# Elevational shift in seed plant distributions in China's mountains over the last 70 years

Kuiling Zu, Jonathan Roger Michel Henri Lenoir, Jingyun Fang, Zhiyao Tang, Zehao Shen, Chengjun Ji, Chengyang Zheng, Ao Luo, Wenqi Song, Niklaus Zimmermann, et al.

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► To cite this version:

Kuiling Zu, Jonathan Roger Michel Henri Lenoir, Jingyun Fang, Zhiyao Tang, Zehao Shen, et al. "Elevational shift in seed plant distributions in China's mountains over the last 70 years." Global Ecology and Biogeography, 2023, 32 (7), pp.1098-1112. 10.1111/geb.13692. hal-04262552

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Title page

Title:

Elevational shift in seed plant distributions in China's mountains over
the last 70 years

Running Title : Elevational shifts of plants in China

List of Authors:

Kuiling Zu¹,², Jonathan Lenoir³, Jingyun Fang¹,⁴, Zhiyao Tang¹, Zehao Shen¹, Chengjun Ji¹, Chengyang Zheng¹, Ao Luo¹, Wenqi Song¹, Niklaus E. Zimmermann⁵, Loïc Pellissier⁵,⁶, Zhiheng Wang¹*

**Institutional affiliations:**

¹Institute of Ecology, and Key Laboratory for Earth Surface Processes of the Ministry of Education, College of Urban and Environmental Sciences, Peking University, Beijing 100871, China

²Key Laboratory of National Forestry and Grassland Administration on Forest Ecosystem Protection and Restoration of Poyang Lake Watershed, College of Forestry, Jiangxi Agricultural University, Nanchang 330045, China

³UMR CNRS 7058 "Ecologie et Dynamique des Systèmes Anthropisés" (EDYSAN), Université de Picardie Jules Verne, 1 Rue des Louvels, 80000 Amiens, France

⁴College of Ecology and Environmental Sciences, Yunnan University, Chenggong, Kunming 650500, China

⁵Landscape Ecology, Institute of Terrestrial Ecosystems, Department of Environmental Systems Science, ETH Zürich, Zürich, Switzerland

⁶Unit of Land Change Science, Swiss Federal Research Institute WSL, Birmensdorf, Switzerland

Author for correspondence:

*Email: zhiheng.wang@pku.edu.cn

Telephone: 010-62745157
**Acknowledgements:**

We gratefully thank the data supports by the National Specimen Information
Infrastructure (NSII). We also thank Nawal Shrestha and Yaoqi Li for discussion and
help improving the manuscript. This work was supported by the Strategic Priority
Research Program of Chinese Academy of Sciences (XDB31000000), the Natural
Science Foundation of Jiangxi, China (20224BAB213033), Jiangxi Provincial
Department of Education Science and Technology Research Project (GJJ2200433), the
National Key Research Development Program of China (2018YFA0606104;
#2017YFA0605101) and the National Natural Science Foundation of China
(#32125026, #31988102).

**Biosketch:**

Kuiling Zu is a teacher at Jiangxi Agricultural University, and she did postdoctoral work at Peking University, and interested in macroecology and botany, currently studying the impacts of global changes on plant resources. Her current research interest is to explore the response of species distributions and phenology to climate change.
1 **Abstract**

2 **Aim:** Significant changes in species elevational ranges in mountains have been repeatedly documented, yet the direction, magnitude and drivers of these shifts remain controversial.

3 Presently, there is still lacking evidence about the general nature of species elevational range shifts in eastern Eurasia in response to anthropogenic climate change. By using historical specimen records and recent field observations for 735 seed plant species across 29 China's mountains, we assessed changes in species' elevational centroids and their drivers.

4 **Location:** China.

5 **Time period:** 1950 - 2018.

6 **Major taxa studied:** Seed plant species.

10 **Methods:** The elevation records of all sampled occurrences in each mountain during the two time periods were estimated, and the null models were developed to test the sampling bias. Ecological niche models (ENMs) were used to evaluate the relative importance of climate factors in constraining each species distribution. Generalized linear models (GLMs) to test the relationships between the centroid elevational range shifts of species and different divers.

15 **Results:** We found that 54% of the species shifting upward and 46% downhill. However, species' elevational shifts significantly differed among species and mountains. Herbaceous and lowland species moved upward faster than woody and high-elevation species. Species in temperate mountains and in mountains with taller elevational gradients moved upward, while species in subtropical mountains and in mountains with shorter elevational gradients moved downward. Precipitation changes experienced by species, species' climatic adaptations, several species' functional traits and mountain size all contributed to explain the magnitude of species' centroid elevational range shifts.

20 **Main conclusions:** Our results highlight complex biodiversity redistribution of seed plants across Chinese mountains, not necessarily conforming to the trend of species upward shifts in elevation. Changes in precipitation regimes may blur the simplistic assumption of isotherm tracking. This study fills an important geographic shortfall for our understanding of biodiversity redistribution under anthropogenic climate change.

25 **KEYWORDS**

30 **biodiversity redistribution, climate change, climatic niche, elevational gradients, macroecology,** precipitation, upslope range shifts, velocity of species range shift

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32 **1 INTRODUCTION**

Anthropogenic climate change is a global threat that has the potential to impact species distributions and ecosystem functioning (Pecl et al., 2017). To better understand the possible impacts of climate change on species distributions, studies have examined the responses of individual species in different mountains and taxonomic groups worldwide (Chen et al., 2009; Crimmins et al., 2011; Freeman et al., 2018; Lenoir et al., 2008; Moritz, 2008; Rumpf et al., 2019; Telwala, 2013; Zu et al., 2021). These original reports collectively provide strong evidence that most species moved upward as climate warmed, while some moved downward or did not move (Lenoir et al., 2010; Maggini et al., 2011). Indeed, based on original studies and meta-analyses strictly or partly focusing on species elevational range shifts, we know that: (1) not all species are moving upward; (2) regional differences in species responses exist; and (3) each biotic response depends on local (and sometimes idiosyncratic) drivers (Chen et al., 2009; Freeman et al., 2021; Lenoir et al., 2020; Parmesan, 2003; Vittasse et al., 2021). Although a few recent studies assessed the elevational changes in species distributions (Wen et al., 2017; Zu et al., 2021) and treelines (Liang et al., 2016; Shi et al., 2020) where data is scarce, there is still lacking evidence about the general nature of species elevational range shifts in eastern Eurasia in response to recent climate change. Hence, filling this geographical gap can help reaching a truly global assessment on biodiversity redistribution in mountain ecosystems worldwide (Feeley et al., 2017; Lenoir & Svenning, 2013; Lenoir & Svenning, 2015).

Mountain regions are biodiversity hotspots and their complex and diverse environments provided refugia under past climatic fluctuations since the Quaternary (López-Pujol et al., 2011; Sandel et al., 2011). Recent studies indicated that warming rate in the European Alps is projected to be 1.6 to 2.5 times faster than the average warming rate observed over the northern hemisphere during the 20th century (Vittasse et al., 2021). For most regions of China, temperatures have been observed to have increased by a rate of 0.05°C/yr between 1961 and 2004 (Wang et al., 2010). More importantly, Nogués-Bravo et al. (2007) predicted that air temperature would increase by 0.30 to 0.45°C per decade until 2100. Temperature has long been considered as the most important driver limiting plants' and animals' distributions along elevational gradients in mountain ecosystems (Grinnell, 1917). However, the general and
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61 overly simplistic hypothesis that increasing temperature is the main determinant pushing species to shift their elevational range upwards ignores the potential compounding effect of precipitation changes (Crimmins et al., 2011) or other abiotic as well as biotic factors (Lenoir et al., 2010). In line with this result, Grytnes et al. (2014) showed that climate warming alone had low explanatory power to explain the observed elevational range shifts of alpine plants across European mountains. However, the relative effects of global warming and precipitation changes on species range shifts in mountains remain controversial. Moreover, whether the response of different mountain plants to climate change is consistent across mountains and which factors influence species' elevational redistributions under climate change remains largely unknown for most Asian mountains.

71 The impact of changes in abiotic (e.g., temperature and precipitation) is not enough to explain the direction and magnitude of species elevational range shifts. Indeed, species intrinsic abilities to cope with these changes matter too. Overall, species with different life forms may respond differently to the same amount of change in environmental conditions (Stewart & Dalen, 2010). For instance, Lenoir et al. (2008) reported that short-lived herbaceous plant species shifted farther upward than long-lived woody plants (trees and shrubs), which can be attributed to the differences in life cycles and the respective capacity to migrate through effective seed dispersal and successful seed germination. Moreover, recent analyses focusing on mountain regions in Europe and worldwide suggests that low-elevation species shifted upward faster than high-elevation species (Mamantov et al., 2021; Rumpf et al., 2018). Consequently, the velocity of elevational range shifts in response to climate change varied among different life forms and across elevations. Inconsistent responses of different species to climate change imply that species interactions are likely to change over multiple trophic levels (Vitasse et al., 2021), thus affecting the structures and functioning of mountain ecosystems at different elevations. Hence, more data are needed on species elevational range shifts to forecast these future disruptions within existing ecological networks, especially for understudied regions such as Asian mountains.

88 Herbarium specimens are sustainable and persistent, providing accurate information on species distributions during both past and present collecting times (Holmes et al., 2016; Page
90 & Riccardi, 2015). The collection density of herbarium specimens in China's mountains is considered intensive and complete according to the database of Chinese vascular plant distributions (Yang et al., 2014). Therefore, herbarium specimens are a key resource to improve our understanding of the impacts of climate change on species distributions and redistributions through space and time (James et al., 2018). In this study, we combined old herbarium specimens with recent botanical field surveys to investigate changes in the elevational distributions of 735 seed plant species across 29 mountains in China. We aimed to: (1) assess the direction and magnitude of species centroid elevational shifts along the elevational gradients in China's mountains since the 1950s, assuming a general trend in seed plant species in China to shift upward in elevation (Zu et al., 2021); (2) explore the main determinants of the observed changes in range centroids across the studied mountains, assuming that temperature changes won't be the main or only driver (Grytnes et al., 2014); (3) compare the magnitude of centroid range shifts between herbaceous and woody plants, assuming greater magnitudes of range shifts for short-lived herbs (Lenoir et al., 2008); and (4) compare the difference in centroid range shifts between low- and high-elevation species, expecting greater magnitudes of range shifts for low-elevation species (Mamantov et al., 2021; Rumpf et al., 2018).

## 2 MATERIALS AND METHODS

### 2.1 The study area

The study area covers 29 mountain sites in China (Figure 1; Table S1), which are mainly distributed across 18 distinct mountain ranges, including the Qionglai, Bayankela, Hengduan, Changbai, Yanshan, Wuling, Lvliang, Daba, Wuyi, Dalou, Miaoling, Lushan, Nanling, Himalayan, Qinling, Tianshan, Taihang, and Aertai Mountain Range. These mountain sites are distributed across 15 provinces and 3 climatic zones (Plateau, Temperate and Subtropics) within China.

There are 12 mountain sites that are located in the climatic zone of Qinghai-Tibet Plateau in western China. The highest mountain is Mt. Namjagbarwa, which is located at the eastern edge of the Himalayan Mountain Range in the southeast of the Qinghai-Tibet Plateau, reaching its highest elevation at 7782 m. Eight mountain sites are located in the Hengduan Mountain Range, the core biodiversity hotspot in south central China (Myers, 2000). The tallest mountain in that
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mountainous area is Mt. Gongga with an elevation peaking at 7556 m (Zu et al., 2021). The

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Hengduan Mountain Range is affected by the Indian and Pacific Ocean monsoon circumfluence,

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resulting in a distinct wet season during summer and autumn and a dry season during winter

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and spring. This region is characterized by a sharp elevational gradient from approximately 70

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to 7556 m, offering an exceptionally long and diverse range of topographical and climatic

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conditions for studying species elevational range shifts in response to climate change (Wu et

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al., 2013). The Qionglai Mountain Range is adjacent to the Hengduan Mountain Range, which

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is located in the Sichuan province. Two mountain sites belong to the Qionglai Mountain Range:

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the Siguniang (SGN) and Balang (BL) site, with the highest elevation being 6250 m and 5072

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m, respectively.

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There are 10 mountain sites that are located in the climatic zone of the subtropics in southern

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China. The lowest mountain ranges are the Lushan and Maoer Mountain Range in southeast

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China, with the highest elevations being 1474 m and 2142 m, respectively. Mt. Hualong (HL)

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and Mt. Shennongjia (SNJ) are located in the Daba Mountain Range, which constitute the main

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body of China's North-South Transitional Zone (Zhang et al., 2021). Mont Fanjing (FJ), Mt.

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Fenghuang (FH) and Mt. Leigong (LG) are located in the Guizhou province while Mt. Jinfo

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(JF) is located at the northeast of the Dalou Mountain Range with the highest elevation being

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2251 m.

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Our study region also includes seven mountain sites that are located in the temperature

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climate zone in the northern part of China. Mont Taibai is the summit of the Qinling Mountain

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Range and the highest mountain in eastern China – reaching its highest elevation at 3767 m

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(Zhang & Chen, 2014). Noteworthy, our study region also covers the Changbai Mountain

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Range which is located in the northeast of China and is situated on the border between China

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and North Korea (Bai et al., 2008). The forested areas in this mountain range is distributed

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below 2000 m and the typical zonal vegetation is a mixed forest made of Korea pines and broad-

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leaved tree species.

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## 2.2 Data on seed plant species

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Plant distribution data across the 29 mountain sites in China were obtained from: (1) historical

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specimens collected during the past 70 years and available through museum and herbarium
collections as well as (2) original field surveys and resurveys carried out during the past 20 years. Herbarium records were extracted from the National Specimen Information Infrastructure (NSII, http://www.nsii.org.cn, accessed on November 11, 2019) and the Chinese Virtual Herbarium (CVH, http://www.cvh.ac.cn). In total, 10918077 herbarium specimens were extracted for all seed plant species in China, covering about 34000 species. Besides, between 2000 and 2019, we carried out original botanical surveys in: the Changbai Mountain Range; the Xiaowutai Mountain Range; the Gongga Mountain Range; the Taibai Mountain Range; and in several other mountain ranges of smaller extents. Botanical surveys were carried out in vegetation plots of 600 m² (20 m × 30 m), which were established along elevational transects from low to high elevations with an elevational interval of about 50 m between two subsequent vegetation plots. The vegetation plots selected for each elevation represented the zonal vegetation types in the areas. In each vegetation plot, we also measured the diameter at breast height (DBH) of all tree individuals with a DBH > 3 cm (Fang et al., 2012).

To ensure data quality, we cleaned the plant species distribution data as following. As a first step, we extracted the botanical specimen data of each county of the corresponding mountain site, counted the number of plant specimens in each site and determined the searched area within each mountain site. Then, we cleaned up the geographical names and verified the accuracy of plant distribution points within each mountain site. We excluded duplicate specimens and specimens lacking information on their exact geographical locations, and only included records with correct information (e.g., elevation record and collection year). Finally, over 5000 complete records were selected across the 29 mountain sites. The total number of herbarium specimens we selected for each mountain site are shown in Table 1. As a second step, we standardized the species names of all records based on the Catalogue of Life (COL, http://www.catalogueoflife.org/) which was supplemented with the plant list (TPL, http://www.theplantlist.org/) as well as the Taxonomic Name Resolution Service (TNRS, http://tnrs.plantcollaborative.org/). As a third step, in order to assess changes in species' elevational distributions between the historical and present times, we divided our dataset of species occurrence records into two distinct periods: 1950 - 1988 vs. 1989 - 2018. The threshold time of 1988 / 1989 was chosen because mean annual temperature in China started to
significantly increase at this time (Wang et al., 2010). Finally, for statistical reason, we selected only seed plant species with at least 10 unique occurrence records for each of the two studied periods within a given mountain site. The number of the selected seed plant species considered per site is shown in Table 1 and Figure S1. The occurrence records for the selected seed plant species per site during the first period (1950 - 1988) and the second period (1989 - 2018) is displayed in Figure S2. The centroid elevation per species was calculated in each mountain range during the two time periods, using the average summary statistic across all sampled occurrences during a given time period, within a given mountain range and for a given focal species. Because the centroid elevation represents the distribution center of the populations for each species, and therefore it can represent the elevation shifts.

## 2.3 Climate data

The climate data (daily temperature and precipitation data covering the period 1951 - 2015 and available from the Geographic Data Sharing Infrastructure, College of Urban and Environmental Science, Peking University: http://geodata.pku.edu.cn) for all 29 mountain sites were extracted from 54 standard meteorological stations across China (Figure S3). These 54 meteorological stations are representative of the macroclimatic conditions experienced within our study areas. For each of the 29 studied mountain sites, we first calculated the mean annual temperature (MAT, °C), the total amount of annual precipitations (AP, mm) and the annual actual evapotranspiration (AET, mm). These three climatic variables were computed for each year during the entire period of 1951 - 2015. Then, we computed the mean of each of the three variables for the periods 1951 - 1988 and 1989 - 2015 separately before estimating the mean changes in MAT, AP and AET between the two periods. The climatic variable AET was calculated following the Thornthwaite and Hare formula (Huntley, 1993), based on the mean monthly temperature and precipitation data. The climatic time series across the 54 meteorological stations showed clear trends of climate warming and precipitation decrease but a less clear trend of change in AET between the two studied periods (Figure S4, S5, and S6). In addition to that, we also computed seasonal changes by focusing on the change in temperatures and precipitations during summer (May - July) and winter (November - Next January) separately and for each of the 29 mountain sites (Figure S7).
## 2.4 Complementary drivers of changes in species' centroid elevational position

Besides climatic drivers, being the usual culprits responsible for species range shifts under anthropogenic climate change, we extracted information on additional potential drivers to explain the variation we found in species' centroid elevational range shifts of seed plants. First, we downloaded a spatial layer of the human footprint index (HI) at 1km resolution from the Last of the Wild (v2, LWP-2, 2005; https://sedac.ciesin.columbia.edu) and extracted HI values across our 29 mountain sites. Then, we extracted data on atmospheric nitrogen deposition from on a national dataset providing atmospheric inorganic nitrogen deposition across China from 1996 to 2015 (spatial raster layers at 1 km × 1 km resolution) (Jia et al., 2019). From this raster layer, we calculated the mean nitrogen deposition across each of the 29 mountain sites.

Previous studies suggest that mountain size, in terms of the length of the tallest elevational gradient within a given mountain range, could significantly influence the distribution of treelines and snowlines in southeastern Eurasia (Yao & Zhang, 2014). In order to assess the impact of the mountain size on the observed species' centroid elevational range shifts, we used the difference between the highest and lowest elevation (i.e., the elevation range) within each of the 29 mountain sites to represent the site-specific mountain size. The highest elevation of each mountain were obtained from Wang et al. 2004, and the lowest elevation of each mountain was obtained from literature collections (Table S2).

We also compiled data on several plant species functional traits, including: lifeform (herbaceous vs woody plants), the main reproductive system (SexSyst), and fruit type. These trait data were obtained from Flora of China (http://www.iplant.cn/foc), Flora Reipublicae Popularis Bulgaricae (http://www. iplant.cn/frps) and other provincial and local floras in China.

## 2.5 Data analysis

We calculated the centroid elevational position of each species in each of the 29 mountain sites during each time period separately (1950 - 1988 vs. 1989 - 2018) by averaging the elevation records of all sampled occurrences of a given species within a given mountain and during a given time period. The shift in the species' centroid elevation position within each mountain site was computed as the raw difference in the species' centroid position, for the focal mountain site, between the baseline (1950 - 1988) and resurvey period (1989 - 2018): resurvey minus
baseline position. Pairwise Student *t* tests were used to test the significance in the mean difference in species' centroid elevational position between the two studied time periods, for each mountain site separately. We also calculated the proportion of species shifting upward and downward for each mountain site separately. To test whether there is a sampling bias caused by a tendency that current researchers would sample at higher or lower elevations than in the past, we built null models to estimate the null expectation of the elevation change in the centroid position of a given species, for each of the 29 mountains separately, under the actual sampling effort per mountain site. The specific methods and steps to build the null models is the same as the one we used in a previous study focusing on one of the study site: Mt. Gongga (Zu et al., 2021). To test the influence of species lifeform on species' centroid elevational range shifts, we used Student *t* tests to compare the mean elevation shifts of herbaceous and woody plants, for each mountain separately.

In order to estimate whether climate changes influence species elevational shifts, we estimated the climate changes (Change in MAT and Change in AP) experienced by each species. To assess the effects of several climatic drivers constraining each species distribution and thus potentially explaining the observed magnitude of species' centroid elevational range shifts, we first collected data on the global distribution of each of the 735 studied species and then rendered an ecological niche modelling (ENM) approach. To do so, we used the MaxEnt algorithm to estimate the relative importance of six climatic variables in explaining each species distribution. We followed the exact same methodology as described in Zu et al. (2021) for one study site on Mt. Gongga. In short, the global distribution ranges of species were obtained from online databases including: the Global Biodiversity Information Facility (GBIF, http://www.gbif.org/); the Chinese Virtual Herbarium (CVH, http://www.cvh.ac.cn); the National Specimen Information Infrastructure (NSII, http://www.nsii.org.cn); RAINBIO (Dauby et al., 2016); and the Botanical Information and Ecology Network (BIEN, http://bien.nceas.ucsb.edu/bien/bindata/), etc. The climate data were from WorldClim (https://www.worldclim.org/data/worldclim21.html). Then, once the relative importance of each of the above-mentioned climatic variables were estimated from the ENM approach, we used generalized linear models (GLMs) to test the strength of the relationship between the magnitude of species' centroid elevational range shifts and the relative importance of each of the six studied climatic drivers. In addition to these six explanatory variables, we also tested the
influence of HI and atmospheric nitrogen deposition. Finally, we also assessed the relationship between species' centroid elevational range shifts and the historical mean elevation position (cf. baseline position) for each species in the 29 mountain sites. All analyses were conducted in R version 3.5.3 (R Core Team, 2019).

# 3 RESULTS

## 3.1 Changes in species' centroid elevational position in different mountains

Between the periods 1950-1988 and 1989-2018, we found that 54% (± 21%) of the studied seed plant species shifted the centroids of their elevational ranges upward across the 29 mountain sites. However, the proportion of species with upward centroid elevational shifts significantly varied among mountain sites. Specifically, 17 of the 29 mountain sites had higher proportions of species shifting upward (69% ± 9%, on average), while 10 mountain sites had lower proportions of species shifting upward (29 ± 10%, on average) (Figure 1). The two remaining mountain sites (Mt. Huanggang and Mt. Tiangeer) had even proportions of species shifting upward and downward (Figure 1; Table 1). After accounting for the difference in sampling efforts between time periods, the proportions of species shifting upward vs. downward across the 29 mountain sites remained similar (Table S3). These results indicate that most plant species in the 29 studied mountain sites have shifted their elevational distributions during the past 70 years, and that these elevational range shifts were not the result of a sampling bias.

## 3.2 The driving factors of species' centroid elevational range shifts

We found that precipitation changes as experienced by species (i.e., the relative importance of changes in AP between 1951 - 1988 and 1989 - 2015), mean climatic conditions (i.e., the relative importance of MAT, AP and AET during 1951 - 2015), species' intrinsic characteristics (i.e., elevational range size, fruit type, family and genus) and mountain sites' properties (climatic region, longitude and mountain size) all significantly contributed to explain the magnitude of species' centroid elevational range shifts (Table 2). These variables collectively explained 66% (R² = 0.66, p < 0.001) of the total variance in species' centroid elevational range shifts across all mountains.

Specifically, the mean magnitude of species' centroid elevational range shifts were negatively correlated with mean annual temperature and changes in summer and winter
precipitation, while positively correlated with mountain size (Figure 1 and Figure S8). In other words, species tend to move further upward in mountains located in colder regions, within tall mountain ranges and in mountains located in drier regions.

The climatic adaptation of a species was significantly correlated with its shift in centroid elevation. Specifically, the centroid elevation shifts of species were positively correlated with the climatic adaptation to mean temperature of coldest quarter (MTCQ) and precipitation seasonality (PS), but negatively correlated with the climatic adaptation to precipitation of warmest quarter (PWQ) (Figure 2). Moreover, species with differing fruit types showed different elevational shifts (Figure S9a). In different climate regions, species centroid elevational shifts differed. For instance, species in subtropics regions tended to shift downward, while those inhabiting the Qinghai-Tibet Plateau tended to upward shifts on average (Figure S9b). The centroid elevation shifts of species were negatively correlated with longitude, and positively correlated with the relative elevation (Figure 2).

## 3.3 Differences in elevational shifts of herbaceous and woody plants

The magnitude of species' centroid elevational range shifts differed between herbaceous and woody plants. Herbs tended to move upward in the Qinghai-Tibet Plateau, while woody plants tended to move upward in the Temperate zone (Figure S9c). As mean annual temperature decreases, both herbaceous and woody plants shifted further upward (Figure S8). In cold mountains, upward range shifts of herbs (+90 m, on average) was greater than that of woody plants (+74 m, on average). In warm mountains, downward range shift of herbs (-170 m, on average) was greater than that of woody plants (-50 m, on average) (Figure 3). Both the elevational range shifts of herbs and woody plants were influenced by the change in precipitation seasonality. However, the elevational shifts of herbs were also significantly influenced by mountain size (Figure S8).

## 3.4 The relationship between species' centroid elevational range shifts and the historical mean elevation of species distribution

We found that species' centroid elevational range shifts were negatively correlated with the historical mean elevation position of species in nine out of the 29 studied mountain sites (Table S4). The mountains with the strongest negative correlation values are mainly Mt. Xiaowutai
(XWT, 0.55) and Mt. Lushan (LS, 0.49) (Figure 4). In these mountains, the high-elevation species tended to shift downward, while the low-elevation species tended to shift upward, resulting in a mid-elevation concentration. The results in these mountains support the hypotheses of a negative relationship between the magnitude and direction of elevational range shifts and the historical position of species along the studied elevational gradients, suggesting that lowland species move upward while alpine species move downward. In contrast, the remaining 20 mountains sites did not support this hypothesis (Table S4).

4 DISCUSSION

We assessed the changes in species' range centroid along elevational gradients using historical and recent (1950 - 1988 vs. 1989 - 2018) occurrence records for 735 seed plant species across 29 mountains in China and explored the drivers of these changes in species distribution over time. Similarly to previous meta-analyses (Chen et al., 2011; Lenoir et al., 2020; Mamantov et al., 2021), we found that not all studied species are moving upward along the studied elevational gradients of our 29 mountain sites in China. Yet, we found a much larger proportion of species shifting their centroid elevation position downward in comparison with these former meta-analyses reporting not more than 20% of species shifting downward against 46% based on our findings. Moreover, we found that herbaceous plant species shifted their centroid elevation further upward, by +16 m on average, than woody plants did, which is consistent with a former study reporting faster rates of upward range shifts for herbs than for woody plants in French mountain forests (Lenoir et al., 2008). Centroid shifts show negative relationships with the average elevation of species in 23 mountains, suggesting that lowland species have shifted upward farther than high-elevation species. Overall, our results suggest that species in mountains are not necessarily moving upward following and according to the rather simplistic isotherm-tracking expectation. Instead, species may demonstrate more complex elevational redistribution in response to environmental changes. The elevational redistributions we found in our study are not entirely due to climate change (e.g. climate warming), but are additionally influenced by other factors such as, among others described below, changes in precipitation regimes, which supports former findings (Crimmins et al., 2011; Grytnes et al., 2014). Our results indicate that several factors contribute to the elevational changes of species in Chinese
mountains: (1) climate change and species' climatic adaptations; (2) mountain properties; (3) the functional traits of different species.

## 4.1 Effect of climate change and species' climatic adaptations on elevational shifts

Climatic conditions in Chinese mountains have significantly changed over the past 70 years, which has influenced species' elevational distributions. Our results indicate that the amount of climate change as experienced by seed plant species significantly influenced the direction and magnitude of species' centroid elevational range shifts. This finding is consistent with previous studies reporting that changes in temperature and precipitation regimes play an important role in species' elevational redistributions (Chen et al., 2009; Grytnes et al., 2014). Previous studies suggested that the overall trend of upward range shifts of forest plant species in France is mainly driven by climate warming (Lenoir et al., 2008), while the overall trend of downward range shifts of woody plants in California is chiefly influenced by precipitation changes (Crimmins et al., 2011). However, our study suggests that temperature changes as experienced by seed plant species do not necessarily play an important role in explaining the overall trend of elevational range shifts in Chinese mountains. In contrast, changes in annual precipitation between the two studied periods and as experienced by species had a significant negative effect on species elevational range shifts (Figure S8). Our findings regarding the insignificant effect of temperature changes over time on the magnitude of species' centroid elevational range shifts might stem from a spatial mismatch between climate as measured by weather stations and climate as experienced by living organisms. Indeed, air temperatures as measured by standardized weather stations normally differ from the actual surface temperatures that low-stature alpine plants may experience in-situ, inside their habitats (Daniel & Körner, 2011). This suggests that macroclimatic conditions as measured by meteorological stations may to some extent lead to low accuracy in predicting species redistributions. Recent studies analyzed multi-decadal understory microclimate dynamics in European forests and provided evidence that responses of forest community to climate change are most closely related to microclimate rather than macroclimate changes (Zellweger et al., 2019). This indicates that local microclimate may hold the key to understand biotic responses, such as species elevational range shifts, to anthropogenic climate change (Alexander et al., 2018; De Frenne et al., 2013; Devictor et al.,
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382 2012).

383 In this study, lowland species tended to shift upward faster than high-elevation species in
384 23 mountains. This result is consistent with a recent finding that most lowland species across
385 20 montane transects moved upward farther than the high-elevation species (Mamantov et al.,
386 2021). Our results together with previous studies seem to indicate a general trend for species
elevational redistribution to be more pronounced at low elevations than at high elevations in
387 response to anthropogenic climate change. In general, the increase in temperature conditions
388 over time is stronger in highland regions than in the lowlands (Pepin et al. 2015), and the
389 stronger warming and precipitation decrease have negative effects on plant nutrient use and
390 resorption efficiencies for the lowland species (Zu et al., 2021). This might be one of the reasons
391 for the stronger upward shifts of lowland species. Moreover, lowland plant species normally
392 have narrower thermal tolerances (including temperature and precipitation) so that they need to
393 migrate farther upward to track their thermal niche (Smith, 2018).
394

395 Interestingly, we found that the adaptation of species to climate has significant effects on
396 the elevational changes of species. Specifically, the adaptation of species to winter temperature
397 and precipitation seasonality was positively correlated with species elevational changes, and
398 hence significantly contributed to the upward shifts of species. For instance, the global
distribution ranges of two lowland species, *Sorbus discolor* and *Cerastium arvense*, were
399 strongly influenced by both winter temperature and precipitation seasonality, and their
400 elevation distribution ranges shifted 72.5 m and 128.9 m upslope, respectively. In contrast, we
401 found that the adaptation of species to higher summer precipitation was negatively related to
402 the elevation shifts, suggesting that species having strong adaptability to the summer
403 precipitation tends to shifts downhill along the elevation gradients. For instance, the global
distribution range of *Paederia foetida* and *Pericampylus glaucus* in subtropical mountains were
404 dominated by summer precipitation, and their elevation range shifted downhill by 177.4 m and
405 94.9 m, respectively. Plant species in these mountains appear to track climatic water balance
406 by shifting their elevational distributions downhill. This is consistent with the previous finding
407 that change in precipitation leads to downward shifts of plant species in California (Crimmins
408 et al., 2011).
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Our results indicated that some plant species showed no or little elevational shifts.

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Previous studies suggest that the local persistence of plant populations may be the result of

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microenvironment on species persistence. Micro-environmental conditions including

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topography, soil, hydrological characteristics, and microclimate could create a multitude of

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microhabitats offering refugial persistence to some plant species (Rabasa et al., 2013; Daniel &

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Körner, 2011). The non-existent or slow migration of some plant species might be also the

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reason that these species having limited ability to expand into newly climatically suitable

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habitat due to dispersal and/or recruitment limitation (Feeley, 2012). If this condition continues

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into the future, these species with little or on elevational shifts will face risks of extinctions due

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to reduced habit areas and population sizes under future climate changes.

421

## 4.2 Effect of mountain properties on elevational shifts of species

422

Our results indicate that in addition to the effects of climate change and species' climate

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adaptation on species elevational redistributions, the size of the mountains (i.e. measured as the

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difference between the highest and lowest elevations of a mountain) plays an important role in

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determining species' elevational distribution shifts. The mean magnitude of upward shifts and

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the proportions of species with upward shifts were both higher in larger mountains (Figure 2).

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For instance, the mountains in the Qinghai-Tibet Plateau have higher elevational range (i.e.

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larger mountains) compared with the mountains in other regions, and the upward movement of

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the species are larger. Studies indicated that the plant species' optimum elevation movement

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along the elevational gradient since 1970s was significantly influenced by the mountain

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topography and summit heights in European Alps (Holmes et al., 2016). Based on studies of

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516 forest lines in the Northern Hemisphere, a recent meta-analysis indicates that mountains

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size explains 48.9% of variations in the distribution of forest lines, which also supports our

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finding that mountains size influences the redistribution of woody plants in response to climate

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changes (Yao & Zhang, 2014; Zhao et al., 2014). Our results together with previous findings

436

suggest that higher mountains may provide more suitable habitats for species to move upward.

437

Moreover, our results showed that species in temperate mountains tended to shift upward,

438

while species in warm and wet mountains (i.e. those in subtropical regions) tended to shift

439

downward. In line with our results, a recent study also demonstrated downward shifts of
centroid elevations of plant species living in subtropical mountains (O'Sullivan & Jump, 2020).

In subtropical regions, the low seasonal temperature variation reduces overlap in thermal regimes between lowlands and high altitudes, and species tend to have narrow adaptations to temperatures (Shrestha et al., 2018). Lowland species may be restricted to lower elevations because of their limited tolerance to cold stress (Heatwole et al., 1969), while high-elevation species are generally not limited by warm temperature and are therefore able to move to lowland regions (Ghalambor et al., 2006). This explained the results that the high-elevation species in subtropical mountains (e.g. EM, SNJ, BL, HG, JF, LS) tended to shifts downward. Besides, the interaction among species (e.g. species competition), topography and anthropogenic influences may also contribute to the downward elevation shifts of plant species in subtropical mountains (Bhatta, 2018). Recent studies indicate that in subtropical mountains, high-elevation species cannot survive the lowlands because they are often outcompeted by the more competitive lowland species, not due to physiological limitations (Mamantov et al., 2021; Zu et al., 2021).

If climate warming reduces the competitive fitness of lowland species, then higher elevation species tend to shift downslope more easily (Rabasa et al., 2013). How species' elevational redistributions may influence species interactions in subtropical regions requires urgent attention.

## 4.3 Effect of functional traits on elevational shifts of species

Our finding demonstrates that the elevational shifts of species in response to climate change may vary among species with different functional traits (including life forms and fruit types).

First, several previous studies have demonstrated that the distributions of species with different lifeforms may respond to climate change differently. For example, studies on plants' elevational distributions in the French Alps in Europe (Lenoir et al., 2008) and the Mt. Gongga in China (Zu et al., 2021) and on the forest communities in Switzerland (Küchler, 2014) all showed that herbaceous species had faster upslope shifts than trees and shrubs. In contrast, a recent review on species elevational shifts in the European Alps showed that the optimum elevations of long-lived woody plants were shifting upward at the same rate with non-alpine herbaceous plants (Vitasse et al., 2021). These findings suggest that the conclusions on the responses of woody and herbaceous plants remain controversial. In this study, our result showed that the direction
and magnitude of elevational shifts in China' mountains differed between life forms. We found that herbaceous plants shifted their elevational centroids significantly faster than woody plants in 15 mountains. The reason for these differences between the elevational shifts of woody and herbaceous plant species in response to climate change may be partly due to the fact that different lifeforms have different climate adaptation and physiological properties (Grytnes et al., 2014). Generally, the woody plants are long-lived species and have deeper roots, and they seem to be less sensitive to the soil water (Wang et al., 2020). On the contrary, herbaceous species have shorter life cycles and shallower roots than woody species, and they tend to be more sensitive to precipitation changes (Lande, 1993).

The rate and distance of elevational shifts of species may also be influenced by other functional traits in addition to lifeform (Alexander et al., 2018; Matteodo et al., 2016). In this study, we found that species with different fruit types shifted their centroid elevations in different trends and with different magnitudes. For instance, the species with samara fruits (e.g., *Acer oliverianum* and *Betula albosinensis*) shifted their centroid elevations upward, while the species with caryopsis fruits (e.g., *Eulalia speciose* and *Bromus remotiflorus*) shifted their centroid elevations downhill. This is because samara fruits are easily dispersed by wind, while caryopsis fruits tend to drop to the ground directly or beneath the parent plants. Moreover, species elevational shifts also significantly different between different genera and families. This is likely because different genera and families normally have different functional traits, climatic adaptation and evolutionary history (Li et al., 2017; Losos, 2011). These results suggest that species distributions of different taxonomic groups may respond differently to environmental changes, and hence may have different risks of local extinction under future environmental change scenarios. In this study, there are only 20 gymnosperm species for which we had sufficient data while we had good time series for 715 angiosperm species in total. Indeed, gymnosperm and angiosperm species have very different traits and climate adaptation, hence we also assumed that the magnitude of elevation range shifts will be different between both taxonomic groups. Yet, we found no significant difference in the magnitude of mean elevational range shifts between gymnosperm and angiosperm species (t = 1.1132, df = 19.754, p-value = 0.279), most likely due to the extremely low sample size for gymnosperms relative to
498 angiosperms.

499 Finally, we suggest to consider both regional and species characteristics in future researches related to elevational range shifts. We recommend to focus on the taxon that is more sensitive to climate change and in fragile biodiversity regions. For species that tend to shift upward, those living at high-elevational regions will face more climatic and competitive pressures than lowland species, and thus should be given much more attentions under future climate warming. On the contrary, for species that tend to shift downhill, the lowland species and their natural habitats are more vulnerable to climate change and intensive human pressure (e.g. tourism and recreation). Compared with traditional suggestions to protect or restore species habitats, we recommend to protect dispersal corridors of species to improve their dispersal processes along the elevational or latitudinal gradients. Our study provides essential scientific references for managing protected areas in China's mountain regions.

510 COMPETING INTEREST STATEMENT

511 The authors declare no competing interests.

512 DATA AVAILABILITY STATEMENT

513 The data used in the analyses of the paper are uploaded to the Dryad website (https://doi.org/10.5061/dryad.r7sqv9sj0). The data of study regions and the total number of herbarium specimens are provided in the Supporting Information (Table S1); The climate data that support this study are available from the Geographic Data Sharing Infrastructure, College of Urban and Environmental Science, Peking University (http://geodata.pku.edu.cn), and WorldClim (https://www.worldclim.org/data/worldclim21.html). The distribution data of plant species were from the National Specimen Information Infrastructure (NSII, http://www.nsii.org.cn), the Chinese Virtual Herbarium (CVH, http://www.cvh.ac.cn), Global Biodiversity Information Facility (GBIF, http://www.gbif.org/), and the Botanical Information and Ecology Network (BIEN, http://bien.nceas.ucsb.edu/bien/biendata/). The human footprint data is from the Last of the Wild (https://sedac.ciesin.columbia.edu).

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TABLES AND FIGURES

<table><caption>TABLE 1. Summary information on the 29 studied mountain sites and the average range shifts of the centroid elevational position across all species occurring in a given mountain site.</caption><thead><tr><th>Mountain range</th><th>Mount</th><th>Abbr.</th><th>Lat (°N)</th><th>Long (°E)</th><th>Total_num</th><th>N</th><th>Up-ratio</th><th>Shift(m)</th></tr></thead><tbody><tr><td>Qionglai(邛崃山)</td><td>Balang(巴郎山)</td><td>BL</td><td>31.167</td><td>102.917</td><td>13502</td><td>114</td><td>0.42</td><td>-33.12</td></tr><tr><td>Bayankela(巴颜喀拉山脉)</td><td>Bayankela(巴颜喀拉山)</td><td>BYKL</td><td>34.204</td><td>97.659</td><td>11936</td><td>18</td><td>0.61</td><td>50.21</td></tr><tr><td>Hengduan(横断山脉)</td><td>Baimang(白茫茫雪山)</td><td>BMX</td><td>28.183</td><td>99.117</td><td>13705</td><td>21</td><td>0.67</td><td>53.94</td></tr><tr><td>Changbai(长白山脉)</td><td>Baitou(白头山)</td><td>BT</td><td>41.983</td><td>128.083</td><td>13498</td><td>29</td><td>0.59</td><td>17.56</td></tr><tr><td>Yanshan(燕山山脉)</td><td>Dongling(东灵山)</td><td>DL</td><td>39.733</td><td>115.5</td><td>14267</td><td>13</td><td>0.38</td><td>-86.32</td></tr><tr><td>Qionglai(邛崃山脉)</td><td>Emei(峨眉山)</td><td>EM</td><td>29.6</td><td>103.5</td><td>5451</td><td>25</td><td>0.8</td><td>440.46</td></tr><tr><td>Wuling(武陵山脉)</td><td>Fanjing(梵净山)</td><td>FJ</td><td>27.917</td><td>108.767</td><td>7146</td><td>22</td><td>0.41</td><td>-415.29</td></tr><tr><td>Wuling(武陵山脉)</td><td>Fenghuang(凤凰山)</td><td>FH</td><td>28.11</td><td>108.68</td><td>8188</td><td>5</td><td>0.6</td><td>-37.16</td></tr><tr><td>Hengduan(横断山脉)</td><td>Gaoligong(高黎贡山)</td><td>GLG</td><td>25.267</td><td>98.783</td><td>11047</td><td>35</td><td>0.2</td><td>-173.28</td></tr><tr><td>Hengduan(横断山脉)</td><td>Gongga(贡嘎山)</td><td>GG</td><td>29.6</td><td>101.883</td><td>27669</td><td>83</td><td>0.64</td><td>135.74</td></tr><tr><td>Lvliang(吕梁山脉)</td><td>Guandi(关帝山孝文山)</td><td>GD</td><td>37.8</td><td>110.8</td><td>7123</td><td>13</td><td>0.69</td><td>15.73</td></tr><tr><td>Hengduan(横断山脉)</td><td>Haba(哈巴雪山)</td><td>HB</td><td>27.45</td><td>100.117</td><td>5503</td><td>7</td><td>0.57</td><td>92.84</td></tr><tr><td>Daba(大巴山脉)</td><td>Hualong(化龙山)</td><td>HL</td><td>32.054</td><td>109.419</td><td>5387</td><td>5</td><td>0.8</td><td>116.47</td></tr><tr><td>Wuyi(武夷山脉)</td><td>Huanggang(黄岗山)</td><td>HG</td><td>27.8</td><td>117.6</td><td>11821</td><td>54</td><td>0.48</td><td>15.46</td></tr><tr><td>Dalou(大娄山脉)</td><td>Jinfo(金佛山)</td><td>JF</td><td>29.25</td><td>108.333</td><td>40841</td><td>231</td><td>0.4</td><td>-57.85</td></tr><tr><td>Hengduan(横断山脉)</td><td>Kagebo(卡格薄峰)</td><td>KGB</td><td>28.449</td><td>98.575</td><td>5372</td><td>38</td><td>0.66</td><td>97.76</td></tr><tr><td>Hengduan(横断山脉)</td><td>Laojun(老君山)</td><td>LJ</td><td>26.703</td><td>99.75</td><td>6254</td><td>16</td><td>0.19</td><td>-179.05</td></tr><tr><td>Miaoling(苗岭)</td><td>Leigong(雷公山)</td><td>LG</td><td>26.417</td><td>108.25</td><td>6276</td><td>5</td><td>0.8</td><td>-26.2</td></tr><tr><td>Lushan(庐山)</td><td>Lushan(庐山)</td><td>LS</td><td>29.55</td><td>115.967</td><td>19261</td><td>31</td><td>0.16</td><td>-218.86</td></tr><tr><td>Nanling(南岭)</td><td>Maoer(猫儿山)</td><td>ME</td><td>25.918</td><td>110.42</td><td>7105</td><td>22</td><td>0.36</td><td>-144.65</td></tr><tr><td>Himalayas(喜马拉雅山脉)</td><td>Nanjiabawa(南迦巴瓦峰)</td><td>NJBW</td><td>29.633</td><td>95.067</td><td>5037</td><td>6</td><td>0.67</td><td>-14.38</td></tr><tr><td>Hengduan(横断山脉)</td><td>Queer(雀儿山)</td><td>QE</td><td>31.77</td><td>99.05</td><td>6251</td><td>13</td><td>0.69</td><td>346.99</td></tr><tr><td>Daba(大巴山脉)</td><td>Shennongjia(神农架)</td><td>SNJ</td><td>31.667</td><td>110.583</td><td>59131</td><td>61</td><td>0.59</td><td>134.8</td></tr><tr><td>Qionglai(邛崃山脉)</td><td>Siguniang(四姑娘山)</td><td>SGN</td><td>31</td><td>103.217</td><td>6412</td><td>10</td><td>0.9</td><td>155.53</td></tr><tr><td>Qinling(秦岭)</td><td>Taibai(太白山)</td><td>TB</td><td>33.95</td><td>107.75</td><td>8295</td><td>46</td><td>0.72</td><td>196.38</td></tr><tr><td>Tianshan(天山山脉)</td><td>Tiangeer(天格尔峰)</td><td>TGE</td><td>43.168</td><td>86.623</td><td>8717</td><td>18</td><td>0.5</td><td>18.87</td></tr><tr><td>Hengduan(横断山脉)</td><td>Wuliang(无量山)</td><td>WL</td><td>24.45</td><td>100.733</td><td>6850</td><td>7</td><td>0.71</td><td>153.52</td></tr><tr><td>Taihan(太行山脉)</td><td>Xiaowutai(小五台山)</td><td>XWT</td><td>39.967</td><td>114.833</td><td>5360</td><td>12</td><td>0.17</td><td>-60.97</td></tr><tr><td>Aertai(阿尔泰山脉)</td><td>Youyi(友谊峰)</td><td>YYF</td><td>49.1</td><td>87.817</td><td>5706</td><td>6</td><td>0.33</td><td>1.59</td></tr></tbody></table>

Mount: the name of each of the 29 studied mountain site; Total_num: the total number of herbarium specimens for each of the 29 mountain sites. N: the total number of selected seed plant species in each of the 29 mountain sites; Up-ratio: the percentage of species shifting upward for each mountain site; Shift: the mean range shift of the centroid elevational position across all species of a given mountain site and between the two study periods (1950-1988 vs. 1980-2018).
<table><caption>TABLE 2 The multiple regression model evaluating the relationships between species elevational shifts and climate change experienced by species, the climatic adaptation of species, species properties and mountain properties</caption><thead><tr><th>Type</th><th>Variation</th><th>Df</th><th>Sum Sq</th><th>Mean Sq</th><th>F</th><th>p</th></tr></thead><tbody><tr><td rowspan="4">Climate change</td><td>Change in tem</td><td>1</td><td>37051</td><td>37051</td><td>0.43</td><td>0.51</td></tr><tr><td>Change in pre</td><td>1</td><td>345512</td><td>345512</td><td>3.99</td><td>0.04*</td></tr><tr><td>Importance of TS</td><td>1</td><td>3304</td><td>3304</td><td>0.04</td><td>0.85</td></tr><tr><td>Importance of MTWQ</td><td>1</td><td>46407</td><td>46407</td><td>0.54</td><td>0.46</td></tr><tr><td rowspan="6">Climatic adaption</td><td>Importance of MTCQ</td><td>1</td><td>414136</td><td>414136</td><td>4.78</td><td>0.03*</td></tr><tr><td>Importance of PS</td><td>1</td><td>578382</td><td>578382</td><td>6.68</td><td>0.01*</td></tr><tr><td>Importance of PWQ</td><td>1</td><td>357892</td><td>357892</td><td>4.13</td><td>0.04*</td></tr><tr><td>Importance of PCQ</td><td>1</td><td>75940</td><td>75940</td><td>0.88</td><td>0.35</td></tr><tr><td>Elevation</td><td>1</td><td>75603</td><td>75603</td><td>0.87</td><td>0.35</td></tr><tr><td>Range</td><td>1</td><td>294868</td><td>294868</td><td>3.40</td><td>0.07.</td></tr><tr><td rowspan="10">Species properties</td><td>Life form</td><td>2</td><td>94746</td><td>47373</td><td>0.55</td><td>0.58</td></tr><tr><td>SexSyst</td><td>2</td><td>348638</td><td>174319</td><td>2.01</td><td>0.13</td></tr><tr><td>Fruit type</td><td>13</td><td>1903992</td><td>146461</td><td>1.69</td><td>0.06.</td></tr><tr><td>Family</td><td>109</td><td>15435053</td><td>141606</td><td>1.64</td><td>&lt;0.001***</td></tr><tr><td>Genus</td><td>224</td><td>31068140</td><td>138697</td><td>1.60</td><td>&lt;0.001***</td></tr><tr><td>Re-elevation</td><td>1</td><td>3235916</td><td>3235916</td><td>37.37</td><td>&lt;0.001***</td></tr><tr><td>Latitude</td><td>1</td><td>5789</td><td>5789</td><td>0.07</td><td>0.80</td></tr><tr><td>Longitude</td><td>1</td><td>498763</td><td>498763</td><td>5.76</td><td>0.02*</td></tr><tr><td>Climate regions</td><td>2</td><td>533050</td><td>266525</td><td>3.08</td><td>0.04*</td></tr><tr><td>Mountain properties</td><td>Mountain</td><td>23</td><td>8981946</td><td>390519</td><td>4.51</td><td>&lt;0.001***</td></tr><tr><td></td><td>Residuals</td><td>452</td><td>39143600</td><td>86601</td><td></td><td></td></tr></tbody></table>

Change in tem: the temperature changes experienced by each species; Change in pre: the precipitation changes experienced by each species; TS: temperature seasonality; MTWQ: mean temperature of warmest quarter; MTCQ: mean temperature of coldest quarter; PS: Precipitation seasonality; PWQ: precipitation of warmest quarter; PCQ: precipitation of coldest quarter. .p<0.1, *, p < 0.05, **, p < 0.01, ***, p < 0.001.
<figure><img src="image_3.png" /><figcaption>FIGURE 1 (a) The locations of the 29 study mountains and the ratio of species experiencing upward and downhill elevational shifts in these mountains. (b, c) The changes in the average magnitude of species' elevational shifts as functions of the average temperature (b) and size (c) of the mountains. In (a), blue bars indicated the ratio of species with upward shifts, and red bars indicated the ratios of species with downhill shifts. In (b) and (c), blue circles indicated the average magnitude of upward shifts, red triangles indicated the magnitude of downhill shifts, and white square indicated non-significant elevational shifts. The abbreviations of mountain names are shown in Table 1</figcaption></figure>

<figure><img src="image_4.png" /></figure>

<figure><img src="image_5.png" /></figure>
<figure><img src="image_6.png" /><figcaption>FIGURE 2 Relationships between the changes in optimum elevations and the climatic adaptation of species. The climatic adaptation was represented by the importance of climate factors in determining the global distribution ranges of the studied species (see Methods for details). MTCQ: mean temperature of coldest quarter; PS: Precipitation seasonality; PWQ: precipitation of warmest quarter</figcaption></figure>
<figure><img src="image_7.png" /><figcaption>FIGURE 3 The average magnitude of elevational shifts (a) and relative species percentage of upward shifts (b) for herbaceous and woody plant species shown along the gradient of mean temperature of mountains. The abbreviations of mountain names are shown in Table 1</figcaption></figure>
<figure><img src="image_8.png" /><figcaption>FIGURE 4 The relationship between the centroid elevation shifts and elevational distributions of species in different mountains. The x-axes represent the historical centroid elevation of species, and the y-axes represent the changes in the centroid elevation between the two studied periods. The 95% confidence intervals of the regression lines are shown in grey. The abbreviations of mountain names are shown in Table 1</figcaption></figure>
## Supplemental Figures and Tables

**Figure S1:** The species numbers studied in each mountains.

**Figure S2:** Elevational distribution of the selected species occurrence records through time for each mountain.

**Figure S3:** The standard meteorological stations were used in this study.

**Figure S4:** Change in temperature between historical (1950 - 1988) and present time (1989 - 2015) periods at long-term meteorological stations in China's mountains.

**Figure S5:** Change in precipitations between historical (1950 - 1988) and present time (1989 - 2015) periods at long-term meteorological stations in China's mountains.

**Figure S6:** Change in AET between historical (1950 - 1988) and present time (1989 - 2015) periods at long-term meteorological stations in China's mountains.

**Figure S7:** Change in summer/winter temperatures and precipitations between historical (1950 - 1988) and present time (1989 - 2015) periods at long-term meteorological stations in China's mountains.

**Figure S8:** Relationships between changes in optimum elevations and the importance of climate factors.

**Figure S9:** Relationships of the changes in optimum elevations with the functional traits of species, and the elevational shifts of herbs and woody plants in different climate regions (c). The functional traits included fruit types (a), and climate regions (b).

**Table S1:** The mountain sites located in China.

**Tables S2:** The lowest elevational data from literature.

**Table S3:** The random shifts in the species' centroid elevation calculated after the
resampling of the historical and current survey records for each mountain.

**Table S4:** The relationship between elevational shifts and elevation. The red indicates the significant relationship.
Supplementary Materials for

Elevational shift in seed plant distributions in China's mountains over
the last 70 years

For Peer Review
## Supplemental Figures and Tables

**Figure S1:** The species numbers studied in each mountains.

**Figure S2:** Elevational distribution of the selected species occurrence records through time for each mountain.

**Figure S3:** The standard meteorological stations were used in this study.

**Figure S4:** Change in temperature between historical (1950 - 1988) and present time (1989 - 2015) periods at long-term meteorological stations in China's mountains.

**Figure S5:** Change in precipitations between historical (1950 - 1988) and present time (1989 - 2015) periods at long-term meteorological stations in China's mountains.

**Figure S6:** Change in AET between historical (1950 - 1988) and present time (1989 - 2015) periods at long-term meteorological stations in China's mountains.

**Figure S7:** Change in summer/winter temperatures and precipitations between historical (1950 - 1988) and present time (1989 - 2015) periods at long-term meteorological stations in China's mountains.

**Figure S8:** Relationships between changes in optimum elevations and the importance of climate factors.

**Figure S9:** Relationships of the changes in optimum elevations with the functional traits of species, and the elevational shifts of herbs and woody plants in different climate regions (c). The functional traits included fruit types (a), and climate regions (b).

**Table S1:** The mountain sites located in China.

**Tables S2:** The lowest elevational data from literature.

**Table S3:** The random shifts in the species' centroid elevation calculated after the
resampling of the historical and current survey records for each mountain.

**Table S4:** The relationship between elevational shifts and elevation. The red indicates the significant relationship.
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<figure><img src="image_9.png" /><figcaption>Figure S1: The species numbers studied in each mountains.</figcaption></figure>
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<figure><img src="image_24.png" /></figure>
<figure><img src="image_25.png" /><figcaption>Figure S2: Elevational distribution of the selected species occurrence records through time for each mountain.</figcaption></figure>

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<figure><img src="image_26.png" /><figcaption>Figure S3: The standard meteorological stations were used in this study.</figcaption></figure>
<figure><img src="image_27.png" /></figure>

<figure><img src="image_28.png" /><figcaption>Figure S4: Change in temperature between historical (1950 - 1988) and present time (1989 - 2015) periods at long-term meteorological stations in</figcaption></figure>
<figure><img src="image_29.png" /><figcaption>China's mountains.</figcaption></figure>

<figure><img src="image_30.png" /></figure>
Figure S5: Change in precipitations between historical (1950 - 1988) and present time (1989 - 2015) periods at long-term meteorological stations in China's mountains.
<figure><img src="image_31.png" /></figure>

<figure><img src="image_32.png" /><figcaption>Figure S6: Change in AET between historical (1950 - 1988) and present time (1989 - 2015) periods at long-term meteorological stations in China's mountains.</figcaption></figure>
A. Temperature rises gradually

<figure><img src="image_33.png" /></figure>

B. Temperature rises gradually

<figure><img src="image_34.png" /></figure>

C. Precipitations rises gradually

<figure><img src="image_35.png" /></figure>

D. Precipitations rises gradually

<figure><img src="image_36.png" /><figcaption>Figure S7: Change in summer/winter temperatures and precipitations between historical (1950 - 1988) and present time (1989 - 2015) periods at</figcaption></figure>
long-term meteorological stations in China' mountains.
<figure><img src="image_37.png" /></figure>

Figure S8: Relationships between changes in optimum elevations and the importance of climate factors. The x-axes are MAT, MAP, AET, changes in MAT, MAP, AET, relative elevation(Reele), Seasonal changes in MAT(CTsummer and CTwinter) and Seasonal changes in MAP(CPsummer and CPwinter), atmospheric nitrogen deposition(ND), and human footprint index (HI). The Y-axes represent changes in optimum elevation for all
species, the shifts of herb plants and the shifts of woody.
<figure><img src="image_38.png" /></figure>

Figure S9: Relationships of the changes in optimum elevations with the functional traits of species, and the elevational shifts of herbs and woody plants in different climate regions (c). The functional traits included fruit types (a), and climate regions (b).
<table><caption>Table S1: The mountain sites located in China.</caption><thead><tr><th>Mountain range</th><th>Mount</th><th>Abbr.</th><th>Lat (°N)</th><th>Long (°E)</th><th>The highest elevation</th><th>Reelevation(max elevation-min elevation)</th><th>Province</th><th>climate region</th></tr></thead><tbody><tr><td>Qionglai(邛崃山)</td><td>Balang(巴郎山)</td><td>BL</td><td>31.167</td><td>102.917</td><td>5072</td><td>3072</td><td>Sichuan</td><td>Plateau</td></tr><tr><td>Bayankela(巴颜喀拉山脉)</td><td>Bayankela(巴颜喀拉山)</td><td>BYKL</td><td>34.204</td><td>97.659</td><td>5267</td><td>1667</td><td>Qinghai</td><td>Plateau</td></tr><tr><td>Hengduan(横断山脉)</td><td>Baimang(白茫雪山)</td><td>BMX</td><td>28.183</td><td>99.117</td><td>5430</td><td>3500</td><td>Yunnan</td><td>Plateau</td></tr><tr><td>Changbai(长白山脉)</td><td>Baitou(白头山)</td><td>BT</td><td>41.983</td><td>128.083</td><td>2691</td><td>1591</td><td>Jilin</td><td>Temperate</td></tr><tr><td>Yanshan(燕山山脉)</td><td>Dongling(东灵山)</td><td>DL</td><td>39.733</td><td>115.5</td><td>2303</td><td>1803</td><td>Beijing</td><td>Temperate</td></tr><tr><td>Qionglai(邛崃山脉)</td><td>Emei(峨眉山)</td><td>EM</td><td>29.6</td><td>103.5</td><td>3099</td><td>2700</td><td>Sichuan</td><td>Subtropics</td></tr><tr><td>Wuling(武陵山脉)</td><td>Fanjing(梵净山)</td><td>FJ</td><td>27.917</td><td>108.767</td><td>2493</td><td>2000</td><td>Guizhou</td><td>Subtropics</td></tr><tr><td>Wuling(武陵山脉)</td><td>Fenghuang(凤凰山)</td><td>FH</td><td>28.11</td><td>108.68</td><td>2572</td><td>1572</td><td>Guizhou</td><td>Subtropics</td></tr></tbody></table>
<table><tr><td>Hengduan(横断山脉)</td><td>Gaoligong(高黎贡山)</td><td>GLG</td><td>25.267</td><td>98.783</td><td>3374</td><td>3000</td><td>Yunnan</td><td>Plateau</td></tr><tr><td>Hengduan(横断山脉)</td><td>Gongga(贡嘎山)</td><td>GG</td><td>29.6</td><td>101.883</td><td>7556</td><td>6466</td><td>Sichuan</td><td>Plateau</td></tr><tr><td>Lvliang(吕梁山脉)</td><td>Guandi(关帝山孝文山)</td><td>GD</td><td>37.8</td><td>110.8</td><td>2831</td><td>2051</td><td>Shanxi</td><td>Temperate</td></tr><tr><td>Hengduan(横断山脉)</td><td>Haba(哈巴雪山)</td><td>HB</td><td>27.45</td><td>100.117</td><td>5396</td><td>3846</td><td>Yunnan</td><td>Plateau</td></tr><tr><td>Daba(大巴山脉)</td><td>Hualong(化龙山)</td><td>HL</td><td>32.054</td><td>109.419</td><td>2917</td><td>2537</td><td>Shanxi</td><td>Subtropics</td></tr><tr><td>Wuyi(武夷山脉)</td><td>Huanggang(黄岗山)</td><td>HG</td><td>27.8</td><td>117.6</td><td>2158</td><td>1808</td><td>Fujian</td><td>Subtropics</td></tr><tr><td>Dalou(大娄山脉)</td><td>Jinfo(金佛山)</td><td>JF</td><td>29.25</td><td>108.333</td><td>2251</td><td>1911</td><td>Chongqing</td><td>Subtropics</td></tr><tr><td>Hengduan(横断山脉)</td><td>Kagebo(卡格薄峰)</td><td>KGB</td><td>28.449</td><td>98.575</td><td>6740</td><td>4700</td><td>Yunnan</td><td>Plateau</td></tr><tr><td>Hengduan(横断山脉)</td><td>Laojun(老君山)</td><td>LJ</td><td>26.703</td><td>99.75</td><td>4247</td><td>2697</td><td>Yunnan</td><td>Plateau</td></tr><tr><td>Miaoling(苗岭)</td><td>Leigong(雷公山)</td><td>LG</td><td>26.417</td><td>108.25</td><td>2179</td><td>1500</td><td>Guizhou</td><td>Subtropics</td></tr><tr><td>Lushan(庐山)</td><td>Lushan(庐山)</td><td>LS</td><td>29.55</td><td>115.967</td><td>1474</td><td>1140</td><td>Jiangxi</td><td>Subtropics</td></tr><tr><td>Nanling(南岭)</td><td>Maoer(猫儿山)</td><td>ME</td><td>25.918</td><td>110.42</td><td>2141.5</td><td>1862</td><td>Guangxi</td><td>Subtropics</td></tr></table>
<table><tr><td>Himalayas(喜马拉雅山脉)</td><td>Nanjiabawa(南迦巴瓦峰)</td><td>NJBW</td><td>29.633</td><td>95.067</td><td>7782</td><td>5682</td><td>Xizang</td><td>Plateau</td></tr><tr><td>Hengduan(横断山脉)</td><td>Queer(雀儿山)</td><td>QE</td><td>31.77</td><td>99.05</td><td>6168</td><td>4000</td><td>Sichuan</td><td>Plateau</td></tr><tr><td>Daba(大巴山脉)</td><td>Shennongjia(神农架)</td><td>SNJ</td><td>31.667</td><td>110.583</td><td>3106</td><td>2706</td><td>Hubei</td><td>Subtropics</td></tr><tr><td>Qionglai(邛崃山脉)</td><td>Siguniang(四姑娘山)</td><td>SGN</td><td>31</td><td>103.217</td><td>6250</td><td>4050</td><td>Sichuan</td><td>Plateau</td></tr><tr><td>Qinling(秦岭)</td><td>Taibai(太白山)</td><td>TB</td><td>33.95</td><td>107.75</td><td>3767</td><td>3000</td><td>Shanxi</td><td>Temperate</td></tr><tr><td>Tianshan(天山山脉)</td><td>Tiangeer(天格尔峰)</td><td>TGE</td><td>43.168</td><td>86.623</td><td>4562</td><td>3000</td><td>Xinjiang</td><td>Temperate</td></tr><tr><td>Hengduan(横断山脉)</td><td>Wuliang(无量山)</td><td>WL</td><td>24.45</td><td>100.733</td><td>3254</td><td>2200</td><td>Yunnan</td><td>Plateau</td></tr><tr><td>Taihan(太行山脉)</td><td>Xiaowutai(小五台山)</td><td>XWT</td><td>39.967</td><td>114.833</td><td>2870</td><td>2070</td><td>Hebei</td><td>Temperate</td></tr><tr><td>Aertai(阿尔泰山脉)</td><td>Youyi(友谊峰)</td><td>YYF</td><td>49.1</td><td>87.817</td><td>4374</td><td>2000</td><td>Xinjiang</td><td>Temperate</td></tr></table>

“Reelevation (max elevation-min elevation)” means the relative elevation, and it is the value of the highest elevation minus the lowest elevation.
<table><caption>Tables S2: The lowest elevational data from literature.</caption>
  <tr>
    <td>Cao W, Li Y. Vertical pattern of flora from Changbai Mountain, China .Journal of Beijing Forestry University. 2008, 30(4): 53-58.</td>
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  <tr>
    <td>Du J, Hao M, Tian Z, et al. A study on the plant diversity of Xiaowutai Mountain district. Herbei Journal of forestry and orchard research. 2016, 31(01): 88-93.</td>
  </tr>
  <tr>
    <td>Fu Z. Floristic and geographical components of seed plants in Hualong Mountain, southern Shaanxi Province. Geographical research. 1997, (02): 73-81.</td>
  </tr>
  <tr>
    <td>Gao LL, Yang HJ, Liu GZ, Wang CH, Yang GP, Cai CT. Diversity of medicinal vascular plant resources in Wuliangshan National Nature Reserve , Yunnan , China. Plant Science Journal, 2018, 36(2): 203-210.</td>
  </tr>
  <tr>
    <td>Gu HY, Li CH. Biodiversity and flora of the mixed evergreen and deciduous broadleaved forest in Emei. Bulletin of Botanical Research. 2006, (05): 618-623.</td>
  </tr>
  <tr>
    <td>Gu HY, Li CH. Preliminarily Study on the Pteridophytes Flora of Emei Mountain. Acta Bot. Boreal. Occident. Sin. 2008, (02): 2381-2387.</td>
  </tr>
  <tr>
    <td>Hu XX, Zheng WC, Li JQ, et al. Tianpei Guan1* Preliminary survey on mammal and bird diversity at Siguniang Mountains National Nature Reserve, Sichuan, China. Biodiversity Science. 2018, 26 (12): 1325-1331.</td>
  </tr>
  <tr>
    <td>Li HW, Zhao YF. Plant diversity of Baimaxueshan National Nature Reserve. Guihaia. 2007, (01): 71-76.</td>
  </tr>
  <tr>
    <td>Li J, Teng CD. Vertical distribution of vegetation in Guandi Mountain. Journal of Shanxi University (Natural Science Edition). 1979, (01): 144-149.</td>
  </tr>
  <tr>
    <td>Liu JL, Chen K, Wang JY, Chen WH. Comparative Study on Seed Plant Flora between Southern and Northern Gaoligong Mountains. Journal of Tropical and Subtropical Botany. 2016, 24(3): 327-332.</td>
  </tr>
  <tr>
    <td>Liu JY, Liu JJ, Han QS. Pteridophyte flora in Taibaishan Nature Reserve of Shaanxi Province. Guihaia, 2014, 34(1): 77-83.</td>
  </tr>
  <tr>
    <td>Ou ZY, Pang SL, He F, et al. Variation and Correlation of Leaf Functional Traits of Common Plants at Main Peak of Mao'er Moutain, Guangxi. Guangxi Forestry Science. 2021,50(05):557-562.</td>
  </tr>
  <tr>
    <td>Peng H. The Endemism in the Flora of Seed Plants in Mt. Wuliangshan. Acta Botanica Yunnanica. 1997, (01): 3-16.</td>
  </tr>
  <tr>
    <td>Peng TL, Tie J. Wild plant resources in Shennongjia Nature Reserve and its protection measures. Journal of Changzhi University. 2010, 27(05): 23-26.</td>
  </tr>
  <tr>
    <td>Song XT, Shao XM, Sun Y, et al. Research on the bryoflora of Dongling Mountain , Beijing , China. Plant Science Journal, 2018, 36(4): 554-561.</td>
  </tr>
  <tr>
    <td>Wang J. Study on the Flora of Spermatophyte in Xinluhai Nature Reserve of the Queer Mountain. Journal of Anhui Agri. Sci. 2012, 40(21): 11020-11021.</td>
  </tr>
  <tr>
    <td>Wang LG, Xu C. Preliminary Study on Plant Resources Magnoliaceae, Wuyi Mountain National Nature Reserve, Fujian Province. Fujian Forestry Science and Technology. 2010, 37(02): 90-93.</td>
  </tr>
</table>
Wang PL, Zhong AW, Liang TJ, Chen BH. Investigation and Conservation of Germplasm Resources of Wild seed plants in Lushan Mountain. Guizhou Agricultural Sciences. 2016, 44(04): 12-16.

Wu RX, Xiong KN, Li R. Characteristics of spermatophyte flora of Fanjing Mountain and its phytogeographical significance. Guihaia, 2017, 37(10): 1348-1354.

Wu XJ, Yu CY. Vegetation Distribution Characteristics and Diversity Conservation in Habaxueshan Provincial Nature Reserve. Forest Inventory and Planning, 2020, 45(1): 47-51.

Wu YH. The Floristic Characteristics in the Region of Bayan Har Mountains. Acta Botanica Yunnanica. 2004, (06): 587-603.

Yang HY, Luo XQ, Yang F. Algae flora of Leigong Mountain Nature Reserve of Guizhou. Guihaia. 2014, 34(04): 570-574.

Yi SR, Huang Y. Preliminary Study on Seed Flora in Jinfo Mountain Nature Reserve. Acta Botanica Sinica Northwest China, 2004, (01): 83-93.

Yu WWG, Zhu LT, Wang SE, et al. Study on the flora of Changbai Mountain. Journal of Shandong Normal University (Natural Science Edition), 2006, (02): 115-117.

Zhang HB. Investigation on the Construction Status of Laojun Mountain National Park in Lijiang. Yunnan Social Sciences, 2013, (03): 96-99.

Zhao Q, Zhang Y, Yuan L, et al. Preliminary investigation on medicinal fungi resources in Laojunshan, Yunnan Province. Chinese Journal of Microbiology, 2006, (04): 85-88.

Zu KL, Luo A, Shrestha N, et al. Altitudinal biodiversity patterns of seed plants along Gongga Mountain in the southeastern Qinghai-Tibetan Plateau. Ecology and Evolution, 2019, 9:9586-9596.
<table><caption>Table S3: The random shifts in the species' centroid elevation calculated after the resampling of the historical and current survey records for each mountain.</caption><thead><tr><th rowspan="2">Mountain range</th><th rowspan="2">Mount</th><th rowspan="2">Abbr.</th><th colspan="3">downward species</th><th colspan="3">upward species</th></tr><tr><th>t</th><th>df</th><th>p</th><th>t</th><th>df</th><th>p</th></tr></thead><tbody><tr><td>Qionglai(邛崃山)</td><td>Balang(巴郎山)</td><td>BL</td><td>5.7924</td><td>63</td><td>2.39E-07</td><td>-12.142</td><td>49</td><td>&lt; 2.2e-16</td></tr><tr><td>Bayankela(巴颜喀拉山脉)</td><td>Bayankela(巴颜喀拉山)</td><td>BYKL</td><td>2.5854</td><td>5</td><td>0.04911</td><td>-5.4648</td><td>11</td><td>0.0001964</td></tr><tr><td>Hengduan(横断山脉)</td><td>Baimang(白茫雪山)</td><td>BMX</td><td>5.26</td><td>6</td><td>0.001901</td><td>-1.8095</td><td>13</td><td>0.09354</td></tr><tr><td>Changbai(长白山脉)</td><td>Baitou(白头山)</td><td>BT</td><td>1.7506</td><td>12</td><td>0.1055</td><td>-3.4488</td><td>4</td><td>0.02608</td></tr><tr><td>Yanshan(燕山山脉)</td><td>Dongling(东灵山)</td><td>DL</td><td>1.7625</td><td>6</td><td>0.1285</td><td>-5.1134</td><td>5</td><td>0.003728</td></tr><tr><td>Qionglai(邛崃山脉)</td><td>Emei(峨眉山)</td><td>EM</td><td>4.8982</td><td>4</td><td>0.008055</td><td>-2.6158</td><td>19</td><td>0.017</td></tr><tr><td>Wuling(武陵山脉)</td><td>Fanjing(梵净山)</td><td>FJ</td><td>6.2574</td><td>12</td><td>0.00004208</td><td>-3.8442</td><td>8</td><td>0.004918</td></tr><tr><td>Wuling(武陵山脉)</td><td>Fenghuang(凤凰山)</td><td>FH</td><td>4.7051</td><td>1</td><td>0.1333</td><td>-0.6637</td><td>2</td><td>0.5752</td></tr><tr><td>Hengduan(横断山脉)</td><td>Gaoligong(高黎贡山)</td><td>GLG</td><td>2.9785</td><td>17</td><td>0.008432</td><td>-3.3971</td><td>6</td><td>0.01455</td></tr><tr><td>Hengduan(横断山脉)</td><td>Gongga(贡嘎山)</td><td>GG</td><td>10.84</td><td>18</td><td>&lt; 0.001</td><td>-3.97</td><td>52</td><td>0.0002</td></tr><tr><td>Lvliang(吕梁山脉)</td><td>Guandi(关帝山孝文山)</td><td>GD</td><td>5.4114</td><td>3</td><td>0.01238</td><td>-5.804</td><td>8</td><td>0.0004033</td></tr><tr><td>Hengduan(横断山脉)</td><td>Haba(哈巴雪山)</td><td>HB</td><td>2.6428</td><td>2</td><td>0.1183</td><td>-1.5516</td><td>3</td><td>0.2186</td></tr><tr><td>Daba(大巴山脉)</td><td>Hualong(化龙山)</td><td>HL</td><td>—</td><td>—</td><td>—</td><td>3.603</td><td>3</td><td>0.03668</td></tr><tr><td>Wuyi(武夷山脉)</td><td>Huanggang(黄岗山)</td><td>HG</td><td>6.5888</td><td>27</td><td>4.56E-07</td><td>-6.1212</td><td>25</td><td>2.127E-06</td></tr><tr><td>Dalou(大娄山脉)</td><td>Jinfo(金佛山)</td><td>JF</td><td>4.1557</td><td>133</td><td>0.0000577</td><td>-21.055</td><td>96</td><td>&lt; 2.2e-16</td></tr><tr><td>Hengduan(横断山脉)</td><td>Kagebo(卡格薄峰)</td><td>KGB</td><td>5.0916</td><td>12</td><td>0.0002655</td><td>-5.4549</td><td>24</td><td>0.0000132</td></tr><tr><td>Hengduan(横断山脉)</td><td>Laojun(老君山)</td><td>LJ</td><td>1.8963</td><td>12</td><td>0.08225</td><td>-3.5976</td><td>2</td><td>0.06933</td></tr></tbody></table>
<table><tr><td>Miaoling(苗岭)</td><td>Leigong(雷公山)</td><td>LG</td><td>—</td><td></td><td></td><td>-1.9746</td><td>3</td><td>0.1428</td></tr><tr><td>Lushan(庐山)</td><td>Lushan(庐山)</td><td>LS</td><td>0.8541</td><td>25</td><td>0.4012</td><td>-26.477</td><td>4</td><td>0.00001209</td></tr><tr><td>Nanling(南岭)</td><td>Maoer(猫儿山)</td><td>ME</td><td>3.6141</td><td>13</td><td>0.003146</td><td>-6.0956</td><td>7</td><td>0.0004932</td></tr><tr><td>Himalayas(喜马拉雅山脉)</td><td>Nanjiabawa(南迦巴瓦峰)</td><td>NJBW</td><td>1.7692</td><td>1</td><td>0.3275</td><td>-5.8342</td><td>3</td><td>0.01003</td></tr><tr><td>Hengduan(横断山脉)</td><td>Queer(雀儿山)</td><td>QE</td><td>7.7699</td><td>3</td><td>0.004435</td><td>-3.668</td><td>8</td><td>0.006328</td></tr><tr><td>Daba(大巴山脉)</td><td>Shennongjia(神农架)</td><td>SNJ</td><td>3.8511</td><td>20</td><td>0.0009962</td><td>-9.1901</td><td>39</td><td>2.65E-11</td></tr><tr><td>Qionglai(邛崃山脉)</td><td>Siguniang(四姑娘山)</td><td>SGN</td><td>-0.478</td><td>8</td><td>0.6456</td><td>—</td><td></td><td></td></tr><tr><td>Qinling(秦岭)</td><td>Taibai(太白山)</td><td>TB</td><td>7.7445</td><td>9</td><td>0.00002866</td><td>7.7175</td><td>35</td><td>4.64E-09</td></tr><tr><td>Tianshan(天山山脉)</td><td>Tiangeer(天格尔峰)</td><td>TGE</td><td>8.9414</td><td>8</td><td>0.00001944</td><td>-2.1882</td><td>8</td><td>0.06009</td></tr><tr><td>Hengduan(横断山脉)</td><td>Wuliang(无量山)</td><td>WL</td><td>1.4907</td><td>1</td><td>0.3762</td><td>-2.5967</td><td>4</td><td>0.06026</td></tr><tr><td>Taihan(太行山脉)</td><td>Xiaowutai(小五台山)</td><td>XWT</td><td>0.1616</td><td>9</td><td>0.8752</td><td>-5.2865</td><td>1</td><td>0.119</td></tr><tr><td>Aertai(阿尔泰山脉)</td><td>Youyi(友谊峰)</td><td>YYF</td><td>4.3688</td><td>3</td><td>0.02218</td><td>-3.6829</td><td>1</td><td>0.1688</td></tr></table>
<table><caption>Table S4: The relationship between elevational shifts and elevation. The red indicate the significant relationship.</caption><thead><tr><th>Mountain range</th><th>Mount</th><th>Abbr.</th><th>slope</th><th>r2</th><th>p</th></tr></thead><tbody><tr><td>Qionglai(邛崃山)</td><td>Balang(巴郎山)</td><td>BL</td><td>-0.23</td><td>0.06</td><td>0.01</td></tr><tr><td>Bayankela(巴颜喀拉山脉)</td><td>Bayankela(巴颜喀拉山)</td><td>BYKL</td><td>-0.33</td><td>0.11</td><td>0.18</td></tr><tr><td>Hengduan(横断山脉)</td><td>Baimang(白茫雪山)</td><td>BMX</td><td>-0.09</td><td>0.03</td><td>0.50</td></tr><tr><td>Changbai(长白山脉)</td><td>Baitou(白头山)</td><td>BT</td><td>-0.12</td><td>0.03</td><td>0.40</td></tr><tr><td>Yanshan(燕山山脉)</td><td>Dongling(东灵山)</td><td>DL</td><td>-0.29</td><td>0.08</td><td>0.36</td></tr><tr><td>Qionglai(邛崃山脉)</td><td>Emei(峨眉山)</td><td>EM</td><td>-0.73</td><td>0.39</td><td>0.00</td></tr><tr><td>Wuling(武陵山脉)</td><td>Fanjing(梵净山)</td><td>FJ</td><td>0.34</td><td>0.05</td><td>0.44</td></tr><tr><td>Wuling(武陵山脉)</td><td>Fenghuang(凤凰山)</td><td>FH</td><td>0.17</td><td>0.11</td><td>0.58</td></tr><tr><td>Hengduan(横断山脉)</td><td>Gaoligong(高黎贡山)</td><td>GLG</td><td>0.03</td><td>0.00</td><td>0.87</td></tr><tr><td>Hengduan(横断山脉)</td><td>Gongga(贡嘎山)</td><td>GG</td><td>-0.13</td><td>0.05</td><td>0.03</td></tr><tr><td>Lvliang(吕梁山脉)</td><td>Guandi(关帝山孝文山)</td><td>GD</td><td>-0.41</td><td>0.26</td><td>0.08</td></tr><tr><td>Hengduan(横断山脉)</td><td>Haba(哈巴雪山)</td><td>HB</td><td>-0.42</td><td>0.24</td><td>0.26</td></tr><tr><td>Daba(大巴山脉)</td><td>Hualong(化龙山)</td><td>HL</td><td>0.15</td><td>0.27</td><td>0.37</td></tr><tr><td>Wuyi(武夷山脉)</td><td>Huanggang(黄岗山)</td><td>HG</td><td>-0.36</td><td>0.13</td><td>0.01</td></tr><tr><td>Dalou(大娄山脉)</td><td>Jinfo(金佛山)</td><td>JF</td><td>-0.25</td><td>0.22</td><td>0.00</td></tr><tr><td>Hengduan(横断山脉)</td><td>Kagebo(卡格薄峰)</td><td>KGB</td><td>-0.06</td><td>0.02</td><td>0.47</td></tr><tr><td>Hengduan(横断山脉)</td><td>Laojun(老君山)</td><td>LJ</td><td>-0.11</td><td>0.02</td><td>0.58</td></tr><tr><td>Miaoling(苗岭)</td><td>Leigong(雷公山)</td><td>LG</td><td>-2.28</td><td>0.73</td><td>0.06</td></tr><tr><td>Lushan(庐山)</td><td>Lushan(庐山)</td><td>LS</td><td>-0.52</td><td>0.49</td><td>0.00</td></tr><tr><td>Nanling(南岭)</td><td>Maoer(猫儿山)</td><td>ME</td><td>-0.37</td><td>0.08</td><td>0.21</td></tr><tr><td>Himalayas(喜马拉雅山脉)</td><td>Nanjiabawa(南迦巴瓦峰)</td><td>NJBW</td><td>-0.01</td><td>0.00</td><td>0.96</td></tr><tr><td>Hengduan(横断山脉)</td><td>Queer(雀儿山)</td><td>QE</td><td>0.50</td><td>0.19</td><td>0.14</td></tr><tr><td>Daba(大巴山脉)</td><td>Shennongjia(神农架)</td><td>SNJ</td><td>-0.54</td><td>0.32</td><td>0.00</td></tr><tr><td>Qionglai(邛崃山脉)</td><td>Siguniang(四姑娘山)</td><td>SGN</td><td>-0.09</td><td>0.02</td><td>0.68</td></tr><tr><td>Qinling(秦岭)</td><td>Taibai(太白山)</td><td>TB</td><td>-0.45</td><td>0.29</td><td>0.00</td></tr><tr><td>Tianshan(天山山脉)</td><td>Tiangeer(天格尔峰)</td><td>TGE</td><td>0.20</td><td>0.04</td><td>0.41</td></tr><tr><td>Hengduan(横断山脉)</td><td>Wuliang(无量山)</td><td>WL</td><td>-0.01</td><td>0.00</td><td>0.99</td></tr><tr><td>Taihan(太行山脉)</td><td>Xiaowutai(小五台山)</td><td>XWT</td><td>-0.44</td><td>0.55</td><td>0.02</td></tr><tr><td>Aertai(阿尔泰山脉)</td><td>Youyi(友谊峰)</td><td>YYF</td><td>-0.39</td><td>0.58</td><td>0.08</td></tr></tbody></table>