7ba1e864c177bc74eede594b5f0c05ac pmc_10476895.pdf a4efcf47c364476bd84793917b03a39f307065f6 pmc_10476895.pdf dff599dc65231cafa7354d42accee5f8d72132defea44f0b8d5f9e7fbcf260cd pmc_10476895.pdf Title: Warming-induced phenological mismatch between trees and shrubs explains high-elevation forest expansion Subject: DOI: 10.1093/nsr/nwad182 National Science Review, 10, 0, 26-06-2023. Abstract: Despite the importance of species interaction in modulating the range shifts of plants, little is known about the responses of coexisting life forms to a warmer climate. Here, we combine long-term monitoring of cambial phenology in sympatric trees and shrubs at two treelines of the Tibetan Plateau, with a meta-analysis of ring-width series from 344 shrubs and 575 trees paired across 11 alpine treelines in the Northern Hemisphere. Under a spring warming of +1°C, xylem resumption advances by 2–4 days in trees, but delays by 3–8 days in shrubs. The divergent phenological response to warming was due to shrubs being 3.2 times more sensitive than trees to chilling accumulation. Warmer winters increased the thermal requirement for cambial reactivation in shrubs, leading to a delayed response to warmer springs. Our meta-analysis confirmed such a mechanism across continental scales. The warming-induced phenological mismatch may give a competitive advantage to trees over shrubs, which would provide a new explanation for increasing alpine treeline shifts under the context of climate change. Keywords: chilling, climate warming, forcing, treeline shift, xylogenesis Author: Li Xiaoxia, Liang Eryuan, Camarero J. Julio, Rossi Sergio, Zhang Jingtian, Zhu Haifeng, Fu Yongshuo H., Sun Jian, Wang Tao, Piao Shilong, Peñuelas Josep Creator: OUP Producer: Acrobat Distiller 10.0.0 (Windows) CreationDate: Tue Jul 25 08:27:36 2023 CEST ModDate: Tue Jul 25 08:28:13 2023 CEST Custom Metadata: yes Metadata Stream: yes Tagged: yes UserProperties: no Suspects: no Form: none JavaScript: no Pages: 10 Encrypted: no Page size: 595.276 x 782.362 pts Page rot: 0 File size: 866662 bytes Optimized: yes PDF version: 1.3 name type encoding emb sub uni object ID ------------------------------------ ----------------- ---------------- --- --- --- --------- FNFEDN+ArnoPro-Bold Type 1C WinAnsi yes yes no 1510 0 FNFECM+UniversLTStd-BoldCn Type 1C WinAnsi yes yes no 1513 0 FNFCDG+HelveticaNeueLTStd-MdCn Type 1C WinAnsi yes yes no 1516 0 FNFDCB+MTSY Type 1C Custom yes yes yes 1519 0 FNFCDD+UniversLTStd-LightCn Type 1C Custom yes yes yes 1524 0 FNFCDF+UniversLTStd-LightCnObl Type 1C WinAnsi yes yes no 1529 0 FNFECN+ArnoPro-Regular Type 1C Custom yes yes yes 1532 0 FNFCDE+dummy-space Type 1C WinAnsi yes yes no 1537 0 FNFFKH+Arial-ItalicMT Type 1C WinAnsi yes yes no 11 0 FNFFKG+Symbol Type 1C WinAnsi yes yes no 14 0 FNFFJF+ArialMT Type 1C WinAnsi yes yes no 17 0 FNFGKC+RMTMI Type 1C Custom yes yes yes 401 0 FNFGLC+ArnoPro-Italic Type 1C WinAnsi yes yes no 406 0 FNGEDK+MTEX Type 1C Custom yes yes no 242 0 FNGEJJ+ArnoPro-BoldItalic Type 1C Custom yes yes yes 261 0 Jhove (Rel. 1.28.0, 2023-05-18) Date: 2025-03-12 02:43:39 CET RepresentationInformation: pmc_10476895.pdf ReportingModule: PDF-hul, Rel. 1.12.4 (2023-03-16) LastModified: 2025-03-11 02:07:55 CET Size: 866662 Format: PDF Version: 1.3 Status: Well-Formed and valid SignatureMatches: PDF-hul MIMEtype: application/pdf Profile: Linearized PDF, Tagged PDF PDFMetadata: Objects: 1619 FreeObjects: 1 IncrementalUpdates: 1 DocumentCatalog: ViewerPreferences: HideToolbar: false HideMenubar: false HideWindowUI: false FitWindow: false CenterWindow: false DisplayDocTitle: true NonFullScreenPageMode: UseNone Direction: L2R ViewArea: CropBox ViewClip: CropBox PrintArea: CropBox PageClip: CropBox PageLayout: TwoColumn PageMode: UseOutlines Language: English Outlines: Item: Title: INTRODUCTION Destination: section*.2 Item: Title: RESULTS Destination: section*.4 Children: Item: Title: Phenological mismatch at the Tibetan Plateau Destination: section*.6 Item: Title: Phenological mismatch in the Northern Hemisphere Destination: section*.8 Item: Title: DISCUSSION Destination: section*.10 Item: Title: METHODS Destination: section*.12 Children: Item: Title: Study sites and species Destination: section*.14 Item: Title: Cambial phenology and temperature Destination: section*.16 Item: Title: Growth-climate relationship of shrubs and trees at alpine treelines Destination: section*.18 Item: Title: Modeling the cambial phenology of shrubs and trees across alpine treelines Destination: section*.20 Item: Title: Assessment of temperature effects on the cambial phenology of trees and shrubs Destination: section*.22 Item: Title: DATA AVAILABILITY Destination: section*.24 Item: Title: SUPPLEMENTARY DATA Destination: section*.26 Item: Title: ACKNOWLEDGEMENTS Destination: section*.28 Item: Title: FUNDING Destination: section*.30 Item: Title: AUTHOR CONTRIBUTIONS Destination: section*.32 Item: Title: REFERENCES Destination: section*.34 Info: Title: Warming-induced phenological mismatch between trees and shrubs explains high-elevation forest expansion Author: Li Xiaoxia, Liang Eryuan, Camarero J. Julio, Rossi Sergio, Zhang Jingtian, Zhu Haifeng, Fu Yongshuo H., Sun Jian, Wang Tao, Piao Shilong, Peñuelas Josep Subject: DOI: 10.1093/nsr/nwad182 National Science Review, 10, 0, 26-06-2023. Abstract: Despite the importance of species interaction in modulating the range shifts of plants, little is known about the responses of coexisting life forms to a warmer climate. Here, we combine long-term monitoring of cambial phenology in sympatric trees and shrubs at two treelines of the Tibetan Plateau, with a meta-analysis of ring-width series from 344 shrubs and 575 trees paired across 11 alpine treelines in the Northern Hemisphere. Under a spring warming of +1°C, xylem resumption advances by 2–4 days in trees, but delays by 3–8 days in shrubs. The divergent phenological response to warming was due to shrubs being 3.2 times more sensitive than trees to chilling accumulation. Warmer winters increased the thermal requirement for cambial reactivation in shrubs, leading to a delayed response to warmer springs. Our meta-analysis confirmed such a mechanism across continental scales. The warming-induced phenological mismatch may give a competitive advantage to trees over shrubs, which would provide a new explanation for increasing alpine treeline shifts under the context of climate change. 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Author(s) 2023. Published by Oxford University Press on behalf of China Science Publishing & Media Ltd. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (https://creativecommons.org/licenses/by/4.0/), which permits unrestricted reuse, distribution, and reproduction in any medium, provided the original work is properly cited. All Rights Reserved application/pdf doi:10.1093/nsr/nwad182 © The Author(s) 2023. Published by Oxford University Press on behalf of China Science Publishing & Media Ltd. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (https://creativecommons.org/licenses/by/4.0/), which permits unrestricted reuse, distribution, and reproduction in any medium, provided the original work is properly cited. Li Xiaoxia Liang Eryuan Camarero J. Julio Rossi Sergio Zhang Jingtian Zhu Haifeng Fu Yongshuo H. Sun Jian Wang Tao Piao Shilong Peñuelas Josep Oxford University Press DOI: 10.1093/nsr/nwad182 National Science Review, 10, 0, 26-06-2023. Abstract: Despite the importance of species interaction in modulating the range shifts of plants, little is known about the responses of coexisting life forms to a warmer climate. Here, we combine long-term monitoring of cambial phenology in sympatric trees and shrubs at two treelines of the Tibetan Plateau, with a meta-analysis of ring-width series from 344 shrubs and 575 trees paired across 11 alpine treelines in the Northern Hemisphere. Under a spring warming of +1°C, xylem resumption advances by 2–4 days in trees, but delays by 3–8 days in shrubs. The divergent phenological response to warming was due to shrubs being 3.2 times more sensitive than trees to chilling accumulation. Warmer winters increased the thermal requirement for cambial reactivation in shrubs, leading to a delayed response to warmer springs. Our meta-analysis confirmed such a mechanism across continental scales. The warming-induced phenological mismatch may give a competitive advantage to trees over shrubs, which would provide a new explanation for increasing alpine treeline shifts under the context of climate change. chilling climate warming forcing treeline shift xylogenesis Warming-induced phenological mismatch between trees and shrubs explains high-elevation forest expansion Li Xiaoxia, Liang Eryuan, Camarero J. 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