Distribution, species richness, and relative importance of different plant life forms across drylands in China

doi:10.1016/j.pld.2024.09.007

Abstract

Abstract: Studies on plant diversity are usually based on the total number of species in a community. However, few studies have examined species richness (SR) of different plant life forms in a community along large-scale environmental gradients. Particularly, the relative importance (RIV) of different plant life forms in a community and how they vary with environmental variables are still unclear. To fill these gaps, we determined plant diversity of ephemeral plants, annual herbs, perennial herbs, and woody plants from 187 sites across drylands in China. The SR patterns of herbaceous plants, especially perennial herbs, and their RIV in plant communities increased with increasing precipitation and soil nutrient content; however, the RIV of annual herbs was not altered along these gradients. The SR and RIV of ephemeral plants were affected mainly by precipitation seasonality. The SR of woody plants had a unimodal relationship with air temperature and exhibited the highest RIV and SR percentage in plant communities under the harshest environments. An obvious shift emerged in plant community composition, SR and their critical impact factors at 238.5 mm of mean annual precipitation (MAP). In mesic regions (> 238.5 mm), herbs were the dominant species, and the SR displayed a relatively slow decreasing rate with increasing aridity, which was mediated mainly by MAP and soil nutrients. In arid regions (< 238.5 mm), woody plants were the dominant species, and the SR displayed a relatively fast decreasing rate with increasing aridity, which was mediated mainly by climate variables, especially precipitation. Our findings highlight the importance of comparative life form studies in community structure and biodiversity, as their responses to gradients differed substantially on a large scale.

Key words: Drylands, Environmental gradients, Plant life forms, Relative importance, Species richness, Threshold

Shuran Yao, Weigang Hu, Mingfei Ji, Abraham Allan Degen, Qiajun Du, Muhammad Adnan Akram, Yuan Sun, Ying Sun, Yan Deng, Longwei Dong, Haiyang Gong, Qingqing Hou, Shubin Xie, Xiaoting Wang, Jinzhi Ran, Bernhard Schmid, Qinfeng Guo, Karl J. Niklas, Jianming Deng. Distribution, species richness, and relative importance of different plant life forms across drylands in China[J]. Plant Diversity, 2025, 47(02): 273-281.

References

Adler, P.B., and Levine, J.M. 2007. Contrasting relationships between precipitation and species richness in space and time. Oikos 116, 221-232. https://doi.org/10.1111/j.2006.0030-1299.15327.x.
Berdugo, M., Kefi, S., Soliveres, S., et al. 2017. Plant spatial patterns identify alternative ecosystem multifunctionality states in global drylands. Nat. Ecol. Evol. 1, 0003. https://doi.org/10.1038/s41559-016-0003.
Berdugo, M., Maestre, F.T., Kefi, S., et al. 2019. Aridity preferences alter the relative importance of abiotic and biotic drivers on plant species abundance in global drylands. J. Ecol. 107, 190-202. https://doi.org/10.1111/1365-2745.13006.
Blunch, N. 2008. Introduction to structural equation modeling using SPSS and AMOS. SAGE Publications Ltd. https://doi.org/10.4135/9781446249345.
Borvka, L., Vacek, O., and Jehlika, J. 2005. Principal component analysis as a tool to indicate the origin of potentially toxic elements in soils. Geoderma 128, 289-300, https://doi.org/10.1016/j.geoderma.2005.04.010.
Chen, R., Ran, J., Huang, H., et al. 2019. Life history strategies drive size-dependent biomass allocation patterns of dryland ephemerals and shrubs. Ecosphere 10, e02709. https://doi.org/10.1002/ecs2.2709.
Currie, D.J., Mittelbach, G.G., Cornell, H.V., et al. 2004. Predictions and tests of climate-based hypotheses of broad-scale variation in taxonomic richness. Ecol. Lett. 7, 1121-1134. https://doi.org/10.1111/j.1461-0248.2004.00671.x.
Curtis, J.T., and McIntosh, R.P. 1951. An upland forest continuum in the prairie-forest border region of Wisconsin. Ecology 32, 476-496. https://doi.org/10.2307/1931725.
De'ath, G., and Fabricius, K.E. 2000. Classification and regression trees: a powerful yet simple technique for ecological data analysis. Ecology 81, 3178-3192. https://doi.org/10.1890/0012-9658(2000)081[3178:CARTAP]2.0.CO;2.
Delgado-Baquerizo, M., Maestre, F.T., Gallardo, A., et al. 2013. Decoupling of soil nutrient cycles as a function of aridity in global drylands. Nature 502, 672-676. https://doi.org/10.1038/nature12670.
Deng, J., Wang, G., Morris, E., et al. 2006. Plant mass-density relationship along a moisture gradient in north-west China. J. Ecol. 94, 953-958. https://doi.org/10.1111/j.1365-2745.2006.01141.x.
Deng, J., Li, T., Wang, G., et al. 2008. Trade-offs between the metabolic rate and population density of plants. PLoS One 3, e1799. https://doi.org/10.1371/journal.pone.0001799.
Dong, L., Sun, Y., Ran, J., et al. 2022. Ecosystem organic carbon storage and their drivers across the drylands of China. Catena 214, 106280. https://doi.org/10.1016/j.catena.2022.106280.
Dornelas, M., Chase, J.M., Gotelli, N.J., et al. 2023. Looking back on biodiversity change: lessons for the road ahead. Phil. Trans. R. Soc. B. 37820220199. http://doi.org/10.1098/rstb.2022.0199.
Facelli, J.M., Chesson, P., and Barnes, N. 2005. Differences in seed biology of annual plants in arid lands: a key ingredient of the storage effect. Ecology 86, 2998-3006. https://doi.org/10.1890/05-0304.
Fan, L., Tang, L., Wu, L., et al. 2014. The limited role of snow water in the growth and development of ephemeral plants in a cold desert. J. Veg. Sci. 25, 681-690. https://doi.org/10.1111/jvs.12121.
Fang, J., Shen, Z., Tang, Z., et al. 2012. Forest community survey and the structural characteristics of forests in China. Ecography 35, 1059-1071. https://doi.org/10.1111/j.1600-0587.2013.00161.x.
Fang, J., Yu, G., Liu, L., et al. 2018. Climate change, human impacts, and carbon sequestration in China. Proc. Natl Acad. Sci. USA. 115, 4015-4020. https://doi.org/10.1073/pnas.1700304115.
Fick, S.E., and Hijmans, R.J. 2017. WorldClim 2: new 1-km spatial resolution climate surfaces for global land areas. Int. J. Climatol. 37, 4302-4315. https://doi.org/10.1002/joc.5086.
Fong, Y., Huang, Y., Gilbert, P.B., et al. 2017. Chngpt: threshold regression model estimation and inference. BMC Bioinformatics 18, 454. https://doi.org/10.1186/s12859-017-1863-x.
Francis, A.P., and Currie, D.J. 2003. A globally consistent richness-climate relationship for angiosperms. The American Naturalist 161, 523-536. https://doi.org/10.1086/368223.
Hageer, Y., Esperon-Rodriguez, M., Baumgartner, J.B., et al. 2017. Climate, soil or both? Which variables are better predictors of the distributions of Australian shrub species? PeerJ. 5, 1-22. https://doi.org/10.7717/peerj.3446.
Harrison, S.P., Gornish, E.S., and Copeland, S. 2015. Climate-driven diversity loss in a grassland community. Proc. Natl Acad. Sci. USA 112, 8672-8677. https://doi.org/10.1073/pnas.1502074112.
Hu, W., Ran, J., Dong, L., et al. 2021. Aridity-driven shift in biodiversity-soil multifunctionality relationships. Nat. Commun. 12, 1-15. https://doi.org/10.1038/s41467-021-25641-0.
Hu, W., Hou, Q., Delgado-Baquerizo, M., et al. 2022. Continental-scale niche differentiation of dominant topsoil archaea in drylands. Environ. Microbiol. 24, 5483-5497. https://doi.org/10.1111/1462-2920.16099.
Huang, J., Li, Y., Fu, C., et al. 2017. Dryland climate change: recent progress and challenges. Rev. Geophys. 55, 719-778. https://doi.org/10.1002/2016RG000550.
Husakova, I., Weiner, J., and Munzbergova, Z. 2018. Species traits and shoot-root biomass allocation in 20 dry-grassland species. J. Plant Ecol. 11, 273-285. https://doi.org/10.1093/jpe/rtw143.
Hooper, D.U., Adair, E.C., Cardinale, B.J., et al. 2012. A global synthesis reveals biodiversity loss as a major driver of ecosystem change. Nature 486, 105-108. https://doi.org/10.1038/nature11118.
Jia, Y., Sun, Y., Zhang, T., et al. 2020. Elevated precipitation alters the community structure of spring ephemerals by changing dominant species density in Central Asia. Ecol. Evol. 10, 2196-2212. https://doi.org/10.1002/ece3.6057.
Kuznetsova, A., Brockhoff, P.B., and Christensen, R.H.B. 2017. lmerTest package: tests in linear mixed effects models. J. Stat. Softw. 82, 1-26. https://doi.org/10.18637/jss.v082.i13.
Ladwig, L.M., Ratajczak, Z.R., Ocheltree, T.W., et al. 2016. Beyond arctic and alpine: the influence of winter climate on temperate ecosystems. Ecology 97, 372-382. https://doi.org/10.1890/15-0153.1.
Le Bagousse-Pinguet, Y., Gross, N., Maestre, F.T., et al. 2017. Testing the environmental filtering concept in global drylands. J. Ecol. 105, 1058-1069. https://doi.org/10.1111/1365-2745.12735.
Li, X., Song, G., Hui, R., et al. 2017. Precipitation and topsoil attributes determine the species diversity and distribution patterns of crustal communities in desert ecosystems. Plant Soil 420, 163-175. https://doi.org/10.1007/s11104-017-3385-8.
Li, Y., Zhang, X., and Lv, G. 2019. Phylogeography of Ixiolirion songaricum, a spring ephemeral species endemic to Northwest China. Plant Syst. Evol. 305, 205-221. https://doi.org/10.1007/s00606-018-1563-7.
Li, C., Fu, B., Wang, S., et al. 2021. Drivers and impacts of changes in China’s drylands. Nat. Rev. Earth Environ. 2, 858-873. https://doi.org/10.1038/s43017-021-00226-z.
Liu, Y., Su, X. Shrestha, N., et al. 2018. Effects of contemporary environment and Quaternary climate change on drylands plant diversity differ between growth forms. Ecography 42, 334-345. https://doi.org/10.1111/ecog.03698.
Maestre, F.T., Quero, J.L., Gotelli, N.J., et al. 2012. Plant species richness and ecosystem multifunctionality in global drylands. Science 335, 214-218. https://doi.org/10.1126/science.1215442.
Meng, H., Gao, X., Huang, J., et al. 2015. Plant phylogeography in arid Northwest China: Retrospectives and perspectives. J. Syst. Evol. 53, 33-46. https://doi.org/10.1111/jse.12088.
Ochoa-Hueso, R., Eldridge, D.J., Delgado-Baquerizo, M., et al. 2018. Soil fungal abundance and plant functional traits drive fertile island formation in global drylands. J. Ecol. 106, 242-253. https://doi.org/10.1111/1365-2745.12871.
Palpurina, S., Wagner, V., von Wehrden, H., et al. 2017. The relationship between plant species richness and soil pH vanishes with increasing aridity across Eurasian dry grasslands. Glob. Ecol. Biogeogr. 26, 425-434. https://doi.org/10.1111/geb.12549.
Parmesan, C. 2006. Ecological and evolutionary responses to recent climate change. Annu. Rev. Ecol. Evol. Syst. 37, 637-669. https://doi.org/10.1146/ecolsys.2006.37.issue-110.1146/annurev.ecolsys.37.091305.110100.
Reed, S.C., Coe, K.K., Sparks, J.P., et al. 2012. Changes to dryland rainfall result in rapid moss mortality and altered soil fertility. Nat. Clim. Chang. 2, 752-755. https://doi.org/10.1038/nclimate1596.
Roberts, D.W. 2020. Comparison of distance-based and model-based ordinations. Ecology 101, e02908. https://doi.org/10.1002/ecy.2908.
Simova, I., Ordonez, A., and Storch, D. 2023. The dynamics of the diversity-energy relationship during the last 21,000 years. Glob. Ecol. Biogeogr. 32, 707-718. https://doi.org/10.1111/geb.13649.
Saiz, H., Gomez-Gardenes, J., Borda, J.P., et al. 2018. The structure of plant spatial association networks is linked to plant diversity in global drylands. J. Ecol. 106, 1443-1453. https://doi.org/10.1111/1365-2745.12935.
Schimel, D.S. 2010. Drylands in the earth system. Science 327, 418-419. https://doi.org/10.1126/science.1184946.
Spaak, J.W., Baert, J.M., Baird, D.J., et al. 2017. Shifts of community composition and population density substantially affect ecosystem function despite invariant richness. Ecol. Lett. 20, 1315-1324. https://doi.org/10.1111/ele.12828.
Sun, Y., Sun, Y., Yao, S., et al. 2021. Impact of climate change on plant species richness across drylands in China: From past to present and into the future. Ecol. Indic. 132, 108288. https://doi.org/10.1016/j.ecolind.2021.108288.
Tang, Z., Wang, Z., Zheng, C., et al. 2006. Biodiversity in China's mountains. Front. Ecol. Environ. 4, 347-352. https://doi.org/10.1890/1540-9295(2006)004[0347:BICM]2.0.CO;2.
Tao, Y., Qiu, D., Gong, Y., et al. 2022. Leaf-root-soil N:P stoichiometry of ephemeral plants in a temperate desert in Central Asia. J. Plant Res. 135, 55-67. https://doi.org/10.1007/s10265-021-01355-8.
Therneau, T.M., and Atkinson, E.J. 2019. An introduction to recursive partitioning using the RPART routines. Rochester, M.N: Division of biomedical statistics and informatics, Mayo Clinic; 2018. https://cran.r-project.org/web/packages/rpart/vignettes/longintro.pdf.
Ulrich, W., Soliveres, S., Maestre, F.T., et al. 2014. Climate and soil attributes determine plant species turnover in global drylands. J. Biogeogr. 41, 2307-2319. https://doi.org/10.1111/jbi.12377.
Wang, C., Wang, S., Fu, B., et al. 2017. Precipitation gradient determines the tradeoff between soil moisture and soil organic carbon, total nitrogen, and species richness in the Loess Plateau, China. Sci. Total Environ. 575, 1538-1545. https://doi.org/10.1016/j.scitotenv.2016.10.047.
Wang, Z., Fang, J., Tang, Z., et al. 2010. Patterns, determinants and models of woody plant diversity in China. Proc. R. Soc. B-Biol. Sci. 278, 2122-2132. https://doi.org/10.1098/rspb.2010.1897.
Wright, I.J., Reich, P.B., Westoby, M., et al. 2004. The worldwide leaf economics spectrum. Nature 428, 821-827. https://doi.org/10.1038/nature02403.
Xu, X., Wang, Z., Rahbek, C., et al. 2016. Geographical variation in the importance of water and energy for oak diversity. J. Biogeogr. 43, 279-288. https://doi.org/10.1111/jbi.12620.
Yao, S., Akram, M.A., Hu, W., et al. 2021. Effects of water and energy on plant diversity along the aridity gradient across dryland in China. Plants 10, 636. https://doi.org/10.3390/plants10040636.
Zhang, R., Liu, T., Zhang, J., et al. 2015. Spatial and environmental determinants of plant species diversity in a temperate desert. J. Plant Ecol. 9, 124-131. https://doi.org/10.1093/jpe/rtv053.
Zhu, J., Yu, J., Wang, P., et al. 2013. Distribution patterns of groundwater-dependent vegetation species diversity and their relationship to groundwater attributes in northwestern China. Ecohydrology 6, 191-200. https://doi.org/10.1002/eco.1258.

[1]	Miao Liu, Tiancai Zhou, Quansheng Fu. Leaf nitrogen and phosphorus are more sensitive to environmental factors in dicots than in monocots, globally [J]. Plant Diversity, 2024, 46(06): 804-811.
[2]	Wei Wang, Kun Xin, Yujun Chen, Yuechao Chen, Zhongmao Jiang, Nong Sheng, Baowen Liao, Yanmei Xiong. Spatio-temporal variation of water salinity in mangroves revealed by continuous monitoring and its relationship to floristic diversity [J]. Plant Diversity, 2024, 46(01): 134-143.
[3]	Carlos A. Vargas, Marius Bottin, Tiina Sarkinen, James E. Richardson, Marcela Celis, Boris Villanueva, Adriana Sanchez. How to fill the biodiversity data gap: Is it better to invest in fieldwork or curation? [J]. Plant Diversity, 2024, 46(01): 39-48.
[4]	Hong Qian, Jian Zhang, Meichen Jiang. Global patterns of taxonomic and phylogenetic diversity of flowering plants:Biodiversity hotspots and coldspots [J]. Plant Diversity, 2023, 45(03): 265-271.
[5]	Romina Vidal-Russell, Mariana Tadey, Romana Urfusová, Tomáš Urfus, Cintia Paola Souto. Evolutionary importance of the relationship between cytogeography and climate: New insights on creosote bushes from North and South America [J]. Plant Diversity, 2022, 44(05): 492-498.
[6]	Wen-Jing Fang, Qiong Cai, Qing Zhao, Cheng-Jun Ji, Jiang-Ling Zhu, Zhi-Yao Tang, Jing-Yun Fang. Species richness patterns and the determinants of larch forests in China [J]. Plant Diversity, 2022, 44(05): 436-444.
[7]	Li-Shen Qian, Hong-Hua Shi, Xiao-Kun Ou, Hang Sun. Elevational patterns of functional diversity and trait of Delphinium (Ranunculaceae) in Hengduan Mountains, China [J]. Plant Diversity, 2022, 44(01): 20-29.
[8]	Wen-Yun Chen, Tao Su. Asian monsoon shaped the pattern of woody dicotyledon richness in humid regions of China [J]. Plant Diversity, 2020, 42(03): 148-154.
[9]	Kflay Gebrehiwot, Sebsebe Demissew, Zerihun Woldu, Mekbib Fekadu, Temesgen Desalegn, Ermias Teferi. Elevational changes in vascular plants richness, diversity, and distribution pattern in Abune Yosef mountain range, Northern Ethiopia [J]. Plant Diversity, 2019, 41(04): 220-228.
[10]	Mouldi Gamoun, Azaiez Ouled Belgacem, Mounir Louhaichi. Diversity of desert rangelands of Tunisia [J]. Plant Diversity, 2018, 40(05): 217-225.
[11]	Jie Gao, Peng Zhang, Xing Zhang, Yanhong Liu. Multi-scale analysis on species diversity within a 40-ha old-growth temperate forest [J]. Plant Diversity, 2018, 40(02): 45-49.
[12]	Uma Shankar, Amit Kumar Tripathi. Rainforests north of the Tropic of Cancer: Physiognomy, floristics and diversity in ‘lowland rainforests’ of Meghalaya, India [J]. Plant Diversity, 2017, 39(01): 20-36.
[13]	Yahuang Luo, Jie Liu, Shaolin Tan, Marc W. Cadotte, Kun Xu, Lianming Gao, Dezhu Li. Trait variation and functional diversity maintenance of understory herbaceous species coexisting along an elevational gradient in Yulong Mountain, Southwest China [J]. Plant Diversity, 2016, 38(06): 303-311.
[14]	HUANG Man-Rong-, GUO Wei. Altitudinal Gradients of Lichen Species Richness in Tibet, China [J]. Plant Diversity, 2012, 34(2): 192-198.
[15]	David A.GALBRAITH, Natalie E.IWANYCKI, Brechann V.McGOEY, Jamie McGREGOR. The Evolving Role of Botanical Gardens and Natural Areas: A Floristic Case Study from Royal Botanical Gardens, Canada [J]. Plant Diversity, 2011, 33(01): 123-131.