How does plant diversity drive productivity in China’s drylands: Through complementary or mass ratio effects?

doi:10.1016/j.pld.2026.04.011

Plant Diversity ›› 2026, Vol. 48 ›› Issue (04): 766-781.DOI: 10.1016/j.pld.2026.04.011

• Articles • Previous Articles Next Articles

How does plant diversity drive productivity in China’s drylands: Through complementary or mass ratio effects?

Hao Guo^a,b,c, Hong-Hu Meng^d, Jin-Fei Yin^e, Tong Li^a,b,c, Ben-Feng Yin^a,b,c, Xiao-Bing Zhou^a,b,c, Yuan-Ming Zhang^a,b,c

a. State Key Laboratory of Desert and Oasis Ecology, Key Laboratory of Ecological Safety and Sustainable Development in Arid Lands, Xinjiang Institute of Ecology and Geography, Chinese Academy of Sciences, Urumqi 830011, China;
b. Xinjiang Key Laboratory of Biodiversity Conservation and Application in Arid Lands, Xinjiang Institute of Ecology and Geography, Chinese Academy of Sciences, Urumqi 830011, China;
c. Xinjiang Field Scientific Observation Research Station of Tianshan Wild Fruit Forest Ecosystem, Yili Botanical Garden, Xinjiang Institute of Ecology and Geography, Chinese Academy of Sciences, Urumqi 830011, China;
d. Plant Phylogenetics and Conservation Group, Center for Integrative Conservation, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Kunming 650223, China;
e. School of Life Sciences, Hebei University, Baoding 071002, China

Received:2025-11-26 Revised:2026-04-29 Accepted:2026-04-30 Online:2026-05-12 Published:2026-07-25
Contact: Xiao-Bing Zhou,E-mail:zhouxb@ms.xjb.ac.cn;Yuan-Ming Zhang,E-mail:ymzhang@ms.xjb.ac.cn
Supported by:
This study was also supported by the Leading Talents in Sci-Technological Innovation Project of the “Tianshan Talent” Training Plan of Xinjiang Uygur Autonomous Region (2022TSYCLJ0058), the Natural Science Foundation of Xinjiang Uygur Autonomous Region (2022D01D83), the Tianchi Talent Introduction Project of Xinjiang Uygur Autonomous Region (E4350113), the National Natural Science Foundation of China (42471072), and the Xinjiang Tianshan Talent Project (2022TSYCCX0007).

How does plant diversity drive productivity in China’s drylands: Through complementary or mass ratio effects?

Hao Guo^a,b,c, Hong-Hu Meng^d, Jin-Fei Yin^e, Tong Li^a,b,c, Ben-Feng Yin^a,b,c, Xiao-Bing Zhou^a,b,c, Yuan-Ming Zhang^a,b,c

a. State Key Laboratory of Desert and Oasis Ecology, Key Laboratory of Ecological Safety and Sustainable Development in Arid Lands, Xinjiang Institute of Ecology and Geography, Chinese Academy of Sciences, Urumqi 830011, China;
b. Xinjiang Key Laboratory of Biodiversity Conservation and Application in Arid Lands, Xinjiang Institute of Ecology and Geography, Chinese Academy of Sciences, Urumqi 830011, China;
c. Xinjiang Field Scientific Observation Research Station of Tianshan Wild Fruit Forest Ecosystem, Yili Botanical Garden, Xinjiang Institute of Ecology and Geography, Chinese Academy of Sciences, Urumqi 830011, China;
d. Plant Phylogenetics and Conservation Group, Center for Integrative Conservation, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Kunming 650223, China;
e. School of Life Sciences, Hebei University, Baoding 071002, China

通讯作者: Xiao-Bing Zhou,E-mail:zhouxb@ms.xjb.ac.cn;Yuan-Ming Zhang,E-mail:ymzhang@ms.xjb.ac.cn
基金资助:
This study was also supported by the Leading Talents in Sci-Technological Innovation Project of the “Tianshan Talent” Training Plan of Xinjiang Uygur Autonomous Region (2022TSYCLJ0058), the Natural Science Foundation of Xinjiang Uygur Autonomous Region (2022D01D83), the Tianchi Talent Introduction Project of Xinjiang Uygur Autonomous Region (E4350113), the National Natural Science Foundation of China (42471072), and the Xinjiang Tianshan Talent Project (2022TSYCCX0007).

Abstract

Abstract: Dryland biodiversity-productivity relationships remain poorly resolved. Specifically, the environmental conditions governing the shift between complementarity and mass ratio mechanisms remain unclear, limiting the effectiveness of restoration and management strategies. To address this gap, the aim of this study was to investigate the geographical patterns of diversity and biomass production in herbaceous communities along a 2100-km precipitation gradient in North China. We studied how α- and β-diversity affect community-wide productivity using linear mixed-effects models and piecewise structural equation models, along with rolling-window change-point analyses. In arid regions, biomass productivity was primarily driven by interspecific niche complementarity, where higher functional diversity (FD(α)) enhanced resource-use efficiency. However, in semi-arid regions, productivity was regulated by the mass ratio effect, specifically through the traits of dominant species, including community weighted mean height and specific leaf area, as these species exploited broader resource spectra with increasing water availability. A critical mechanistic shift occurred at a mean annual precipitation (MAP) threshold of ∼168 mm (95% CI: 152-171 mm; p < 0.001). Below this threshold, productivity was driven by diversity-mediated complementarity and stress tolerant strategies. Conversely, as MAP surpassed 168 mm, the system transitioned to mass ratio control, coincident with a shift toward competitive strategies. Overall, our study provides empirical evidence to guide dryland management: prioritising the maintenance of functional diversity in arid communities, while emphasising dominant-trait optimisation (plant height and specific leaf area) in semi-arid communities to maximise aboveground biomass.

Key words: Biodiversity, Aboveground biomass (AGB), Complementarity effect, Mass ratio effect, CSR strategy, Threshold effect

摘要： Dryland biodiversity-productivity relationships remain poorly resolved. Specifically, the environmental conditions governing the shift between complementarity and mass ratio mechanisms remain unclear, limiting the effectiveness of restoration and management strategies. To address this gap, the aim of this study was to investigate the geographical patterns of diversity and biomass production in herbaceous communities along a 2100-km precipitation gradient in North China. We studied how α- and β-diversity affect community-wide productivity using linear mixed-effects models and piecewise structural equation models, along with rolling-window change-point analyses. In arid regions, biomass productivity was primarily driven by interspecific niche complementarity, where higher functional diversity (FD(α)) enhanced resource-use efficiency. However, in semi-arid regions, productivity was regulated by the mass ratio effect, specifically through the traits of dominant species, including community weighted mean height and specific leaf area, as these species exploited broader resource spectra with increasing water availability. A critical mechanistic shift occurred at a mean annual precipitation (MAP) threshold of ∼168 mm (95% CI: 152-171 mm; p < 0.001). Below this threshold, productivity was driven by diversity-mediated complementarity and stress tolerant strategies. Conversely, as MAP surpassed 168 mm, the system transitioned to mass ratio control, coincident with a shift toward competitive strategies. Overall, our study provides empirical evidence to guide dryland management: prioritising the maintenance of functional diversity in arid communities, while emphasising dominant-trait optimisation (plant height and specific leaf area) in semi-arid communities to maximise aboveground biomass.

关键词: Biodiversity, Aboveground biomass (AGB), Complementarity effect, Mass ratio effect, CSR strategy, Threshold effect

Hao Guo, Hong-Hu Meng, Jin-Fei Yin, Tong Li, Ben-Feng Yin, Xiao-Bing Zhou, Yuan-Ming Zhang. How does plant diversity drive productivity in China’s drylands: Through complementary or mass ratio effects?[J]. Plant Diversity, 2026, 48(04): 766-781.

References

[1] Adeleye, M.A., Haberle, S.G., Gallagher, R., et al., 2023. Changing plant functional diversity over the last 12,000 years provides perspectives for tracking future changes in vegetation communities. Nat. Ecol. Evol. 7, 224-235. https://doi.org/10.1038/s41559-022-01943-4.
[2] Amyntas, A., Berti, E., Gauzens, B., et al., 2023. Niche complementarity among plants and animals can alter the biodiversity-ecosystem functioning relationship. Funct. Ecol. 37, 2652-2665. https://doi.org/10.1111/1365-2435.14419.
[3] Aragon, L., Messier, J., Atuesta-Escobar, N., et al., 2023. Tropical shrubs living in an extreme environment show convergent ecological strategies but divergent ecophysiological strategies. Ann. Bot. 131, 491-502. https://doi.org/10.1093/aob/mcad002.
[4] Balazs, K.R., Munson, S.M., Butterfield, B.J., 2022. Functional composition of plant communities mediates biomass effects on ecosystem service recovery across an experimental dryland restoration network. Funct. Ecol. 36, 2317-2330. https://doi.org/10.1111/1365-2435.14129.
[5] Berdugo, M., Delgado-Baquerizo, M., Soliveres, S., et al., 2020. Global ecosystem thresholds driven by aridity. Science 367, 787-790. https://doi.org/10.1126/science.aay5958.
[6] Bertness, M.D., Callaway, R.M., 1994. Positive interactions in communities. Trends Ecol. Evol. 9, 191-193. https://doi.org/10.1016/0169-5347(94)90088-4.
[7] Botta-Dukat, Z., Czucz, B., 2016. Testing the ability of functional diversity indices to detect trait convergence and divergence using individual-based simulation. Methods Ecol. Evol. 7, 114-126. https://doi.org/10.1111/2041-210x.12450.
[8] Brandt, M., Rasmussen, K., Penuelas, J., et al., 2017. Human population growth offsets climate-driven increase in woody vegetation in sub-Saharan Africa. Nat. Ecol. Evol. 1, 0081. https://doi.org/10.1038/s41559-017-0081.
[9] Bricca, A., Carranza, M.L., Varricchione, M., et al., 2021. Exploring plant functional diversity and redundancy of mediterranean high-mountain habitats in the Apennines. Diversity 13, 466. https://doi.org/10.3390/d13100466.
[10] Bruelheide, H., Dengler, J., Purschke, O., et al., 2018. Global trait-environment relationships of plant communities. Nat. Ecol. Evol. 2, 1906-1917. https://doi.org/10.1038/s41559-018-0699-8.
[11] Butterfield, B.J., Cavieres, L.A., Callaway, R.M., et al., 2013. Alpine cushion plants inhibit the loss of phylogenetic diversity in severe environments. Ecol. Lett. 16, 478-486. https://doi.org/10.1111/ele.12070.
[12] Caccianiga, M., Luzzaro, A., Pierce, S., et al., 2006. The functional basis of a primary succession resolved by CSR classification. Oikos 112, 10-20. https://doi.org/10.1111/j.0030-1299.2006.14107.x.
[13] Cadotte, M.W., Carscadden, K., Mirotchnick, N., 2011. Beyond species: Functional diversity and the maintenance of ecological processes and services. J. Appl. Ecol. 48, 1079-1087. https://doi.org/10.1111/j.1365-2664.2011.02048.x.
[14] Catano, C.P., Questad, E.J., Cordell, S., et al., 2023. Community assembly history alters relationships between biodiversity and ecosystem functions during restoration. Ecology 104, e3910. https://doi.org/10.1002/ecy.3910.
[15] Cavender-Bares, J., Schneider, F.D., Santos, M.J., et al., 2022. Integrating remote sensing with ecology and evolution to advance biodiversity conservation. Nat. Ecol. Evol. 6, 506-519. https://doi.org/10.1038/s41559-022-01702-5.
[16] Chitale, V., Tripathi, P., Behera, M.D., et al., 2012. On the relationships among diversity, productivity and climate from an Indian tropical ecosystem: a preliminary investigation. Biodivers. Conserv. 21, 1177-1197. https://doi.org/10.1007/s10531-012-0247-9.
[17] Coe, K., Sparks, J., 2014. Physiology-based prognostic modeling of the influence of changes in precipitation on a keystone dryland plant species. Oecologia 176, 933-942. https://doi.org/10.1007/s00442-014-3067-7.
[18] Connolly, J., Cadotte, M., Brophy, C., et al., 2011. Phylogenetically diverse grasslands are associated with pairwise interspecific processes that increase biomass. Ecology 92, 1385-1392. https://doi.org/10.1890/10-2270.1.
[19] Craven, D., Eisenhauer, N., Pearse, W.D., et al., 2018. Multiple facets of biodiversity drive the diversity-stability relationship. Nat. Ecol. Evol. 2, 1579-1587. https://doi.org/10.1038/s41559-018-0647-7.
[20] Davis, M.K., Cook, E., Collins, S., et al., 2015. Top-down vs. bottom-up regulation of herbaceous primary production and composition in an arid, urbanizing ecosystem. J. Arid Environ. 116, 103-114. https://doi.org/10.1016/j.jaridenv.2015.01.018.
[21] Delgado-Baquerizo, M., Eldridge, D.J., Maestre, F.T., et al., 2017. Climate legacies drive global soil carbon stocks in terrestrial ecosystems. Sci. Adv. 3, e1602008. https://doi.org/10.1126/sciadv.1501101.
[22] Delgado-Baquerizo, M., Reich, P.B., Trivedi, C., et al., 2020. Multiple elements of soil biodiversity drive ecosystem functions across biomes. Nat. Ecol. Evol. 4, 210-220. https://doi.org/10.1038/s41559-019-1084-y.
[23] Diaz, S., Cabido, M., 2001. Vive la difference: plant functional diversity matters to ecosystem processes. Trends Ecol. Evol. 16, 646-655. https://doi.org/10.1016/s0169-5347(01)02283-2.
[24] Diaz, S., Kattge, J., Cornelissen, J.H.C., et al., 2016. The global spectrum of plant form and function. Nature 529, 167-171. https://doi.org/10.1038/nature16489.
[25] Diaz, S., Lavorel, S., de Bello, F., et al., 2007. Incorporating plant functional diversity effects in ecosystem service assessments. Proc. Natl. Acad. Sci. U. S. A. 104, 20684-20689. https://doi.org/10.1073/pnas.0704716104.
[26] Eskelinen, A., Harrison, S., Tuomi, M., 2012. Plant traits mediate consumer and nutrient control on plant community productivity and diversity. Ecology 93, 2705-2718. https://doi.org/10.1890/12-0393.1.
[27] Fargione, J., Tilman, D., Dybzinski, R., et al., 2007. From selection to complementarity: shifts in the causes of biodiversity-productivity relationships in a long-term biodiversity experiment. Proc. R. Soc. B 274, 871-876. https://doi.org/10.1098/rspb.2006.0351.
[28] Felton, A., Shriver, R., Bradford, J., et al., 2021. Biotic versus abiotic controls on temporal sensitivity of primary production to precipitation across North American drylands. New Phytol. 231, 2150-2161. https://doi.org/10.1111/nph.17543.
[29] Firn, J., McGree, J.M., Harvey, E., et al., 2019. Leaf nutrients, not specific leaf area, are consistent indicators of elevated nutrient inputs. Nat. Ecol. Evol. 3, 400-406. https://doi.org/10.1038/s41559-018-0790-1.
[30] Flynn, D.F.B., Mirotchnick, N., Jain, M., et al., 2011. Functional and phylogenetic diversity as predictors of biodiversity-ecosystem functioning relationships. Ecology 92, 1573-1581. https://doi.org/10.1890/10-1245.1.
[31] Fridley, J., Grime, J.P., 2010. Community and ecosystem effects of intraspecific genetic diversity in grassland microcosms of varying species diversity. Ecology 91, 2272-2283. https://doi.org/10.1890/09-1240.1.
[32] Galand, P., Salter, I., Kalenitchenko, D., 2015. Ecosystem productivity is associated with bacterial phylogenetic distance in surface marine waters. Mol. Ecol. 24, 5785-5795. https://doi.org/10.1111/mec.13347.
[33] Gallagher, R.V., Falster, D.S., Maitner, B.S., et al., 2020. Open Science principles for accelerating trait-based science across the Tree of Life. Nat. Ecol. Evol. 4, 294-303. https://doi.org/10.1038/s41559-020-1109-6.
[34] Garcia-Palacios, P., Maestre, F.T., Gallardo, A., 2011. Soil nutrient heterogeneity modulates ecosystem responses to changes in the identity and richness of plant functional groups. J. Ecol. 99, 551-562. https://doi.org/10.1111/j.1365-2745.2010.01765.x.
[35] Grace, J., Anderson, T.M., Smith, M.D., et al., 2007. Does species diversity limit productivity in natural grassland communities? Ecol. Lett. 10, 680-689. https://doi.org/10.1111/j.1461-0248.2007.01058.x.
[36] Granda, E., Gazol, A., Camarero, J.J., 2018. Functional diversity differently shapes growth resilience to drought for co-existing pine species. J. Veg. Sci. 29, 265-275. https://doi.org/10.1111/jvs.12617.
[37] Grime, J.P., 1973. Competitive exclusion in herbaceous vegetation. Nature 242, 344-347. https://doi.org/10.1038/242344a0.
[38] Grime, J.P., 1977. Evidence for the existence of three primary strategies in plants and its relevance to ecological and evolutionary theory. Am. Nat. 111, 1169-1194. https://doi.org/10.1086/283244.
[39] Grime, J.P., 1998. Benefits of plant diversity to ecosystems: immediate, filter and founder effects. J. Ecol. 86, 902-910. https://doi.org/10.1046/j.1365-2745.1998.00306.x.
[40] Grossman, J.J., Cavender-Bares, J., Hobbie, S., et al., 2017. Species richness and traits predict overyielding in stem growth in an early-successional tree diversity experiment. Ecology 98, 2601-2614. https://doi.org/10.1002/ecy.1958.
[41] Grunzweig, J.M., De Boeck, H.J., Rey, A., et al., 2022. Dryland mechanisms could widely control ecosystem functioning in a drier and warmer world. Nat. Ecol. Evol. 6, 1064-1076. https://doi.org/10.1038/s41559-022-01779-y.
[42] Guo, H., Zhou, X., Tao, Y., et al., 2023b. More effect of ephemeral plant species diversity on aboveground biomass than functional diversity and functional composition. Ecol. Res. 38, 828-841. https://doi.org/10.1111/1440-1703.12411.
[43] Guo, H., Zhou, X.B., Tao, Y., et al., 2023a. Perennial herb diversity contributes more than annual herb diversity to multifunctionality in dryland ecosystems of North-western China. Front. Plant Sci. 14, 1099110. https://doi.org/10.3389/fpls.2023.1099110.
[44] Gusewell, S., 2004. N:P ratios in terrestrial plants: variation and functional significance. New Phytol. 164, 243-266. https://doi.org/10.1111/j.1469-8137.2004.01192.x.
[45] Hamilton, M.A., Murray, B.R., Cadotte, M.W., et al., 2005. Life-history correlates of plant invasiveness at regional and continental scales. Ecol. Lett. 8, 1066-1074. https://doi.org/10.1111/j.1461-0248.2005.00809.x.
[46] Hodgson, J.G., Montserrat-Marti, G., Charles, M., et al., 2011. Is leaf dry matter content a better predictor of soil fertility than specific leaf area? Ann. Bot. 108, 1337-1345. https://doi.org/10.1093/aob/mcr225.
[47] Hu, W., Ran, J., Dong, L., et al., 2021. Aridity-driven shift in biodiversity-soil multifunctionality relationships. Nat. Commun. 12, 5350. https://doi.org/10.1038/s41467-021-25641-0.
[48] Huang, K., Xia, J., Wang, Y., et al., 2018. Enhanced peak growth of global vegetation and its key mechanisms. Nat. Ecol. Evol. 2, 1897-1905. https://doi.org/10.1038/s41559-018-0714-0.
[49] Huxman, T.E., Snyder, K.A., Tissue, D., et al., 2004. Precipitation pulses and carbon fluxes in semiarid and arid ecosystems. Oecologia 141, 254-268. https://doi.org/10.1007/s00442-004-1682-4.
[50] Isbell, F., Tilman, D., Reich, P.B., et al., 2019. Deficits of biodiversity and productivity linger a century after agricultural abandonment. Nat. Ecol. Evol. 3, 1533-1538. https://doi.org/10.1038/s41559-019-1012-1.
[51] Jabot, F., Pottier, J., 2012. A general modelling framework for resource-ratio and CSR theories of plant community dynamics. J. Ecol. 100, 1296-1302. https://doi.org/10.1111/j.1365-2745.2012.02024.x.
[52] Jin, Y., Qian, H., 2022. V.PhyloMaker2: An updated and enlarged R package that can generate very large phylogenies for vascular plants. Plant Divers. 44, 335-339. https://doi.org/10.1016/j.pld.2022.05.005.
[53] Jing, X., Sanders, N.J., Shi, Y., et al., 2015. The links between ecosystem multifunctionality and above-and belowground biodiversity are mediated by climate. Nat. Commun. 6, 8159. https://doi.org/10.1038/ncomms9159.
[54] Kahmen, A., Renker, C., Unsicker, S., et al., 2006. Niche complementarity for nitrogen: an explanation for the biodiversity and ecosystem functioning relationship? Ecology 87, 1244-1255. https://doi.org/10.1890/0012-9658(2006)87[1244:ncfnae]2.0.co;2.
[55] Klironomos, J., McCune, J.L., Hart, M., et al., 2000. The influence of arbuscular mycorrhizae on the relationship between plant diversity and productivity. Ecol. Lett. 3, 137-141. https://doi.org/10.1046/j.1461-0248.2000.00131.x.
[56] Knapp, A.K., Smith, M.D., 2001. Variation among biomes in temporal dynamics of aboveground primary production. Science 291, 481-484. https://doi.org/10.1126/science.291.5503.481.
[57] Koerner, S.E., Smith, M.D., Burkepile, D.E., et al., 2018. Change in dominance determines herbivore effects on plant biodiversity. Nat. Ecol. Evol. 2, 1925-1932. https://doi.org/10.1038/s41559-018-0696-y.
[58] La Pierre, K.J., Blumenthal, D.M., Brown, C.S., et al., 2016. Drivers of variation in aboveground net primary productivity and plant community composition differ across a broad precipitation gradient. Ecosystems 19, 521-533. https://doi.org/10.1007/s10021-015-9949-7.
[59] LaManna, J.A., Belote, R.T., Burkle, L.A., et al., 2017. Negative density dependence mediates biodiversity-productivity relationships across scales. Nat. Ecol. Evol. 1, 1107-1115. https://doi.org/10.1038/s41559-017-0225-4.
[60] Lambers, H., 2021. Phosphorus Acquisition and Utilization in Plants. Annu. Rev. Plant Biol. 72, 17-44. https://doi.org/10.1146/annurev-arplant-102720-125738.
[61] Laughlin, D., Moore, M.M., 2009. Climate-induced temporal variation in the productivity-diversity relationship. Oikos 118, 897-902. https://doi.org/10.1111/j.1600-0706.2009.17514.x.
[62] Le Bagousse-Pinguet, Y., Soliveres, S., Gross, N., et al., 2019. Phylogenetic, functional, and taxonomic richness have both positive and negative effects on ecosystem multifunctionality. Proc. Natl. Acad. Sci. U. S. A. 116, 8419-8424. https://doi.org/10.1073/pnas.1815727116.
[63] Leps, J., de Bello, F., Lavorel, S., et al., 2006. Quantifying and interpreting functional diversity of natural communities: practical considerations. J. Veg. Sci. 17, 55-74. https://doi.org/10.1111/j.1654-1103.2006.tb02424.x.
[64] Liao, H., Li, D., Zhou, T., et al., 2021. The role of functional strategies in global plant distribution. Ecography 44, 493-503. https://doi.org/10.1111/ecog.05476.
[65] Liu, J., Zhang, X., Song, F., et al., 2015. Explaining maximum variation in productivity requires phylogenetic diversity and single functional traits. Ecology 96, 176-183. https://doi.org/10.1890/14-1034.1.
[66] Loranger, H., Weisser, W., Ebeling, A., et al., 2013. Invertebrate herbivory increases along an experimental gradient of grassland plant diversity. Oecologia 174, 183-193. https://doi.org/10.1007/s00442-013-2741-5.
[67] Loreau, M., 2000. Biodiversity and ecosystem functioning: recent theoretical advances. Oikos 91, 3-17. https://doi.org/10.1034/j.1600-0706.2000.910101.x.
[68] Loreau, M., Hector, A., 2001. Partitioning selection and complementarity in biodiversity experiments. Nature 412, 72-76. https://doi.org/10.1038/35083573.
[69] Luo, W., Muraina, T.O., Griffin-Nolan, R.J., et al., 2023. Responses of a semiarid grassland to recurrent drought are linked to community functional composition. Ecology 104, e3920. https://doi.org/10.1002/ecy.3920.
[70] Luo, Y., Peng, Q., Li, K., et al., 2021. Patterns of nitrogen and phosphorus stoichiometry among leaf, stem and root of desert plants and responses to climate and soil factors in Xinjiang, China. Catena 199, 105100. https://doi.org/10.1016/j.catena.2020.105100.
[71] Ma, W., He, J.S., Yang, Y., et al., 2010. Environmental factors covary with plant diversity-productivity relationships among Chinese grassland sites. Glob. Ecol. Biogeogr. 19, 233-243. https://doi.org/10.1111/j.1466-8238.2009.00508.x.
[72] Maestre, F.T., Benito, B.M., Berdugo, M., et al., 2021. Biogeography of global drylands. New Phytol. 231, 540-558. https://doi.org/10.1111/nph.17395.
[73] Maestre, F.T., Biancari, L., Chen, N., et al., 2024. Research needs on the biodiversity-ecosystem functioning relationship in drylands. npj Biodivers. 3, 12. https://doi.org/10.1038/s44185-024-00046-6.
[74] 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.
[75] Mahaut, L., Fort, F., Violle, C., et al., 2019. Multiple facets of diversity effects on plant productivity: Species richness, functional diversity, species identity and intraspecific competition. Funct. Ecol. 34, 287-298. https://doi.org/10.1111/1365-2435.13473.
[76] Mason-Jones, K., Robinson, S.L., Veen, G.F., et al., 2022. Microbial storage and its implications for soil ecology. ISME J. 16, 617-629. https://doi.org/10.1038/s41396-021-01110-w.
[77] McDowell, N., Pockman, W.T., Allen, C.D., et al., 2008. Mechanisms of plant survival and mortality during drought: Why do some plants survive while others succumb to drought? New Phytol. 178, 719-739. https://doi.org/10.1111/j.1469-8137.2008.02436.x.
[78] Medeiros, A.S., Scaloppi, J.C., Damasceno, E.S., et al., 2023. Arbuscular mycorrhizal fungi communities shaped by host-plant affect the outcome of plant-soil feedback in dryland restoration. J. Appl. Ecol. 60, 507-518. https://doi.org/10.1111/1365-2664.14330.
[79] Meng, H.H., Gao, X.Y., Huang, J.F., 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.
[80] Miller, J.E., Damschen, E.I., Ives, A.R., 2019. Functional traits and community composition: A comparison among community-weighted means, weighted correlations, and multilevel models. Methods Ecol. Evol. 10, 415-425. https://doi.org/10.1111/2041-210x.13119.
[81] Mokany, K., Ash, J., Roxburgh, S., 2008. Functional identity is more important than diversity in influencing ecosystem processes in a temperate native grassland. J. Ecol. 96, 884-893. https://doi.org/10.1111/j.1365-2745.2008.01395.x.
[82] Montes, N., Maestre, F., Ballini, C., et al., 2008. On the relative importance of the effects of selection and complementarity as drivers of diversity-productivity relationships in Mediterranean shrublands. Oikos 117, 1345-1350. https://doi.org/10.1111/j.0030-1299.2008.16910.x.
[83] Mori, A.S., 2018. Environmental controls on the causes and functional consequences of tree species diversity. J. Ecol. 106, 113-125. https://doi.org/10.1111/1365-2745.12851.
[84] Morris, E., Caruso, T., Buscot, F., et al., 2014. Choosing and using diversity indices: insights for ecological applications from the German Biodiversity Exploratories. Ecol. Evol. 4, 3514-3524. https://doi.org/10.1002/ece3.1155.
[85] Munoz, F., Violle, C., Cheptou, P.O., 2016. CSR ecological strategies and plant mating systems: outcrossing increases with competitiveness but stress-tolerance is related to mixed mating. Oikos 125, 1296-1303. https://doi.org/10.1111/oik.02328.
[86] Niklaus, P.A., Baruffol, M., He, J.S., et al., 2017. Can niche plasticity promote biodiversity-productivity relationships through increased complementarity? Ecology 98, 1104-1116. https://doi.org/10.1002/ecy.1748.
[87] Norris, C., Hobson, P., Ibisch, P., 2011. Microclimate and vegetation function as indicators of forest thermodynamic efficiency. J. Appl. Ecol. 49, 562-570. https://doi.org/10.1111/j.1365-2664.2011.02084.x.
[88] Oliveira, B.F., Machac, A., Costa, G.C., et al., 2016. Species and functional diversity accumulate differently in mammals. Glob. Ecol. Biogeogr. 25, 1119-1130. https://doi.org/10.1111/geb.12471.
[89] Pan, Y.L., Tang, H.P., Liu, D., 2025. Geographical patterns and drivers of plant productivity and species diversity in the Qinghai-Tibet Plateau. Plant Divers. 48, 908–919.
[90] Paquette, A., Messier, C., 2011. The effect of biodiversity on tree productivity: from temperate to boreal forests. Glob. Ecol. Biogeogr. 20, 170-180. https://doi.org/10.1111/j.1466-8238.2010.00592.x.
[91] Parolari, A., Goulden, M., Bras, R., 2015. Ecohydrological controls on grass and shrub above-ground net primary productivity in a seasonally dry climate. Ecohydrology 8, 1572-1583. https://doi.org/10.1002/eco.1605.
[92] Penuelas, J., Ciais, P., Canadell, J.G., et al., 2017. Shifting from a fertilization-dominated to a warming-dominated period. Nat. Ecol. Evol. 1, 1438-1445. https://doi.org/10.1038/s41559-017-0274-8.
[93] Perez-Harguindeguy, N., Diaz, S., Garnier, E., et al., 2013. New handbook for standardised measurement of plant functional traits worldwide. Aust. J. Bot. 61, 167-234. https://doi.org/10.1071/bt12225.
[94] Pierce, S., Negreiros, D., Cerabolini, B.E., et al., 2017. A global method for calculating plant CSR ecological strategies applied across biomes world-wide. Funct. Ecol. 31, 444-457. https://doi.org/10.1111/1365-2435.12722.
[95] Pointing, S., Belnap, J., 2012. Microbial colonization and controls in dryland systems. Nat. Rev. Microbiol. 10, 551-562. https://doi.org/10.1038/nrmicro2831.
[96] Prado-Junior, J.A., Schiavini, I., Vale, V.S., et al., 2016. Conservative species drive biomass productivity in tropical dry forests. J. Ecol. 104, 817-827. https://doi.org/10.1111/1365-2745.12543.
[97] Prieto, I., Armas, C., Pugnaire, F.I., 2012. Water release through plant roots: New insights into its consequences at the plant and ecosystem level. New Phytol. 193, 87-103. https://doi.org/10.1111/j.1469-8137.2011.04039.x.
[98] Reich, P.B., 2014. The world-wide ‘fast-slow’ plant economics spectrum: a traits manifesto. J. Ecol. 102, 275-301. https://doi.org/10.1111/1365-2745.12211.
[99] Ricotta, C., Dalle Fratte, M., Pierce, S., et al., 2023. Quantifying the extent of plant functional specialization using Grime’s CSR strategies. Ecol. Indic. 148, 110066. https://doi.org/10.1016/j.ecolind.2023.110066.
[100] Rio, M., Pretzsch, H., Ruiz-Peinado, R., et al., 2017. Species interactions increase the temporal stability of community productivity in Pinus sylvestris-Fagus sylvatica mixtures across Europe. J. Ecol. 105, 746-763. https://doi.org/10.1111/1365-2745.12727.
[101] Rosado, B.H.P., Mattos, E.A., 2017. On the relative importance of CSR ecological strategies and integrative traits to explain species dominance at local scales. Funct. Ecol. 31, 1969-1974. https://doi.org/10.1111/1365-2435.12894.
[102] Roscher, C., Thein, S., Schmid, B., et al., 2008. Complementary nitrogen use among potentially dominant species in a biodiversity experiment varies between two years. J. Ecol. 96, 477-488. https://doi.org/10.1111/j.1365-2745.2008.01353.x.
[103] Rosenfield, M.F., Mueller, S.C., Overbeck, G.E., 2019. Short gradient, but distinct plant strategies: The CSR scheme applied to subtropical forests. J. Veg. Sci. 30, 984-993. https://doi.org/10.1111/jvs.12787.
[104] Ruppert, J., Harmoney, K., Henkin, Z., et al., 2015. Quantifying drylands' drought resistance and recovery: the importance of drought intensity, dominant life history and grazing regime. Glob. Change Biol. 21, 1258-1270. https://doi.org/10.1111/gcb.12777.
[105] Schob, C., Armas, C., Guler, M., et al., 2013. Variability in functional traits mediates plant interactions along stress gradients. J. Ecol. 101, 753-762. https://doi.org/10.1111/1365-2745.12062.
[106] Shamrikova, E.V., Kondratenok, B.M., Tumanova, E.A., et al., 2022. Transferability between soil organic matter measurement methods for database harmonization. Geoderma 412, 115547. https://doi.org/10.1016/j.geoderma.2021.115547.
[107] Shen, H., Zhang, J., Peng, L., et al., 2023. Spatial patterns and climatic factors influence the branch xylem anatomical traits of Reaumuria soongarica in the desert region of northwestern China. Environ. Exp. Bot. 210, 105338. https://doi.org/10.1016/j.envexpbot.2023.105338.
[108] Soliveres, S., Smit, C., Maestre, F.T., 2015. Moving forward on facilitation research: Response to changing environments and effects on the diversity, functioning and evolution of plant communities. Biol. Rev. 90, 297-313. https://doi.org/10.1111/brv.12110.
[109] Su, Y.G., Liu, J., Zhang, Y.M., et al., 2021. More drought leads to a greater significance of biocrusts to soil multifunctionality. Funct. Ecol. 35, 989-1000. https://doi.org/10.1111/1365-2435.13761.
[110] Taugourdeau, S., Diedhiou, A., Fassinou, C., et al., 2022. Estimating herbaceous aboveground biomass in Sahelian rangelands using Structure from Motion data collected on the ground and by UAV. Ecol. Evol. 12, e8867. https://doi.org/10.1002/ece3.8867.
[111] Teichert, N., Lepage, M., Chevillot, X., et al., 2018. Environmental drivers of taxonomic, functional and phylogenetic diversity (alpha, beta and gamma components) in estuarine fish communities. J. Biogeogr. 45, 406-417. https://doi.org/10.1111/jbi.13133.
[112] Thompson, P.L., Davies, T.J., Gonzalez, A., 2015. Ecosystem functions across trophic levels are linked to functional and phylogenetic diversity. PLoS One 10, e0117595. https://doi.org/10.1371/journal.pone.0117595.
[113] Tilman, D., 1996. Biodiversity: Population versus ecosystem stability. Ecology 77, 350-363. https://doi.org/10.2307/2265614.
[114] Tilman, D., Isbell, F., Cowles, J.M., 2014. Biodiversity and ecosystem functioning. Annu. Rev. Ecol. Evol. Syst. 45, 471-493. https://doi.org/10.1146/annurev-ecolsys-120213-091917.
[115] Ulrich, W., Soliveres, S., Maestre, F., 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.
[116] Urbansky, E.T., 2001. Total organic carbon analyzers as tools for measuring carbonaceous matter in natural waters. J. Environ. Monit. 3, 102-112. https://doi.org/10.1039/b006564l.
[117] Valencia, E., Maestre, F.T., Le Bagousse-Pinguet, Y., et al., 2015. Functional diversity enhances the resistance of ecosystem multifunctionality to aridity in Mediterranean drylands. New Phytol. 206, 660-671. https://doi.org/10.1111/nph.13268.
[118] Venail, P., Vives, M.J., 2013. Phylogenetic distance and species richness interactively affect the productivity of bacterial communities. Ecology 94, 2529-2536. https://doi.org/10.1890/12-2002.1.
[119] Villeger, S., Mason, N.W., Mouillot, D., 2008. New multidimensional functional diversity indices for a multifaceted framework in functional ecology. Ecology 89, 2290-2301. https://doi.org/10.1890/07-1206.1.
[120] Wang, H., Zhang, R., Cai, Y., et al., 2022a. Ecological uniqueness and the determinants in arid desert ecosystems of Northwest China. Glob. Ecol. Conserv. 34, e02005. https://doi.org/10.1016/j.gecco.2022.e02005.
[121] Wang, L., Jiao, W., MacBean, N., et al., 2022b. Dryland productivity under a changing climate. Nat. Clim. Chang. 12, 981-994. https://doi.org/10.1038/s41558-022-01499-y.
[122] Wehn, S., Taugourdeau, S., Johansen, L., et al., 2017. Effects of abandonment on plant diversity in semi-natural grasslands along soil and climate gradients. J. Veg. Sci. 28, 838-847. https://doi.org/10.1111/jvs.12543.
[123] Williams, L.J., Paquette, A., Cavender-Bares, J., et al., 2017. Spatial complementarity in tree crowns explains overyielding in species mixtures. Nat. Ecol. Evol. 1, 0063. https://doi.org/10.1038/s41559-016-0063.
[124] Wilson, P.J., Thompson, K.E.N., Hodgson, J.G., 1999. Specific leaf area and leaf dry matter content as alternative predictors of plant strategies. New Phytol. 143, 155-162. https://doi.org/10.1046/j.1469-8137.1999.00427.x.
[125] Wonkka, C., Lafon, C., Hutton, C., et al., 2013. A CSR classification of tree life history strategies and implications for ice storm damage. Oikos 122, 209-222. https://doi.org/10.1111/j.1600-0706.2012.20346.x.
[126] 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.
[127] Xie, J., Wang, Z., Li, Y., 2022. Stomatal opening ratio mediates trait coordinating network adaptation to environmental gradients. New Phytol. 235, 907-922. https://doi.org/10.1111/nph.18189.
[128] Yan, P., Zhang, J., He, N., et al., 2023. Functional diversity and soil nutrients regulate the interannual variability in gross primary productivity. J. Ecol. 111, 1307-1318. https://doi.org/10.1111/1365-2745.14082.
[129] Yao, S., Hu, W., Ji, M., et al., 2025. Distribution, species richness, and relative importance of different plant life forms across drylands in China. Plant Divers. 47, 273–281.
[130] Yao, H., Peng, H., Hong, B., et al., 2022. Seasonal and diurnal variation in ecosystem respiration and environmental controls from an alpine wetland in arid northwest China. J. Plant Ecol. 15, 933-946. https://doi.org/10.1093/jpe/rtac050.
[131] Yuan, Z., Ali, A., Ruiz-Benito, P., et al., 2020. Above-and below-ground biodiversity jointly regulate temperate forest multifunctionality along a local-scale environmental gradient. J. Ecol. 108, 2012-2024. https://doi.org/10.1111/1365-2745.13378.
[132] Zhang, A., Yin, J., Zhang, Y., et al., 2023. Plants alter their aboveground and belowground biomass allocation and affect community-level resistance in response to snow cover change in Central Asia, Northwest China. Sci. Total Environ. 902, 166059. https://doi.org/10.1016/j.scitotenv.2023.166059.
[133] Zhang, Q., Buyantuev, A., Li, F.Y., et al., 2017. Functional dominance rather than taxonomic diversity and functional diversity mainly affects community aboveground biomass in the Inner Mongolia grassland. Ecol. Evol. 7, 1605-1615. https://doi.org/10.1002/ece3.2778.
[134] Zhang, Y., Chen, H.Y.H., Reich, P., 2012. Forest productivity increases with evenness, species richness and trait variation: a global meta-analysis. J. Ecol. 100, 742-749. https://doi.org/10.1111/j.1365-2745.2011.01944.x.
[135] Zheng, L.T., Chen, H.Y., Hautier, Y., et al., 2022. Functionally diverse tree stands reduce herbaceous diversity and productivity via canopy packing. Funct. Ecol. 36, 950-961. https://doi.org/10.1111/1365-2435.14009.
[136] Zomer, R.J., Xu, J., Trabucco, A., 2022. Version 3 of the global aridity index and potential evapotranspiration database. Sci. Data 9, 409. https://doi.org/10.1038/s41597-022-01493-1.
[137] Zuo, X., Li, X., Yue, P., et al., 2022. Drought-driven shifts in relationships between plant biodiversity and productivity in temperate steppes. Funct. Ecol. 36, 2917-2928. https://doi.org/10.1111/1365-2435.14219.

How does plant diversity drive productivity in China’s drylands: Through complementary or mass ratio effects?

How does plant diversity drive productivity in China’s drylands: Through complementary or mass ratio effects?

PDF (PC)

Knowledge

Abstract

Cite this article

share this article

References

Related Articles 15

Recommended Articles

Metrics

Comments

[1]	Jithu K. Jose, Saranya K.R.L. Humboldtia Vahl - An under-utilised, under-researched, and vulnerable tree genus endemic to the Western Ghats-Sri Lanka biodiversity hotspot [J]. Plant Diversity, 2026, 48(02): 433-437.
[2]	Zhi-Qiang Shen, Xian-Hui Zhu, Ming-Qiang Wang, Ming Ni, Wen-Da Cheng, Wei Lin, Cheng-Jin Chu, You-Shi Wang. Effects of neighborhood diversity on herbivory in early-stage forests in southern China [J]. Plant Diversity, 2026, 48(02): 381-388.
[3]	Ke-Ping Ma. Hangzhou strategic action plan charts a decade for biosphere reserves [J]. Plant Diversity, 2026, 48(01): 216-218.
[4]	Fei-Fei Li, Qiang Hao, Xia Cui, Ruo-Zhu Lin, Bin-Sheng Luo, Jin-Shuang Ma. Global invasive alien plant management lists: Assessing current practices and adapting to new demands [J]. Plant Diversity, 2025, 47(04): 666-680.
[5]	Mustaqeem Ahmad, Ya-Huang Luo (罗亚皇), Sonia Rathee, Robert A. Spicer, Jian Zhang (张健), Moses C. Wambulwa, Guang-Fu Zhu (朱光福), Marc W. Cadotte, Zeng-Yuan Wu (吴增源), Shujaul Mulk Khan, Debabrata Maity, De-Zhu Li (李德铢), Jie Liu (刘杰). Multifaceted plant diversity patterns across the Himalaya: Status and outlook [J]. Plant Diversity, 2025, 47(04): 529-543.
[6]	Ziwei Chen, Dongsheng Zhao, Siqi Deng, Yu Zhu, Ke Wang, Shunsheng Wang, Du Zheng. Resistance of plant diversity to road disturbance: Evidence from the Tibetan Plateau [J]. Plant Diversity, 2025, 47(03): 394-402.
[7]	Jian Zhang, Hong Qian, Xinyang Wang. An online version and some updates of R package U.Taxonstand for standardizing scientific names in plant and animal species [J]. Plant Diversity, 2025, 47(01): 166-168.
[8]	José Luiz Alves Silva, Alexandre Souza, Angela Pierre Vitória. Detection of functional diversity gradients and their geoclimatic filters is sensitive to data types (occurrence vs. abundance) and spatial scales (sites vs. regions) [J]. Plant Diversity, 2024, 46(06): 732-743.
[9]	Ling-Yun Wu, Shuang-Quan Huang, Ze-Yu Tong. Elevational and temporal patterns of pollination success in distylous and homostylous buckwheats (Fagopyrum) in the Hengduan Mountains [J]. Plant Diversity, 2024, 46(05): 661-670.
[10]	Yanwei Guan, Yongru Wu, Zheng Cao, Zhifeng Wu, Fangyuan Yu, Haibin Yu, Tiejun Wang. Island biogeography theory and the habitat heterogeneity jointly explain global patterns of Rhododendron diversity [J]. Plant Diversity, 2024, 46(05): 565-574.
[11]	Xu Chen, Haining Lu, Zhengru Ren, Yuqiu Zhang, Ruoxuan Liu, Yunhai Zhang, Xingguo Han. Reproductive height determines the loss of clonal grasses with nitrogen enrichment in a temperate grassland [J]. Plant Diversity, 2024, 46(02): 256-264.
[12]	Lin Lin, Xiao-Long Jiang, Kai-Qi Guo, Amy Byrne, Min Deng. Climate change impacts the distribution of Quercus section Cyclobalanopsis (Fagaceae), a keystone lineage in East Asian evergreen broadleaved forests [J]. Plant Diversity, 2023, 45(05): 552-568.
[13]	Karla J.P. Silva-Souza, Maíra G. Pivato, Vinícius C. Silva, Ricardo F. Haidar, Alexandre F. Souza. New patterns of the tree beta diversity and its determinants in the largest savanna and wetland biomes of South America [J]. Plant Diversity, 2023, 45(04): 369-384.
[14]	Thant Sin Aung, Alice C. Hughes, Phyo Kay Khine, Bo Liu, Xiao-Li Shen, Ke-Ping Ma. Patterns of floristic inventory and plant collections in Myanmar [J]. Plant Diversity, 2023, 45(03): 302-308.
[15]	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.