Journal of Mountain Science

, Volume 11, Issue 5, pp 1182–1191 | Cite as

Species richness, diversity and density of understory vegetation along disturbance gradients in the Himalayan conifer forest

  • Kesang WangchukEmail author
  • András Darabant
  • Prem Bahadur Rai
  • Maria Wurzinger
  • Werner Zollitsch
  • Georg Gratzer


We investigated whether species richness, diversity and density of understory herbaceous plants differed along logging (gap) and grazing (primarily by cattle) disturbance gradients, and sought to identify drivers of richness, diversity and density of understory vegetation of logged sites. A factorial experiment was conducted in the mixed conifer forest of Gidakom in Western Bhutan. Levels of the logging treatment included small (0.15–0.24 ha), medium (0.25–0.35 ha) and large (0.36–1.31 ha) gaps. The grazing treatment included grazed (primarily by cattle) and ungrazed (where herbivores were excluded by a fence) plots nested within each gap. Data were collected from 12 gaps (4 replicates at each level of logging) using the point intercept method. Shannon Weaver Diversity and Margalef’s indices were used to estimate species diversity and describe species richness, respectively. Soil samples were analyzed for pH and nutrients. The interaction effect of logging and grazing was significant (p≤0.001) only on species diversity. Relative to ungrazed areas, species diversity was significantly higher (0.01≤p≤0.05) in medium grazed gaps. Under grazed conditions, soil P was negatively correlated with gap size and species diversity. While species diversity was positively correlated (0.01≤p≤0.05) with soil N in grazed plots species richness was positively correlated (0.001≤p≤0.01) with soil N in ungrazed plots. Relative density of Yushania microphylla and Carex nubigena were higher under ungrazed conditions. Our study suggests that the combined effect of cattle grazing and logging results in higher species diversity of understory vegetation in medium and grazed gaps in mixed conifer forests of Bhutan,whereas increase or decrease in relative density of major species is determined primarily by the independent effects of grazing and logging. From management perspective, forest managers must refrain from creating large gaps to avoid loss of nutrients (mainly P and N), which may eventually affect tree regeneration. Managers intending to maintain understory vegetation diversity must consider the combined effects of grazing and logging, ensuring low to moderate grazing pressure.


Forest gap Grazing Relative density Soil nutrients Species diversity Species richness 


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  1. Adams PW, Boyle JR (1982) Soil fertility changes following clear-cut and whole tree logging and burning in Central Michigan. Soil Science Society of America Journal 46: 638–640. DOI: 10.2136/sssaj1982.03615995004600030037xCrossRefGoogle Scholar
  2. Bakker ES, Ritchie ME, Olff H, et al. (2006) Herbivore impact on grassland plant diversity depends on habitat productivity and herbivore size. Ecology letters 9: 780–788. DOI: 10.1111/j.1461-0248.2006.00925.xCrossRefGoogle Scholar
  3. Blackhall M, Raffaele E, Veblen TT (2008) Cattle affect early post-fire regeneration in a Nothofagus dombeyiAustrocedrus chilensis mixed forest in Northern Patagonia, Argentina. Biological Conservation 141: 2251–2261. DOI:10.1016/j.biocon.2008.06.016CrossRefGoogle Scholar
  4. Bray RM, Kurtz LT (1945) Determination of total, organic and available forms of phosphorus in soils. Soil Science 59: 39–45.CrossRefGoogle Scholar
  5. Brosofske KD, Chen J, Crow TR (2001) Understory vegetation and site factors: implications for a managed Wisconsin landscape. Forest Ecology and Management 146: 75–87. DOI: 10.1016/S0378-1127(00)00447-3CrossRefGoogle Scholar
  6. Canham CD, Burbank DH (1994) Causes and consequences of resource heterogeneity in forests: interspecific variation in light transmission by canopy trees. Canadian Journal of Forest Research 24: 337–349. DOI: 10.1139/x94-046CrossRefGoogle Scholar
  7. Chen Y, Li Y, Zhao X, et al. (2012) Effects of grazing exclusion on soil properties and on ecosystem carbon and nitrogen storage in a sandy rangeland of Inner Mongolia, Northern China. Environmental Management 50: 622–632. DOI: 10.1007/s00267-012-9919-1CrossRefGoogle Scholar
  8. Connell JH (1978) Diversity in tropical rain forests and coral reefs. Science 199: 1302–1310.CrossRefGoogle Scholar
  9. Darabant A, Rai PB, Gratzer G (2005) Interactive effects of biotic and abiotic factors on Tsuga dumosa seedling establishment in silvicultural group gaps. University of Natural Resources and Applied Life Sciences, Department of Forest and Soil Sciences, Institute of Forest Ecology, Peter Jordan Strasse 82, A-1190 Wien, Austria.Google Scholar
  10. Darabant A, Rai PB, Tenzin K, et al. (2007) Cattle grazing facilitates tree regeneration in a conifer forest with palatable bamboo understory. Forest Ecology and Management 252: 73–83. DOI: 10.1016/j.foreco.2007.06.018CrossRefGoogle Scholar
  11. Dorji S (2004) Natural regeneration at cable crane logged sites in the mixed conifer belt of Gidakom Forest Management Unit, Thimphu, Bhutan. MSc Dissertation, University of Natural Resources and Life Sciences, Vienna, Austria.Google Scholar
  12. Everson CS, Clarke GPY (1987) A comparison of six methods of botanical analysis in the montane grasslands of NATAL. Plant Ecology 73: 47–51. DOI: 10.1007/BF00031850CrossRefGoogle Scholar
  13. Foster BL, Gross KL (1998) Species richness in successional grassland: Effects of nitrogen enrichment and plant litter. Ecology 79: 2593–2602. DOI: 10.1890/0012-9658(1998)079 [2593:SRIASG]2.0.CO;2CrossRefGoogle Scholar
  14. Gilliam FS (2007) The ecological significance of the herbaceous layer in temperate forest ecosystems. BioScience 57: 845–858. DOI: 10.1641/B571007CrossRefGoogle Scholar
  15. Gratzer G, Rai PB, Glatzel G (1999) The influence of the bamboo Yushania microphylla on regeneration of Abies densa in central Bhutan. Canadian Journal of Forest Research 29: 1518–1527. DOI: 10.1139/x99-125CrossRefGoogle Scholar
  16. Gray AN, Spies TA (1996) Gap size, within gap position and canopy structure effects on conifer seedling establishment. Journal of Ecology 84: 635–645. DOI: 10.2307/2261327CrossRefGoogle Scholar
  17. Grime JP (1973) Competitive exclusion in herbaceous vegetation. Nature 242: 344–347.CrossRefGoogle Scholar
  18. Guebel DV, Nudel BC, Giulietti AM (1991) A simple and rapid micro-Kjeldahl method for total nitrogen analysis. Biotechnology Techniques 5: 427–430.CrossRefGoogle Scholar
  19. Harrison M, Evans JR, Moore A (2008) Grazing induced changes in light interception and radiation-use efficiency of winter wheat. Proceedings of 14th Agronomy Conference 2008, 21–25 September 2008, Adelaide, South Australia.Google Scholar
  20. Haynes RJ, Williams PH (1993) Nutrient cycling and soil fertility in the grazed pasture ecosystem. Advances in Agronomy 49: 119–199. DOI:10.1016/S0065-2113(08)60794-4CrossRefGoogle Scholar
  21. IUSS GW (2006) World reference base for soil resources 2006. World Soil Resources Report No. 103, FAO, Rome, Italy.Google Scholar
  22. Jewell PL, Käuferle D, Güsewell S, et al. (2007) Redistribution of phosphorus by cattle on a traditional mountain pasture in the Alps. Agriculture Ecosystems and Environment 122: 377–386. DOI: 10.1016/j.agee.2007.02.012CrossRefGoogle Scholar
  23. Kneeshaw DD, Bergeron Y (1998) Canopy gap characteristics and tree replacement in the southeastern boreal forest. Ecology 79: 783–794. DOI: 10.1890/0012-9658(1998)079[0783:CGCATR]2.0.CO;2CrossRefGoogle Scholar
  24. Kreutzweiser, David P, Hazlett, et al. (2008) Logging impacts on the biogeochemistry of boreal forest soils and nutrient export to aquatic systems: A review. Environmental Review 16: 157–179. DOI: 10.1139/A08-006CrossRefGoogle Scholar
  25. Kuijper DPJ (2011) Lack of natural control mechanisms increases wildlife forestry conflict in managed temperate European forest systems. European Journal of Forest Research 130: 895–909. DOI: 10.1007/s10342-011-0523-3CrossRefGoogle Scholar
  26. Kyriazopoulos AP, Sklavou P, Nastis AS, Papanastasis VP (2009) Interactions between grazing behavior and plant community structure in shrubland and their consequences on desertification. Options Mediterraneennes A/no. 85.Google Scholar
  27. Landau S, Everitt BS (2004) A handbook of statistical analysis using SPSS. Chapman and Hall/CRC, CRC Press Company.Google Scholar
  28. Lavorel S, McIntyre S, Landsberg J, Forbes TDA (1997) Plant functional classifications: from general groups to specific groups based on response to disturbance. Trends in Ecology and Evolution 12: 474–478. DOI: 10.1016/S0169-5347(97)01219-6CrossRefGoogle Scholar
  29. Margalef R (1958) Temporal succession and spatial heterogeneity in phytoplankton. In: Buzzati Traverso (ed), Perspectives in marine biology. pp 323–347.Google Scholar
  30. Marini L, Scotton M, Klimek S, et al. (2007) Effects of local factors on plant species richness and composition of alpine meadows. Agriculture Ecosystems and Environment 119: 281–288. DOI: 10.1016/j.agee.2006.07.015CrossRefGoogle Scholar
  31. McEvoy PM, Flexen M, McAdam JH (2006) The effects of livestock grazing on ground flora in broadleaf woodlands in Northern Ireland. Forest Ecology and Management 225: 39–50. DOI: 10.1016/j.foreco.2005.12.026CrossRefGoogle Scholar
  32. McLean EO (1982). Soil pH and lime requirement. In: Page, A. L., Miller, R. H., Keeney, D. R. (Eds.), Methods of soil analysis Part 2 — Chemical and microbiological properties (2nd Ed.), Agronomy 9: 199–223.Google Scholar
  33. McNaughton SJ (1993) Grasses and grazers, science and management. Ecological Applications 3: 17–20. DOI: 10.2307/1941782CrossRefGoogle Scholar
  34. Medina-Roldán E, PazFerreiro J, Bardgett RD (2012) Grazing exclusion affects soil and plant communities, but has no impact on soil carbon storage in an upland grassland. Agriculture Ecosystems and Environment 149: 118–123. DOI: 10.1016/j.agee.2011.12.012CrossRefGoogle Scholar
  35. Merino A, Edeso JM, González MJ, Marauri P (1998) Soil properties in a hilly area following different logging management practices. Forest Ecology and Management 103: 235–246. DOI: PI1 SO378-1 127(97)00229-6CrossRefGoogle Scholar
  36. Milchunas DG, Lauenroth WK (1993) Quantitative effects of grazing on vegetation and soils over a global range of environments. Ecological Monograph 63: 327–366. DOI: 10.2307/2937150CrossRefGoogle Scholar
  37. Modrý M, Hubený D, Rejšek K (2004) Differential response of naturally regenerated European shade tolerant tree species to soil type and light availability. Forest Ecology and Management 188:185–195. DOI: 10.1016/j.foreco.2003.07.029CrossRefGoogle Scholar
  38. Moe SR, Wegge P (2008) Effects of deposition of deer dung on nutrient redistribution and on soil and plant nutrients on intensively grazed grasslands in lowland Nepal. Ecological Research 23: 227–234. DOI: 10.1007/s11284-007-0367-yCrossRefGoogle Scholar
  39. Moktan MR, Gratzer G, Richards WH, et al. (2009) Regeneration of mixed conifer forests under group tree selection harvest management in western Bhutan Himalayas. Forest Ecology and Management 257: 2121–2132. DOI: 10.1016/j.foreco.2009.02.022CrossRefGoogle Scholar
  40. Moore MR, Vankat JL (1986) Responses of the herb layer to the gap dynamics of a mature beech-maple forest. American Midland Naturalist 115: 336–347.CrossRefGoogle Scholar
  41. Norbu L (2002) Grazing management in broadleaf forests. Journal of Bhutan Studies 7: 99–129.Google Scholar
  42. Nuttle T, Royo AA, Adams MB, Carson WP (2013) Historic disturbance regimes promote tree diversity only under low browsing regimes in eastern deciduous forest. Ecological Monographs 83: 3–17. DOI: 10.1890/11-2263.1CrossRefGoogle Scholar
  43. Ohsawa M, Wangda P, Kitazawa T, et al. (2002) Secondary succession and soil development in tseri-farming system, Shemgang, Southern Bhutan. In: Ohsawa, M. (Ed.), Life Zone Ecology of the Bhutan Himalaya III. pp 125–143.Google Scholar
  44. Palviainen M, Finér L, Laiho R, et al. (2010) Carbon and nitrogen release from decomposing Scots pine, Norway spruce and silver birch stumps. Forest Ecology and Management 259: 390–398. DOI: 10.1016/j.foreco.2009.10.034CrossRefGoogle Scholar
  45. Pedreira CGS, Sollenberger LE, Mislevy P (2000) Botanical composition, light interception, and carbohydrate reserve status of grazed ‘Florakirk’ bermudagrass. Agronomy Journal 92: 194–199. DOI: 10.2134/agronj2000.922194xCrossRefGoogle Scholar
  46. Phuntsho Y (2012) Gidakom forest management unit management plan 2012–2022. Forest Resources and Management Division, Department of Forests and Park Services, Ministry of Agriculture, Thimphu, Bhutan.Google Scholar
  47. Prober SM, Standish RJ, Wiehl G (2011) After the fence: Vegetation and topsoil condition in grazed, fenced and benchmark eucalypt woodlands of fragmented agricultural landscapes. Australian Journal of Botany 59: 369–381. DOI: 0.1071/BT11026Google Scholar
  48. Proulx M, Mazumder A (1998) Reversal of grazing impact on plant species richness in nutrient poor vs. nutrient — rich ecosystems. Ecology 79: 2581–2592. DOI: 10.1890/0012-9658(1998)079[2581:ROGIOP]2.0.CO;2Google Scholar
  49. Pyakälä J (2003) Effects of restoration with cattle grazing on plant species composition and richness of semi natural grasslands. Biodiversity and Conservation 12: 2211–2226. DOI: 10.1023/A:1024558617080CrossRefGoogle Scholar
  50. Rich CI (1965) Elemental analysis by flame emission photometry. In: Black CA, Evans DD, White J (eds.), Methods of Soil Analysis, Madison, Wisconsin, American Society of Agronomy. pp 849–864.Google Scholar
  51. Royo AA, Collins R, Adams MB, et al. (2010) Pervasive interactions between ungulate browsers and disturbance regimes promote temperate forest herbaceous diversity. Ecology 91: 93–105. DOI: 10.1890/08-1680.1CrossRefGoogle Scholar
  52. Schultz NL, Morgan JW, Lunt ID (2011) Effects of grazing exclusion on plant species richness and phytomass accumulation vary across a regional productivity gradient. Journal of Vegetation Science 22: 130–142. DOI: 10.1111/j.1654-1103.2010.01235.xCrossRefGoogle Scholar
  53. Shannon CE, Weaver W (1949) The mathematical theory of communication. Urbana, IL: University of Illinois Press. pp 177.Google Scholar
  54. Slesak RA, Schoenholtzb SH, Harringtonc TB (2011) Soil carbon and nutrient pools in Douglas fir plantations 5 years after manipulating biomass and competing vegetation in the Pacific Northwest. Forest Ecology and Management 262: 1722–1728. DOI: 10.1016/j.foreco.2011.07.021CrossRefGoogle Scholar
  55. Sparks TH, Greatorex-Davies JN, Mountford JO, et al. (1996) The effects of shade on the plant communities of rides in plantation woodland and implications for butterfly conservation. Forest Ecology and Management 80: 197–207. DOI: SSDI 0378-1127(95)03639-3CrossRefGoogle Scholar
  56. Tasker EM, Bradstock RA (2006) Influence of cattle grazing practices on forest understory structure in north-eastern New South Wales. Austral Ecology 31: 490–502. DOI: 10.1111/j.1442-9993.2006.01597.xCrossRefGoogle Scholar
  57. Tenzin K (2010) Regeneration and gap dynamics in mixed conifer forests of the Bhutan Himalayas. PhD Dissertation, University of Natural Resources and Life Sciences, Vienna, Austria.Google Scholar
  58. Tessema ZK, de Boer WF, Baars RMT, Prins HHT (2011) Changes in soil nutrients, vegetation structure and herbaceous biomass in response to grazing in a semi-arid savanna of Ethiopia. Journal of Arid Environment 75: 662–670. DOI: 10.1016/j.jaridenv.2011.02.004CrossRefGoogle Scholar
  59. Thomas SC, Halpern CB, Falk DA, et al. (1999) Plant diversity in managed forests: Understory responses to thinning and fertilization. Ecological Applications 9: 864–879. DOI: 10.1890/1051-0761(1999)009[0864:PDIMFU]2.0.CO;2CrossRefGoogle Scholar
  60. Tilman D (1982) Resource competition and community structure. Princeton University Press, Princeton, NJ.Google Scholar
  61. Tilman D, Wedin D, Knops JMH (1996) Productivity and sustainability influenced by biodiversity in grassland ecosystems. Nature 379: 718–720. DOI: 10.1038/379718a0CrossRefGoogle Scholar
  62. Tshering P (2005) The interaction of grazing and competition in the conifer forests of Bhutan. MSc Dissertation, University of Natural Resources and Life Sciences, Vienna, Austria.Google Scholar
  63. Walker BH (1970) An evaluation of eight methods of botanical analysis on grasslands in Rhodesia. Journal of Applied Ecology 7: 403–416. DOI: 10.2307/2401967CrossRefGoogle Scholar
  64. Wangchuk K (2007) Natural regeneration ecology of mixed conifer forests in Western Bhutan. MSc Dissertation, University of Natural Resources and Life Sciences, Vienna, Austria.Google Scholar
  65. Wangchuk K, Wurzinger M, Darabant A, et al. (2014) The changing face of cattle raising and forest grazing in the Bhutan Himalaya. Mountain Research and Development 34: 131–138. DOI: 10.1659/MRD-Journal-D-13-00021.1CrossRefGoogle Scholar
  66. Whigham DF (2004) Ecology of woodland herbs in temperate deciduous forests. Annual Review of Ecology and Evolution System 35: 583–621. DOI: 10.1146/annurev.ecolsys.35.021103.105708CrossRefGoogle Scholar
  67. Worm B, Lotze HK, Hillebrand H, Sommer U (2002) Consumer versus resource control of species diversity and ecosystem functioning. Nature 417:848–851. DOI: 10.1038/nature00830CrossRefGoogle Scholar
  68. Wright JP, Flecker AS, Jones CG (2003) Local vs. landscape controls on plant species richness in beaver meadows. Ecology 84: 3162–3173. DOI: 10.1890/02-0598CrossRefGoogle Scholar
  69. Yokoyama S, Shibata EI (1998) The effects of sika deer browsing on the biomass and morphology of a dwarf bamboo, Sasa nipponica, in Mt. Ohdaigahara, central Japan. Forest Ecology and Management 103: 49–56. DOI:PII S0378-1127 97 00175-8CrossRefGoogle Scholar

Copyright information

© Science Press, Institute of Mountain Hazards and Environment, CAS and Springer-Verlag Berlin Heidelberg 2014

Authors and Affiliations

  • Kesang Wangchuk
    • 1
    • 2
    Email author
  • András Darabant
    • 2
    • 3
  • Prem Bahadur Rai
    • 2
  • Maria Wurzinger
    • 1
  • Werner Zollitsch
    • 1
  • Georg Gratzer
    • 3
  1. 1.Department of Sustainable Agricultural Systems, Division of Livestock SciencesUniversity of Natural Resources and Life SciencesViennaAustria
  2. 2.Renewable Natural Resources Research and Development CenterBumthangBhutan
  3. 3.Department of Forest and Soil Sciences, Institute of Forestry EcologyUniversity of Natural Resources and Life SciencesViennaAustria

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