<?xml version="1.0" encoding="UTF-8"?>
<!DOCTYPE article PUBLIC "-//NLM//DTD Journal Publishing with OASIS Tables v3.0 20080202//EN" "https://jats.nlm.nih.gov/nlm-dtd/publishing/3.0/journalpub-oasis3.dtd">
<article xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:oasis="http://docs.oasis-open.org/ns/oasis-exchange/table" xml:lang="en" dtd-version="3.0" article-type="research-article">
  <front>
    <journal-meta><journal-id journal-id-type="publisher">BG</journal-id><journal-title-group>
    <journal-title>Biogeosciences</journal-title>
    <abbrev-journal-title abbrev-type="publisher">BG</abbrev-journal-title><abbrev-journal-title abbrev-type="nlm-ta">Biogeosciences</abbrev-journal-title>
  </journal-title-group><issn pub-type="epub">1726-4189</issn><publisher>
    <publisher-name>Copernicus Publications</publisher-name>
    <publisher-loc>Göttingen, Germany</publisher-loc>
  </publisher></journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.5194/bg-23-5943-2026</article-id><title-group><article-title>Saturation effect of background temperature and aridity on vegetation phenological sensitivity to urban warming</article-title><alt-title>Saturation effect of background temperature</alt-title>
      </title-group>
      <contrib-group>
        <contrib contrib-type="author" equal-contrib="yes" corresp="no" rid="aff1">
          <name><surname>Zhenzhen</surname><given-names>Zhang</given-names></name>
          
        <ext-link>https://orcid.org/0000-0002-5663-3267</ext-link></contrib>
        <contrib contrib-type="author" equal-contrib="yes" corresp="no" rid="aff1">
          <name><surname>Xinxin</surname><given-names>Hu</given-names></name>
          
        </contrib>
        <contrib contrib-type="author" corresp="no" rid="aff1">
          <name><surname>Yongqi</surname><given-names>Zhang</given-names></name>
          
        </contrib>
        <contrib contrib-type="author" corresp="no" rid="aff2">
          <name><surname>Shufen</surname><given-names>Cui</given-names></name>
          
        </contrib>
        <contrib contrib-type="author" corresp="no" rid="aff3">
          <name><surname>Liheng</surname><given-names>Sun</given-names></name>
          
        </contrib>
        <contrib contrib-type="author" corresp="yes" rid="aff1">
          <name><surname>Xingwen</surname><given-names>Lin</given-names></name>
          <email>linxw@zjnu.edu.cn</email>
        <ext-link>https://orcid.org/0000-0002-4189-7366</ext-link></contrib>
        <contrib contrib-type="author" corresp="no" rid="aff1">
          <name><surname>Chaofan</surname><given-names>Wu</given-names></name>
          
        </contrib>
        <contrib contrib-type="author" corresp="no" rid="aff1">
          <name><surname>Zhaoyang</surname><given-names>Zhang</given-names></name>
          
        </contrib>
        <contrib contrib-type="author" corresp="no" rid="aff1">
          <name><surname>Yuanjian</surname><given-names>Chen</given-names></name>
          
        </contrib>
        <contrib contrib-type="author" corresp="no" rid="aff4">
          <name><surname>Qingqing</surname><given-names>Wen</given-names></name>
          
        </contrib>
        <aff id="aff1"><label>1</label><institution>College of Geography and Environmental Sciences, Zhejiang Normal University, Jinhua 321004, China</institution>
        </aff>
        <aff id="aff2"><label>2</label><institution>College of Business, Lishui University, Lishui 323200, China</institution>
        </aff>
        <aff id="aff3"><label>3</label><institution>Faculty of Management, Universiti Teknologi Malaysia, Johor Bahru 81310, Malaysia</institution>
        </aff>
        <aff id="aff4"><label>4</label><institution>The Administration Center of Jinhua Wucheng Nanshan Provincial Nature Reserve, Jinhua 321000, China</institution>
        </aff><author-comment content-type="econtrib"><p>These authors contributed equally to this work.</p></author-comment>
      </contrib-group>
      <author-notes><corresp id="corr1">Lin Xingwen (linxw@zjnu.edu.cn)</corresp></author-notes><pub-date><day>28</day><month>August</month><year>2026</year></pub-date>
      
      <volume>23</volume>
      <issue>16</issue>
      <fpage>5943</fpage><lpage>5959</lpage>
      <history>
        <date date-type="received"><day>16</day><month>January</month><year>2026</year></date>
           <date date-type="rev-request"><day>4</day><month>March</month><year>2026</year></date>
           <date date-type="rev-recd"><day>9</day><month>August</month><year>2026</year></date>
           <date date-type="accepted"><day>10</day><month>August</month><year>2026</year></date>
      </history>
      <permissions>
        <copyright-statement>Copyright: © 2026 Zhang Zhenzhen et al.</copyright-statement>
        <copyright-year>2026</copyright-year>
      <license license-type="open-access"><license-p>This work is licensed under the Creative Commons Attribution 4.0 International License. To view a copy of this licence, visit <ext-link ext-link-type="uri" xlink:href="https://creativecommons.org/licenses/by/4.0/">https://creativecommons.org/licenses/by/4.0/</ext-link></license-p></license></permissions><self-uri xlink:href="https://bg.copernicus.org/articles/23/5943/2026/bg-23-5943-2026.html">This article is available from https://bg.copernicus.org/articles/23/5943/2026/bg-23-5943-2026.html</self-uri><self-uri xlink:href="https://bg.copernicus.org/articles/23/5943/2026/bg-23-5943-2026.pdf">The full text article is available as a PDF file from https://bg.copernicus.org/articles/23/5943/2026/bg-23-5943-2026.pdf</self-uri>
      <abstract><title>Abstract</title>

      <p id="d2e186">Quantifying how background temperature and aridity constrain phenological sensitivity to urban warming requires identifying critical thermal thresholds and hydrological boundaries driving spatial heterogeneity. This study assessed urban warming effects on vegetation phenology across 293 Chinese cities during 2010–2020. Urban-rural disparities (<inline-formula><mml:math id="M1" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS, <inline-formula><mml:math id="M2" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS) and temperature sensitivity (<inline-formula><mml:math id="M3" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-SOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>, <inline-formula><mml:math id="M4" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-EOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>) were quantified using satellite metrics. Results showed that urban heat islands advanced SOS by 12.06 d and delayed EOS by 9.86 d relative to rural areas. A non-linear saturation effect was detected: phenological sensitivity to urban warming peaked at 4 °C (spring) and 6 °C (autumn), and weakened significantly beyond the saturation thresholds of 12.5 °C (SOS) and 4 °C (EOS). Aridity acted as a negative regulator that weakened warming-induced phenological effects across most aridity index (AI) ranges and reversed them within 1.4 <inline-formula><mml:math id="M5" display="inline"><mml:mo>&lt;</mml:mo></mml:math></inline-formula> AI <inline-formula><mml:math id="M6" display="inline"><mml:mo>&lt;</mml:mo></mml:math></inline-formula> 2.0. Combined, LST and AI explained 75.05 % and 76.21 % of spatial variance in SOS and EOS responses, respectively. Conceptually, these findings challenge the linear paradigm of warming-driven phenological shifts by demonstrating finite physiological responses to urban warming. Ecologically, this study highlights the coupled heat-aridity constraints on urban vegetation, providing key implications for climate-adaptive urban ecological planning.</p>
  </abstract>
    </article-meta>
  </front>
<body>
      

<sec id="Ch1.S1" sec-type="intro">
  <label>1</label><title>Introduction</title>
      <p id="d2e255">Vegetation phenology, a critical aspect of ecosystem dynamics, exhibits significant sensitivity to the coupling of thermal and moisture variations (Badeck et al., 2004; Yu et al., 2003). While recent studies have shown that urban warming advances the start of the growing season (SOS) and delays the end of the growing season (EOS) (Li et al., 2017; Zhou et al., 2016), it is increasingly recognized that these responses are not solely temperature-driven but are fundamentally constrained by the background aridity of the locale. Thus, the phenological effects of the Urban Heat Island (UHI) effect must be evaluated through a dual lens of heat and water availability (Jeong et al., 2019; Buyantuyev and Wu, 2012; Crawford et al., 2025). The cascading effects of these shifts extend beyond mere phenological alterations, contributing to a range of ecological and environmental challenges (Zhou et al., 2016; Qiu et al., 2017; Ding et al., 2020).</p>
      <p id="d2e258">However, the degree to which warming promotes phenological changes has been widely debated (Chmielewski and Rötzer, 2001; Ding et al., 2020; Yao et al., 2017). Urban environments offer a unique “natural laboratory” for testing these phenological theories. Unlike experimental warming plots, the urban heat island (UHI) effect represents a sustained, large-scale warming scenario that has persisted for decades (Grimm et al., 2008). This allows researchers to observe long-term vegetation adaptation and identify thresholds that are difficult to capture in short-term manipulative experiments. Moreover, the vast geographic distribution of cities across climatic gradients provides a natural experimental framework to test how identical warming magnitudes (UHI) interact with varying background climates (Fig. 1). Empirical evidence from the Northern Hemisphere consistently shows that cities in cold high-latitude regions exhibit more advanced SOS and delayed EOS in response to urban warming (Dallimer et al., 2016; Gazal et al., 2008; Jia et al., 2021; Li et al., 2021; Meng et al., 2020). While these studies offer valuable evidence regarding spatial heterogeneity, the non-linear dynamics of these responses – specifically, the saturation thresholds where urban warming-induced phenological effects diminish, and how background aridity modulates these thresholds – remain inadequately quantified across climatic gradients. Identifying such thresholds is critical for predicting urban ecosystem stability under continuous global warming scenarios.</p>
      <p id="d2e261">The magnitude of urban-induced phenological shifts is not uniform. Background climate heterogeneity constrains vegetation responses to additional urban warming: phenological advancements are strong in cold regions but weaken as baseline temperatures rise, implying that phenological sensitivity is a variable state conditioned by local thermal conditions. In ecology, the “saturation effect” describes the phenomenon whereby stimulus efficacy declines beyond a threshold, rooted in tolerance curve theory (Angilletta, 2009): organismal physiological functions increase with environmental drivers until reaching physiological saturation, beyond which no further enhancement occurs (Ketola and Kristensen, 2017). Plant physiological processes (photosynthesis, respiration, transpiration) do not increase linearly with temperature; they slow down or even decrease beyond a thermal threshold (Sage and Kubien, 2007; Ge et al., 2021). Meta-analysis has shown greater positive growth responses to warming in colder ecosystems (Rustad et al., 2001; Büntgen et al., 2019). We therefore hypothesize that the phenological effects of urban warming (advanced SOS, delayed EOS) gradually approach saturation in warmer regions, where additional UHI warming no longer promotes phenological shifts. This hypothesis remains to be tested in urban ecosystems.</p>
      <p id="d2e264">Drought disrupts temperature-driven phenological responses (Peng et al., 2019; Yuan et al., 2020). Urban vegetation is particularly vulnerable to moisture deficits due to the “Urban Dry Island” effect (reduced soil moisture and elevated evaporation in built environments) and impervious surfaces (Zhang et al., 2019). UHI warming combined with high aridity elevates vapor pressure deficit and water demand, triggering defensive mechanisms (such as abscisic acid (ABA) accumulation and protein degradation) (Brodribb and McAdam, 2013) that offset thermal advancements. Drought delays spring SOS and advances autumn EOS by inducing water stress and early dormancy (Brodribb and McAdam, 2013; Čehulić et al., 2019). However, in urban environments, this drought effect is compounded by the Urban Dry Island and impervious surfaces, further attenuating phenological sensitivity to extra warming, diminishing the positive impacts of UHI; this effect is stronger in warmer cities where plants approach thermal limits.</p>
      <p id="d2e268">Consequently, we hypothesize that background aridity (AI) acts as a critical negative regulator of the thermal saturation threshold. Specifically, we predict that as background aridity increases, the “saturation point” at which warming-induced phenological effects (such as advanced SOS or delayed EOS) plateau will occur at lower LST values. This implies that the saturation effect is not a fixed thermal threshold but a dynamic state conditioned by water stress, leading to a more rapid decline in phenological sensitivity in arid urban environments.</p>
      <p id="d2e271">Furthermore, phenological sensitivity to environmental drivers is species-specific and modulated by functional traits. Broad-leaf and coniferous species show divergent responses to warming and drought due to differences in leaf morphology and hydraulic architecture (Zheng et al., 2022). However, whether the saturation effect and “aridity-driven inhibition” are consistent across urban vegetation types remains unclear.</p>
      <p id="d2e274">As a rapidly urbanizing country with diverse climates, China faces severe UHI challenges (He et al., 2017). UHI intensity averages 0.9 <inline-formula><mml:math id="M7" display="inline"><mml:mo>±</mml:mo></mml:math></inline-formula> 1.1 °C but varies drastically across regions (Li et al., 2021). Previous urban phenology studies have largely focused on linear temperature sensitivity, and have overlooked non-linear saturation effects and aridity interactions. No study has quantified critical thermal/arid saturation thresholds or clarified the coupling mechanism between pre-season temperature (<inline-formula><mml:math id="M8" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>, phenological sensitivity) and land surface temperature (LST, background climate) in urban ecosystems. China's large climatic gradients and rapid urbanization make it an ideal platform to test the saturation hypothesis and aridity modulation.</p>
      <p id="d2e291">Investigating plant phenological responses to urbanization across China's diverse climates would improve our understanding of vegetation acclimation capacity. However, how background temperature and aridity interact to determine the thresholds and “saturation” of phenological sensitivity to urban warming remains largely unquantified at the macro-scale. Addressing this gap is essential for predicting the stability of urban carbon sinks under future climate scenarios. To this end, the primary objectives of this study were to: (1) explore the spatial distribution of plant phenological responses to the urban heat island effect throughout mainland China; (2) quantify the saturation effect caused by the spatial variability of phenological responses correlated with background temperature across the cold-hot gradient; and (3) assess whether the phenological effects of urban warming would be counteracted in drought-affected areas.</p>
</sec>
<sec id="Ch1.S2">
  <label>2</label><title>Materials and Methods</title>
<sec id="Ch1.S2.SS1">
  <label>2.1</label><title>Study area</title>
      <p id="d2e309">In our study, we conducted an analysis covering a total of 293 prefecture-level cities in China (excluding Taiwan, autonomous prefectures, and other special administrative regions), based on data from the year 2020 (<uri>http://xzqh.mca.gov.cn/map</uri>, last access: 19 August 2026) (Fig. 1). We hypothesized that the urban warming effect is most pronounced in urban centers and gradually diminishes towards rural areas (Zhang et al., 2004). To determine the urbanization intensity of each city, nighttime light data from NPP-VIIRS (Table 1) were used (Elvidge et al., 2017), employing the comparison method. The optimal segmentation threshold was established based on the statistical yearbook (Fan et al., 2019). Moreover, urban built-up areas were identified using the comparison method and annual average data derived from NPP-VIIRS nighttime light data between 2013 and 2020. All data processing procedures were performed using ArcGIS 10.2. We created ten buffer zones extending to a radius of 20 km from each city center, with each zone representing a 2 km annulus (Fig. 1).</p>
      <p id="d2e315">This 20 km boundary was chosen because our preliminary analysis revealed that the urban warming footprint on phenological metrics (SOS and EOS) gradually diminishes to near zero beyond 15 km (Zhang et al., 2004; Figs. 1 and 2). Thus, defining the outermost 20 km buffer zone as the “Rural” baseline ensures a background environment largely free of urban heat island interference. The subscript (10) refers to the 10th buffer zone at 20 km from the urban center. Equation notations use a standardized buffer index (0 for urban center, 10 for rural reference) to avoid confusion between distance and buffer number.</p>

      <fig id="F1"><label>Figure 1</label><caption><p id="d2e320">City centers (red area) and buffer zones (blue area) of the 293 prefecture-level cities in China. The map, generated using ArcGIS version 10.2 (<uri>https://www.esri.com/about/newsroom/arcwatch/the-best-of-arcgis-10-2/</uri>, last access: 19 August 2026), features administrative boundaries sourced from the Ministry of Civil Affairs of the People's Republic of China (<uri>http://xzqh.mca.gov.cn/map</uri>, last access: 19 August 2026). Source: Esri, TomTom, FAO, USGS, and the GIS User Community; Ministry of Civil Affairs of the People's Republic of China <inline-formula><mml:math id="M9" display="inline"><mml:mo>|</mml:mo></mml:math></inline-formula> Powered by Esri.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/5943/2026/bg-23-5943-2026-f01.png"/>

        </fig>

</sec>
<sec id="Ch1.S2.SS2">
  <label>2.2</label><title>Vegetation phenology extraction</title>
      <p id="d2e350">Phenology data spanning the years 2010 to 2020 were derived from MODIS Enhanced Vegetation Index (EVI) 16 d composite data with a spatial resolution of 250 m (MOD13Q1) and processed with TIMESAT (Jönsson and Eklundh, 2004). To mitigate challenges such as cloud cover and snow, the raw EVI time series data were smoothed using the Savitzky-Golay (S-G) filter method (Cai et al., 2017). Subsequently, a dynamic threshold technique was employed to extract the phenology data, utilizing a seasonal amplitude of 20 %, a parameter proven effective for phenology extraction in urban environments (Zhou et al., 2016; Buyantuyev and Wu, 2012; Cong et al., 2012; White et al., 2009). The datasets underwent preprocessing and quality control to eliminate pixels with minimal vegetation EVI (e.g., bodies of water and fully paved surfaces). To exclude unrealistic phenological values, we retained only SOS and EOS values within valid ranges (30–180 d for SOS and 210–360 d for EOS), consistent with previous studies (Zhang et al., 2006). To reduce the influence of outlier years, we averaged the phenology data for each pixel over 2010–2020. After applying these quality control thresholds and land-cover filtering criteria, approximately 48.5 % of the initial pixels within the study buffer zones were excluded.</p>
      <p id="d2e353">Furthermore, we employed land cover data from the MCD12Q1 dataset, which has a 500 m resolution, to mitigate the impact of agricultural practices on the mainland during our analysis. The MCD12Q1 dataset categorizes land into 17 distinct classes. These include various types of forested areas such as Evergreen Coniferous Forests, Deciduous Needle-leaf Forests, Evergreen Broad-leaf Forests, Deciduous Broad-leaf Forests, Mixed Forests, as well as Closed Shrub lands, Open Shrub lands, Savannas, Woody Savannas, Grasslands, and Croplands.</p>
      <p id="d2e356">Given the frequent human-induced disturbances in rural croplands and the regular manual pruning of urban grasslands, we excluded savannas, woody savannas, grasslands, and croplands from our analysis. The remaining land types were consolidated into broader categories: broad-leaf forests (BF, deciduous and evergreen broadleaf forests), coniferous forests (CF, deciduous and evergreen coniferous forests), mixed forests (MF), and shrubs (closed and open shrublands). This classification approach allowed us to focus our analysis on land types that are less influenced by human activities.</p>

<table-wrap id="T1" specific-use="star"><label>Table 1</label><caption><p id="d2e363">Data inventory used in this study.</p></caption><oasis:table frame="topbot"><oasis:tgroup cols="5">
     <oasis:colspec colnum="1" colname="col1" align="left"/>
     <oasis:colspec colnum="2" colname="col2" align="left"/>
     <oasis:colspec colnum="3" colname="col3" align="left"/>
     <oasis:colspec colnum="4" colname="col4" align="left"/>
     <oasis:colspec colnum="5" colname="col5" align="left"/>
     <oasis:thead>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1">Data type</oasis:entry>
         <oasis:entry colname="col2">Resolution</oasis:entry>
         <oasis:entry colname="col3">Layer</oasis:entry>
         <oasis:entry colname="col4">Data sources</oasis:entry>
         <oasis:entry colname="col5">Year</oasis:entry>
       </oasis:row>
     </oasis:thead>
     <oasis:tbody>
       <oasis:row>
         <oasis:entry colname="col1">MOD13Q1</oasis:entry>
         <oasis:entry colname="col2">250 m/16 d</oasis:entry>
         <oasis:entry colname="col3">EVI</oasis:entry>
         <oasis:entry colname="col4"><uri>https://search.earthdata.nasa.gov</uri></oasis:entry>
         <oasis:entry colname="col5">2010–2020</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">MOD11A2</oasis:entry>
         <oasis:entry colname="col2">1000 m/8 d</oasis:entry>
         <oasis:entry colname="col3">LST_Day LST_Night</oasis:entry>
         <oasis:entry colname="col4"><uri>https://search.earthdata.nasa.gov</uri></oasis:entry>
         <oasis:entry colname="col5">2010–2020</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">MOD16A2</oasis:entry>
         <oasis:entry colname="col2">500 m/8 d</oasis:entry>
         <oasis:entry colname="col3">PET</oasis:entry>
         <oasis:entry colname="col4"><uri>https://search.earthdata.nasa.gov</uri></oasis:entry>
         <oasis:entry colname="col5">2010–2020</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">MCD12Q1</oasis:entry>
         <oasis:entry colname="col2">500 m</oasis:entry>
         <oasis:entry colname="col3">Land Cover</oasis:entry>
         <oasis:entry colname="col4"><uri>https://search.earthdata.nasa.gov</uri></oasis:entry>
         <oasis:entry colname="col5">2010–2020</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">VIIRS</oasis:entry>
         <oasis:entry colname="col2">500 m</oasis:entry>
         <oasis:entry colname="col3">yearly average</oasis:entry>
         <oasis:entry colname="col4"><uri>https://eogdata.mines.edu</uri></oasis:entry>
         <oasis:entry colname="col5">2013–2020</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Precipitation data</oasis:entry>
         <oasis:entry colname="col2">1000 m</oasis:entry>
         <oasis:entry colname="col3">monthly average</oasis:entry>
         <oasis:entry colname="col4"><uri>http://www.geodata.cn</uri></oasis:entry>
         <oasis:entry colname="col5">2010–2020</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">China's urban zoning data</oasis:entry>
         <oasis:entry colname="col2"/>
         <oasis:entry colname="col3"/>
         <oasis:entry colname="col4"><uri>https://www.resdc.cn</uri></oasis:entry>
         <oasis:entry colname="col5">2020</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Station climates</oasis:entry>
         <oasis:entry colname="col2"/>
         <oasis:entry colname="col3">monthly average</oasis:entry>
         <oasis:entry colname="col4"><uri>http://data.cma.cn/en</uri></oasis:entry>
         <oasis:entry colname="col5">2010–2020</oasis:entry>
       </oasis:row>
     </oasis:tbody>
   </oasis:tgroup></oasis:table><table-wrap-foot><p id="d2e366">All URLs were last accessed on 19 August 2026.</p></table-wrap-foot></table-wrap>

</sec>
<sec id="Ch1.S2.SS3">
  <label>2.3</label><title>Pre-season temperature and background climates</title>
      <p id="d2e567">Building upon the methodology of a previous study (Piao et al., 2006), we defined pre-season temperature (<inline-formula><mml:math id="M10" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>) as the mean temperature for the three months preceding phenological onset. To align the temperature data with the phenological data, the average temperature dataset was resampled to a 250 m resolution using the nearest neighbor interpolation technique. To evaluate the performance of the downscaling procedure, the observed long-term monthly temperature and precipitation across China were obtained from the National Meteorological Information Center of China (<uri>http://data.cma.cn/en</uri>, last access: 19 August 2026). This dataset included observations from 496 national weather stations during 2010 to 2020. Upon conducting a fitting analysis with station data, we observed that the coefficient of determination (<inline-formula><mml:math id="M11" display="inline"><mml:mrow><mml:msup><mml:mi>R</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>) for monthly mean temperature and remote sensing data reached 0.93, while that for monthly precipitation and remote sensing data achieved 0.85. These results indicated that the downscaled meteorological data sufficiently met the requirements of this study.</p>
      <p id="d2e591">Background climate was characterized using land surface temperature (LST) and the aridity index (AI) as indicators. The LST and AI data for each pixel were averaged from 2010 to 2020. Specifically, MODIS LST datasets exhibiting an absolute bias of less than 1 K were selected as optimal for our investigation, in line with previous studies (Gow et al., 2016; Pablos et al., 2016; Qiao et al., 2013; Wan, 2008). Collection 6 LST products from the Terra satellite (MOD11A2, 8 d composite, 1 km spatial resolution, covering the period from 2010 to 2020) were used. Additionally, daily temperature (<inline-formula><mml:math id="M12" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>) was calculated as the mean of daytime and nighttime temperatures, following the methodology outlined by Kang et al. (2016). AI was expressed as:

            <disp-formula id="Ch1.E1" content-type="numbered"><label>1</label><mml:math id="M13" display="block"><mml:mrow><mml:mi mathvariant="normal">AI</mml:mi><mml:mo>=</mml:mo><mml:mstyle displaystyle="true"><mml:mfrac style="display"><mml:mi mathvariant="normal">PET</mml:mi><mml:mi>P</mml:mi></mml:mfrac></mml:mstyle></mml:mrow></mml:math></disp-formula>

          where PET denotes the annual potential evapotranspiration and <inline-formula><mml:math id="M14" display="inline"><mml:mi>P</mml:mi></mml:math></inline-formula> denotes the annual precipitation. The PET values spanning from 2010 to 2020 were extracted from the MOD16A2 monthly synthetic product, characterized by an 8 d temporal resolution and a spatial resolution of 500 m (Mu et al., 2011), as delineated in Table 1. The annual precipitation (<inline-formula><mml:math id="M15" display="inline"><mml:mi>P</mml:mi></mml:math></inline-formula>) dataset was derived from the monthly precipitation records of China encompassing the timeframe from 1901 to 2020, at a spatial resolution of 1 km. Subsequently, both PET and <inline-formula><mml:math id="M16" display="inline"><mml:mi>P</mml:mi></mml:math></inline-formula> datasets were resampled to a spatial resolution of 250 m to facilitate the ensuing spatial analysis (Peng, 2020).</p>
      <p id="d2e638">To robustly quantify these responses, we defined the distinct analytical roles of two critical thermal dimensions: pre-season air temperature (<inline-formula><mml:math id="M17" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>) and land surface temperature (LST). As the direct biological driver of plant physiology (e.g., chilling requirements), <inline-formula><mml:math id="M18" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula> was used exclusively to quantify the pure biological phenological sensitivity (<inline-formula><mml:math id="M19" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mi>t</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>) and identify true physiological thresholds. In contrast, as the standard metric for quantifying the urban heat island (UHI) effect, LST was used to characterize the spatial “background climate” of each city across the continent.</p>
      <p id="d2e666">Because pre-season air temperature (<inline-formula><mml:math id="M20" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>) is the direct driver of phenological sensitivity, we first derived the biological thresholds using <inline-formula><mml:math id="M21" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula> (Fig. 5). However, to address our core objective of quantifying how background LST shapes these phenological shifts, we defined the quantitative relationship between the background LST and <inline-formula><mml:math id="M22" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>. We fitted the relationship between LST and <inline-formula><mml:math id="M23" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula> across all cities. To minimize data matching biases during this conversion, we used the spatiotemporally aligned 10-year climatological means (2010–2020) for both variables at a unified 250 m resolution. We observed a highly robust linear correlation (Fig. S4). Based on this high goodness-of-fit, we applied the empirical conversion formula between the two variables to mathematically replace <inline-formula><mml:math id="M24" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula> with LST in the sensitivity curves (Wan, 2008). This empirical translation allowed us to systematically convert the <inline-formula><mml:math id="M25" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>-driven physiological thresholds into the specific LST-determined background climate thresholds (12.5 °C for SOS and 4 °C for EOS), explicitly projecting pure biological limits onto the urban map. Although <inline-formula><mml:math id="M26" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula> and LST represent different physical properties, they are highly correlated in urban settings (<inline-formula><mml:math id="M27" display="inline"><mml:mrow><mml:msup><mml:mi>R</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup><mml:mo>=</mml:mo><mml:mn mathvariant="normal">0.93</mml:mn></mml:mrow></mml:math></inline-formula> for monthly mean temperature), confirming the reliability of this empirical conversion between physiological response and climatic background.</p>
</sec>
<sec id="Ch1.S2.SS4">
  <label>2.4</label><title>Urbanization effect on phenology</title>
      <p id="d2e742">The phenology dates of each vegetation type were extracted in each buffer zone. Finally, the mean phenology in each buffer zone was calculated as a weighted mean of the phenology of each vegetation type. To ensure comparability in phenological analyses across buffer zones, we first identified the common vegetation types present in all zones and excluded the non-shared types. We then calculated the phenological means for these common types within each buffer zone and averaged these values to determine the overall mean vegetation phenology for the entire buffer zone, as shown in Eqs. (2) and (3).

                <disp-formula specific-use="gather" content-type="numbered"><mml:math id="M28" display="block"><mml:mtable displaystyle="true"><mml:mlabeledtr id="Ch1.E2"><mml:mtd><mml:mtext>2</mml:mtext></mml:mtd><mml:mtd><mml:mrow><mml:mstyle displaystyle="true" class="stylechange"/><mml:msub><mml:mi mathvariant="normal">SOS</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mo>=</mml:mo><mml:mstyle displaystyle="true"><mml:mfrac style="display"><mml:mrow><mml:mo>∑</mml:mo><mml:msub><mml:mi mathvariant="normal">SOS</mml:mi><mml:mrow><mml:mi>i</mml:mi><mml:mi>j</mml:mi></mml:mrow></mml:msub></mml:mrow><mml:mi>n</mml:mi></mml:mfrac></mml:mstyle></mml:mrow></mml:mtd></mml:mlabeledtr><mml:mlabeledtr id="Ch1.E3"><mml:mtd><mml:mtext>3</mml:mtext></mml:mtd><mml:mtd><mml:mrow><mml:mstyle class="stylechange" displaystyle="true"/><mml:msub><mml:mi mathvariant="normal">EOS</mml:mi><mml:mi>i</mml:mi></mml:msub><mml:mo>=</mml:mo><mml:mstyle displaystyle="true"><mml:mfrac style="display"><mml:mrow><mml:mo>∑</mml:mo><mml:msub><mml:mi mathvariant="normal">EOS</mml:mi><mml:mrow><mml:mi>i</mml:mi><mml:mi>j</mml:mi></mml:mrow></mml:msub></mml:mrow><mml:mi>n</mml:mi></mml:mfrac></mml:mstyle></mml:mrow></mml:mtd></mml:mlabeledtr></mml:mtable></mml:math></disp-formula>

          SOS<sub><italic>i</italic></sub> and EOS<sub><italic>i</italic></sub> arethe mean SOS and EOS of the <inline-formula><mml:math id="M31" display="inline"><mml:mi>i</mml:mi></mml:math></inline-formula>th buffer zone; SOS<sub><italic>i</italic><italic>j</italic></sub> and EOS<sub><italic>i</italic><italic>j</italic></sub> are the mean SOS and EOS of the <inline-formula><mml:math id="M34" display="inline"><mml:mi>j</mml:mi></mml:math></inline-formula>th vegetation type in the <inline-formula><mml:math id="M35" display="inline"><mml:mi>i</mml:mi></mml:math></inline-formula>th buffer zone; <inline-formula><mml:math id="M36" display="inline"><mml:mi>n</mml:mi></mml:math></inline-formula> is the number of shared vegetation types across all buffer zones.</p>
      <p id="d2e880">To investigate the impact of urbanization on phenology, mean values of start of season (SOS) and end of season (EOS) in each buffer zone were analyzed during 2010–2020, and fitted with respect to their distance from the urban center. In this study, the outermost buffer zone (20 km from the urban center) was defined as the rural reference region, and the urban center was defined as the urban region. This delineation was justified by the observation that phenological change rates approached zero at 15 km and beyond (Fig. S1), and similar patterns were observed for all four vegetation types (Fig. S2). Following prior investigations, the phenological sensitivity (<inline-formula><mml:math id="M37" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mi>t</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>) to temperature was quantified by calculating the partial correlation coefficient between <inline-formula><mml:math id="M38" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula> and the phenology dates (<inline-formula><mml:math id="M39" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-SOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> and <inline-formula><mml:math id="M40" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-EOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>) (Meng et al., 2020). The phenological difference between urban and rural regions was calculated to quantify the magnitude of the urban warming effect on phenology:

                <disp-formula specific-use="gather" content-type="numbered"><mml:math id="M41" display="block"><mml:mtable displaystyle="true"><mml:mlabeledtr id="Ch1.E4"><mml:mtd><mml:mtext>4</mml:mtext></mml:mtd><mml:mtd><mml:mrow><mml:mstyle displaystyle="true" class="stylechange"/><mml:mi mathvariant="normal">Δ</mml:mi><mml:mi mathvariant="normal">SOS</mml:mi><mml:mo>=</mml:mo><mml:msub><mml:mi mathvariant="normal">SOS</mml:mi><mml:mrow><mml:mo>(</mml:mo><mml:mn mathvariant="normal">0</mml:mn><mml:mo>)</mml:mo></mml:mrow></mml:msub><mml:mo>-</mml:mo><mml:msub><mml:mi mathvariant="normal">SOS</mml:mi><mml:mrow><mml:mo>(</mml:mo><mml:mn mathvariant="normal">10</mml:mn><mml:mo>)</mml:mo></mml:mrow></mml:msub></mml:mrow></mml:mtd></mml:mlabeledtr><mml:mlabeledtr id="Ch1.E5"><mml:mtd><mml:mtext>5</mml:mtext></mml:mtd><mml:mtd><mml:mrow><mml:mstyle class="stylechange" displaystyle="true"/><mml:mi mathvariant="normal">Δ</mml:mi><mml:mi mathvariant="normal">EOS</mml:mi><mml:mo>=</mml:mo><mml:msub><mml:mi mathvariant="normal">EOS</mml:mi><mml:mrow><mml:mo>(</mml:mo><mml:mn mathvariant="normal">0</mml:mn><mml:mo>)</mml:mo></mml:mrow></mml:msub><mml:mo>-</mml:mo><mml:msub><mml:mi mathvariant="normal">EOS</mml:mi><mml:mrow><mml:mo>(</mml:mo><mml:mn mathvariant="normal">10</mml:mn><mml:mo>)</mml:mo></mml:mrow></mml:msub></mml:mrow></mml:mtd></mml:mlabeledtr></mml:mtable></mml:math></disp-formula>

          where SOS<sub>(0)</sub> and SOS<sub>(10)</sub> are the mean SOS of the urban center (buffer index 0) and the 10th buffer zone (located 20 km away), respectively. Similarly, EOS<sub>(0)</sub> and EOS<sub>(10)</sub> are the mean EOS of the urban center and the 10th buffer zone. Biologically, a negative <inline-formula><mml:math id="M46" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS indicates an advanced start of the season (spring) in the urban center relative to the rural area, while a positive <inline-formula><mml:math id="M47" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS indicates a delayed end of the season (autumn senescence) in the urban center.</p>
      <p id="d2e1076">Similarly, the difference in phenological sensitivity to temperature between urban and rural regions was defined as:

                <disp-formula specific-use="gather" content-type="numbered"><mml:math id="M48" display="block"><mml:mtable displaystyle="true"><mml:mlabeledtr id="Ch1.E6"><mml:mtd><mml:mtext>6</mml:mtext></mml:mtd><mml:mtd><mml:mrow><mml:mstyle displaystyle="true" class="stylechange"/><mml:mi mathvariant="normal">Δ</mml:mi><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-SOS</mml:mtext></mml:mrow></mml:msub><mml:mo>=</mml:mo><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-SOS(0)</mml:mtext></mml:mrow></mml:msub><mml:mo>-</mml:mo><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-SOS(10)</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:mtd></mml:mlabeledtr><mml:mlabeledtr id="Ch1.E7"><mml:mtd><mml:mtext>7</mml:mtext></mml:mtd><mml:mtd><mml:mrow><mml:mstyle class="stylechange" displaystyle="true"/><mml:mi mathvariant="normal">Δ</mml:mi><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-EOS</mml:mtext></mml:mrow></mml:msub><mml:mo>=</mml:mo><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-EOS(0)</mml:mtext></mml:mrow></mml:msub><mml:mo>-</mml:mo><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-EOS(10)</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:mtd></mml:mlabeledtr></mml:mtable></mml:math></disp-formula>

          where <inline-formula><mml:math id="M49" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-SOS(0)</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> and <inline-formula><mml:math id="M50" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-SOS(10)</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> are the mean <inline-formula><mml:math id="M51" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-SOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> of the urban center (buffer index 0) and the 10th buffer zone (located 20 km away), respectively. Similarly, <inline-formula><mml:math id="M52" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-EOS(0)</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> andR<sub><italic>t</italic>-EOS(10)</sub> are the mean <inline-formula><mml:math id="M54" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-EOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> of the urban center and the 10th buffer zone.</p>
</sec>
<sec id="Ch1.S2.SS5">
  <label>2.5</label><title>Background climate determines phenological responses</title>
      <p id="d2e1251">To elucidate the distinct impacts of LST and AI on urban and rural components, we performed a partial correlation analysis. Specifically, the average LST or AI values from 2010 to 2020 for each of the 293 cities were correlated with their respective <inline-formula><mml:math id="M55" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS, <inline-formula><mml:math id="M56" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS, <inline-formula><mml:math id="M57" display="inline"><mml:mrow><mml:mi mathvariant="normal">Δ</mml:mi><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-SOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>, and <inline-formula><mml:math id="M58" display="inline"><mml:mrow><mml:mi mathvariant="normal">Δ</mml:mi><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-EOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> values, while controlling for the influence of the other climatic factor.</p>
      <p id="d2e1300">Path analysis (PA) (Doncaster, 2007) was used to examine both the direct and indirect effects of background climate variables (LST and AI) on phenological responses (SOS and EOS), as well as their corresponding discrepancies (<inline-formula><mml:math id="M59" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS and <inline-formula><mml:math id="M60" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS) and rates of change (<inline-formula><mml:math id="M61" display="inline"><mml:mrow><mml:mi mathvariant="normal">Δ</mml:mi><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-SOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> and <inline-formula><mml:math id="M62" display="inline"><mml:mrow><mml:mi mathvariant="normal">Δ</mml:mi><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-EOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>).</p>
      <p id="d2e1349">In the path analysis, we first assessed model fit using fit indices, including the chi-square (<inline-formula><mml:math id="M63" display="inline"><mml:mrow><mml:msup><mml:mi mathvariant="italic">χ</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>) statistic, Comparative Fit Index (CFI), and Root Mean Square Error of Approximation (RMSEA). Then, path coefficients were derived for both the direct and indirect effects of each independent variable on the dependent variable from the model. Squared path coefficients were used to estimate the respective direct and indirect contributions of each independent variable to the dependent variable. The sum of these direct and indirect contributions represents the total contribution, that is, the relative contribution.</p>
</sec>
<sec id="Ch1.S2.SS6">
  <label>2.6</label><title>Validation of phenology “saturation effect”</title>
      <p id="d2e1371">To determine if there was a temperature saturation effect on vegetation phenology, we conducted a model comparison. Rather than assuming a single relationship a priori, we evaluated the fit of phenology versus pre-season temperature using competing models: a linear model, a quadratic model, a segmented (piecewise) model, and a logistic model. We specifically selected the logistic function as our primary model because quadratic models biologically imply a symmetric reversal (a continuous steep decline) rather than a plateau, and piecewise models assume an abrupt, angular change in biological rates, which is less physiologically realistic than the smooth, asymptotic transition captured by the logistic curve (Pinheiro and Bates, 2000). The asymptote of the logistic function mathematically represents the biological “saturation point” – the maximum physiological limit of thermal responses.</p>
      <p id="d2e1374">Statistically, the model comparison results (Table 2) confirmed that the logistic model consistently outperforms the other three formulations for both spring SOS and autumn EOS, providing quantitative support for the non-linear saturation effect interpretation. For SOS, the logistic model yields an AIC value 2778 units lower than the linear model and 1424 units lower than the piecewise model, along with an <inline-formula><mml:math id="M64" display="inline"><mml:mrow><mml:msup><mml:mi>R</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula> of 0.37, which is approximately 19 % higher than that of the piecewise model. For EOS, the logistic model still achieves the lowest AIC and highest <inline-formula><mml:math id="M65" display="inline"><mml:mrow><mml:msup><mml:mi>R</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>, but the performance improvement is more modest: its <inline-formula><mml:math id="M66" display="inline"><mml:mrow><mml:msup><mml:mi>R</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula> (0.40) is only marginally higher than that of the piecewise model (0.39), consistent with a weaker but stable saturation pattern.</p>

<table-wrap id="T2" specific-use="star"><label>Table 2</label><caption><p id="d2e1413">Performance comparison of four competing models describing phenological responses to pre-season temperature.</p></caption><oasis:table frame="topbot"><oasis:tgroup cols="6">
     <oasis:colspec colnum="1" colname="col1" align="left"/>
     <oasis:colspec colnum="2" colname="col2" align="left"/>
     <oasis:colspec colnum="3" colname="col3" align="right"/>
     <oasis:colspec colnum="4" colname="col4" align="right"/>
     <oasis:colspec colnum="5" colname="col5" align="right"/>
     <oasis:colspec colnum="6" colname="col6" align="right"/>
     <oasis:thead>
       <oasis:row rowsep="1">

         <oasis:entry colname="col1">Phase</oasis:entry>

         <oasis:entry colname="col2">Model</oasis:entry>

         <oasis:entry colname="col3">AIC</oasis:entry>

         <oasis:entry colname="col4">BIC</oasis:entry>

         <oasis:entry colname="col5"><inline-formula><mml:math id="M67" display="inline"><mml:mrow><mml:msup><mml:mi>R</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula></oasis:entry>

         <oasis:entry colname="col6">RMSE</oasis:entry>

       </oasis:row>
     </oasis:thead>
     <oasis:tbody>
       <oasis:row>

         <oasis:entry rowsep="1" colname="col1" morerows="3">Spring_SOS</oasis:entry>

         <oasis:entry colname="col2">Linear</oasis:entry>

         <oasis:entry colname="col3">104 203.62</oasis:entry>

         <oasis:entry colname="col4">104 218.97</oasis:entry>

         <oasis:entry colname="col5">0.25</oasis:entry>

         <oasis:entry colname="col6">26.52</oasis:entry>

       </oasis:row>
       <oasis:row>

         <oasis:entry colname="col2">Quadratic</oasis:entry>

         <oasis:entry colname="col3">103 636.33</oasis:entry>

         <oasis:entry colname="col4">103 659.35</oasis:entry>

         <oasis:entry colname="col5">0.27</oasis:entry>

         <oasis:entry colname="col6">26.05</oasis:entry>

       </oasis:row>
       <oasis:row>

         <oasis:entry colname="col2">Piecewise</oasis:entry>

         <oasis:entry colname="col3">102 849.67</oasis:entry>

         <oasis:entry colname="col4">102 880.36</oasis:entry>

         <oasis:entry colname="col5">0.31</oasis:entry>

         <oasis:entry colname="col6">25.41</oasis:entry>

       </oasis:row>
       <oasis:row rowsep="1">

         <oasis:entry colname="col2">Logistic</oasis:entry>

         <oasis:entry colname="col3">101 425.33</oasis:entry>

         <oasis:entry colname="col4">101 456.02</oasis:entry>

         <oasis:entry colname="col5">0.37</oasis:entry>

         <oasis:entry colname="col6">24.29</oasis:entry>

       </oasis:row>
       <oasis:row>

         <oasis:entry colname="col1" morerows="3">Autumn_EOS</oasis:entry>

         <oasis:entry colname="col2">Linear</oasis:entry>

         <oasis:entry colname="col3">93 252.94</oasis:entry>

         <oasis:entry colname="col4">93 268.34</oasis:entry>

         <oasis:entry colname="col5">0.23</oasis:entry>

         <oasis:entry colname="col6">17.39</oasis:entry>

       </oasis:row>
       <oasis:row>

         <oasis:entry colname="col2">Quadratic</oasis:entry>

         <oasis:entry colname="col3">90 241.5</oasis:entry>

         <oasis:entry colname="col4">90 264.6</oasis:entry>

         <oasis:entry colname="col5">0.36</oasis:entry>

         <oasis:entry colname="col6">15.86</oasis:entry>

       </oasis:row>
       <oasis:row>

         <oasis:entry colname="col2">Piecewise</oasis:entry>

         <oasis:entry colname="col3">89 309.4</oasis:entry>

         <oasis:entry colname="col4">89 340.2</oasis:entry>

         <oasis:entry colname="col5">0.39</oasis:entry>

         <oasis:entry colname="col6">15.41</oasis:entry>

       </oasis:row>
       <oasis:row>

         <oasis:entry colname="col2">Logistic</oasis:entry>

         <oasis:entry colname="col3">85 678.49</oasis:entry>

         <oasis:entry colname="col4">85 709.14</oasis:entry>

         <oasis:entry colname="col5">0.40</oasis:entry>

         <oasis:entry colname="col6">15.22</oasis:entry>

       </oasis:row>
     </oasis:tbody>
   </oasis:tgroup></oasis:table></table-wrap>

      <p id="d2e1625">We then extracted phenological information from all vegetation pixels across the country and generated scatter plots of phenology versus pre-season temperature. Next, we fitted logistic growth curves to these scatter plots and evaluated the assumption using the goodness-of-fit metrics, p-value and <inline-formula><mml:math id="M68" display="inline"><mml:mrow><mml:msup><mml:mi>R</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>. Subsequently, we applied the same logistic curve to characterize the phenological responses of urban and rural vegetation in the 293 different cities against their pre-season temperatures, testing whether the phenological response to temperature in urban areas also follows a logistic growth pattern. Finally, we replaced the pre-season temperature with the background temperature LST to determine how the phenological saturation effect is regulated by the background temperature.</p>
</sec>
</sec>
<sec id="Ch1.S3">
  <label>3</label><title>Results</title>
<sec id="Ch1.S3.SS1">
  <label>3.1</label><title>Spatial distribution of urban-rural phenological differences</title>
      <p id="d2e1655">Among the 293 cities analyzed, 84 % (<inline-formula><mml:math id="M69" display="inline"><mml:mrow><mml:mi>N</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">246</mml:mn></mml:mrow></mml:math></inline-formula>) exhibited an advanced SOS in urban areas compared to rural regions, while 82 % (<inline-formula><mml:math id="M70" display="inline"><mml:mrow><mml:mi>N</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">241</mml:mn></mml:mrow></mml:math></inline-formula>) showed a delayed EOS (Fig. S3). Spatially, cities located in high-latitude regions generally demonstrated more negative <inline-formula><mml:math id="M71" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS values than those in low-latitude regions (Fig. 2), indicating that urban warming advanced spring phenology most strongly in northern and western China (Fig. 2a, b). This pattern is consistent with the conventional understanding of earlier SOS in colder regions. Additionally, cities in coastal provinces exhibited larger <inline-formula><mml:math id="M72" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS values than inland cities (Fig. 2).</p>
      <p id="d2e1696">A strong positive correlation was observed between <inline-formula><mml:math id="M73" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS and the rural rate of SOS change (<inline-formula><mml:math id="M74" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-SOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>), suggesting that cities with more negative <inline-formula><mml:math id="M75" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-SOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> values exhibited larger urban-rural disparities (Fig. 3). Notably, cities with lower temperatures tended to have more negative <inline-formula><mml:math id="M76" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-SOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> and <inline-formula><mml:math id="M77" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS values.</p>
      <p id="d2e1755"><inline-formula><mml:math id="M78" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS also showed a significant increasing trend with latitude (Fig. 2), with northern cities displaying more positive <inline-formula><mml:math id="M79" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS values, indicating a delayed autumn phenology in urban areas (Fig. 2c, d). In contrast, the majority of cities in southern regions demonstrated negative <inline-formula><mml:math id="M80" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS values, signifying an advancement in autumn phenology. Furthermore, cities in the coastal provinces also exhibited larger <inline-formula><mml:math id="M81" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS values.</p>
      <p id="d2e1786">Similarly, a positive correlation was observed between <inline-formula><mml:math id="M82" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS and the rural rate of EOS change (<inline-formula><mml:math id="M83" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-EOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>). Positive and negative <inline-formula><mml:math id="M84" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS values were predominantly found in cold and warm regions, respectively (Fig. 3). Moreover, when <inline-formula><mml:math id="M85" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS <inline-formula><mml:math id="M86" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 0, both the absolute value of <inline-formula><mml:math id="M87" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS and <inline-formula><mml:math id="M88" display="inline"><mml:mrow><mml:mi mathvariant="normal">Δ</mml:mi><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-EOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> increased with temperature, while for <inline-formula><mml:math id="M89" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS <inline-formula><mml:math id="M90" display="inline"><mml:mo>&lt;</mml:mo></mml:math></inline-formula> 0, the absolute value of <inline-formula><mml:math id="M91" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS and <inline-formula><mml:math id="M92" display="inline"><mml:mrow><mml:mi mathvariant="normal">Δ</mml:mi><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-EOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> decreased with temperature.</p>

      <fig id="F2" specific-use="star"><label>Figure 2</label><caption><p id="d2e1895">Spatial distributions of urban-rural disparities in SOS (<inline-formula><mml:math id="M93" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS) <bold>(a, b)</bold> and in EOS (<inline-formula><mml:math id="M94" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS) <bold>(c, d)</bold> across mainland China. Mean land surface temperature (LST) is shown in grayscale for the study area, and the data reflect the 10-year mean for 2010–2020 for each city. <bold>(e)</bold> depicts the linear correlation between <inline-formula><mml:math id="M95" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS and latitude (<inline-formula><mml:math id="M96" display="inline"><mml:mrow><mml:msup><mml:mi>R</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup><mml:mo>=</mml:mo><mml:mn mathvariant="normal">0.35</mml:mn></mml:mrow></mml:math></inline-formula>, <inline-formula><mml:math id="M97" display="inline"><mml:mrow><mml:mi>p</mml:mi><mml:mo>&lt;</mml:mo><mml:mn mathvariant="normal">0.01</mml:mn></mml:mrow></mml:math></inline-formula>); <bold>(f)</bold> depicts the linear correlation between <inline-formula><mml:math id="M98" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS and latitude (<inline-formula><mml:math id="M99" display="inline"><mml:mrow><mml:msup><mml:mi>R</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup><mml:mo>=</mml:mo><mml:mn mathvariant="normal">0.46</mml:mn></mml:mrow></mml:math></inline-formula>, <inline-formula><mml:math id="M100" display="inline"><mml:mrow><mml:mi>p</mml:mi><mml:mo>&lt;</mml:mo><mml:mn mathvariant="normal">0.01</mml:mn></mml:mrow></mml:math></inline-formula>); <bold>(g)</bold> compares <inline-formula><mml:math id="M101" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS between coastal and non-coastal provinces (<sup>*</sup> <inline-formula><mml:math id="M103" display="inline"><mml:mrow><mml:mi>p</mml:mi><mml:mo>&lt;</mml:mo><mml:mn mathvariant="normal">0.05</mml:mn></mml:mrow></mml:math></inline-formula>); <bold>(h)</bold> compares <inline-formula><mml:math id="M104" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS between coastal and non-coastal provinces (<sup>**</sup> <inline-formula><mml:math id="M106" display="inline"><mml:mrow><mml:mi>p</mml:mi><mml:mo>&lt;</mml:mo><mml:mn mathvariant="normal">0.01</mml:mn></mml:mrow></mml:math></inline-formula>). The map, generated using ArcGIS version 10.2, features administrative boundaries sourced from the Ministry of Civil Affairs of the People's Republic of China (<uri>http://xzqh.mca.gov.cn/map</uri>, last access: 19 August 2026). Source: Esri, TomTom, FAO, USGS, and the GIS User Community; Ministry of Civil Affairs of the People's Republic of China <inline-formula><mml:math id="M107" display="inline"><mml:mo>|</mml:mo></mml:math></inline-formula> Powered by Esri.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/5943/2026/bg-23-5943-2026-f02.png"/>

        </fig>

      <fig id="F3" specific-use="star"><label>Figure 3</label><caption><p id="d2e2078">Relationship between urban-rural phenological difference and temperature sensitivity across 293 Chinese cities. <bold>(a)</bold> Urban-rural difference in start of growing season (<inline-formula><mml:math id="M108" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS) versus temperature sensitivity of SOS (<inline-formula><mml:math id="M109" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-SOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>). <bold>(b)</bold> Urban-rural difference in end of growing season (<inline-formula><mml:math id="M110" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS) versus temperature sensitivity of EOS (<inline-formula><mml:math id="M111" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-EOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>). Each point represents one of the 293 cities analyzed, colored by land surface temperature (LST, °C). Red solid lines indicate linear regression fits.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/5943/2026/bg-23-5943-2026-f03.png"/>

        </fig>

</sec>
<sec id="Ch1.S3.SS2">
  <label>3.2</label><title>Phenological responses to LST</title>
      <p id="d2e2144">As we compared urban and rural regions, logistic trends were observed for both SOS and EOS in relation to LST (Fig. 4). In urban areas where LST was below 12 °C, the SOS occurred notably earlier compared to rural areas, while urban areas exhibited a delayed EOS when LST was below 18.5 °C. At the low-LST (cold) end, SOS plateaued at 101.3 d in urban areas and 129.7 d in rural areas, while the EOS stabilized at 321.3 d for urban areas and 310.8 d for rural areas. In contrast, when LST exceeded the specific thresholds for urban-rural differences (LST <inline-formula><mml:math id="M112" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 18 °C for <inline-formula><mml:math id="M113" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS and LST <inline-formula><mml:math id="M114" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 18.5 °C for <inline-formula><mml:math id="M115" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS), the phenological disparities between urban and rural areas were no longer significant and sometimes even reversed (Fig. 4c, d). As LST increased, the urban and rural SOS stabilized at 71.7 d (Fig. 4a), and the urban and rural EOS stabilized at 327.4   and 332.4 d, respectively. Fitted curves of <inline-formula><mml:math id="M116" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS and <inline-formula><mml:math id="M117" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS against LST revealed a positive and a negative relationship, respectively, intersecting at LST <inline-formula><mml:math id="M118" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 18  and 18.5 °C, indicating a shift in SOS and EOS dynamics with increasing temperatures beyond these thresholds.</p>

      <fig id="F4" specific-use="star"><label>Figure 4</label><caption><p id="d2e2199">Relationship between background temperature (LST) and phenology (SOS and EOS) in urban and rural areas <bold>(a, b)</bold> for 2010 to 2020, and between <inline-formula><mml:math id="M119" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS and LST and between <inline-formula><mml:math id="M120" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS and LST <bold>(c, d)</bold>.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/5943/2026/bg-23-5943-2026-f04.png"/>

        </fig>

      <fig id="F5" specific-use="star"><label>Figure 5</label><caption><p id="d2e2230">Relationships between pre-season temperature (<inline-formula><mml:math id="M121" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>) and SOS <inline-formula><mml:math id="M122" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> EOS <bold>(a, b)</bold>, and the corresponding first-order derivatives with respect to temperature <bold>(c, d)</bold>, representing phenological sensitivity to temperature.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/5943/2026/bg-23-5943-2026-f05.png"/>

        </fig>

      <p id="d2e2260">To determine whether the temperature sensitivity influenced the logistic trends of phenology with LST, we conducted a nationwide fitting of phenology to <inline-formula><mml:math id="M123" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula> and derived logistic curves for SOS and EOS (Fig. 5). The first-order derivatives of the fitted curves, representing the phenological sensitivity to <inline-formula><mml:math id="M124" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>, revealed warming-induced phenological effects on SOS and EOS, ranging from <inline-formula><mml:math id="M125" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>3.5–10 °C and 2.0–13.5 °C, respectively. The highest sensitivity was observed at <inline-formula><mml:math id="M126" display="inline"><mml:mrow><mml:mi>T</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">4</mml:mn></mml:mrow></mml:math></inline-formula> °C for SOS and <inline-formula><mml:math id="M127" display="inline"><mml:mrow><mml:mi>T</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">6</mml:mn></mml:mrow></mml:math></inline-formula> °C for EOS. Notably, a decreasing trend for EOS was observed when <inline-formula><mml:math id="M128" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula> <inline-formula><mml:math id="M129" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 15 °C, corresponding to the advanced EOS when <inline-formula><mml:math id="M130" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula> <inline-formula><mml:math id="M131" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 18.5 °C (Fig. 5d).</p>
      <p id="d2e2337">To assess the temperature sensitivity of phenological variations between urban and rural settings across vegetation types, we established regression models of <inline-formula><mml:math id="M132" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS and <inline-formula><mml:math id="M133" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS against LST for each type (Fig. 6). Results showed that shrublands had the highest temperature sensitivity for both metrics, whereas CF displayed the lowest. The temperature response of MF and BF did not differ significantly.</p>

      <fig id="F6" specific-use="star"><label>Figure 6</label><caption><p id="d2e2356">Relationships between LST and <inline-formula><mml:math id="M134" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS <inline-formula><mml:math id="M135" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> <inline-formula><mml:math id="M136" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS across the 293 cities for four vegetation types. The inset subplots compare the mean phenological differences among vegetation groups. BF: Broad-leaf forest, CF: Coniferous forest, MF: Mixed forest, Shrubs: Shrublands.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/5943/2026/bg-23-5943-2026-f06.png"/>

        </fig>

</sec>
<sec id="Ch1.S3.SS3">
  <label>3.3</label><title>Quantify the LST determined phenological sensitivity</title>
      <p id="d2e2394">To obtain LST-determined phenological sensitivity, the <inline-formula><mml:math id="M137" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula> in the first-order derivative curves shown in Fig. 5c and d was converted to LST based on the strong correlation between <inline-formula><mml:math id="M138" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula> and LST (Fig. S4). Three distinct LST thresholds were identified: phenological date stabilization thresholds, urban-rural disparity reversal thresholds, and phenological sensitivity saturation thresholds. Specifically, phenological date stabilization and urban-rural disparity reversal occur at LST <inline-formula><mml:math id="M139" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 12 °C (SOS) <inline-formula><mml:math id="M140" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> 18.5 °C (EOS), while phenological sensitivity saturates at LST <inline-formula><mml:math id="M141" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 12.5 °C (SOS) <inline-formula><mml:math id="M142" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> 4 °C (EOS). Based on the first-order derivatives, we identified the specific thresholds for phenological sensitivity. We observed that when background LST exceeded 12.5 °C for spring and 4 °C for autumn, the temperature sensitivity (<inline-formula><mml:math id="M143" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mi>t</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>) began to weaken significantly. This indicates a saturation of the physiological response rate, even though the absolute urban-rural difference (<inline-formula><mml:math id="M144" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS <inline-formula><mml:math id="M145" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> <inline-formula><mml:math id="M146" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS) might still persist until higher temperatures are reached.</p>

      <fig id="F7" specific-use="star"><label>Figure 7</label><caption><p id="d2e2474">Schematic diagram of the three scenarios that likely occurred in urban (red dots) and rural (blue dots) regions for SOS (I: LST<inline-formula><mml:math id="M147" display="inline"><mml:mrow><mml:msub><mml:mi/><mml:mrow><mml:mi mathvariant="normal">R</mml:mi><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msub><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mo>&lt;</mml:mo></mml:mrow></mml:math></inline-formula> LST<inline-formula><mml:math id="M148" display="inline"><mml:mrow><mml:msub><mml:mi/><mml:mrow><mml:mi mathvariant="normal">U</mml:mi><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msub><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mo>&lt;</mml:mo></mml:mrow></mml:math></inline-formula> 12.5 °C; II: 12.5 °C <inline-formula><mml:math id="M149" display="inline"><mml:mo>&lt;</mml:mo></mml:math></inline-formula> LST<inline-formula><mml:math id="M150" display="inline"><mml:mrow><mml:msub><mml:mi/><mml:mrow><mml:mi mathvariant="normal">R</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:mrow></mml:msub><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mo>&lt;</mml:mo></mml:mrow></mml:math></inline-formula> LST<sub>U2</sub>; III: LST<inline-formula><mml:math id="M152" display="inline"><mml:mrow><mml:msub><mml:mi/><mml:mrow><mml:mi mathvariant="normal">R</mml:mi><mml:mn mathvariant="normal">3</mml:mn></mml:mrow></mml:msub><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mo>&lt;</mml:mo></mml:mrow></mml:math></inline-formula> 12.5 °C <inline-formula><mml:math id="M153" display="inline"><mml:mo>&lt;</mml:mo></mml:math></inline-formula> LST<sub>U3</sub>) and EOS (I: LST<inline-formula><mml:math id="M155" display="inline"><mml:mrow><mml:msub><mml:mi/><mml:mrow><mml:mi mathvariant="normal">R</mml:mi><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msub><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mo>&lt;</mml:mo></mml:mrow></mml:math></inline-formula> LST<inline-formula><mml:math id="M156" display="inline"><mml:mrow><mml:msub><mml:mi/><mml:mrow><mml:mi mathvariant="normal">U</mml:mi><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msub><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mo>&lt;</mml:mo></mml:mrow></mml:math></inline-formula> 4 °C; II: 4 °C <inline-formula><mml:math id="M157" display="inline"><mml:mo>&lt;</mml:mo></mml:math></inline-formula> LST<inline-formula><mml:math id="M158" display="inline"><mml:mrow><mml:msub><mml:mi/><mml:mrow><mml:mi mathvariant="normal">R</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:mrow></mml:msub><mml:mspace width="0.125em" linebreak="nobreak"/><mml:mo>&lt;</mml:mo></mml:mrow></mml:math></inline-formula> LST<sub>U2</sub>; III: LST<inline-formula><mml:math id="M160" display="inline"><mml:mrow><mml:msub><mml:mi/><mml:mrow><mml:mi mathvariant="normal">R</mml:mi><mml:mn mathvariant="normal">3</mml:mn></mml:mrow></mml:msub><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mo>&lt;</mml:mo></mml:mrow></mml:math></inline-formula> 4 °C <inline-formula><mml:math id="M161" display="inline"><mml:mo>&lt;</mml:mo></mml:math></inline-formula> LST<sub>U3</sub>). Scenario I: when LST was less than 12.5 °C (4 °C) for SOS (EOS), higher LST was associated with heightened temperature sensitivity, resulting in urban regions (U<sub>1</sub>) exhibiting higher sensitivity than rural regions (<inline-formula><mml:math id="M164" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mn mathvariant="normal">1</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>) (Fig. 5). Scenario II: when LST exceeded 12.5 °C (4 °C) for SOS (EOS), temperature sensitivity weakened as LST increased, yet urban regions (U<sub>2</sub>) still exhibited higher sensitivity than rural regions (<inline-formula><mml:math id="M166" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msub></mml:mrow></mml:math></inline-formula>) (Fig. 5). Scenario III: when urban LST surpassed 12.5 °C (4 °C) while rural LST remained below 12.5 °C (4 °C), the difference in sensitivity between urban and rural regions was inconclusive.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/5943/2026/bg-23-5943-2026-f07.png"/>

        </fig>

</sec>
<sec id="Ch1.S3.SS4">
  <label>3.4</label><title>Phenological responses to aridity</title>
      <p id="d2e2738">To characterize how phenological responses vary with background aridity, we fitted linear regressions of <inline-formula><mml:math id="M167" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-SOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>, <inline-formula><mml:math id="M168" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-EOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>, <inline-formula><mml:math id="M169" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS, and <inline-formula><mml:math id="M170" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS against the aridity index (AI) (Fig. 8). The zero-crossing points of the fitted regression lines (i.e., the AI value at which the regression-predicted response variable equals zero) were used to demarcate contrasting phenological response regimes. Because these response variables are signed, these zero-crossing points separate opposing phenological patterns (e.g., from warming-induced advancement to warming-induced delay of SOS).</p>
      <p id="d2e2783"><inline-formula><mml:math id="M171" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-SOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> was positively related to AI (Fig. 8a), with its fitted regression line crossing zero at AI <inline-formula><mml:math id="M172" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 1.7. When AI <inline-formula><mml:math id="M173" display="inline"><mml:mo>&lt;</mml:mo></mml:math></inline-formula> 1.7, the absolute value of <inline-formula><mml:math id="M174" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-SOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> decreased with increasing AI, indicating that increasing aridity weakened the temperature sensitivity of SOS. In contrast, when AI <inline-formula><mml:math id="M175" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 1.7, most urban areas exhibited positive <inline-formula><mml:math id="M176" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-SOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> (resulting in delayed SOS), with higher AI values corresponding to greater <inline-formula><mml:math id="M177" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-SOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>, suggesting that severe drought markedly delayed the onset of SOS.</p>
      <p id="d2e2863"><inline-formula><mml:math id="M178" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-EOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> was negatively related to AI (Fig. 8c), with its fitted regression line crossing zero at AI <inline-formula><mml:math id="M179" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 1.4. When AI <inline-formula><mml:math id="M180" display="inline"><mml:mo>&lt;</mml:mo></mml:math></inline-formula> 1.4, most <inline-formula><mml:math id="M181" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-EOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> values were positive and decreased with AI, i.e., intensified drought counteracted the EOS-delaying effect induced by urban warming. When AI <inline-formula><mml:math id="M182" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 1.4, most cities exhibited negative <inline-formula><mml:math id="M183" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-EOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>, and higher AI values corresponded to more negative <inline-formula><mml:math id="M184" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-EOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>. That is, extreme drought reversed the warming-delaying effect on EOS.</p>

      <fig id="F8" specific-use="star"><label>Figure 8</label><caption><p id="d2e2946">Relationships between the aridity index (AI) and <bold>(a)</bold> the temperature sensitivity of SOS (<inline-formula><mml:math id="M185" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-SOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>), <bold>(b)</bold> the urban-rural difference in SOS (<inline-formula><mml:math id="M186" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS), <bold>(c)</bold> the temperature sensitivity of EOS (<inline-formula><mml:math id="M187" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-EOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>), and <bold>(d)</bold> the urban-rural difference in EOS (<inline-formula><mml:math id="M188" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS). Dashed lines mark the AI thresholds (AI <inline-formula><mml:math id="M189" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 1.7, 1.4, and 2.0 in panels <bold>a</bold>, <bold>c</bold>, and <bold>d</bold>, respectively) at which the fitted regression lines cross zero.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/5943/2026/bg-23-5943-2026-f08.png"/>

        </fig>

      <p id="d2e3027">When urban-rural phenological differences were fitted to AI, <inline-formula><mml:math id="M190" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS was positively related to AI (Fig. 8b), indicating that increasing aridity progressively reduced the advanced green-up typically induced by urban warming. <inline-formula><mml:math id="M191" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS was negatively related to AI (Fig. 8d), with its fitted regression line crossing zero at AI <inline-formula><mml:math id="M192" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 2.0. Notably, negative <inline-formula><mml:math id="M193" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS tended to appear when AI <inline-formula><mml:math id="M194" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 2, suggesting that under extreme drought, urban EOS occurred earlier than rural EOS.</p>
      <p id="d2e3065">The same relationships were evident across all four vegetation types (Fig. 9): increasing AI was associated with a reduced advance of SOS (less negative <inline-formula><mml:math id="M195" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS) and an earlier urban EOS (more negative <inline-formula><mml:math id="M196" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS). Notably, <inline-formula><mml:math id="M197" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS for CF showed less sensitivity to AI than the other vegetation types. Both BF and MF demonstrated lower sensitivities to AI for both <inline-formula><mml:math id="M198" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS and <inline-formula><mml:math id="M199" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS than shrublands.</p>

      <fig id="F9" specific-use="star"><label>Figure 9</label><caption><p id="d2e3105">Relationships between AI and urban-rural phenological differences (<inline-formula><mml:math id="M200" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS, <inline-formula><mml:math id="M201" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS) across different vegetation types. <bold>(a)</bold> Scatter plot of <inline-formula><mml:math id="M202" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS versus AI with linear regression fits for each vegetation type. <bold>(b)</bold> Scatter plot of <inline-formula><mml:math id="M203" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS versus AI with linear regression fits for each vegetation type.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/5943/2026/bg-23-5943-2026-f09.png"/>

        </fig>

      <fig id="F10"><label>Figure 10</label><caption><p id="d2e3151">Venn diagrams illustrating the relative contributions of AI and LST to <inline-formula><mml:math id="M204" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS, <inline-formula><mml:math id="M205" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS, <inline-formula><mml:math id="M206" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-SOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>, and <inline-formula><mml:math id="M207" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-EOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> across 293 Chinese cities. Note: Values in each section represent the percentage of variance independently explained by AI or LST and their shared explanatory power in the overlapping region, and the total variance explained by both factors is labeled below each panel.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/5943/2026/bg-23-5943-2026-f10.png"/>

        </fig>

      <p id="d2e3203">To further quantify the individual and combined contributions of LST and AI to phenological variations, a variance decomposition analysis was conducted (Legendre and Legendre, 2012; Fig. 10). The Venn diagram revealed that AI and LST explained 75.05 %, 76.21 %, 71.72 % and 82.14 % of the variance in <inline-formula><mml:math id="M208" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS, <inline-formula><mml:math id="M209" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS, <inline-formula><mml:math id="M210" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-SOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>, and <inline-formula><mml:math id="M211" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-EOS,</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> respectively (Fig. 10), and LST always contributed more to these phenological indicators. Additionally, the interaction effect of LST and AI ranged from 3.29 % to 5.66 %.</p>
</sec>
</sec>
<sec id="Ch1.S4">
  <label>4</label><title>Discussion</title>
<sec id="Ch1.S4.SS1">
  <label>4.1</label><title>Overall urbanization effects on plant phenology in China</title>
      <p id="d2e3264">Our multi-city assessment confirms a predominant urban-rural phenological divergence across China, characterized by advanced SOS and delayed EOS within a concentrated 10 km urbanization footprint. This consistent spatial pattern confirms that urban warming acts as a primary driver of phenological shifts. Beyond mere thermal forcing, this convergence across 293 cities reflects the profound impact of urban microclimates on plant life cycles. The observed 10 km threshold is particularly significant as it suggests a localized but intense ecological footprint that necessitates specialized urban greening management strategies, especially in high-density northern cities where these shifts are most pronounced.</p>
      <p id="d2e3267">Spatially, the pronounced north-south gradient in <inline-formula><mml:math id="M212" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS and <inline-formula><mml:math id="M213" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS across China, as shown in Fig. 2, closely mirrors the spatial pattern of LST, reflecting stronger temperature sensitivity of vegetation phenology in northern regions than in the south. This finding aligns with previous research documenting greater phenological shifts in northern areas (Jia et al., 2021; Yang et al., 2025), and further highlights that background climate acts as a key modulator of urban phenological impacts. The more pronounced urban warming effects observed in coastal provinces relative to inland cities (Fig. 2) likely stem from the buffering effect of the maritime climate on seasonal temperature constraints, which interacts with urban heat islands to amplify urban-rural phenological divergence. Moreover, the strong correlations between <inline-formula><mml:math id="M214" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS <inline-formula><mml:math id="M215" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> <inline-formula><mml:math id="M216" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS, temperature sensitivity indicators (<inline-formula><mml:math id="M217" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-SOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> and <inline-formula><mml:math id="M218" display="inline"><mml:mrow><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-EOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>), LST, and AI, as presented in Fig. 3, confirm that background temperature and aridity are critical drivers shaping the magnitude of urbanization-induced phenological shifts across the 293 cities, rather than just passive correlates, validating the core hypothesis of this study.</p>
      <p id="d2e3334">Notably, the pattern of urban warming's effects on vegetation phenology was consistent among the four vegetation types (Fig. S2). This consistency could be attributed to the ecological convergence of plants coexisting in shared local habitats within each city (Wang et al., 2016). Ecologically, the consistent 10 km effective footprint validates that urban environments function as distinct microclimatic islands. This spatial boundary conceptually highlights that urban ecological engineering and greening strategies must account for highly localized, rather than regional, thermal forcings.</p>
</sec>
<sec id="Ch1.S4.SS2">
  <label>4.2</label><title>Background temperature related phenological response variance</title>
      <p id="d2e3345">When we quantified phenological responses to temperature across the climatic gradient, clear logistic trends were observed for both SOS and EOS (Fig. 5). Consistent with this logistic trajectory, we observed that in urban areas, the growth rate for SOS <inline-formula><mml:math id="M219" display="inline"><mml:mo>/</mml:mo></mml:math></inline-formula> EOS began to plateau when temperatures exceeded 4 °C/6 °C, respectively (Fig. 5). Given the significant correlation between LST and <inline-formula><mml:math id="M220" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula> (Fig. S4), we anticipated a diminished sensitivity in phenology with higher LST (Fig. 7). These findings lend credibility to the “saturation” effect hypothesis, suggesting that the accelerating influence of warmth on vegetation phenology reported in earlier studies has its limits (Badeck et al., 2004; Li et al., 2021). Indeed, plants in southern regions exhibit not just a dampened phenological response but also a decelerated growth rate when faced with increasing temperatures (Li et al., 2021; Büntgen et al., 2019). As illustrated by the first-order derivatives in Fig. 5c and d, the phenological sensitivity to temperature does not increase linearly, but peaks at <inline-formula><mml:math id="M221" display="inline"><mml:mrow><mml:mi>T</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">4</mml:mn></mml:mrow></mml:math></inline-formula> °C for SOS and <inline-formula><mml:math id="M222" display="inline"><mml:mrow><mml:mi>T</mml:mi><mml:mo>=</mml:mo><mml:mn mathvariant="normal">6</mml:mn></mml:mrow></mml:math></inline-formula> °C for EOS before rapidly declining. Specifically, the stabilization of urban SOS at 101.3 d and rural SOS at 129.7 d when LST was low (Fig. 4a) further demonstrated that plants reach a required physiological threshold that cannot be bypassed by additional thermal forcing. Collectively, these results highlight that threshold-based non-linear models rather than simple linear relationships are more suitable for predicting phenological responses under scenarios of continuous environmental warming, especially in urban settings. Conceptually, these saturation thresholds challenge the traditional linear paradigm of “warming-driven phenological advancement”. By identifying explicit thermal boundaries, our framework establishes that the phenological response of urban vegetation to additional warming is bounded.</p>
      <p id="d2e3386">The observation of temperature saturation for the SOS in our study likely stems from the chilling requirements for vernalization (Kim et al., 2009; Hanninen et al., 2019). Typically, vegetation needs to undergo a period of exposure to low chilling temperatures to break dormancy. This mechanism is consistent with our spatial observations in Fig. 2a, where cities in lower latitudes (warmer regions) exhibited significantly smaller <inline-formula><mml:math id="M223" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS values. This suggests that in these warmer regions, the higher background LST may fail to satisfy the necessary chilling requirements, leading to the diminished phenological sensitivity we quantified in Fig. 3. Moreover, an earlier onset of greening increases the risk of damage from freezing events, posing a survival threat to plants (Duan et al., 2011; Wang et al., 2015). This phenomenon explains the delayed SOS in approximately 16 % of the cities under study, mainly distributed in low-latitude regions (Fig. 4).</p>
      <p id="d2e3396">Regarding the EOS, a moderate increase in temperature can enhance the activity of photosynthetic enzymes (Shi et al., 2014), slow down the degradation of chlorophyll (Fracheboud et al., 2009), reduce the risk of frost exposure in autumn, and increase the potential for growth and photosynthetic production, ultimately promoting the extension of the growing season. However, the positive effects of increased temperature can be offset by the negative impacts of heat waves and excessive thermal stress (Mathur et al., 2014). Our data in Fig. 5b and d provide direct evidence for this: when pre-season temperature (<inline-formula><mml:math id="M224" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula>) exceeded 15 °C, a decreasing trend for EOS was observed. This aligns with the intersection point identified in Fig. 4d at LST <inline-formula><mml:math id="M225" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 18.5 °C, where the urban-rural difference in EOS reverses, confirming that excessive heat in urban centers triggers premature leaf senescence rather than further delaying it. In fact, when <inline-formula><mml:math id="M226" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula> <inline-formula><mml:math id="M227" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 15 °C, EOS tends to be reduced, with an increased sensitivity to temperature changes (Estiarte and Peñuelas, 2015; Fig. 5b).</p>
      <p id="d2e3427">Despite the existence of “saturation” effects, lower background temperatures do not necessarily correspond to higher phenological sensitivity (Fig. 4b, c). For SOS, when <inline-formula><mml:math id="M228" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula> <inline-formula><mml:math id="M229" display="inline"><mml:mo>&lt;</mml:mo></mml:math></inline-formula> 4 °C, the phenological sensitivity approached zero with decreased <inline-formula><mml:math id="M230" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula> (Fig. 5c), which led to a stable SOS of 120 d (Fig. 5a, c). In fact, phenological events such as bud burst could not be triggered when the threshold for high forcing temperatures was not reached (Hanninen et al., 2019). For EOS, when <inline-formula><mml:math id="M231" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula> <inline-formula><mml:math id="M232" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 6 °C, the increased <inline-formula><mml:math id="M233" display="inline"><mml:mi>T</mml:mi></mml:math></inline-formula> sensitivity produces more EOS advancement in warmer regions. This reversal may be associated with abscisic acid synthesis induced by low temperatures, which accelerates the progress of defoliation.</p>
      <p id="d2e3474">Beyond thermal controls alone, our findings regarding the aridity-driven modulation of phenological responses provide a critical geographic context that complements previous observations. While earlier studies primarily emphasized thermal forcing as the dominant driver of urban phenology (Meng et al., 2020), our results demonstrate that AI acts as a significant “environmental filter” that can override or even reverse warming-induced phenological effects. As shown in Figs. 8 and 10, the high explanatory power of AI (17.33 % for <inline-formula><mml:math id="M234" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS) suggests that in water-limited regions of northern and western China, the physiological stress induced by drought prevents plants from utilizing the additional heat provided by the Urban Heat Island (UHI) effect. This phenomenon is consistent with the “resource limitation” theory, where plant development is constrained by the most limiting resource (water; Harpole et al., 2011), regardless of optimal temperatures. Compared to coastal cities where water is abundant, the weakened warming-induced phenological effects in arid inland cities highlight the need for regionally differentiated urban irrigation strategies to maintain ecological productivity. However, when LST <inline-formula><mml:math id="M235" display="inline"><mml:mo>&lt;</mml:mo></mml:math></inline-formula> 12.5 °C, warming regions exhibited higher urban-rural differences in <inline-formula><mml:math id="M236" display="inline"><mml:mrow><mml:mi mathvariant="normal">Δ</mml:mi><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-SOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula>, which contradicted the weakened urban-rural difference in <inline-formula><mml:math id="M237" display="inline"><mml:mrow><mml:mi mathvariant="normal">Δ</mml:mi><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-SOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> reported in the previous study. For EOS, the warming-induced phenological effects only occurred when 0 <inline-formula><mml:math id="M238" display="inline"><mml:mo>&lt;</mml:mo></mml:math></inline-formula> LST <inline-formula><mml:math id="M239" display="inline"><mml:mo>&lt;</mml:mo></mml:math></inline-formula> 18 °C. Moreover, the weakened sensitivity was only detected at LST <inline-formula><mml:math id="M240" display="inline"><mml:mo>&gt;</mml:mo></mml:math></inline-formula> 4 °C. The weakened <inline-formula><mml:math id="M241" display="inline"><mml:mrow><mml:mi mathvariant="normal">Δ</mml:mi><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-SOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> and <inline-formula><mml:math id="M242" display="inline"><mml:mrow><mml:mi mathvariant="normal">Δ</mml:mi><mml:msub><mml:mi>R</mml:mi><mml:mrow><mml:mi>t</mml:mi><mml:mtext>-EOS</mml:mtext></mml:mrow></mml:msub></mml:mrow></mml:math></inline-formula> reported in the USA may reflect the fact that the LST range there was mostly within 8–22 °C.</p>
</sec>
<sec id="Ch1.S4.SS3">
  <label>4.3</label><title>AI related phenological response variance</title>
      <p id="d2e3585">In addition to background temperature (LST), the aridity index (AI) drives significant spatial variation in urban-rural phenological disparities (<inline-formula><mml:math id="M243" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS, <inline-formula><mml:math id="M244" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS). Spatially, the coastal cities experienced more pronounced warming-induced phenological effects (Fig. 2). Our statistical decomposition in Fig. 10 further elucidates this aridity-driven modulation, showing that AI and LST together explain more than 75 % of the variance in <inline-formula><mml:math id="M245" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS and <inline-formula><mml:math id="M246" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS. Notably, the fact that AI explains 17.33 % of the variation in <inline-formula><mml:math id="M247" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>EOS (Fig. 10c) supports our inference that water availability is a critical limiting factor in autumn. This is further substantiated by the positive correlation between <inline-formula><mml:math id="M248" display="inline"><mml:mi mathvariant="normal">Δ</mml:mi></mml:math></inline-formula>SOS and AI in Fig. 8b, which quantifies how increasing aridity progressively offsets the advanced green-up typically induced by urban warming. These results highlight that the warming-induced phenological effects on vegetation phenology will be weakened in those drought-dominated regions. In fact, the delayed SOS has already been demonstrated in previous studies (Ji et al., 2021; Yu et al., 2003).</p>
      <p id="d2e3631">Water deficit inhibits leaf growth via chemical signals (Knauer et al., 2017), and urbanization exacerbates drought stress (Zhang et al., 2019; Pollastrini et al., 2019), further weakening or even reversing the positive effects of urban warming on phenology. Furthermore, in water-stressed regions, temperatures are already close to the optimum for photosynthesis, thus fewer positive physiological effects are expected (Peñuelas et al., 2004). The weakened temperature sensitivity in drought-prone cities is likely to lead to less extension of the growing season.</p>
      <p id="d2e3634">Notably, the influence of AI on autumn phenology response was higher than that on spring phenology response (Fig. 10). During spring leaf development, plants may use water stored in their bodies to support the growth of new leaves (Shi et al., 2023). Supporting evidence comes from the synchrony between plant water storage and leaf sprouting for the boreal and temperate forests (Tian et al., 2018). In this case, plants may have relatively little demand for external water supply, especially in some wetter areas. By comparison, in summer and autumn, plant transpiration is higher, and water demand is greater (Wu et al., 2022), so the influence of drought severity on plant phenology is more pronounced. The overarching ecological implication here is a profound vulnerability: as background climates dry under global change, the compounding stress of UHI and background aridity may lead to “phenological compression”. In arid urban ecosystems, this moisture deficit effectively neutralizes the anticipated carbon sequestration benefits of thermally extended growing seasons, rendering them highly sensitive to future climate extremes.</p>
</sec>
<sec id="Ch1.S4.SS4">
  <label>4.4</label><title>Phenological response variation among different vegetation types</title>
      <p id="d2e3646">The phenological response to LST and AI varies significantly across vegetation types, reflecting the influence of distinct ecological strategies. Our results (Figs. 6 and 9) demonstrate that shrublands are the most sensitive to both warming and aridity, likely due to their opportunistic resource utilization strategy and higher leaf turnover rates (Xiong et al., 2024). In contrast, coniferous forests exhibited a muted response to both drivers (Malla et al., 2023). While our analysis relies on broad land-cover categories and does not directly measure species-specific functional traits, these discrepancies likely reflect generalized divergent ecological strategies among the vegetation groups.</p>
      <p id="d2e3649">This insensitivity is substantiated by our AI-sensitivity analysis (Fig. 9a), suggesting that the conservative hydraulic traits of conifers – such as needle morphology and lower stomatal conductance (Sperry et al., 2002; Brodribb and McAdam, 2013) – act as a buffer against the “aridity-driven inhibition” that more severely affects broadleaf species. Based on generalized ecological principles, this might be associated with typical drought-tolerant adaptations found in many conifers (e.g., needle morphology and stricter stomatal control), which could limit the potential positive impact of warming on their phenology. In contrast, the heightened sensitivity of shrublands observed in our dataset aligns with literature suggesting more opportunistic resource utilization strategies in shrubs (Xiong et al., 2024).</p>
      <p id="d2e3652">By considering these functional differences, our study provides a more nuanced understanding of urban ecosystem resilience, suggesting that urban greening programs should prioritize drought-tolerant coniferous or mixed forest structures in regions where the warming-saturation effect is most pronounced. However, we note that these functional interpretations remain broad generalizations; future studies incorporating field-based, species-level trait measurements are required to mechanistically validate these large-scale categorical patterns. Recognizing these trait-mediated responses is a crucial conceptual step for urban forestry. It indicates that building climate-resilient urban ecosystems requires shifting from generic greening to deploying specific functional traits – such as the conservative hydraulic strategies of conifers – to buffer against the coupled heat-aridity saturation effects.</p>
</sec>
</sec>
<sec id="Ch1.S5" sec-type="conclusions">
  <label>5</label><title>Conclusions</title>
      <p id="d2e3665">Our study used remote sensing data to investigate the impact of urbanization on vegetation phenology across 293 Chinese cities from 2010 to 2020. Urban heat islands consistently advanced the start of the growing season (SOS) by 12.06 d and delayed the end of the growing season (EOS) by 9.86 d relative to rural areas. However, the magnitude of these shifts varied markedly with background land surface temperature (LST) and aridity index (AI), indicating that warming-induced phenological responses are not universally distributed but are constrained by local climatic context.</p>
      <p id="d2e3668">We identified a non-linear saturation effect in phenological sensitivity to urban warming. Temperature sensitivity peaked at 4 °C (spring) and 6 °C (autumn), and weakened significantly beyond the saturation thresholds of 12.5 °C for SOS and 4 °C for EOS. Within the effective ranges of 3.5 to 17.5 °C (SOS) and 0–18 °C (EOS), urban warming exerted pronounced positive effects on phenological shifts; outside these ranges, sensitivity approached zero. Notably, extreme high temperatures reversed these positive phenological effects on EOS, causing weakly delayed or even advanced senescence in warm urban regions.</p>
      <p id="d2e3671">Aridity emerged as a critical negative regulator of warming-induced phenological effects. Across most AI ranges, increasing aridity attenuated the positive effects of UHI on both SOS and EOS. Within the intermediate aridity band of 1.4 <inline-formula><mml:math id="M249" display="inline"><mml:mo>&lt;</mml:mo></mml:math></inline-formula> AI <inline-formula><mml:math id="M250" display="inline"><mml:mo>&lt;</mml:mo></mml:math></inline-formula> 2.0, warming-induced phenological effects were entirely reversed. Consequently, coastal cities with lower AI exhibited substantially larger urban-rural phenological disparities than arid inland cities, where coupled heat-aridity stress compressed the effective growing season.</p>
      <p id="d2e3688">Collectively, these findings challenge the linear assumption that urban warming universally extends the growing season. Instead, they demonstrate that phenological responses to UHI are bounded by finite physiological limits and modulated by water availability. For urban ecological planning, our results imply that greening strategies must be climate-specific: drought-tolerant vegetation types should be prioritized in arid regions where saturation occurs at lower thermal thresholds, whereas coastal and northern cities can more effectively harness the phenological effects of UHI for growing season extension. Future research incorporating species-level functional traits and process-based phenology models will be essential to refine these thresholds and project urban ecosystem dynamics under continued global warming.</p>
</sec>

      
      </body>
    <back><notes notes-type="dataavailability"><title>Data availability</title>

      <p id="d2e3695">The data supporting the findings of this study are available in the public-domain resources listed in Table 1.</p>
  </notes><notes notes-type="authorcontribution"><title>Author contributions</title>

      <p id="d2e3701">Lin Xingwen: Conceptualization, Methodology, Funding Acquisition; Zhang Zhenzhen: Data curation, Writing – original draft preparation; Hu Xinxin: Methodology, Formal analysis, Validation, Writing – review and editing; Zhang Yongqi: Visualization, Investigation; Sun Liheng, Wu Chaofan and Cui Shufen: Supervision; Zhang Zhaoyang, Chen Yuanjian: Software, Validation; Wen Qingqing: Funding Acquisition.</p>
  </notes><notes notes-type="competinginterests"><title>Competing interests</title>

      <p id="d2e3707">The contact author has declared that none of the authors has any competing interests.</p>
  </notes><notes notes-type="disclaimer"><title>Disclaimer</title>

      <p id="d2e3713">Publisher's note: Copernicus Publications remains neutral with regard to jurisdictional claims made in the text, published maps, institutional affiliations, or any other geographical representation in this paper. The authors bear the ultimate responsibility for providing appropriate place names. Views expressed in the text are those of the authors and do not necessarily reflect the views of the publisher.</p>
  </notes><ack><title>Acknowledgements</title><p id="d2e3719">We appreciate the valuable suggestions from the Associate Editor and the anonymous reviewers, which substantially improved this manuscript. We also acknowledge the computing support provided by our university and the members of our research group for their assistance throughout this work.</p></ack><notes notes-type="financialsupport"><title>Financial support</title>

      <p id="d2e3724">This study was supported by Jinhua scientific and technological projects (grant numbers 2026-3-108, 2024-3-002), Open Research Project of Innovation Center of Yangtze River Delta, Zhejiang University (grant number: 2025KFKY004).</p>
  </notes><notes notes-type="reviewstatement"><title>Review statement</title>

      <p id="d2e3730">This paper was edited by Sara Vicca and reviewed by two anonymous referees.</p>
  </notes><ref-list>
    <title>References</title>

      <ref id="bib1.bib1"><label>1</label><mixed-citation>Angilletta, M. J.: Thermal Adaptation: A Theoretical and Empirical Synthesis, Oxford University Press, Oxford, <ext-link xlink:href="https://doi.org/10.1093/acprof:oso/9780198570875.001.1" ext-link-type="DOI">10.1093/acprof:oso/9780198570875.001.1</ext-link>, 2009.</mixed-citation></ref>
      <ref id="bib1.bib2"><label>2</label><mixed-citation>Badeck, F.-W., Bondeau, A., Böttcher, K., Doktor, D., Lucht, W., Schaber, J., and Sitch, S.: Responses of spring phenology to climate change, New Phytol., 162, 295–309, <ext-link xlink:href="https://doi.org/10.1111/j.1469-8137.2004.01059.x" ext-link-type="DOI">10.1111/j.1469-8137.2004.01059.x</ext-link>, 2004.</mixed-citation></ref>
      <ref id="bib1.bib3"><label>3</label><mixed-citation>Brodribb, T. J. and McAdam, S. A. M.: Abscisic acid mediates a divergence in the drought response of two conifers, Plant Physiol., 162, 1370–1377, <ext-link xlink:href="https://doi.org/10.1104/pp.113.217877" ext-link-type="DOI">10.1104/pp.113.217877</ext-link>, 2013.</mixed-citation></ref>
      <ref id="bib1.bib4"><label>4</label><mixed-citation>Buyantuyev, A. and Wu, J.: Urbanization diversifies land surface phenology in arid environments: Interactions among vegetation, climatic variation, and land use pattern in the Phoenix metropolitan region, USA, Landsc. Urban Plan., 105, 149–159, <ext-link xlink:href="https://doi.org/10.1016/j.landurbplan.2011.12.013" ext-link-type="DOI">10.1016/j.landurbplan.2011.12.013</ext-link>, 2012.</mixed-citation></ref>
      <ref id="bib1.bib5"><label>5</label><mixed-citation>Büntgen, U., Krusic, P. J., Piermattei, A., Coomes, D. A., Esper, J., Myglan, V. S., Kirdyanov, A. V., Camarero, J. J., Crivellaro, A., and Körner, C.: Limited capacity of tree growth to mitigate the global greenhouse effect under predicted warming, Nat. Commun., 10, 2171, <ext-link xlink:href="https://doi.org/10.1038/s41467-019-10174-4" ext-link-type="DOI">10.1038/s41467-019-10174-4</ext-link>, 2019.</mixed-citation></ref>
      <ref id="bib1.bib6"><label>6</label><mixed-citation>Cai, Z., Jönsson, P., Jin, H., and Eklundh, L.: Performance of smoothing methods for reconstructing NDVI time-Series and estimating vegetation phenology from MODIS Data, Remote Sens., 9, 1271, <ext-link xlink:href="https://doi.org/10.3390/rs9121271" ext-link-type="DOI">10.3390/rs9121271</ext-link>, 2017.</mixed-citation></ref>
      <ref id="bib1.bib7"><label>7</label><mixed-citation>Čehulić, I., Sever, K., Katičić Bogdan, I., Jazbec, A., S̆kvorc, Z̆., and Bogdan, S.: Drought Impact on Leaf Phenology and Spring Frost Susceptibility in a <italic>Quercus robur L.</italic> Provenance Trial, Forests, 10, 50, <ext-link xlink:href="https://doi.org/10.3390/f10010050" ext-link-type="DOI">10.3390/f10010050</ext-link>, 2019.</mixed-citation></ref>
      <ref id="bib1.bib8"><label>8</label><mixed-citation>Chmielewski, F.-M. and Rötzer, T.: Response of tree phenology to climate change across Europe, Agr. Forest Meteorol., 108, 101–112, <ext-link xlink:href="https://doi.org/10.1016/S0168-1923(01)00233-7" ext-link-type="DOI">10.1016/S0168-1923(01)00233-7</ext-link>, 2001.</mixed-citation></ref>
      <ref id="bib1.bib9"><label>9</label><mixed-citation>Cong, N., Piao, S., Chen, A., Wang, X., Lin, X., Chen, S., Han, S., Zhou, G., and Zhang, X.: Spring vegetation green-up date in China inferred from SPOT NDVI data: A multiple model analysis, Agr. Forest Meteorol., 165, 104–113, <ext-link xlink:href="https://doi.org/10.1016/j.agrformet.2012.06.009" ext-link-type="DOI">10.1016/j.agrformet.2012.06.009</ext-link>, 2012.</mixed-citation></ref>
      <ref id="bib1.bib10"><label>10</label><mixed-citation>Crawford, B., Kelsey, K., Ibsen, P., Rees, A., and Charobee, A.: Intra-urban variations in land surface phenology in a semi-arid environment, Environ. Res. Lett., 20, 014036, <ext-link xlink:href="https://doi.org/10.1088/1748-9326/ad9759" ext-link-type="DOI">10.1088/1748-9326/ad9759</ext-link>, 2025.</mixed-citation></ref>
      <ref id="bib1.bib11"><label>11</label><mixed-citation>Dallimer, M., Tang, Z., Gaston, K. J., and Davies, Z. G.: The extent of shifts in vegetation phenology between rural and urban areas within a human-dominated region, Ecol. Evol., 6, 1942–1953, <ext-link xlink:href="https://doi.org/10.1002/ece3.1990" ext-link-type="DOI">10.1002/ece3.1990</ext-link>, 2016.</mixed-citation></ref>
      <ref id="bib1.bib12"><label>12</label><mixed-citation>Ding, H., Xu, L., Elmore, A. J., Ma, X., and Li, X.: Vegetation phenology influenced by rapid urbanization of The Yangtze Delta region, Remote Sens., 12, 1783, <ext-link xlink:href="https://doi.org/10.3390/rs12111783" ext-link-type="DOI">10.3390/rs12111783</ext-link>, 2020.</mixed-citation></ref>
      <ref id="bib1.bib13"><label>13</label><mixed-citation>Doncaster, C. P.: Structural Equation Modeling and Natural Systems, Fish Fish., 8, 368–369, <ext-link xlink:href="https://doi.org/10.1111/j.1467-2979.2007.00260.x" ext-link-type="DOI">10.1111/j.1467-2979.2007.00260.x</ext-link>, 2007.</mixed-citation></ref>
      <ref id="bib1.bib14"><label>14</label><mixed-citation> Duan, J., Zhang, Q.-B., Lv, L., and Zhang, C.: Regional-scale winter-spring temperature variability and chilling damage dynamics over the past two centuries in southeastern China, Clim. Dynam., 39, 919–928, 2011.</mixed-citation></ref>
      <ref id="bib1.bib15"><label>15</label><mixed-citation>Elvidge, C. D., Baugh, K., Zhizhin, M., Hsu, F. C., and Ghosh, T.: VIIRS night-time lights, Int. J. Remote Sens., 38, 5860–5879, <ext-link xlink:href="https://doi.org/10.1080/01431161.2017.1342050" ext-link-type="DOI">10.1080/01431161.2017.1342050</ext-link>, 2017.</mixed-citation></ref>
      <ref id="bib1.bib16"><label>16</label><mixed-citation>Estiarte, M. and Peñuelas, J.: Alteration of the phenology of leaf senescence and fall in winter deciduous species by climate change: effects on nutrient proficiency, Glob. Change Biol., 21, 1005–1017, <ext-link xlink:href="https://doi.org/10.1111/gcb.12804" ext-link-type="DOI">10.1111/gcb.12804</ext-link>, 2015.</mixed-citation></ref>
      <ref id="bib1.bib17"><label>17</label><mixed-citation>Fan, J., He, H., Hu, T., Zhang, P., Yu, X., and Zhou, Y.: Estimation of landscape pattern changes in BRICS from 1992 to 2013 using DMSP-OLS NTL images, J. Indian Soc. Remote Sens., 47, 725–735, <ext-link xlink:href="https://doi.org/10.1007/s12524-019-00963-1" ext-link-type="DOI">10.1007/s12524-019-00963-1</ext-link>, 2019.</mixed-citation></ref>
      <ref id="bib1.bib18"><label>18</label><mixed-citation>Fracheboud, Y., Luquez, V., Bjorken, L., Sjodin, A., Tuominen, H., and Jansson, S.: The control of autumn senescence in European aspen, Plant Physiol., 149, 1982–1991, <ext-link xlink:href="https://doi.org/10.1104/pp.108.133249" ext-link-type="DOI">10.1104/pp.108.133249</ext-link>, 2009.</mixed-citation></ref>
      <ref id="bib1.bib19"><label>19</label><mixed-citation>Gazal, R., White, M. A., Gillies, R., Rodemaker, E. L. I., Sparrow, E., and Gordon, L.: GLOBE students, teachers, and scientists demonstrate variable differences between urban and rural leaf phenology, Glob. Change Biol., 14, 1568–1580, <ext-link xlink:href="https://doi.org/10.1111/j.1365-2486.2008.01602.x" ext-link-type="DOI">10.1111/j.1365-2486.2008.01602.x</ext-link>, 2008.</mixed-citation></ref>
      <ref id="bib1.bib20"><label>20</label><mixed-citation>Ge, W., Han, J., Zhang, D., and Wang, F.: Divergent impacts of droughts on vegetation phenology and productivity in the Yungui Plateau, southwest China, Ecol. Indic., 127, 107743, <ext-link xlink:href="https://doi.org/10.1016/j.ecolind.2021.107743" ext-link-type="DOI">10.1016/j.ecolind.2021.107743</ext-link>, 2021.</mixed-citation></ref>
      <ref id="bib1.bib21"><label>21</label><mixed-citation>Gow, L. J., Barrett, D. J., Renzullo, L. J., Phinn, S. R., and O'Grady, A. P.: Characterising groundwater use by vegetation using a surface energy balance model and satellite observations of land surface temperature, Environ. Modell. Softw., 80, 66–82, <ext-link xlink:href="https://doi.org/10.1016/j.envsoft.2016.02.021" ext-link-type="DOI">10.1016/j.envsoft.2016.02.021</ext-link>, 2016.</mixed-citation></ref>
      <ref id="bib1.bib22"><label>22</label><mixed-citation>Grimm, N. B., Faeth, S. H., Golubiewski, N. E., Redman, C. L., Wu, J., Bai, X., and Briggs, J. M.: Global change and the ecology of cities, Science, 319, 756–760, <ext-link xlink:href="https://doi.org/10.1126/science.1150195" ext-link-type="DOI">10.1126/science.1150195</ext-link>, 2008.</mixed-citation></ref>
      <ref id="bib1.bib23"><label>23</label><mixed-citation>Hanninen, H., Kramer, K., Tanino, K., Zhang, R., Wu, J., and Fu, Y. H.: Experiments are necessary in process-based tree phenology modelling, Trends Plant Sci., 24, 199–209, <ext-link xlink:href="https://doi.org/10.1016/j.tplants.2018.11.006" ext-link-type="DOI">10.1016/j.tplants.2018.11.006</ext-link>, 2019.</mixed-citation></ref>
      <ref id="bib1.bib24"><label>24</label><mixed-citation>Harpole, W. S., Ngai, J. T., Cleland, E. E., Seabloom, E. W., Borer, E. T., Bracken, M. E., Elser, J. J., Gruner, D. S., Hillebrand, H., Shurin, J. B., and Smith, J. E.: Nutrient co-limitation of primary producer communities, Ecol. Lett., 14, 852–862, <ext-link xlink:href="https://doi.org/10.1111/j.1461-0248.2011.01651.x" ext-link-type="DOI">10.1111/j.1461-0248.2011.01651.x</ext-link>, 2011.</mixed-citation></ref>
      <ref id="bib1.bib25"><label>25</label><mixed-citation>He, C., Gao, B., Huang, Q., Ma, Q., and Dou, Y.: Environmental degradation in the urban areas of China: Evidence from multi-source remote sensing data, Remote Sens. Environ., 193, 65–75, <ext-link xlink:href="https://doi.org/10.1016/j.rse.2017.02.027" ext-link-type="DOI">10.1016/j.rse.2017.02.027</ext-link>, 2017.</mixed-citation></ref>
      <ref id="bib1.bib26"><label>26</label><mixed-citation>Jeong, S.-J., Park, H., Ho, C.-H., and Kim, J.: Impact of urbanization on spring and autumn phenology of deciduous trees in the Seoul Capital Area, South Korea, Int. J. Biometeorol., 63, 627–637, <ext-link xlink:href="https://doi.org/10.1007/s00484-018-1610-7" ext-link-type="DOI">10.1007/s00484-018-1610-7</ext-link>, 2019.</mixed-citation></ref>
      <ref id="bib1.bib27"><label>27</label><mixed-citation>Ji, S., Ren, S., Li, Y., Dong, J., Wang, L., Quan, Q., and Liu, J.: Diverse responses of spring phenology to preseason drought and warming under different biomes in the North China Plain, Sci. Total Environ., 766, 144437, <ext-link xlink:href="https://doi.org/10.1016/j.scitotenv.2020.144437" ext-link-type="DOI">10.1016/j.scitotenv.2020.144437</ext-link>, 2021.</mixed-citation></ref>
      <ref id="bib1.bib28"><label>28</label><mixed-citation>Jia, W., Zhao, S., Zhang, X., Liu, S., Henebry, G. M., and Liu, L.: Urbanization imprint on land surface phenology: The urban-rural gradient analysis for Chinese cities, Glob. Change Biol., 27, 2895–2904, <ext-link xlink:href="https://doi.org/10.1111/gcb.15602" ext-link-type="DOI">10.1111/gcb.15602</ext-link>, 2021.</mixed-citation></ref>
      <ref id="bib1.bib29"><label>29</label><mixed-citation>Jönsson, P. and Eklundh, L.: TIMESAT – a program for analyzing time-series of satellite sensor data, Comput. Geosci., 30, 833–845, <ext-link xlink:href="https://doi.org/10.1016/j.cageo.2004.05.006" ext-link-type="DOI">10.1016/j.cageo.2004.05.006</ext-link>, 2004.</mixed-citation></ref>
      <ref id="bib1.bib30"><label>30</label><mixed-citation>Kang, X., Hao, Y., Cui, X., Chen, H., Huang, S., Du, Y., Li, W., Kardol, P., Xiao, X., and Cui, L.: Variability and changes in climate, phenology, and gross primary production of an Alpine wetland ecosystem, Remote Sens., 8, 391, <ext-link xlink:href="https://doi.org/10.3390/rs8050391" ext-link-type="DOI">10.3390/rs8050391</ext-link>, 2016.</mixed-citation></ref>
      <ref id="bib1.bib31"><label>31</label><mixed-citation>Ketola, T. and Kristensen, T. N.: Experimental approaches for testing if tolerance curves are useful for predicting fitness in fluctuating environments, Front. Ecol. Evol., 5, 129, <ext-link xlink:href="https://doi.org/10.3389/fevo.2017.00129" ext-link-type="DOI">10.3389/fevo.2017.00129</ext-link>, 2017.</mixed-citation></ref>
      <ref id="bib1.bib32"><label>32</label><mixed-citation>Kim, D. H., Doyle, M. R., Sung, S., and Amasino, R. M.: Vernalization: winter and the timing of flowering in plants, Annu. Rev. Cell Dev. Biol., 25, 277–299, <ext-link xlink:href="https://doi.org/10.1146/annurev.cellbio.042308.113411" ext-link-type="DOI">10.1146/annurev.cellbio.042308.113411</ext-link>, 2009.</mixed-citation></ref>
      <ref id="bib1.bib33"><label>33</label><mixed-citation>Knauer, J., Zaehle, S., Reichstein, M., Medlyn, B. E., Forkel, M., Hagemann, S., and Werner, C.: The response of ecosystem water‐use efficiency to rising atmospheric CO2 concentrations: sensitivity and large‐scale biogeochemical implications, New Phytol., 213, 1654–1666, <ext-link xlink:href="https://doi.org/10.1111/nph.14288" ext-link-type="DOI">10.1111/nph.14288</ext-link>, 2017.</mixed-citation></ref>
      <ref id="bib1.bib34"><label>34</label><mixed-citation> Legendre, P. and Legendre, L.: Numerical Ecology, 3rd edn., Elsevier, Amsterdam, ISBN 978-0-444-53869-7, 2012.</mixed-citation></ref>
      <ref id="bib1.bib35"><label>35</label><mixed-citation>Li, K., Wang, C., Sun, Q., Rong, G., Tong, Z., Liu, X., and Zhang, J.: Spring phenological sensitivity to climate Change in the northern hemisphere: comprehensive evaluation and driving force analysis, Remote Sens., 13, 1972, <ext-link xlink:href="https://doi.org/10.3390/rs13101972" ext-link-type="DOI">10.3390/rs13101972</ext-link>, 2021.</mixed-citation></ref>
      <ref id="bib1.bib36"><label>36</label><mixed-citation>Li, X., Zhou, Y., Asrar, G. R., Mao, J., Li, X., and Li, W.: Response of vegetation phenology to urbanization in the conterminous United States, Glob. Change Biol., 23, 2818–2830, <ext-link xlink:href="https://doi.org/10.1111/gcb.13562" ext-link-type="DOI">10.1111/gcb.13562</ext-link>, 2017.</mixed-citation></ref>
      <ref id="bib1.bib37"><label>37</label><mixed-citation>Malla, R., Neupane, P. R., and Köhl, M.: Climate change impacts: Vegetation shift of broad-leaved and coniferous forests, Trees For. People, 14, 100457, <ext-link xlink:href="https://doi.org/10.1016/j.tfp.2023.100457" ext-link-type="DOI">10.1016/j.tfp.2023.100457</ext-link>, 2023.</mixed-citation></ref>
      <ref id="bib1.bib38"><label>38</label><mixed-citation>Mathur, S., Agrawal, D., and Jajoo, A.: Photosynthesis: response to high temperature stress, J. Photochem. Photobiol. B, 137, 116–126, <ext-link xlink:href="https://doi.org/10.1016/j.jphotobiol.2014.01.010" ext-link-type="DOI">10.1016/j.jphotobiol.2014.01.010</ext-link>, 2014.</mixed-citation></ref>
      <ref id="bib1.bib39"><label>39</label><mixed-citation>Meng, L., Mao, J., Zhou, Y., Richardson, A. D., Lee, X., Thornton, P. E., Ricciuto, D. M., Li, X., Dai, Y., Shi, X., and Jia, G.: Urban warming advances spring phenology but reduces the response of phenology to temperature in the conterminous United States, P. Natl. Acad. Sci. USA, 117, 4228–4233, <ext-link xlink:href="https://doi.org/10.1073/pnas.1911117117" ext-link-type="DOI">10.1073/pnas.1911117117</ext-link>, 2020.</mixed-citation></ref>
      <ref id="bib1.bib40"><label>40</label><mixed-citation>Mu, Q., Zhao, M., and Running, S. W.: Improvements to a MODIS global terrestrial evapotranspiration algorithm, Remote Sens. Environ., 115, 1781–1800, <ext-link xlink:href="https://doi.org/10.1016/j.rse.2011.02.019" ext-link-type="DOI">10.1016/j.rse.2011.02.019</ext-link>, 2011.</mixed-citation></ref>
      <ref id="bib1.bib41"><label>41</label><mixed-citation>Pablos, M., Martínez-Fernández, J., Piles, M., Sánchez, N., Vall-llossera, M., and Camps, A.: Multi-temporal evaluation of soil moisture and land surface temperature dynamics using in situ and satellite observations, Remote Sens., 8, 587, <ext-link xlink:href="https://doi.org/10.3390/rs8070587" ext-link-type="DOI">10.3390/rs8070587</ext-link>, 2016.</mixed-citation></ref>
      <ref id="bib1.bib42"><label>42</label><mixed-citation>Peng, J., Wu, C., Zhang, X., Wang, X., and Gonsamo, A.: Satellite detection of cumulative and lagged effects of drought on autumn leaf senescence over the Northern Hemisphere, Glob. Change Biol., 25, 2174–2188, <ext-link xlink:href="https://doi.org/10.1111/gcb.14627" ext-link-type="DOI">10.1111/gcb.14627</ext-link>, 2019.</mixed-citation></ref>
      <ref id="bib1.bib43"><label>43</label><mixed-citation>Peng, S.: 1-km monthly precipitation dataset for China (1901–2021), National Tibetan Plateau/Third Pole Environment Data Center [data set], <ext-link xlink:href="https://doi.org/10.5281/zenodo.3185722" ext-link-type="DOI">10.5281/zenodo.3185722</ext-link>, 2020.</mixed-citation></ref>
      <ref id="bib1.bib44"><label>44</label><mixed-citation>Peñuelas, J., Gordon, C., Llorens, L., Nielsen, T., Tietema, A., Beier, C., Bruna, P., Emmett, B., Estiarte, M., and Gorissen, A.: Nonintrusive field experiments show different plant responses to warming and drought among sites, seasons, and species in a north-south European gradient, Ecosystems, 7, 598–612, <ext-link xlink:href="https://doi.org/10.1007/s10021-004-0179-7" ext-link-type="DOI">10.1007/s10021-004-0179-7</ext-link>, 2004.</mixed-citation></ref>
      <ref id="bib1.bib45"><label>45</label><mixed-citation>Piao, S., Fang, J., Zhou, L., Ciais, P., and Zhu, B.: Variations in satellite-derived phenology in China's temperate vegetation, Glob. Change Biol., 12, 672–685, <ext-link xlink:href="https://doi.org/10.1111/j.1365-2486.2006.01123.x" ext-link-type="DOI">10.1111/j.1365-2486.2006.01123.x</ext-link>, 2006.</mixed-citation></ref>
      <ref id="bib1.bib46"><label>46</label><mixed-citation>Pinheiro, J. C. and Bates, D. M.: Mixed-Effects Models in S and S-PLUS, Springer, New York, ISBN 978-0-387-22747-4, <ext-link xlink:href="https://doi.org/10.1007/b98882" ext-link-type="DOI">10.1007/b98882</ext-link>, 2000.</mixed-citation></ref>
      <ref id="bib1.bib47"><label>47</label><mixed-citation>Pollastrini, M., Puletti, N., Selvi, F., Iacopetti, G., and Bussotti, F.: Widespread crown defoliation after a drought and heat wave in the forests of Tuscany (Central Italy) and their recovery – A case study from summer 2017, Front. For. Glob. Change, 2, 74, <ext-link xlink:href="https://doi.org/10.3389/ffgc.2019.00074" ext-link-type="DOI">10.3389/ffgc.2019.00074</ext-link>, 2019.</mixed-citation></ref>
      <ref id="bib1.bib48"><label>48</label><mixed-citation>Qiao, Z., Tian, G., and Xiao, L.: Diurnal and seasonal impacts of urbanization on the urban thermal environment: A case study of Beijing using MODIS data, ISPRS J. Photogramm. Remote Sens., 85, 93–101, <ext-link xlink:href="https://doi.org/10.1016/j.isprsjprs.2013.08.010" ext-link-type="DOI">10.1016/j.isprsjprs.2013.08.010</ext-link>, 2013.</mixed-citation></ref>
      <ref id="bib1.bib49"><label>49</label><mixed-citation>Qiu, T., Song, C., and Li, J.: Impacts of urbanization on vegetation phenology over the past three decades in Shanghai, China, Remote Sens., 9, 970, <ext-link xlink:href="https://doi.org/10.3390/rs9090970" ext-link-type="DOI">10.3390/rs9090970</ext-link>, 2017.</mixed-citation></ref>
      <ref id="bib1.bib50"><label>50</label><mixed-citation>Rustad, L., Campbell, J., Marion, G., Norby, R., Mitchell, M., Hartley, A., Cornelissen, J., Gurevitch, J., and Gcte, N.: A meta-analysis of the response of soil respiration, net nitrogen mineralization, and aboveground plant growth to experimental ecosystem warming, Oecologia, 126, 543–562, <ext-link xlink:href="https://doi.org/10.1007/s004420000544" ext-link-type="DOI">10.1007/s004420000544</ext-link>, 2001.</mixed-citation></ref>
      <ref id="bib1.bib51"><label>51</label><mixed-citation>Sage, R. F. and Kubien, D. S.: The temperature response of C3 and C4 photosynthesis, Plant Cell Environ., 30, 1086–1106, <ext-link xlink:href="https://doi.org/10.1111/j.1365-3040.2007.01682.x" ext-link-type="DOI">10.1111/j.1365-3040.2007.01682.x</ext-link>, 2007.</mixed-citation></ref>
      <ref id="bib1.bib52"><label>52</label><mixed-citation>Shi, C., Sun, G., Zhang, H., Xiao, B., Ze, B., Zhang, N., and Wu, N.: Effects of warming on chlorophyll degradation and carbohydrate accumulation of Alpine herbaceous species during plant senescence on the Tibetan Plateau, PLoS One, 9, e107874, <ext-link xlink:href="https://doi.org/10.1371/journal.pone.0107874" ext-link-type="DOI">10.1371/journal.pone.0107874</ext-link>, 2014.</mixed-citation></ref>
      <ref id="bib1.bib53"><label>53</label><mixed-citation>Shi, W., Li, J., Zhan, H., Yu, L., Wang, C., and Wang, S.: Relation between water storage and photoassimilate accumulation of neosinocalamus affinis with phenology, Forests, 14, 531, <ext-link xlink:href="https://doi.org/10.3390/f14030531" ext-link-type="DOI">10.3390/f14030531</ext-link>, 2023.</mixed-citation></ref>
      <ref id="bib1.bib54"><label>54</label><mixed-citation>Sperry, J. S., Hacke, U. G., Oren, R., and Comstock, J. P.: Water deficits and hydraulic limits to leaf water supply, Plant Cell Environ., 25, 251–263, <ext-link xlink:href="https://doi.org/10.1046/j.0016-8025.2001.00799.x" ext-link-type="DOI">10.1046/j.0016-8025.2001.00799.x</ext-link>, 2002.</mixed-citation></ref>
      <ref id="bib1.bib55"><label>55</label><mixed-citation>Tian, F., Wigneron, J.-P., Ciais, P., Chave, J., Ogée, J., Peñuelas, J., Ræbild, A., Domec, J.-C., Tong, X., Brandt, M., Mialon, A., Rodriguez-Fernandez, N., Tagesson, T., Al-Yaari, A., Kerr, Y., Chen, C., Myneni, R. B., Zhang, W., Ardö, J., and Fensholt, R.: Coupling of ecosystem-scale plant water storage and leaf phenology observed by satellite, Nat. Ecol. Evol., 2, 1428–1435, <ext-link xlink:href="https://doi.org/10.1038/s41559-018-0630-3" ext-link-type="DOI">10.1038/s41559-018-0630-3</ext-link>, 2018.</mixed-citation></ref>
      <ref id="bib1.bib56"><label>56</label><mixed-citation>Wan, Z.: New refinements and validation of the MODIS Land-Surface Temperature/Emissivity products, Remote Sens. Environ., 112, 59–74, <ext-link xlink:href="https://doi.org/10.1016/j.rse.2006.06.026" ext-link-type="DOI">10.1016/j.rse.2006.06.026</ext-link>, 2008.</mixed-citation></ref>
      <ref id="bib1.bib57"><label>57</label><mixed-citation>Wang, C., Cao, R., Chen, J., Rao, Y., and Tang, Y.: Temperature sensitivity of spring vegetation phenology correlates to within-spring warming speed over the Northern Hemisphere, Ecol. Indic., 50, 62–68, <ext-link xlink:href="https://doi.org/10.1016/j.ecolind.2014.11.004" ext-link-type="DOI">10.1016/j.ecolind.2014.11.004</ext-link>, 2015.</mixed-citation></ref>
      <ref id="bib1.bib58"><label>58</label><mixed-citation>Wang, C., Tang, Y., and Chen, J.: Plant phenological synchrony increases under rapid within-spring warming, Sci. Rep., 6, 25460, <ext-link xlink:href="https://doi.org/10.1038/srep25460" ext-link-type="DOI">10.1038/srep25460</ext-link>, 2016.</mixed-citation></ref>
      <ref id="bib1.bib59"><label>59</label><mixed-citation>White, M. A., De Beurs, K. M., Didan, K., Inouye, D. W., Richardson, A. D., Jensen, O. P., O'Keefe, J., Zhang, G., Nemani, R. R., Van Leeuwen, W. J. D., Brown, J. F., De Wit, A., Schaepman, M., Lin, X., Dettinger, M., Bailey, A. S., Kimball, J., Schwartz, M. D., Baldocchi, D. D., Lee, J. T., and Lauenroth, W. K.: Intercomparison, interpretation, and assessment of spring phenology in North America estimated from remote sensing for 1982-2006, Glob. Change Biol., 15, 2335–2359, <ext-link xlink:href="https://doi.org/10.1111/j.1365-2486.2009.01910.x" ext-link-type="DOI">10.1111/j.1365-2486.2009.01910.x</ext-link>, 2009.</mixed-citation></ref>
      <ref id="bib1.bib60"><label>60</label><mixed-citation>Wu, C., Peng, J., Ciais, P., Peñuelas, J., Wang, H., Beguería, S., Black, T. A., Jassal, R. S., Zhang, X., Yuan, W., Liang, E., Wang, X., Hua, H., Liu, R., Ju, W., Fu, Y. H., and Ge, Q.: Increased drought effects on the phenology of autumn leaf senescence, Nat. Clim. Change, 12, 943–949, <ext-link xlink:href="https://doi.org/10.1038/s41558-022-01464-9" ext-link-type="DOI">10.1038/s41558-022-01464-9</ext-link>, 2022. </mixed-citation></ref>
      <ref id="bib1.bib61"><label>61</label><mixed-citation>Xiong, X., Wu, H., Wei, X., and Jiang, M.: Contrasting temperature and light sensitivities of spring leaf phenology between understory shrubs and canopy trees: Implications for phenological escape, Agr. Forest Meteorol., 355, 110144, <ext-link xlink:href="https://doi.org/10.1016/j.agrformet.2024.110144" ext-link-type="DOI">10.1016/j.agrformet.2024.110144</ext-link>, 2024.</mixed-citation></ref>
      <ref id="bib1.bib62"><label>62</label><mixed-citation>Yang, H., Zhang, Q., Zhang, S., Wang, X., and Yu, H.: Joint control of urban expansion and climate change on urban-rural vegetation phenology gradient in 31 cities of China, Front. Ecol. Evol., 13, 1637210, <ext-link xlink:href="https://doi.org/10.3389/fevo.2025.1637210" ext-link-type="DOI">10.3389/fevo.2025.1637210</ext-link>, 2025.</mixed-citation></ref>
      <ref id="bib1.bib63"><label>63</label><mixed-citation>Yao, R., Wang, L., Huang, X., Guo, X., Niu, Z., and Liu, H.: Investigation of Urbanization effects on land Surface phenology in Northeast China during 2001–2015, Remote Sens., 9, 66, <ext-link xlink:href="https://doi.org/10.3390/rs9010066" ext-link-type="DOI">10.3390/rs9010066</ext-link>, 2017.</mixed-citation></ref>
      <ref id="bib1.bib64"><label>64</label><mixed-citation>Yu, F., Price, K. P., Ellis, J., and Shi, P.: Response of seasonal vegetation development to climatic variations in eastern central Asia, Remote Sens. Environ., 87, 42–54, <ext-link xlink:href="https://doi.org/10.1016/S0034-4257(03)00144-5" ext-link-type="DOI">10.1016/S0034-4257(03)00144-5</ext-link>, 2003.</mixed-citation></ref>
      <ref id="bib1.bib65"><label>65</label><mixed-citation>Yuan, M., Zhao, L., Lin, A., Wang, L., Li, Q., She, D., and Qu, S.: Impacts of preseason drought on vegetation spring phenology across the Northeast China Transect, Sci. Total Environ., 738, 140297, <ext-link xlink:href="https://doi.org/10.1016/j.scitotenv.2020.140297" ext-link-type="DOI">10.1016/j.scitotenv.2020.140297</ext-link>, 2020.</mixed-citation></ref>
      <ref id="bib1.bib66"><label>66</label><mixed-citation>Zhang, X., Friedl, M. A., Schaaf, C. B., Strahler, A. H., and Schneider, A.: The footprint of urban climates on vegetation phenology, Geophys. Res. Lett., 31, L12209, <ext-link xlink:href="https://doi.org/10.1029/2004GL020137" ext-link-type="DOI">10.1029/2004GL020137</ext-link>, 2004.</mixed-citation></ref>
      <ref id="bib1.bib67"><label>67</label><mixed-citation>Zhang, X., Friedl, M. A., and Schaaf, C. B.: Global vegetation phenology from Moderate Resolution Imaging Spectro radiometer (MODIS): Evaluation of global patterns and comparison with in situ measurements, J. Geophys. Res.-Biogeo., 111, G04017, <ext-link xlink:href="https://doi.org/10.1029/2006JG000217" ext-link-type="DOI">10.1029/2006JG000217</ext-link>, 2006.</mixed-citation></ref>
      <ref id="bib1.bib68"><label>68</label><mixed-citation>Zhang, X., Chen, N., Sheng, H., Ip, C., Yang, L., Chen, Y., Sang, Z., Tadesse, T., Lim, T. P. Y., Rajabifard, A., Bueti, C., Zeng, L., Wardlow, B., Wang, S., Tang, S., Xiong, Z., Li, D., and Niyogi, D.: Urban drought challenge to 2030 sustainable development goals, Sci. Total Environ., 693, 133536, <ext-link xlink:href="https://doi.org/10.1016/j.scitotenv.2019.07.342" ext-link-type="DOI">10.1016/j.scitotenv.2019.07.342</ext-link>, 2019.</mixed-citation></ref>
      <ref id="bib1.bib69"><label>69</label><mixed-citation>Zheng, W., Liu, Y., Yang, X., and Fan, W.: Spatiotemporal variations of forest vegetation phenology and its response to climate change in northeast China, Remote Sens., 14, 2909, <ext-link xlink:href="https://doi.org/10.3390/rs14122909" ext-link-type="DOI">10.3390/rs14122909</ext-link>, 2022.</mixed-citation></ref>
      <ref id="bib1.bib70"><label>70</label><mixed-citation>Zhou, D., Zhao, S., Zhang, L., and Liu, S.: Remotely sensed assessment of urbanization effects on vegetation phenology in China's 32 major cities, Remote Sens. Environ., 176, 272–281, <ext-link xlink:href="https://doi.org/10.1016/j.rse.2016.02.010" ext-link-type="DOI">10.1016/j.rse.2016.02.010</ext-link>, 2016.</mixed-citation></ref>

  </ref-list></back>
    <!--<article-title-html>Saturation effect of background temperature and aridity on vegetation phenological sensitivity to urban warming</article-title-html>
<abstract-html/>
<ref-html id="bib1.bib1"><label>1</label><mixed-citation>
      
Angilletta, M. J.: Thermal Adaptation: A Theoretical and Empirical Synthesis, Oxford University Press, Oxford, <a href="https://doi.org/10.1093/acprof:oso/9780198570875.001.1" target="_blank">https://doi.org/10.1093/acprof:oso/9780198570875.001.1</a>, 2009.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib2"><label>2</label><mixed-citation>
      
Badeck, F.-W., Bondeau, A., Böttcher, K., Doktor, D., Lucht, W., Schaber, J., and Sitch, S.: Responses of spring phenology to climate change, New Phytol., 162, 295–309, <a href="https://doi.org/10.1111/j.1469-8137.2004.01059.x" target="_blank">https://doi.org/10.1111/j.1469-8137.2004.01059.x</a>, 2004.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib3"><label>3</label><mixed-citation>
      
Brodribb, T. J. and McAdam, S. A. M.: Abscisic acid mediates a divergence in the drought response of two conifers, Plant Physiol., 162, 1370–1377, <a href="https://doi.org/10.1104/pp.113.217877" target="_blank">https://doi.org/10.1104/pp.113.217877</a>, 2013.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib4"><label>4</label><mixed-citation>
      
Buyantuyev, A. and Wu, J.: Urbanization diversifies land surface phenology in arid environments: Interactions among vegetation, climatic variation, and land use pattern in the Phoenix metropolitan region, USA, Landsc. Urban Plan., 105, 149–159, <a href="https://doi.org/10.1016/j.landurbplan.2011.12.013" target="_blank">https://doi.org/10.1016/j.landurbplan.2011.12.013</a>, 2012.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib5"><label>5</label><mixed-citation>
      
Büntgen, U., Krusic, P. J., Piermattei, A., Coomes, D. A., Esper, J., Myglan, V. S., Kirdyanov, A. V., Camarero, J. J., Crivellaro, A., and Körner, C.: Limited capacity of tree growth to mitigate the global greenhouse effect under predicted warming, Nat. Commun., 10, 2171, <a href="https://doi.org/10.1038/s41467-019-10174-4" target="_blank">https://doi.org/10.1038/s41467-019-10174-4</a>, 2019.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib6"><label>6</label><mixed-citation>
      
Cai, Z., Jönsson, P., Jin, H., and Eklundh, L.: Performance of smoothing methods for reconstructing NDVI time-Series and estimating vegetation phenology from MODIS Data, Remote Sens., 9, 1271, <a href="https://doi.org/10.3390/rs9121271" target="_blank">https://doi.org/10.3390/rs9121271</a>, 2017.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib7"><label>7</label><mixed-citation>
      
Čehulić, I., Sever, K., Katičić Bogdan, I., Jazbec, A., S̆kvorc, Z̆., and Bogdan, S.: Drought Impact on Leaf Phenology and Spring Frost Susceptibility in a <i>Quercus robur L.</i> Provenance Trial, Forests, 10, 50, <a href="https://doi.org/10.3390/f10010050" target="_blank">https://doi.org/10.3390/f10010050</a>, 2019.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib8"><label>8</label><mixed-citation>
      
Chmielewski, F.-M. and Rötzer, T.: Response of tree phenology to climate change across Europe, Agr. Forest Meteorol., 108, 101–112, <a href="https://doi.org/10.1016/S0168-1923(01)00233-7" target="_blank">https://doi.org/10.1016/S0168-1923(01)00233-7</a>, 2001.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib9"><label>9</label><mixed-citation>
      
Cong, N., Piao, S., Chen, A., Wang, X., Lin, X., Chen, S., Han, S., Zhou, G., and Zhang, X.: Spring vegetation green-up date in China inferred from SPOT NDVI data: A multiple model analysis, Agr. Forest Meteorol., 165, 104–113, <a href="https://doi.org/10.1016/j.agrformet.2012.06.009" target="_blank">https://doi.org/10.1016/j.agrformet.2012.06.009</a>, 2012.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib10"><label>10</label><mixed-citation>
      
Crawford, B., Kelsey, K., Ibsen, P., Rees, A., and Charobee, A.: Intra-urban variations in land surface phenology in a semi-arid environment, Environ. Res. Lett., 20, 014036, <a href="https://doi.org/10.1088/1748-9326/ad9759" target="_blank">https://doi.org/10.1088/1748-9326/ad9759</a>, 2025.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib11"><label>11</label><mixed-citation>
      
Dallimer, M., Tang, Z., Gaston, K. J., and Davies, Z. G.: The extent of shifts in vegetation phenology between rural and urban areas within a human-dominated region, Ecol. Evol., 6, 1942–1953, <a href="https://doi.org/10.1002/ece3.1990" target="_blank">https://doi.org/10.1002/ece3.1990</a>, 2016.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib12"><label>12</label><mixed-citation>
      
Ding, H., Xu, L., Elmore, A. J., Ma, X., and Li, X.: Vegetation phenology influenced by rapid urbanization of The Yangtze Delta region, Remote Sens., 12, 1783, <a href="https://doi.org/10.3390/rs12111783" target="_blank">https://doi.org/10.3390/rs12111783</a>, 2020.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib13"><label>13</label><mixed-citation>
      
Doncaster, C. P.: Structural Equation Modeling and Natural Systems, Fish Fish., 8, 368–369, <a href="https://doi.org/10.1111/j.1467-2979.2007.00260.x" target="_blank">https://doi.org/10.1111/j.1467-2979.2007.00260.x</a>, 2007.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib14"><label>14</label><mixed-citation>
      
Duan, J., Zhang, Q.-B., Lv, L., and Zhang, C.: Regional-scale winter-spring temperature variability and chilling damage dynamics over the past two centuries in southeastern China, Clim. Dynam., 39, 919–928, 2011.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib15"><label>15</label><mixed-citation>
      
Elvidge, C. D., Baugh, K., Zhizhin, M., Hsu, F. C., and Ghosh, T.: VIIRS night-time lights, Int. J. Remote Sens., 38, 5860–5879, <a href="https://doi.org/10.1080/01431161.2017.1342050" target="_blank">https://doi.org/10.1080/01431161.2017.1342050</a>, 2017.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib16"><label>16</label><mixed-citation>
      
Estiarte, M. and Peñuelas, J.: Alteration of the phenology of leaf senescence and fall in winter deciduous species by climate change: effects on nutrient proficiency, Glob. Change Biol., 21, 1005–1017, <a href="https://doi.org/10.1111/gcb.12804" target="_blank">https://doi.org/10.1111/gcb.12804</a>, 2015.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib17"><label>17</label><mixed-citation>
      
Fan, J., He, H., Hu, T., Zhang, P., Yu, X., and Zhou, Y.: Estimation of landscape pattern changes in BRICS from 1992 to 2013 using DMSP-OLS NTL images, J. Indian Soc. Remote Sens., 47, 725–735, <a href="https://doi.org/10.1007/s12524-019-00963-1" target="_blank">https://doi.org/10.1007/s12524-019-00963-1</a>, 2019.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib18"><label>18</label><mixed-citation>
      
Fracheboud, Y., Luquez, V., Bjorken, L., Sjodin, A., Tuominen, H., and Jansson, S.: The control of autumn senescence in European aspen, Plant Physiol., 149, 1982–1991, <a href="https://doi.org/10.1104/pp.108.133249" target="_blank">https://doi.org/10.1104/pp.108.133249</a>, 2009.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib19"><label>19</label><mixed-citation>
      
Gazal, R., White, M. A., Gillies, R., Rodemaker, E. L. I., Sparrow, E., and Gordon, L.: GLOBE students, teachers, and scientists demonstrate variable differences between urban and rural leaf phenology, Glob. Change Biol., 14, 1568–1580, <a href="https://doi.org/10.1111/j.1365-2486.2008.01602.x" target="_blank">https://doi.org/10.1111/j.1365-2486.2008.01602.x</a>, 2008.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib20"><label>20</label><mixed-citation>
      
Ge, W., Han, J., Zhang, D., and Wang, F.: Divergent impacts of droughts on vegetation phenology and productivity in the Yungui Plateau, southwest China, Ecol. Indic., 127, 107743, <a href="https://doi.org/10.1016/j.ecolind.2021.107743" target="_blank">https://doi.org/10.1016/j.ecolind.2021.107743</a>, 2021.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib21"><label>21</label><mixed-citation>
      
Gow, L. J., Barrett, D. J., Renzullo, L. J., Phinn, S. R., and O'Grady, A. P.: Characterising groundwater use by vegetation using a surface energy balance model and satellite observations of land surface temperature, Environ. Modell. Softw., 80, 66–82, <a href="https://doi.org/10.1016/j.envsoft.2016.02.021" target="_blank">https://doi.org/10.1016/j.envsoft.2016.02.021</a>, 2016.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib22"><label>22</label><mixed-citation>
      
Grimm, N. B., Faeth, S. H., Golubiewski, N. E., Redman, C. L., Wu, J., Bai, X., and Briggs, J. M.: Global change and the ecology of cities, Science, 319, 756–760, <a href="https://doi.org/10.1126/science.1150195" target="_blank">https://doi.org/10.1126/science.1150195</a>, 2008.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib23"><label>23</label><mixed-citation>
      
Hanninen, H., Kramer, K., Tanino, K., Zhang, R., Wu, J., and Fu, Y. H.: Experiments are necessary in process-based tree phenology modelling, Trends Plant Sci., 24, 199–209, <a href="https://doi.org/10.1016/j.tplants.2018.11.006" target="_blank">https://doi.org/10.1016/j.tplants.2018.11.006</a>, 2019.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib24"><label>24</label><mixed-citation>
      
Harpole, W. S., Ngai, J. T., Cleland, E. E., Seabloom, E. W., Borer, E. T., Bracken, M. E., Elser, J. J., Gruner, D. S., Hillebrand, H., Shurin, J. B., and Smith, J. E.: Nutrient co-limitation of primary producer communities, Ecol. Lett., 14, 852–862, <a href="https://doi.org/10.1111/j.1461-0248.2011.01651.x" target="_blank">https://doi.org/10.1111/j.1461-0248.2011.01651.x</a>, 2011.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib25"><label>25</label><mixed-citation>
      
He, C., Gao, B., Huang, Q., Ma, Q., and Dou, Y.: Environmental degradation in the urban areas of China: Evidence from multi-source remote sensing data, Remote Sens. Environ., 193, 65–75, <a href="https://doi.org/10.1016/j.rse.2017.02.027" target="_blank">https://doi.org/10.1016/j.rse.2017.02.027</a>, 2017.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib26"><label>26</label><mixed-citation>
      
Jeong, S.-J., Park, H., Ho, C.-H., and Kim, J.: Impact of urbanization on spring and autumn phenology of deciduous trees in the Seoul Capital Area, South Korea, Int. J. Biometeorol., 63, 627–637, <a href="https://doi.org/10.1007/s00484-018-1610-7" target="_blank">https://doi.org/10.1007/s00484-018-1610-7</a>, 2019.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib27"><label>27</label><mixed-citation>
      
Ji, S., Ren, S., Li, Y., Dong, J., Wang, L., Quan, Q., and Liu, J.: Diverse responses of spring phenology to preseason drought and warming under different biomes in the North China Plain, Sci. Total Environ., 766, 144437, <a href="https://doi.org/10.1016/j.scitotenv.2020.144437" target="_blank">https://doi.org/10.1016/j.scitotenv.2020.144437</a>, 2021.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib28"><label>28</label><mixed-citation>
      
Jia, W., Zhao, S., Zhang, X., Liu, S., Henebry, G. M., and Liu, L.: Urbanization imprint on land surface phenology: The urban-rural gradient analysis for Chinese cities, Glob. Change Biol., 27, 2895–2904, <a href="https://doi.org/10.1111/gcb.15602" target="_blank">https://doi.org/10.1111/gcb.15602</a>, 2021.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib29"><label>29</label><mixed-citation>
      
Jönsson, P. and Eklundh, L.: TIMESAT – a program for analyzing time-series of satellite sensor data, Comput. Geosci., 30, 833–845, <a href="https://doi.org/10.1016/j.cageo.2004.05.006" target="_blank">https://doi.org/10.1016/j.cageo.2004.05.006</a>, 2004.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib30"><label>30</label><mixed-citation>
      
Kang, X., Hao, Y., Cui, X., Chen, H., Huang, S., Du, Y., Li, W., Kardol, P., Xiao, X., and Cui, L.: Variability and changes in climate, phenology, and gross primary production of an Alpine wetland ecosystem, Remote Sens., 8, 391, <a href="https://doi.org/10.3390/rs8050391" target="_blank">https://doi.org/10.3390/rs8050391</a>, 2016.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib31"><label>31</label><mixed-citation>
      
Ketola, T. and Kristensen, T. N.: Experimental approaches for testing if tolerance curves are useful for predicting fitness in fluctuating environments, Front. Ecol. Evol., 5, 129, <a href="https://doi.org/10.3389/fevo.2017.00129" target="_blank">https://doi.org/10.3389/fevo.2017.00129</a>, 2017.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib32"><label>32</label><mixed-citation>
      
Kim, D. H., Doyle, M. R., Sung, S., and Amasino, R. M.: Vernalization: winter and the timing of flowering in plants, Annu. Rev. Cell Dev. Biol., 25, 277–299, <a href="https://doi.org/10.1146/annurev.cellbio.042308.113411" target="_blank">https://doi.org/10.1146/annurev.cellbio.042308.113411</a>, 2009.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib33"><label>33</label><mixed-citation>
      
Knauer, J., Zaehle, S., Reichstein, M., Medlyn, B. E., Forkel, M., Hagemann, S., and Werner, C.: The response of ecosystem water‐use efficiency to rising atmospheric CO2 concentrations: sensitivity and large‐scale biogeochemical implications, New Phytol., 213, 1654–1666, <a href="https://doi.org/10.1111/nph.14288" target="_blank">https://doi.org/10.1111/nph.14288</a>, 2017.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib34"><label>34</label><mixed-citation>
      
Legendre, P. and Legendre, L.: Numerical Ecology, 3rd edn., Elsevier, Amsterdam, ISBN 978-0-444-53869-7, 2012.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib35"><label>35</label><mixed-citation>
      
Li, K., Wang, C., Sun, Q., Rong, G., Tong, Z., Liu, X., and Zhang, J.: Spring phenological sensitivity to climate Change in the northern hemisphere: comprehensive evaluation and driving force analysis, Remote Sens., 13, 1972, <a href="https://doi.org/10.3390/rs13101972" target="_blank">https://doi.org/10.3390/rs13101972</a>, 2021.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib36"><label>36</label><mixed-citation>
      
Li, X., Zhou, Y., Asrar, G. R., Mao, J., Li, X., and Li, W.: Response of vegetation phenology to urbanization in the conterminous United States, Glob. Change Biol., 23, 2818–2830, <a href="https://doi.org/10.1111/gcb.13562" target="_blank">https://doi.org/10.1111/gcb.13562</a>, 2017.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib37"><label>37</label><mixed-citation>
      
Malla, R., Neupane, P. R., and Köhl, M.: Climate change impacts: Vegetation shift of broad-leaved and coniferous forests, Trees For. People, 14, 100457, <a href="https://doi.org/10.1016/j.tfp.2023.100457" target="_blank">https://doi.org/10.1016/j.tfp.2023.100457</a>, 2023.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib38"><label>38</label><mixed-citation>
      
Mathur, S., Agrawal, D., and Jajoo, A.: Photosynthesis: response to high temperature stress, J. Photochem. Photobiol. B, 137, 116–126, <a href="https://doi.org/10.1016/j.jphotobiol.2014.01.010" target="_blank">https://doi.org/10.1016/j.jphotobiol.2014.01.010</a>, 2014.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib39"><label>39</label><mixed-citation>
      
Meng, L., Mao, J., Zhou, Y., Richardson, A. D., Lee, X., Thornton, P. E., Ricciuto, D. M., Li, X., Dai, Y., Shi, X., and Jia, G.: Urban warming advances spring phenology but reduces the response of phenology to temperature in the conterminous United States, P. Natl. Acad. Sci. USA, 117, 4228–4233, <a href="https://doi.org/10.1073/pnas.1911117117" target="_blank">https://doi.org/10.1073/pnas.1911117117</a>, 2020.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib40"><label>40</label><mixed-citation>
      
Mu, Q., Zhao, M., and Running, S. W.: Improvements to a MODIS global terrestrial evapotranspiration algorithm, Remote Sens. Environ., 115, 1781–1800, <a href="https://doi.org/10.1016/j.rse.2011.02.019" target="_blank">https://doi.org/10.1016/j.rse.2011.02.019</a>, 2011.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib41"><label>41</label><mixed-citation>
      
Pablos, M., Martínez-Fernández, J., Piles, M., Sánchez, N., Vall-llossera, M., and Camps, A.: Multi-temporal evaluation of soil moisture and land surface temperature dynamics using in situ and satellite observations, Remote Sens., 8, 587, <a href="https://doi.org/10.3390/rs8070587" target="_blank">https://doi.org/10.3390/rs8070587</a>, 2016.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib42"><label>42</label><mixed-citation>
      
Peng, J., Wu, C., Zhang, X., Wang, X., and Gonsamo, A.: Satellite detection of cumulative and lagged effects of drought on autumn leaf senescence over the Northern Hemisphere, Glob. Change Biol., 25, 2174–2188, <a href="https://doi.org/10.1111/gcb.14627" target="_blank">https://doi.org/10.1111/gcb.14627</a>, 2019.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib43"><label>43</label><mixed-citation>
      
Peng, S.: 1-km monthly precipitation dataset for China (1901–2021), National Tibetan Plateau/Third Pole Environment Data Center [data set], <a href="https://doi.org/10.5281/zenodo.3185722" target="_blank">https://doi.org/10.5281/zenodo.3185722</a>, 2020.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib44"><label>44</label><mixed-citation>
      
Peñuelas, J., Gordon, C., Llorens, L., Nielsen, T., Tietema, A., Beier, C., Bruna, P., Emmett, B., Estiarte, M., and Gorissen, A.: Nonintrusive field experiments show different plant responses to warming and drought among sites, seasons, and species in a north-south European gradient, Ecosystems, 7, 598–612, <a href="https://doi.org/10.1007/s10021-004-0179-7" target="_blank">https://doi.org/10.1007/s10021-004-0179-7</a>, 2004.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib45"><label>45</label><mixed-citation>
      
Piao, S., Fang, J., Zhou, L., Ciais, P., and Zhu, B.: Variations in satellite-derived phenology in China's temperate vegetation, Glob. Change Biol., 12, 672–685, <a href="https://doi.org/10.1111/j.1365-2486.2006.01123.x" target="_blank">https://doi.org/10.1111/j.1365-2486.2006.01123.x</a>, 2006.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib46"><label>46</label><mixed-citation>
      
Pinheiro, J. C. and Bates, D. M.: Mixed-Effects Models in S and S-PLUS, Springer, New York, ISBN 978-0-387-22747-4, <a href="https://doi.org/10.1007/b98882" target="_blank">https://doi.org/10.1007/b98882</a>, 2000.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib47"><label>47</label><mixed-citation>
      
Pollastrini, M., Puletti, N., Selvi, F., Iacopetti, G., and Bussotti, F.: Widespread crown defoliation after a drought and heat wave in the forests of Tuscany (Central Italy) and their recovery – A case study from summer 2017, Front. For. Glob. Change, 2, 74, <a href="https://doi.org/10.3389/ffgc.2019.00074" target="_blank">https://doi.org/10.3389/ffgc.2019.00074</a>, 2019.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib48"><label>48</label><mixed-citation>
      
Qiao, Z., Tian, G., and Xiao, L.: Diurnal and seasonal impacts of urbanization on the urban thermal environment: A case study of Beijing using MODIS data, ISPRS J. Photogramm. Remote Sens., 85, 93–101, <a href="https://doi.org/10.1016/j.isprsjprs.2013.08.010" target="_blank">https://doi.org/10.1016/j.isprsjprs.2013.08.010</a>, 2013.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib49"><label>49</label><mixed-citation>
      
Qiu, T., Song, C., and Li, J.: Impacts of urbanization on vegetation phenology over the past three decades in Shanghai, China, Remote Sens., 9, 970, <a href="https://doi.org/10.3390/rs9090970" target="_blank">https://doi.org/10.3390/rs9090970</a>, 2017.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib50"><label>50</label><mixed-citation>
      
Rustad, L., Campbell, J., Marion, G., Norby, R., Mitchell, M., Hartley, A., Cornelissen, J., Gurevitch, J., and Gcte, N.: A meta-analysis of the response of soil respiration, net nitrogen mineralization, and aboveground plant growth to experimental ecosystem warming, Oecologia, 126, 543–562, <a href="https://doi.org/10.1007/s004420000544" target="_blank">https://doi.org/10.1007/s004420000544</a>, 2001.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib51"><label>51</label><mixed-citation>
      
Sage, R. F. and Kubien, D. S.: The temperature response of C3 and C4 photosynthesis, Plant Cell Environ., 30, 1086–1106, <a href="https://doi.org/10.1111/j.1365-3040.2007.01682.x" target="_blank">https://doi.org/10.1111/j.1365-3040.2007.01682.x</a>, 2007.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib52"><label>52</label><mixed-citation>
      
Shi, C., Sun, G., Zhang, H., Xiao, B., Ze, B., Zhang, N., and Wu, N.: Effects of warming on chlorophyll degradation and carbohydrate accumulation of Alpine herbaceous species during plant senescence on the Tibetan Plateau, PLoS One, 9, e107874, <a href="https://doi.org/10.1371/journal.pone.0107874" target="_blank">https://doi.org/10.1371/journal.pone.0107874</a>, 2014.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib53"><label>53</label><mixed-citation>
      
Shi, W., Li, J., Zhan, H., Yu, L., Wang, C., and Wang, S.: Relation between water storage and photoassimilate accumulation of neosinocalamus affinis with phenology, Forests, 14, 531, <a href="https://doi.org/10.3390/f14030531" target="_blank">https://doi.org/10.3390/f14030531</a>, 2023.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib54"><label>54</label><mixed-citation>
      
Sperry, J. S., Hacke, U. G., Oren, R., and Comstock, J. P.: Water deficits and hydraulic limits to leaf water supply, Plant Cell Environ., 25, 251–263, <a href="https://doi.org/10.1046/j.0016-8025.2001.00799.x" target="_blank">https://doi.org/10.1046/j.0016-8025.2001.00799.x</a>, 2002.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib55"><label>55</label><mixed-citation>
      
Tian, F., Wigneron, J.-P., Ciais, P., Chave, J., Ogée, J., Peñuelas, J., Ræbild, A., Domec, J.-C., Tong, X., Brandt, M., Mialon, A., Rodriguez-Fernandez, N., Tagesson, T., Al-Yaari, A., Kerr, Y., Chen, C., Myneni, R. B., Zhang, W., Ardö, J., and Fensholt, R.: Coupling of ecosystem-scale plant water storage and leaf phenology observed by satellite, Nat. Ecol. Evol., 2, 1428–1435, <a href="https://doi.org/10.1038/s41559-018-0630-3" target="_blank">https://doi.org/10.1038/s41559-018-0630-3</a>, 2018.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib56"><label>56</label><mixed-citation>
      
Wan, Z.: New refinements and validation of the MODIS Land-Surface Temperature/Emissivity products, Remote Sens. Environ., 112, 59–74, <a href="https://doi.org/10.1016/j.rse.2006.06.026" target="_blank">https://doi.org/10.1016/j.rse.2006.06.026</a>, 2008.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib57"><label>57</label><mixed-citation>
      
Wang, C., Cao, R., Chen, J., Rao, Y., and Tang, Y.: Temperature sensitivity of spring vegetation phenology correlates to within-spring warming speed over the Northern Hemisphere, Ecol. Indic., 50, 62–68, <a href="https://doi.org/10.1016/j.ecolind.2014.11.004" target="_blank">https://doi.org/10.1016/j.ecolind.2014.11.004</a>, 2015.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib58"><label>58</label><mixed-citation>
      
Wang, C., Tang, Y., and Chen, J.: Plant phenological synchrony increases under rapid within-spring warming, Sci. Rep., 6, 25460, <a href="https://doi.org/10.1038/srep25460" target="_blank">https://doi.org/10.1038/srep25460</a>, 2016.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib59"><label>59</label><mixed-citation>
      
White, M. A., De Beurs, K. M., Didan, K., Inouye, D. W., Richardson, A. D., Jensen, O. P., O'Keefe, J., Zhang, G., Nemani, R. R., Van Leeuwen, W. J. D., Brown, J. F., De Wit, A., Schaepman, M., Lin, X., Dettinger, M., Bailey, A. S., Kimball, J., Schwartz, M. D., Baldocchi, D. D., Lee, J. T., and Lauenroth, W. K.: Intercomparison, interpretation, and assessment of spring phenology in North America estimated from remote sensing for 1982-2006, Glob. Change Biol., 15, 2335–2359, <a href="https://doi.org/10.1111/j.1365-2486.2009.01910.x" target="_blank">https://doi.org/10.1111/j.1365-2486.2009.01910.x</a>, 2009.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib60"><label>60</label><mixed-citation>
      
Wu, C., Peng, J., Ciais, P., Peñuelas, J., Wang, H., Beguería, S., Black, T. A., Jassal, R. S., Zhang, X., Yuan, W., Liang, E., Wang, X., Hua, H., Liu, R., Ju, W., Fu, Y. H., and Ge, Q.: Increased drought effects on the phenology of autumn leaf senescence, Nat. Clim. Change, 12, 943–949, <a href="https://doi.org/10.1038/s41558-022-01464-9" target="_blank">https://doi.org/10.1038/s41558-022-01464-9</a>, 2022.


    </mixed-citation></ref-html>
<ref-html id="bib1.bib61"><label>61</label><mixed-citation>
      
Xiong, X., Wu, H., Wei, X., and Jiang, M.: Contrasting temperature and light sensitivities of spring leaf phenology between understory shrubs and canopy trees: Implications for phenological escape, Agr. Forest Meteorol., 355, 110144, <a href="https://doi.org/10.1016/j.agrformet.2024.110144" target="_blank">https://doi.org/10.1016/j.agrformet.2024.110144</a>, 2024.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib62"><label>62</label><mixed-citation>
      
Yang, H., Zhang, Q., Zhang, S., Wang, X., and Yu, H.: Joint control of urban expansion and climate change on urban-rural vegetation phenology gradient in 31 cities of China, Front. Ecol. Evol., 13, 1637210, <a href="https://doi.org/10.3389/fevo.2025.1637210" target="_blank">https://doi.org/10.3389/fevo.2025.1637210</a>, 2025.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib63"><label>63</label><mixed-citation>
      
Yao, R., Wang, L., Huang, X., Guo, X., Niu, Z., and Liu, H.: Investigation of Urbanization effects on land Surface phenology in Northeast China during 2001–2015, Remote Sens., 9, 66, <a href="https://doi.org/10.3390/rs9010066" target="_blank">https://doi.org/10.3390/rs9010066</a>, 2017.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib64"><label>64</label><mixed-citation>
      
Yu, F., Price, K. P., Ellis, J., and Shi, P.: Response of seasonal vegetation development to climatic variations in eastern central Asia, Remote Sens. Environ., 87, 42–54, <a href="https://doi.org/10.1016/S0034-4257(03)00144-5" target="_blank">https://doi.org/10.1016/S0034-4257(03)00144-5</a>, 2003.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib65"><label>65</label><mixed-citation>
      
Yuan, M., Zhao, L., Lin, A., Wang, L., Li, Q., She, D., and Qu, S.: Impacts of preseason drought on vegetation spring phenology across the Northeast China Transect, Sci. Total Environ., 738, 140297, <a href="https://doi.org/10.1016/j.scitotenv.2020.140297" target="_blank">https://doi.org/10.1016/j.scitotenv.2020.140297</a>, 2020.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib66"><label>66</label><mixed-citation>
      
Zhang, X., Friedl, M. A., Schaaf, C. B., Strahler, A. H., and Schneider, A.: The footprint of urban climates on vegetation phenology, Geophys. Res. Lett., 31, L12209, <a href="https://doi.org/10.1029/2004GL020137" target="_blank">https://doi.org/10.1029/2004GL020137</a>, 2004.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib67"><label>67</label><mixed-citation>
      
Zhang, X., Friedl, M. A., and Schaaf, C. B.: Global vegetation phenology from Moderate Resolution Imaging Spectro radiometer (MODIS): Evaluation of global patterns and comparison with in situ measurements, J. Geophys. Res.-Biogeo., 111, G04017, <a href="https://doi.org/10.1029/2006JG000217" target="_blank">https://doi.org/10.1029/2006JG000217</a>, 2006.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib68"><label>68</label><mixed-citation>
      
Zhang, X., Chen, N., Sheng, H., Ip, C., Yang, L., Chen, Y., Sang, Z., Tadesse, T., Lim, T. P. Y., Rajabifard, A., Bueti, C., Zeng, L., Wardlow, B., Wang, S., Tang, S., Xiong, Z., Li, D., and Niyogi, D.: Urban drought challenge to 2030 sustainable development goals, Sci. Total Environ., 693, 133536, <a href="https://doi.org/10.1016/j.scitotenv.2019.07.342" target="_blank">https://doi.org/10.1016/j.scitotenv.2019.07.342</a>, 2019.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib69"><label>69</label><mixed-citation>
      
Zheng, W., Liu, Y., Yang, X., and Fan, W.: Spatiotemporal variations of forest vegetation phenology and its response to climate change in northeast China, Remote Sens., 14, 2909, <a href="https://doi.org/10.3390/rs14122909" target="_blank">https://doi.org/10.3390/rs14122909</a>, 2022.

    </mixed-citation></ref-html>
<ref-html id="bib1.bib70"><label>70</label><mixed-citation>
      
Zhou, D., Zhao, S., Zhang, L., and Liu, S.: Remotely sensed assessment of urbanization effects on vegetation phenology in China's 32 major cities, Remote Sens. Environ., 176, 272–281, <a href="https://doi.org/10.1016/j.rse.2016.02.010" target="_blank">https://doi.org/10.1016/j.rse.2016.02.010</a>, 2016.

    </mixed-citation></ref-html>--></article>
