Environmental drivers of herbaceous plant diversity in the understory community of a warm-temperate forest

doi:10.1016/j.pld.2025.01.003

Abstract

Abstract: Herbaceous plants are an essential component of forest diversity and driver of ecosystem processes. However, because the growth forms and life-history strategies of herbaceous plants differ from those of woody plants, it is unclear whether the mechanisms that drive patterns plant diversity and community structure in these two plant groups are the same. In this study, we determined whether herb and woody plant communities have similar patterns and drivers of alpha- and beta-diversity. We compared species richness, distribution, and abundance of herbs to woody seedlings in a 20-ha Donglingshan warm-temperate forest (Donglingshan FDP), China. We also determined whether variation in patterns of species richness and composition are better explained by environmental or spatial variables. Herbaceous plants accounted for 72% of all species (81 herbaceous, 31 woody) recorded. Alpha- and beta-diversity were higher in herbs than in woody seedlings. Although alpha-diversity of herbs and woody seedlings was not correlated across the site, the local-site contributions to beta-diversity for herbs and woody seedlings were negatively correlated. Habitat type explained slightly more variation in herb community composition than in woody seedling composition, with the highest diversity in the low-elevation slope. Environmental variables explained the variation in species richness and composition more in herbaceous plants than in woody seedlings. Our results indicate that different mechanisms drive variation in the herb and woody seedling communities, with herbs exhibiting greater environmental sensitivity and habitat dependence. These findings contribute to the better understanding of herbaceous plant diversity and composition in forest communities.

Key words: Donglingshan FDP, Herbs, Woody species diversity, Environmental filtering, Habitat associations

Tingting Deng, Qingqing Du, Yan Zhu, Simon A. Queenborough. Environmental drivers of herbaceous plant diversity in the understory community of a warm-temperate forest[J]. Plant Diversity, 2025, 47(02): 282-290.

References

Aerts, R., Honnay, O., 2011. Forest restoration, biodiversity and ecosystem functioning. BMC Ecol. 11, 29.
Ali, A., Yan, E.R., Chen, H.Y.H., et al., 2016. Stand structural diversity rather than species diversity enhances aboveground carbon storage in secondary subtropical forests in Eastern China. Biogeosciences 13, 4627–4635.
Anderson, M.J., Crist, T.O., Chase, J.M., et al., 2011. Navigating the multiple meanings of β diversity: a roadmap for the practicing ecologist. Ecol. Lett. 14, 19-28.
Both, S., Fang, T., Martin, B., et al., 2011. Lack of tree layer control on herb layer characteristics in a subtropical forest, China. J. Veg. Sci. 22, 1120-1131.
Braun, E.L., 1950. Deciduous Forests of Eastern North America. The Blakiston Company, Philadelphia, PA, USA.
Burton, J.I., Mladenoff, D.J., Clayton, M.K., et al., 2011. The roles of environmental filtering and colonization in the fine-scale spatial patterning of ground-layer plant communities in north temperate deciduous forests. J. Ecol. 99, 764-776.
Chao, A., 1984. Nonparametric estimation of the number of classes in a population. Scand. J. Stat. 11, 265-270.
Chazdon, R.L., Pearcy, R.W., 1991. The importance of sun-flecks for forest understory plants. Bioscience 41, 760-766.
Chen, Z., 2008. Study on the Diversity and Distribution Pattern of Herbaceous Plants under Parashorea Forest of Xishuangbanna. Shanghai: MS. Chinese Academy of Sciences.
Cielo-Filho, R., Gneri, M.A., Martins, F.R., 2007. Position on slope, disturbance, and tree species coexistence in a seasonal semideciduous forest in SE Brazil. Plant Ecol. 190, 189-203.
Condit, R., 1995. Research in large, long-term tropical forest plots. Trends Ecol. Evol. 10, 18-22.
Cook, J.E., 2015. Structural effects on understory attributes in second-growth forests of northern Wisconsin, USA. For. Ecol. Manag. 347, 188-199.
Crisci, J.V., Katinas, L., Apodaca, M.J., 2020. The end of botany. Trends Plant Sci. 25, 1173–1176.
De'ath, G., 2002. Multivariate regression trees: a new technique for modeling species-environment relationships. Ecology 83, 1105-1117.
Dray, S., Blanchet, F.G., Borcard, D., et al., 2021. Adespatial: Multivariate Multiscale Spatial Analysis. R package version 0.3-14. https://CRAN.R-project.org/package=adespatial.
Eisenhauer, N., Yee, K., Johnson, A.E., et al., 2011. Positive relationship between herbaceous layer diversity and the performance of soil biota in a temperate forest. Soil Biol. Biochem. 43, 462-465.
FAO, UNEP, 2020. The State of the World's Forests 2020. Forests, Biodiversity and People. Rome https://doi.org/10.4060/ca8642en.
Gentry, A.H., Dodson, C., 1987. Contribution of nontrees to species richness of a tropical rainforest. Biotropica 19, 149-156.
Gilliam, F.S., 2006. Response of the herbaceous layer of forest ecosystems to excess nitrogen deposition. J. Ecol. 94, 1176-1191.
Gilliam, F.S., 2007. The ecological significance of the herbaceous layer in temperate forest ecosystems. Bioscience 57, 845-858.
Gilliam, F.S., 2014. The Herbaceous Layer in Forests of Eastern North America. second ed. Oxford University Press, New York, USA.
Grime, J.P., 1973. Competitive exclusion in herbaceous vegetation. Nature 242, 344-347.
Guo, Y., Wang, B., Li, D., et al., 2017. Effects of topography and spatial processes on structuring tree species composition in a diverse heterogeneous tropical karst seasonal rainforest. Flora 231, 21-28.
Hart, S.A., Chen, H.Y., 2008. Fire, logging, and overstory affect understory abundance, diversity, and composition in boreal forest. Ecol. Monogr. 78, 123-140.
Jia, S.H., Wang, X.G., Hao, Z.Q., et al., 2022. The effects of natural enemies on herb diversity in a temperate forest depend on species traits and neighbouring tree composition. J. Ecol. 110, 2615-2627.
Jiang, Z.H., Ma K.M., Liu H.Y., et al., 2018. A trait-based approach reveals the importance of biotic filter for elevational herb richness pattern. J. Biogeogr. 45, 2288-2298.
Johansson, D., 1974. Ecology of vascular epiphytes in West African rain forest. Acta Phytogeogr. Suec. 59, 1-136.
Legendre, P., Borcard, D., Peres-Neto, P.R., 2005. Analyzing beta diversity: partitioning the spatial variation of community composition data. Ecol. Monogr. 75, 435-450.
Legendre, P., Caceres, M. De, 2013. Beta diversity as the variance of community data: dissimilarity coefficients and partitioning. Ecol. Lett. 16, 951-963.
Legendre, P., Gallagher, E.D., 2001. Ecologically meaningful transformations for ordination of species data. Oecologia 129, 271-280.
Legendre, P., Mi, X., Ren, H., et al., 2009. Partitioning beta diversity in a subtropical broad-leaved forest of China. Ecology 90, 663-674.
Lewis, S.L., Lloyd, J., Sitch, S., et al., 2009 Changing ecology of tropical forests: evidence and drivers. Annu. Rev. Ecol. Evol. Syst. 40, 529-549.
Linares-Palomino, R., Cardona, V., Hennig, E.I., et al., 2009. Non-woody life-form contribution to vascular plant species richness in a tropical American forest. Plant Ecol. 201, 87-99.
Liu, H., Xue, D.Y., Sang, W.G., 2014. Species diffusion and niche differentiation of the warm temperate deciduous broad-leaved forest in its functional development process. Chin. Sci. Bull. 59, 2359-2366.
Lutz, J.A., Furniss, T.J., Johnson, D.J., et al., 2018. Global importance of large-diameter trees. Global Ecol. Biogeogr. 27, 849-864.
Lutz, J.A., Larson, A.J., Freund, J.A., et al., 2013. The importance of large-diameter trees to forest structural heterogeneity. PLoS One 8, e82784.
Machado, M.A., De Almeida, J.E.B., 2019. Spatial structure, diversity, and edaphic factors of an area of Amazonian coast vegetation in Brazil. J. Torrey Bot. Soc. 146, 58-68.
Mao, Q., Chen, H., Gurmesa, G. A., et al., 2021. Negative effects of long-term phosphorus additions on understory plants in a primary tropical forest. Sci. Total Environ. 798, 149306.
Matlack, G. R., 1994. Plant species migration in a mixed-history forest landscape in eastern north America. Ecology 75, 1491-1502.
Morales-Hidalgo, D., Oswalt, S.N., Somanathan, E., 2015. Status and trends in global primary forest, protected areas, and areas designated for conservation of biodiversity from the Global Forest Resources Assessment 2015. For. Ecol. Manag. 352, 68-77.
Muller, R.N., 2014. Nutrient Relations of the Herbaceous Layer in Deciduous Forest Ecosystems. Oxford University Press, 13-34.
Murphy, S.J., Salpeter, K., Comita, L.S., 2016. Higher β-diversity observed for herbs over woody plants is driven by stronger habitat filtering in a tropical understory. Ecology 97, 2074-2084.
Nakashizuka, T., 2001. Species coexistence in temperate, mixed deciduous forests. Trends Ecol. Evol. 16, 205-210.
Nilsson, M.C., Wardle, D.A., 2005. Understory vegetation as a forest ecosystem driver: evidence from the northern Swedish boreal forest. Front. Ecol. Environ. 3, 421-428.
Oksanen, J., Blanchet, F.G., Friendly, M., et al., 2022. Vegan: Community Ecology Package. R Packages Version 2.5-7. https://CRAN.R-project.org/package=vegan/. (accessed on 2020-November-28).
Peres-Neto, P.R., Legendre, P., Dray, S., et al., 2006. Variation partitioning of species data matrices: estimation and comparison of fractions. Ecology 87, 2614-2625.
R Core Team, 2021. R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna. https://www.R-project.org/.
Radhamoni, H.V.N., Queenborough, S.A., Arietta, A.Z.A., et al., 2023. Local- and landscape-scale drivers of terrestrial herbaceous plant diversity along a tropical rainfall gradient in western Ghats, India. J. Ecol. 111, 1021-1036.
Reich, P.B., Frelich, L.E., Voldseth, R.A., et al., 2012. Understorey diversity in southern boreal forests is regulated by productivity and its indirect impacts on resource availability and heterogeneity. J. Ecol. 100, 539-545.
Richards, P.W., 1952. The Tropical Rain Forest: an Ecological Study. Cambridge University Press, Cambridge, UK.
Schnitzer, S.A., Carson, W.P., 2000. Have we forgotten the forest because of the trees? Trends Ecol. Evol. 15, 375-376.
Shvidenko, A., Barber, C.V., Persson, R., 2005. Forest and woodland systems. In: Hassan, R., Scholes, R., and Ash, N. (Eds.), Millennium Ecosystem Assessment: Ecosystems and Human Well-Being: Current State and Trends. Cambridge University Press, Cambridge, UK.
Spicer, M.E., Mellor, H., Carson, W.P., 2020. Seeing beyond the trees: a comparison of tropical and temperate plant growth-forms and their vertical distribution. Ecology 101, 1-9.
Spicer, M.E., Radhamoni, H.V.N., Duguid, M.C., et al., 2022. Herbaceous plant diversity in forest ecosystems: patterns, mechanisms, and threats. Plant Ecol. 223, 117-129.
Spurr, S.H., Barnes, B.V., 1973. Forest Ecology. second ed. Wiley.
Standish, R.J., Williams, P.A., Robertson, A.W., et al., 2004. Invasion by a perennial herb increases decomposition rate and alters nutrient availability in warm temperate lowland forest remnants. Biol. Invasions 6, 71-81.
Su, H., Li, G., 2012. Simulating the response of the Quercus mongolica forest ecosystem carbon budget to asymmetric warming. Chin. Sci. Bull. 57, 1544–1552.
Thrippleton, T., Bugmann, H., Kramer-Priewasser, K., et al., 2016. Herbaceous understorey: an overlooked player in forest landscape dynamics? Ecosystems 19, 1240-1254.
Tian, K., Chai P.T., Wang Y.Q., et al., 2023. Species diversity pattern and its drivers of the understory herbaceous plants in a Chinese subtropical forest. Front. Ecol. Evol. 10, 1113742.
Valencia, R., Foster, R.B., Villa, G.R., et al., 2004. Tree species distributions and local habitat variation in the Amazon: large forest plot in eastern Ecuador. J. Ecol. 92, 214-229.
Warren, R.J., Bradford, M.A., 2011. The shape of things to come: woodland herb niche contraction begins during recruitment in mesic forest microhabitat. Proc. Roy. Soc. B-biol. Sci. 278, 1390-1398.
Wiharto, M., Wijaya, M., Hamka, L., et al., 2021. The understory herbaceous vegetation at tropical mountain forest of mount Bawakaraeng, South Sulawesi. J. Phys. 1899, 12002.
Wright, J.S., 2002. Plant diversity in tropical forests: a review of mechanisms of species coexistence. Oecologia 130, 1-14.
Wright, S.J., 1992. Seasonal drought, soil fertility and the species density of tropical forest plant communities. Trends Ecol. Evol. 7, 260-263.
Wu, Y., Li, C., Cheng, C., 2014. Responses of soil organic carbon to long-term understory removal in subtropical Cinnamomum camphora stands. Int. J. Ecol. 4, 1-6.
Xie, J.Y., Chen, L.Z., 1994. Species diversity characteristics of deciduous forests in the warm temperate zone of North China. Acta Ecol. Sin. 14, 337-344.
Xu, S., Su, H., Ren, S., et al., 2023. Functional traits and habitat heterogeneity explain tree growth in a warm temperate forest. Oecologia 203, 371-381.
Zhang, Y.X., Liu, T.R., Guo, J.P., et al., 2021. Changes in the understory diversity of secondary Pinus tabulaeformis forests are the result of stand density and soil properties. Glob. Ecol. Conserv. 28, e01628.

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