Different hydrothermal conditions drive the radial growth pattern of larch (Larix spp.) species on the southeastern Tibetan Plateau

Abstract

The Hengduan Mountains, a biodiversity hotspot on the Tibetan Plateau, are undergoing rapid elevation-dependent warming, profoundly altering alpine forest dynamics. Larix species in this region dominate high-altitude treelines and represent the world’s lowest-latitude natural populations of larch forests. However, climate-growth relationships of Larix species at the whole regional scales have received little attention. To address this gap, we investigated the spatiotemporal variability of radial growth in Larix species and their climatic sensitivity across this region using a network of 26 tree-ring chronologies spanning 1960–2022. Hierarchical clustering identified three distinct geographical clusters (southwestern, central, and northeastern), revealing divergent growth trajectories and climate responses. Results demonstrated that growing-season temperature was the primary climatic driver of Larix radial growth, but its influence varied spatially: southwestern populations correlate strongly with May–August mean temperatures, central populations with May–August mean temperatures, and northeastern populations with April–June maximum temperatures. Each cluster exhibited unique growth trends and thermal sensitivities before and after the rapid warming since 1990. Meanwhile, an inter-individual response divergence became apparent under warming, reflecting intensified competition and microhabitat-scale stress, which highlights the limitations of traditional population-level climate response models that assume a uniform response. Spatial heterogeneity in climate-growth relationships reflected synergistic thermal-hydrological effects and species-specific adaptations, with warming enhancing carbon sequestration in moisture-sufficient areas but threatening high-elevation ecosystems through growth suppression and treelines instability. These findings underscore the need for regionally tailored conservation strategies to address climate-driven ecological imbalances in alpine forests.

Keywords

Larix species / Radial growth / Climate sensitivity / Hengduan Mountains / Dendrochronology

References

[1] Anderson-Teixeira KJ, Miller AD, Mohan JE, Hudiburg TW, Duval BD, DeLucia EH. Altered dynamics of forest recovery under a changing climate. Glob Change Biol, 2013, 19(7): 2001-2021.

[2] Aryal S, Grießinger J, Arsalani M, Meier WJ, Fu PL, Fan ZX, Bräuning A. Insect infestations have an impact on the quality of climate reconstructions using Larix ring-width chronologies from the Tibetan Plateau. Ecol Indic, 2023, 148110124

[3] Beniston M, Stoffel M. Assessing the impacts of climatic change on mountain water resources. Sci Total Environ, 2014, 493: 1129-1137.

[4] Bennett AC, McDowell NG, Allen CD, Anderson-Teixeira KJ. Larger trees suffer most during drought in forests worldwide. Nat Plants, 2015, 115139

[5] Bhattacharya A (2022) Effect of low temperature stress on photosynthesis and allied traits: a review. In: Physiological processes in plants under low temperature stress. Springer Singapore, pp 199–297. https://doi.org/10.1007/978-981-16-9037-2_3

[6] Brokaw N, Busing RT, Brokaw N, Busing RT. Niche versus chance and tree diversity in forest gaps. Trends Ecol Evol, 2000, 15(5): 183-188.

[7] Bunn AG. A dendrochronology program library in R (dplR). Dendrochronologia, 2008, 26(2): 115-124.

[8] Bunn AG. Statistical and visual crossdating in R using the dplR library. Dendrochronologia, 2010, 28(4): 251-258.

[9] Buras A. A comment on the expressed population signal. Dendrochronologia, 2017, 44: 130-132.

[10] Cook ER, Kairiukstis LA (2013) Methods of dendrochronology: applications in the environmental sciences. Springer Science & Business Media.

[11] D’Arrigo R, Wilson R, Liepert B, Cherubini P. On the ‘Divergence problem’ in Northern Forests: a review of the tree-ring evidence and possible causes. Glob Planet Change, 2008, 60(3–4): 289-305.

[12] de S Magalhães JG, Amoroso MM, Larson BC. What evidence exists on the effects of competition on trees’ responses to climate change? A systematic map protocol. Environ Evid, 2021, 10(134

[13] Deng GF, Li MQ, Hao ZX, Shao XM. Responses to climate change of maximum latewood density from Larix speciosa Cheng et law and Abies delavayi franch. in the northwest of Yunnan Province, China. Forests, 2022, 13(5720

[14] Du QQ, Rossi S, Lu XM, Wang YF, Zhu HF, Liang EY, Camarero JJ. Negative growth responses to temperature of sympatric species converge under warming conditions on the southeastern Tibetan Plateau. Trees, 2020, 342): 395-404.

[15] Fan ZX, Bräuning A, Cao KF. Tree-ring based drought reconstruction in the central Hengduan Mountains region (China) since A.D. 1655. Int J Climatol, 2008, 28(14): 1879-1887.

[16] Fan ZX, Bräuning A, Cao KF, Zhu SD. Growth–climate responses of high-elevation conifers in the central Hengduan Mountains, southwestern China. For Ecol Manag, 2009, 258(3): 306-313.

[17] Fan ZX, Bräuning A, Tian QH, Yang B, Cao KF. Tree ring recorded May–August temperature variations since A.D. 1585 in the Gaoligong Mountains, southeastern Tibetan Plateau. Palaeogeogr Palaeoclimatol Palaeoecol, 2010, 296(1–2): 94-102.

[18] Fritts H. Tree rings and climate, 1976, San Diego. Academic Press

[19] Gai XR, Yu DP, Wang SL, Jia X, Wu J, Zhou WM, Zhou L, Dai LM. A review of formation mechanism on the ‘divergence problem’ of tree growth-climate relationship. Chin J Ecol, 2017, 36(11): 3273-3280. in Chinese)

[20] Guiot J. The bootstrapped response function. Tree-Ring Bull, 1991, 51: 39-41

[21] Guo MM, Zhang YD, Liu SR, Gu FX, Wang XC, Li ZS, Shi CM, Fan ZX. Divergent growth between spruce and fir at alpine treelines on the east edge of the Tibetan Plateau in response to recent climate warming. Agric for Meteorol, 2019, 276–277107631

[22] Hamed KH, Ramachandra Rao A. A modified Mann-Kendall trend test for autocorrelated data. J Hydrol, 1998, 204(1–4): 182-196.

[23] Harsch MA, Hulme PE, McGlone MS, Duncan RP. Are treelines advancing? A global meta-analysis of treeline response to climate warming. Ecol Lett, 2009, 12(10): 1040-1049.

[24] Holmes RL (1983) Computer-assisted quality control in tree-ring dating and measurement.

[25] Hu M, Zhao XE, Chen F, Wang SJ, Torbenson MCA, Hou TY, Chen YP. Increasing evidence of an anthropogenic signal in drought variations on the river source areas of southeastern Tibetan Plateau. J Hydrol, 2025, 660133508

[26] IPCC (2021) Climate Change 2021: The Physical Science Basis. Contribution of Working Group I to the Sixth Assessment Report of the Intergovernmental Panel on Climate Change. In. Cambridge University Press.

[27] Jevšenak J. New features in the dendroTools R package: bootstrapped and partial correlation coefficients for monthly and daily climate data. Dendrochronologia, 2020, 63125753

[28] Körner C. Alpine treelines. Springer Basel, 2012

[29] Li ZS, Liu GH, Fu BJ, Hu CJ, Luo SZ, Liu XL, He F. Anomalous temperature–growth response of Abies faxoniana to sustained freezing stress along elevational gradients in China’s Western Sichuan Province. Trees, 2012, 26(4): 1373-1388.

[30] Li MY, Wang L, Fan ZX, Shen CC. Tree-ring density inferred late summer temperature variability over the past three centuries in the Gaoligong Mountains, southeastern Tibetan Plateau. Palaeogeogr Palaeoclimatol Palaeoecol, 2015, 422: 57-64.

[31] Li WQ, Jiang Y, Dong MY, Du EZ, Zhou ZJ, Zhao SD, Xu H. Diverse responses of radial growth to climate across the southern part of the Asian boreal forests in Northeast China. For Ecol Manag, 2020, 458117759

[32] Li ZS, Keyimu M, Fan ZX, Wang XC. Climate sensitivity of conifer growth doesn’t reveal distinct low–high dipole along the elevation gradient in the Wolong National Natural Reserve, SW China. Dendrochronologia, 2020, 61125702

[33] Li T, Peng JF, Au TF, Li JR, Li JB, Zhang Y. Dendroclimatological study ofSabina saltuaria and Abies faxoniana in the mixed forests of the Qionglai Mountains, eastern Tibetan Plateau. J Res, 2024, 35(120

[34] Liang EY, Wang YF, Piao SL, Lu XM, Camarero JJ, Zhu HF, Zhu LP, Ellison AM, Ciais P, Peñuelas J. Species interactions slow warming-induced upward shifts of treelines on the Tibetan Plateau. Proc Natl Acad Sci USA, 2016, 11316): 4380-4385.

[35] Luo WW, Xie CS, Shi SL, Li JJ, Wang GY, Bie XJ. Climatic warming caused a transition in tree growth sensitivity from temperature to moisture conditions: evidence from multi-species tree-ring data in the southeastern Tibetan Plateau. Forests, 2024, 15(2): 241.

[36] Mamet SD, Brown CD, Trant AJ, Laroque CP. Shifting global Larix distributions: northern expansion and southern retraction as species respond to changing climate. J Biogeogr, 2019, 461): 30-44.

[37] McCarthy J. Gap dynamics of forest trees: a review with particular attention to boreal forests. Dossiers Environ, 2001, 9(1): 1-59.

[38] Murtagh F, Legendre P. Ward’s hierarchical agglomerative clustering method: which algorithms implement ward’s criterion?. J Classif, 2014, 31(3): 274-295.

[39] Oliver CD, Larson BC. Forest stand dynamics, 1996, New York. Wiley

[40] Pallardy SG (2008) The woody plant body. In: Physiology of woody plants. Elsevier. pp 9–38. https://doi.org/10.1016/b978-012088765-1.50003-8

[41] Peltier DMP, Ogle K. Tree growth sensitivity to climate is temporally variable. Ecol Lett, 2020, 23(11): 1561-1572.

[42] Pepin NC, Arnone E, Gobiet A, Haslinger K, Kotlarski S, Notarnicola C, Palazzi E, Seibert P, Serafin S, Schöner W, Terzago S, Thornton JM, Vuille M, Adler C. Climate changes and their elevational patterns in the mountains of the world. Rev Geophys, 2022, 60(1e2020RG000730

[43] Peterson DW, Peterson DL. Mountain hemlock growth responds to climatic variability at annual and decadal time scales. Ecology, 2001, 82(12): 3330-3345.

[44] Qi ZH, Liu HY, Wu XC, Hao Q. Climate-driven speedup of alpine treeline forest growth in the Tianshan Mountains, Northwestern China. Glob Change Biol, 2015, 21(2): 816-826.

[45] Rousseeuw PJ. Silhouettes: a graphical aid to the interpretation and validation of cluster analysis. J Comput Appl Math, 1987, 20: 53-65.

[46] Ryhti K, Schiestl-Aalto P, Tang Y, Rinne-Garmston KT, Ding YY, Pumpanen J, Biasi C, Saurer M, Bäck J, Kulmala L. Effects of variable temperature and moisture conditions on respiration and nonstructural carbohydrate dynamics of tree roots. Agric for Meteorol, 2022, 323109040

[47] Schwab N, Bürzle B, Böhner J, Chaudhary RP, Scholten T, Schickhoff U (2021) Environmental drivers of species composition and tree species density of a near-natural central Himalayan treeline ecotone: consequences for the response to climate change. In: Mountain landscapes in transition. Springer International Publishing. pp 349–370. https://doi.org/10.1007/978-3-030-70238-0_13

[48] Shen TQ, Jiang P, Ju Q, Zhao JH, Chen XG, Lin H, Yang B, Tan CH, Zhang Y, Fu XT, Yu ZB. Permafrost on the Tibetan Plateau is degrading: historical and projected trends. J Hydrol, 2024, 628130501

[49] Shi CM, Shen MG, Wu XC, Cheng X, Li XY, Fan TY, Li ZS, Zhang YD, Fan ZX, Shi FZ, Wu GC. Growth response of alpine treeline forests to a warmer and drier climate on the southeastern Tibetan Plateau. Agric for Meteorol, 2019, 264: 73-79.

[50] Shi SL, Liu GH, Li ZS, Ye X. Elevation-dependent growth trends of forests as affected by climate warming in the southeastern Tibetan Plateau. For Ecol Manag, 2021, 498119551

[51] Stokes MA. An introduction to tree-ring dating, 1996University of Arizona Press

[52] Sun Y, Wang LL, Chen J, Duan JP, Shao XM, Chen KL. Growth characteristics and response to climate change of Larix miller tree-ring in China. Sci China Earth Sci, 2010, 53(6): 871-879.

[53] Tian L, Fu WX, Tao Y, Li MY, Wang L. Dynamics of the alpine timberline and its response to climate change in the Hengduan mountains over the period 1985–2015. Ecol Indic, 2022, 135108589

[54] Tiwari A, Huang HS, Zhou ZK. Temperature sensitivity of alpine larch treeline in the southeast of Tibetan Plateau. Bot Orient, 2020, 14: 29-38.

[55] Tiwari A, Adhikari A, Fan ZX, Li SF, Jump AS, Zhou ZK. Himalaya to Hengduan: dynamics of alpine treelines under climate change. Reg Environ Change, 2023, 23(4): 157.

[56] Wang QX, Fan XH, Wang MB. Recent warming amplification over high elevation regions across the globe. Clim Dyn, 2014, 431–2): 87-101.

[57] Wang YF, Pederson N, Ellison AM, Buckley HL, Case BS, Liang EY, Julio Camarero J. Increased stem density and competition may diminish the positive effects of warming at alpine treeline. Ecology, 2016, 97(7): 1668-1679.

[58] Wang SJ, Chen F, Hu M, Chen YP, Cao HH, Yue WP, Zhao XE. Past, present and future changes in the annual streamflow of the Lancang-Mekong River and their driving mechanisms. Sci Total Environ, 2024, 947174707

[59] Wang SJ, Chen F, Chen YP, Torbenson MCA, Esper J, Zhao XE, Hu M, Zhang HL, Yue WP, Cao HH. Greening of Eurasia’s center driven by low-latitude climate warming. For Ecosyst, 2025, 13100330

[60] Wei XX, Wang XQ. Recolonization and radiation in Larix (Pinaceae): evidence from nuclear ribosomal DNA paralogues. Mol Ecol, 2004, 13(10): 3115-3123.

[61] Wookey PA, Aerts R, Bardgett RD, Baptist F, Bråthen KA, Cornelissen JHC, Gough L, Hartley IP, Hopkins DW, Lavorel S, Shaver GR. Ecosystem feedbacks and cascade processes: understanding their role in the responses of Arctic and alpine ecosystems to environmental change. Glob Change Biol, 2009, 15(5): 1153-1172.

[62] Wu ZY, Raven PH, Hong DY. Flora of China (Vol. 4: Pinaceae). Beijing: Science Press; St, 1999, Louis. Missouri Botanical Garden Press

[63] Xing YW, Ree RH. Uplift-driven diversification in the Hengduan Mountains, a temperate biodiversity hotspot. Proc Natl Acad Sci U S A, 2017

[64] Yang B, He MH, Shishov V, Tychkov I, Vaganov E, Rossi S, Ljungqvist FC, Bräuning A, Grießinger J. New perspective on spring vegetation phenology and global climate change based on Tibetan Plateau tree-ring data. Proc Natl Acad Sci U S A, 2017, 114(27): 6966-6971.

[65] Yang L, Cui GS, Zhao WL, Zhang ZM, Luo TX, Zhang L. Sensitivity of radial growth of subalpine conifer trees to climate warming on the southeastern Tibetan Plateau. Glob Ecol Conserv, 2023, 43e02470

[66] Yuan S, Jiang Y, Zhao ZF, Cui MH, Shi DD, Wang SJ, Kang MY. Different trends and divergent responses to climate factors in the radial growth of Abies georgei along elevations in the central Hengduan Mountains. Dendrochronologia, 2023, 80126114

[67] Yue HT, Li JN, Xie SY, Chen H, Tian K, Sun M, Zhang DC, Zhang Y. Relationships between climate variability and radial growth of Larix potaninii at the upper altitudinal limit in central Hengduan Mountain, southwestern China. Forests, 2023, 14(9): 1790.

[68] Yue WP, Chen F, Davi NK, Zhang HL, Chen YP, Zhao XE, Gao ZH. Little Ice Age cooling in the WesternHengduan Mountains, China: a 600-year warm-season temperature reconstruction from tree rings. Clim Dyn, 2024, 62(1): 773-790.

[69] Zang C, Biondi F. Treeclim: an R package for the numerical calibration of proxy-climate relationships. Ecography, 2015, 38(4): 431-436.

[70] Zeng XM, Wei CF, Liu XH, Zhang LN. Qinghai spruce (Picea crassifolia) and Chinese pine (Pinus tabuliformis) show high vulnerability and similar resilience to early-growing-season drought in the Helan Mountains, China. Ecol Indic, 2020, 110105871

[71] Zhang Y, Cao RJ, Yin J, Tian K, Xiao DR, Zhang WG, Yin DC. Radial growth response of major conifers to climate change on Haba Snow Mountain, Southwestern China. Dendrochronologia, 2020, 60125682

[72] Zhang H, Fu PL, Lin PL, Ge S, Yang JQ, Ge-Rong QZ, Fan ZX. Intra-annual radial growth of Abies georgei and Larix potaninii and its responses to environmental factors in the Baima Snow Mountain, Northwest Yunnan, China. Ying Yong Sheng Tai Xue Bao, 2022, 33(11): 2881-2888.

[73] Zhang XQ, Xu XM, Li X, Cui P, Zheng D. A new scheme of climate-vegetation regionalization in the Hengduan Mountains Region. Sci China Earth Sci, 2024, 67(3): 751-768.

[74] Zhao SD, Pederson N, D’Orangeville L, HilleRisLambers J, Boose E, Penone C, Bauer B, Jiang Y, Manzanedo RD. The international tree-ring data bank (ITRDB) revisited. J Biogeogr, 2019, 46(2): 355-368.

[75] Zheng XY, Babst F, Camarero JJ, Li XX, Lu XM, Gao S, Sigdel SR, Wang YF, Zhu HF, Liang EY. Density-dependent species interactions modulate alpine treeline shifts. Ecol Lett, 2024, 274e14403

[76] Zhou Y, Yi YJ, Liu HX, Song J, Jia WF, Zhang SH. Altitudinal trends in climate change result in radial growth variation of Pinus yunnanensis at an arid-hot valley of southwest China. Dendrochronologia, 2022, 71125914

RIGHTS & PERMISSIONS

The Author(s)