75:1065–1073CrossRefGoogle Scholar
  45. Taylor AH, Qin Z (1988b) Regeneration patterns in old-growth Abies-Betula forests in the Wolong natural reserve, Sichuan, China. J Ecol 76:1204–1218CrossRefGoogle Scholar
  46. Taylor AH, Qin Z (1989) Structure and composition of selectively cut and uncut Abies-Tsuga forest in Wolong natural reserve and implications for panda conservation in China. Biol Conserv 47:83–108CrossRefGoogle Scholar
  47. Taylor AH, Qin Z (1993) Bamboo regeneration after flowering in the Wolong giant panda reserve, China. Boil Conserv 63:231–234CrossRefGoogle Scholar
  48. Taylor AH, Reid DG, Qin Z, Hu J (1991) Spatial patterns and environmental associates of bamboo (Bashania fangiana Yi) after mass-flowering in southwestern China. Bull Torr Bot Club 118:247–254CrossRefGoogle Scholar
  49. Tian X (1987) Ecological observation on flowering and regeneration of bamboo forests in Mt. Qinling. Bamboo Res 2:39–43Google Scholar
  50. Tolvanen A, Schroderus J, Henry GHR (2002) Age- and stage-based bud demography of Salix artica under contrasting muskox grazing pressure in the High Arctic. Evol Ecol 15:443–462CrossRefGoogle Scholar
  51. Wei FW, Feng Z, Wang Z, Hu J (2000) Habitat use and separation between the giant panda and the red panda. J Mamm 81:448–455CrossRefGoogle Scholar
  52. Wei FW, Feng Z, Wang Z, Zhou A, Hu J (1999) Use of the nutrients in bamboo by the red panda (Ailurus fulgens). J Zool Lond 248:535–541CrossRefGoogle Scholar
  53. Wilsey B (2002) Clonal plants in a spatially heterogeneous environment: effects of integration on Serengeti grassland response to defoliation and urine-hits from grazing mammals. Plant Ecol 159:15–22CrossRefGoogle Scholar
  54. Yong Y (1989) A preliminary observation of giant pandas in Foping Natural Reserve. The survey of natural resources in Foping Natural Reserve. Foping, ChinaGoogle Scholar
  55. Yong Y, Wang K, Wang T (1994) Movement habits of the giant pandas in Foping. Acta Theriol Sin 14:9–14Google Scholar
  56. Zeigenfuss LC, Singer FJ, Williams SA, Johnson TL (2002) Influences of herbivory and water on willow in elk winter range. J Wildl Manage 66:788–795CrossRefGoogle Scholar

Copyright information

© Springer Science+Business Media, Inc. 2006

Authors and Affiliations

  • Wei Wang
    • 1
    Email author
  • Scott B. Franklin
    • 1
  • John R. Ouellette
    • 2
  1. 1.Department of BiologyUniversity of MemphisMemphisUSA
  2. 2.Department of Research and ConservationMemphis ZooMemphisUSA

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