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Abiotic and biotic drivers of species diversity in understory layers of cold temperate coniferous forests in North China

  • Hui Wang
  • Mengtao Zhang
  • Hongwei Nan
Original Paper
  • 14 Downloads

Abstract

Understory plants are important components of forest ecosystems and play a crucial role in regulating community structures, function realization, and community succession. However, little is known about how abiotic and biotic drivers affect the diversity of understory species in cold temperate coniferous forests in the semiarid climate region of North China. We hypothesized that (1) topographic factors are important environmental factors affecting the distribution and variation of understory strata, and (2) different understory strata respond differently to environmental factors; shrubs may be significantly affected by the overstory stratum, and herbs may be more affected by surface soil conditions. To test these hypotheses, we used the boosted regression tree method to analyze abiotic and biotic environmental factors that influence understory species diversity, using data from 280 subplots across 56 sites in cold temperate coniferous forests of North China. Elevation and slope aspect were the dominant and indirect abiotic drivers affecting understory species diversity, and individual tree size inequality (DBH variation) was the dominant biotic driver of understory species diversity; soil water content was the main edaphic factors affecting herb layers. Elevation, slope aspect, and DBH variation accounted for 36.4, 14.5, and 12.1%, respectively, of shrub stratum diversity. Shrub diversity decreased with elevation within the range of altitude of this study, but increased with DBH variation; shrub diversity was highest on north-oriented slopes. The strongest factor affecting herb stratum species diversity was slope aspect, accounting for 25.9% of the diversity, followed by elevation (15.7%), slope (12.2%), and soil water content (10.3%). The highest herb diversity was found on southeast-oriented slopes and the lowest on northeast-oriented slopes; herb diversity decreased with elevation and soil water content, but increased with slope. The results of the study provide a reference for scientific management and biodiversity protection in cold temperate coniferous forests of North China.

Keywords

Abiotic and biotic drivers Cold temperate coniferous forests North China Semi-arid region Understory species diversity 

Notes

Acknowledgements

We thank all our colleagues and students in our lab for constructive discussion and technical support. We also thank Arshad Ali and anonymous reviewers for helpful comments on an earlier draft of this paper.

Supplementary material

11676_2018_795_MOESM1_ESM.docx (27 kb)
Supplementary material 1 (DOCX 26 kb)

References

  1. Alaback PB, Herman FR (1988) Long-term response of understory vegetation to stand density in Picea-Tsuga forests. Can J For Res 18(12):1522–1530Google Scholar
  2. Alatalo RV (1981) Problems in the measurement of evenness in ecology. Oikos 37(2):199–204Google Scholar
  3. Ali A, Yan ER (2017a) The forest strata-dependent relationship between biodiversity and aboveground biomass within a subtropical forest. For Ecol Manage 401:125–134Google Scholar
  4. Ali A, Yan ER (2017b) Functional identity of overstorey tree height and understorey conservative traits drive aboveground biomass in a subtropical forest. Ecol Ind 83:158–168Google Scholar
  5. Ali A, Yan ER (2018) The mediation roles of intraspecific and interspecific functional trait diversity for linking the response of aboveground biomass to species richness across forest strata in a subtropical forest. Ecol Ind 85:493–501Google Scholar
  6. Ali A, Lohbeck M, Yan ER (2018) Forest strata-dependent functional evenness explains whole-community aboveground biomass through opposing mechanisms. For Ecol Manage 424:439–447Google Scholar
  7. Antos JA (2009) Understory plants in temperate forests. For For Plants I:262–279Google Scholar
  8. Aschehoug ET, Callaway RM (2014) Morphological variability in tree root architecture indirectly affects coexistence among competitors in the understory. Ecology 95(7):1731–1736PubMedGoogle Scholar
  9. Augusto L, Dupouey JL, Ranger J (2003) Effects of tree species on understory vegetation and environmental conditions in temperate forests. Ann For Sci 60(8):823–831Google Scholar
  10. Barbier S, Gosselin F, Balandier P (2008) Influence of tree species on understory vegetation diversity and mechanisms involved: a critical review for temperate and boreal forests. For Ecol Manage 254(1):1–15Google Scholar
  11. Bartels SF, Chen HYH (2013) Interactions between overstorey and understorey vegetation along an overstorey compositional gradient. J Veg Sci 24(3):543–552Google Scholar
  12. Beatty SW, Scholes ODV (1988) Leaf litter effect on plant species composition of deciduous forest treefall pits. Can J For Res 18:553–559Google Scholar
  13. Bennie J, Huntley B, Wiltshire A, Hill MO, Baxter R (2008) Slope, aspect and climate: spatially explicit and implicit models of topographic microclimate in chalk grassland. Ecol Model 216:47–59Google Scholar
  14. Brosofske KD, Chen JQ, Crow TR (2001) Understory vegetation and site factors: implications for a managed Wisconsin landscape. For Ecol Manage 146:75–87Google Scholar
  15. Cai WH, Yang J, Liu ZH, Hu YM, Liu SJ, Jing GZ, Zhao ZF (2012) Controls of post-fire tree recruitment in Great Xing’an Mountains in Heilongjiang Province. Acta Ecol Sin 32(11):3303–3312Google Scholar
  16. Cantón Y, Barrio GD, Solé-Benet A, Lázaro R (2004) Topographic controls on the spatial distribution of ground cover in the Tabernas badlands of SE Spain. CATENA 55(3):341–365Google Scholar
  17. Cavard X, Macdonald SE, Bergeron Y, Chen HYH (2011) Importance of mixedwoods for biodiversity conservation: evidence for understory plants, songbirds, soil fauna, and ectomycorrhizae in northern forests. Doss Environ 19(1):142–161Google Scholar
  18. Chastain RA Jr, Currie WS, Townsend PA (2006) Carbon sequestration and nutrient cycling implications of the evergreen understory layer in Appalachian forests. For Ecol Manage 231(1–3):63–77Google Scholar
  19. Chávez V, Macdonald SE (2010) The influence of canopy patch mosaics on understory plant community composition in boreal mixedwood forest. For Ecol Manage 259(6):1067–1075Google Scholar
  20. Chen H, LégaréS BY (2004) Variation of the understory composition and diversity along a gradient of productivity in Populus tremuloides stands of northern British Columbia. Can J Bot 82(9):1314–1323Google Scholar
  21. Chipman SJ, Johnson EA (2002) Understory vascular plant species diversity in the mixedwood boreal forest of Western Canada. Ecol Appl 12(2):588–601Google Scholar
  22. Chopra K, Kumar P (2004) Forest biodiversity and timber extraction: an analysis of the interaction of market and non-market mechanisms. Ecol Econ 49:135–148Google Scholar
  23. Elith J, Leathwick JR, Hastie T (2008) A working guide to boosted regression trees. J Anim Ecol 77(4):802–813PubMedGoogle Scholar
  24. Ewald J (2000) The influence of coniferous canopies on understorey vegetation and soils in mountain forests of the northern Calcareous. Appl Veg Sci 3:123–134Google Scholar
  25. Gamfeldt L, Snäll T, Bagchi R, Jonsson M, Gustafsson L, Kjellander P, Ruiz-Jaen MC, Fröberg M, Stendahl J, Philipson CD, Mikusiński G, Andersson E, Westerlund B, Andrén H, Moberg F, Moen J, Bengtsson J (2013) Higher levels of multiple ecosystem services are found in forests with more tree species. Nat Commun 4:1340PubMedPubMedCentralGoogle Scholar
  26. Gao JF, Zhang YX (2006) Distributional patterns of species diversity of main plant communities along altitudinal gradient in secondary forest region, Guandi Mountain, China. J For Res 17(2):111–115Google Scholar
  27. Gong X, Brueck H, Giese KM, Zhang L, Sattelmacher B, Lin S (2008) Slope aspect has effects on productivity and species composition of hilly grassland in the Xilin River Basin, Inner Mongolia, China. J Arid Environ 72:483–493Google Scholar
  28. Härdtle W, Gv O, Westphal C (2003) The effects of light and soil conditions on the species richness of the ground vegetation of deciduous forests in northern Germany (Schleswig-Holstein). For Ecol Manage 182(1–3):327–338Google Scholar
  29. Hart AS, Chen YH (2006) Understory vegetation dynamics of north American boreal forests. Crit Rev Plant Sci 25(4):381–397Google Scholar
  30. He MZ, Zheng JG, Li XR, Qian YL (2007) Environmental factors affecting vegetation composition in the Alxa Plateau, China. J Arid Environ 69(3):473–489Google Scholar
  31. Holste EK, Kobe RK, Vriesendorp CF (2011) Seedling growth responses to soil resources in the understory of a wet tropical forest. Ecology 92(9):1828–1838PubMedGoogle Scholar
  32. Huang LJ, Zhu WX, Ren H, Chen HF, Wang J (2012) Impact of atmospheric nitrogen deposition on soil properties and herb-layer diversity in remnant forests along an urban–rural gradient in Guangzhou, southern China. Plant Ecol 213(7):1187–1202Google Scholar
  33. Huo H, Feng Q, Su YH (2014) The influences of canopy species and topographic variables on understory species diversity and composition in coniferous forests. Sci World J 2:252489Google Scholar
  34. Kargar-Chigan H, Javadi SA, Zahedi-Amiri G, Khajeddin SJ, Jafari M (2017) Vegetation composition differentiation and species-environment relationships in the northern part of Isfahan Province, Iran. J Arid Land 9(2):161–175Google Scholar
  35. Koellner T, Hersperger AM, Wohlgemuth T (2004) Rarefaction method for assessing plant species diversity on a regional scale. Ecography 27:532–544Google Scholar
  36. Körner C (2007) The use of “altitude” in ecological research. Trends Ecol Evol 22(11):569–574PubMedGoogle Scholar
  37. Lafrankie JV, Ashton PS, Chuyong GB, Co L, Condit R, Davies SJ, Foster R, Hubbell Stepthen P, Kenfack D, Lagunzad D, Losos EC, Nor NS, Tan S, Thomas TW, Valencia R, Villa G (2006) Contrasting structure and composition of the understory in species-rich tropical rain forests. Ecology 87(9):2298–2305PubMedGoogle Scholar
  38. Légaré S, Bergeron Y, Leduc A, Paré D (2001) Comparison of the understory vegetation in boreal forest types of southwest Quebec. Can J Bot 79(9):1019–1027Google Scholar
  39. Li LP, Wang XP, Zerbe S, Zhang LY, Fang JY (2011) Altitudinal patterns of stand structure and herb layer diversity of Picea schrenkiana forests in the central Tianshan Mountains, Northwest China. J Arid Land 3(4):254–260Google Scholar
  40. Lü XT, Yin JX, Tang JW (2011) Diversity and composition of understory vegetation in the tropical seasonal rain forest of Xishuangbanna, SW China. Rev Biol Trop 59(1):1–9Google Scholar
  41. MacDonald SE, Fenniak TE (2007) Understory plant communities of boreal mixedwood forests in western Canada: natural patterns and response to variable–retention harvesting. For Ecol Manage 242(1):34–48Google Scholar
  42. Maltez-Mouro S, García LV, Marañón T, Freitas H (2005) Understory floristic heterogeneity within a Mediterranean oak forest: How much is explained by edaphic, spatial and overstory components? Rev Biol (Lisboa) 23(3):121–133Google Scholar
  43. Matsushima M, Choi WJ, Chang SX (2012) White spruce foliar δ13C and δ15 N indicate changed soil N availability by understory removal and N fertilization in a 13-year-old boreal plantation. Plant Soil 361(1–2):375–384Google Scholar
  44. Meng XN, Zhao YS, Zheng L, Xin Y (2012) Population structure and understory species diversity of different aged Pinus sylvestris var. mongolica plantations in Nenjiang Sandy Land of Northeast China. Chin J Appl Ecol 23(9):2332–2338Google Scholar
  45. Michalet R, Rolland Ch, Joud D, Gafta D, Callaway RM (2002) Associations between canopy and understory species increase along a rainshadow gradient in the Alps: habitat heterogeneity or facilitation? Plant Ecol 165:145–160Google Scholar
  46. Moeslund JE, Arge L, Bøcher PK, Dalgaard T, Svenning JC (2013) Topography as a driver of local terrestrial vascular plant diversity patterns. Nord J Bot 31:129–144Google Scholar
  47. Mölder A, Bernhardt-Römermann M, Schmidt W (2008) Herb-layer diversity in deciduous forests: raised by tree richness or beaten by beech? For Ecol Manage 256(3):272–281Google Scholar
  48. Negi SP (2011) Diversity patterns of understory woody species in the forests of Belasitsa mountain. University of Twente, BulgariaGoogle Scholar
  49. Nilsson MC, Wardle DA (2005) Understory vegetation as a forest ecosystem driver: evidence from the northern Swedish boreal forest. Front Ecol Environ 3(8):421–428Google Scholar
  50. North M, Oakley B, Fiegener R, Gray A, Barbour M (2005) Influence of light and soil moisture on Sierran mixed-conifer understory communities. Plant Ecol 177:13–24Google Scholar
  51. Økland RH (2000) Population biology of the clonal moss Hylocomium splendens in Norwegian boreal spruce forests. 5. Vertical dynamics of individual shoot segments. Oikos 88(3):449–469Google Scholar
  52. Olivero AM, Hix DM (1998) Influence of aspect and stand age on ground flora of southeastern Ohio forest ecosystems. Plant Ecol 39:177–187Google Scholar
  53. Ou QX, Li HK, Yang Y (2017) Factors affecting the biomass conversion and expansion factor of masson pine in Fujian Province. Acta Ecol Sin 37(17):5756–5764Google Scholar
  54. Pachepsky YA, Timlin DJ, Rawls WJ (2001) Soil water retention as related to topographic variables. Soil Sci Soc Am J 65:1787–1795Google Scholar
  55. Pitman NCA, Terborgh JW, Silman MR, Percy NV, Neill DA, Cerón CE, Palacios WA, Aulestia M (2002) A comparison of tree species diversity in two upper Amazonian forests. Ecology 83(11):3210–3224Google Scholar
  56. Prescott CE, Zabek LM, Staley CL, Kabzems R (2000) Decomposition of broadleaf and needle litter in forests of British Columbia: influences of litter type, forest type, and litter mixtures. Can J For Res 30(11):1742–1750Google Scholar
  57. Puumalainen J (2001) Structural, compositional and functional aspects of forest biodiversity in Europe. Joint FAO/UNECE and JRC paper. Geneva forest and timber discussion papers, vol 22, p 87Google Scholar
  58. Qian H, Klinka K, Sivak B (1997) Diversity of the understory vascular vegetation in 40 year-old and old-growth forest stands on Vancouver Island, British Columbia. J Veg Sci 8:773–780Google Scholar
  59. Qian H, Klinka K, Økland RH, Krestov P, Kayahara GJ (2003) Understorey vegetation in boreal Picea mariana and Populus tremuloides stands in British Columbia. J Veg Sci 14(2):173–184Google Scholar
  60. Reich PB, Bakken P, Carlson D, Frelich LF, Friedman SK, Grigal DF (2001) Influence of logging, fire, and forest type on biodiversity and productivity in southern boreal forests. Ecology 82:2731–2748Google Scholar
  61. Ren XM, Yang GH, Wang DX, Qin XW, Liu ZX, Zhao SX, Bai Y (2012) Effects of environmental factors on species distribution and diversity in an Abies fargesii-Betula utilis mixed forest. Acta Ecol Sin 32(2):605–613Google Scholar
  62. Riegel GM, Miller RF, Krueger WC (1992) Competition for resources between understory vegetation and overstory Pinus ponderosa in northern Oregon. Ecol Appl 2(1):71–85PubMedGoogle Scholar
  63. Samuel MJ, Hart RH (1998) Nitrogen fertilization, botanical composition and biomass production on mixed-grass rangeland. J Range Manag 51(4):408–416Google Scholar
  64. Sanaei A, Chahouki MAZ, Ali A, Jafari M, Azarnivand H (2018) Abiotic and biotic drivers of aboveground biomass in semi-steppe rangelands. Sci Total Environ 615:895–905PubMedGoogle Scholar
  65. Sanjerehei MM, Jafari M, Mataji A, Meybodi NB, Bihamta MR (2013) Influence of environmental factors on distribution of plant species in Nodushan rangelands of Yazd province (Iran). Desert 18:19–26Google Scholar
  66. Schoonover JE, Crim JF (2015) An introduction to soil concepts and the role of soils in watershed management. Journal of Contemporary Water Research and Education 154:21–47Google Scholar
  67. Small CJ, McCarthy BC (2005) Relationship of understory diversity to soil nitrogen, topographic variation, and stand age in an eastern oak forest, USA. Fort Ecol Manag 217(2):229–243Google Scholar
  68. Tchouto MGP, Boer WFD, Wilde JJFED, Maesen LJGVD (2006) Diversity patterns in the flora of the Campo–Ma’an rain forest, Cameroon: do tree species tell it all? Biodivers Conserv 15(4):1353–1374Google Scholar
  69. Tessema ZK, Belay EF (2017) Effect of tree species on understory vegetation, herbaceous biomass and soil nutrients in a semi-arid savanna of Ethiopia. J Arid Environ 139:76–84Google Scholar
  70. Thysell DR, Carey BA (2001) Manipulation of density of Pseudotsuga menziesii canopies: preliminary effects on understory vegetation. Can J For Res 31(9):1513–1525Google Scholar
  71. Valladares F, Ülo N (2008) Shade tolerance, a key plant feature of complex nature and consequences. Ann Rev Ecol Evol Syst 39:237–257Google Scholar
  72. Varga P, Chen HYH, Klinka K (2005) Tree-size diversity between single-and mixedspecies stands in three forest types in western Canada. Can J For Res 35(3):593–601Google Scholar
  73. Wang CK (2006) Biomass allometric equations for 10 co-occurring tree species in Chinese temperate forests. For Ecol Manage 222(1):9–16Google Scholar
  74. Wang SX, Wang XA, Li GQ, Guo H, Zhu ZH (2010) Species diversity and environmental interpretation in the process of community succession in the Ziwu Mountain of Shaanxi Province. Acta Ecol Sin 30(6):1638–1647Google Scholar
  75. Warren RJ II (2008) Mechanisms driving understory evergreen herb distributions across slope aspects: as derived from landscape position. Plant Ecol 198:297–308Google Scholar
  76. Wayman RB, North M (2007) Initial response of mixed-conifer understory plant community to burning and thinning restoration treatments. For Ecol Manage 239:32–44Google Scholar
  77. Weisberg PJ, Shandra O, Becker ME (2013) Landscape influences on recent timberline shifts in the Carpathian Mountains: abiotic influences modulate effects of land-use change. Arct Antarct Alp Res 45(3):404–414Google Scholar
  78. Willinghöfer S, Cicuzza D, Kessler M (2012) Elevational diversity of terrestrial rainforest herbs: when the whole is less than the sum of its parts. Plant Ecol 213(3):407–418Google Scholar
  79. Wu XJ, Chang JG, Yu JX, An Y, Guo JR (2010) Spatial Distribution of Larix principis-rupprech tii-Picea spp. Secondary Forests Along Elevation Gradients on North Facing Slope of Luya Mountain, Shanxi. J Northeast For Univ 38(1):10–14Google Scholar
  80. Wu YC, Li ZC, Cheng CF, Liu RJ (2014) Responses of soil organic carbon to long-term understory removal in subtropical Cinnamomum camphora Stands. Int J Ecol 4:1–6Google Scholar
  81. Xu MH, Ma L, Jia YY, Liu M (2017) Integrating the effects of latitude and altitude on the spatial differentiation of plant community diversity in a mountainous ecosystem in China. PLoS ONE 12(3):e0174231PubMedPubMedCentralGoogle Scholar
  82. Yan HB, Han YZ, Yang XQ, Wang LY, Xiang XY (2010) Spatial distribution patterns and associations of tree species in typical natural secondary mountain forest communities of Northern China. Acta Ecol Sin 30(9):2311–2321Google Scholar
  83. Yang FW, Yu XX, Wang SS, Zhang XM, Wu SH (2007) Interspecific association and niche of forest vegetation in rocky mountain area of North of China. Sci Soil Water Conserv 5(1):60–67Google Scholar
  84. You YM, Xu JY, Cai DX, Liu SR, Zhu HG, Wen YG (2016) Environmental factors affecting plant species diversity of understory plant communities in a Castanopsis hystrix plantation chronosequence in Pingxiang, Guangxi, China. Acta Ecol Sin 36(1):164–172Google Scholar
  85. Yu M, Sun OJ (2013) Effects of forest patch type and site on herb-layer vegetation in a temperate forest ecosystem. For Ecol Manage 300(Supplement 1):14–20Google Scholar
  86. Yuan Z, Ali A, Wang S, Gazol A, Freckleton R, Wang XG, Lin F, Ye J, Zhou L, Hao ZQ, Loreau M (2018) Abiotic and biotic determinants of coarse woody productivity in temperate mixed forests. Sci Total Environ 630:422–431PubMedGoogle Scholar
  87. Zhang JT, Zhang F (2007) Diversity and composition of plant functional groups in mountain forests of the lishan Nature Reserve, North China. Bot Stud 48(3):339–348Google Scholar
  88. Zhang F, Shangguan TL, Anders T (1997) The vertical belts of natural vegetaion partitioning of the Guandi Mountains by using ordered plot clustering, Shanxi, North China. Bull Bot Res 17(4):447–454Google Scholar
  89. Zhang QD, Bi RC, Zhang JT, Nyobe L, Miao YM, Liu XN (2012a) Dynamic analysis on spatial pattern of dominant tree species of cold-temperate coniferous forest in the succession process in the Pangquangou Nature Reserve. Acta Ecol Sin 32(18):5713–5720Google Scholar
  90. Zhang Y, Chen HYH, Reich BP (2012b) Forest productivity increases with evenness, species richness and trait variation: a global meta-analysis. J Ecol 100:742–749Google Scholar
  91. Zhang Y, Chen HYH, Taylor A (2014) Multiple drivers of plant diversity in forest ecosystems. Glob Ecol Biogeogr 23(8):885–893Google Scholar
  92. Zhang Y, Chen HYH, Taylor A (2016a) Aboveground biomass of understorey vegetation has a negligible or negative association with overstorey tree species diversity in natural forests. Glob Ecol Biogeogr 25(2):141–150Google Scholar
  93. Zhang Y, Chen HYH, Taylor A (2016b) Positive species diversity and above-ground biomass relationships are ubiquitous across forest strata despite interference from overstorey trees. Funct Ecol 31:419–426Google Scholar
  94. Zuo XA, Wang SK, Zhao XY, Lian J (2014) Scale dependence of plant species richness and vegetation-environment relationship along a gradient of dune stabilization in Horqin Sandy Land, Northern China. J Arid Land 6(3):334–342Google Scholar

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© Northeast Forestry University and Springer-Verlag GmbH Germany, part of Springer Nature 2018

Authors and Affiliations

  1. 1.College of ForestryShanxi Agricultural UniversityTaiguPeople’s Republic of China

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