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


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.


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



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)


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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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