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\begin{document}

\title{Empirical Evaluation of the Elevational Rapoport's Rule: The Species
Richness Patterns for Vascular Plants on Seorak Mountain, Korea}



\author[1]{Ji-Dong Kim}%
\author[1]{Mi-Hyun Lee}%
\author[1]{Juhyeon Song}%
\author[2]{Seong Yeob Byeon}%
\author[3]{Jeong Eun Lee}%
\author[3]{Ho Jin Kim}%
\author[4]{Seung-Beom Chae}%
\author[3]{Chung Weon Yun}%
\affil[1]{Baekdudaegan National Arboretum}%
\affil[2]{Korea Institute of Arboretum Management}%
\affil[3]{Kongju National University}%
\affil[4]{Korea Forest Research Institute}%


\vspace{-1em}



  
  \date{May 24, 2024}


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\selectlanguage{english}
\begin{abstract}
Research on species abundance patterns and the advanced elevational
Rapoport rule (ERR) has been widespread in recent years; however, for
the temperate mountainous regions in northeast Asia, such research is
lacking. Here, we collected plant species from the Seorak Mountain in
northeast Asia through field surveys. The species were divided into 11
groups according to the life-form types and phytogeography affinities of
each species. The ERR was tested using Steven's method and by examining
the species abundance patterns of each group. The species abundance
patterns revealed a positive multimodal pattern along the elevation
gradient, but phytogeography affinities (increasing trend) and life-form
(unimodal) exhibited different patterns. The elevation gradients (1350 m
for the mean elevation-range relationships), which are affected by the
boundary effect and different life-forms, did not consistently support
the ERR. However, herbs as well as rare, endemic, and red list species
showed consistent support for the ERR, which can be influenced by
phytogeography affinities. Thus, the results from Seorak Mountain showed
that the ERR was not consistent for different plant life-forms in the
same area. The result of our field survey revealed that life-forms in
the plant species did not support ERR, whereas phytogeography affinities
could support and explain ERR.%
\end{abstract}%



\sloppy


Empirical Evaluation of the Elevational Rapoport's Rule: The Species
Richness Patterns for Vascular Plants on Seorak Mountain, Korea

Ji-Dong Kim\textsuperscript{1}, Mi-Hyun Lee\textsuperscript{1}, Ju Hyeon
Song\textsuperscript{1}, Seong Yeob Byeon\textsuperscript{2}, Jeong Eun
Lee\textsuperscript{3}, Ho Jin Kim\textsuperscript{3}, Seung-Beom
Chae\textsuperscript{4} and Chung Weon Yun\textsuperscript{3*}

\textsuperscript{1}Baekdudaegan National Arboretum, Bongwha 36209, Korea

\textsuperscript{2}Korea Institute of Arboretum Management, Sejong
30129, Korea

\textsuperscript{3}Department of Forest Science, Kongju National
University, Yesan 32439, Korea

\textsuperscript{4}Division of Forest Bioinformation, National Institute
of Forest Science, Suwon 16631, Korea

*Corresponding author: cwyun@kongju.ac.kr(Chung-Weon Yun)

Abstract: Research on species abundance patterns and the advanced
elevational Rapoport rule (ERR) has been widespread in recent years;
however, for the temperate mountainous regions in northeast Asia, such
research is lacking. Here, we collected plant species from the Seorak
Mountain in northeast Asia through field surveys. The species were
divided into 11 groups according to the life-form types and
phytogeography affinities of each species. The ERR was tested using
Steven's method and by examining the species abundance patterns of each
group. The species abundance patterns revealed a positive multimodal
pattern along the elevation gradient, but phytogeography affinities
(increasing trend) and life-form (unimodal) exhibited different
patterns. The elevation gradients (1350 m for the mean elevation-range
relationships), which are affected by the boundary effect and different
life-forms, did not consistently support the ERR. However, herbs as well
as rare, endemic, and red list species showed consistent support for the
ERR, which can be influenced by phytogeography affinities. Thus, the
results from Seorak Mountain showed that the ERR was not consistent for
different plant life-forms in the same area. The result of our field
survey revealed that life-forms in the plant species did not support
ERR, whereas phytogeography affinities could support and explain ERR.

Keywords: Elevation gradient, Seorak Mountain, Elevational Rapoport
rule, Species diversity, Phytogeography

Introduction

Biodiversity is expected to decrease globally over time, which will
result in widespread changes to the benefits that it currently provides
and directly impact humans and animals (Sala et al., 2000). In
terrestrial ecosystems, it was found that the main cause of biodiversity
decline is anthropogenic habitat loss due to land use changes (Sala et
al., 2005). At the level of biomes such as tundra, desert, northern
coniferous forests, and cold forests, alterations are caused by
environmental changes rather than by artificial habitat destruction
(Sala et al., 2005).

Among the various theories on biodiversity patterns, Steven's (1992)
elevational Rapoport rule (ERR) remains widely debated (Almeida-Neto et
al., 2006; Colwell \& Hurtt, 1994; Fleishman, Austin \& Weiss, 1998;
Ogwu et al., 2019). Although this approach has attracted the attention
of ecologists and biogeographers worldwide, there is considerable
controversy regarding the ERR because of the high level of variability
supporting the hypothesis. For example, some studies showed strong
support for the ERR (Chan et al., 2016; Feng et al., 2016; Luo et al.,
2011; Rohner et al., 2015; Sanders, 2002), whereas in others, little or
no support was found (Bhattarai \& Vetaas, 2006; Fleishman, Austin \&
Weiss, 1998; Guerrero, Dur\selectlanguage{ngerman}án \& Walter, 2011; Kwon et la., 2014;
Külköylüo\selectlanguage{polish}ğ\selectlanguage{english}lu et al., 2012; McCain \& Bracy Knight, 2013). Altitude range
shifts due to climate change can increase the risk of extinction for
range-restricted species (Elsen \& Tingley, 2015; Freeman \& Freeman,
2014; Harte \& Shaw, 1995; McCain \& Colwell, 2011; Sorte \& Jetz,
2010). Therefore, the accumulation of research and information on ERR in
various regions helps to increase not only our understanding of the
rule, but also the conservation of species with particularly narrow
distributions, thus helping to maintain and promote biodiversity.

An important prediction of the ERR is the positive relationship between
range-size and elevation (Stevens, 1992). The pattern in the altitude
range-size may differ for each taxon (Feng et al., 2016; Gaston, 1996),
suggesting that the range altitude relationship varies depending on the
ecological characteristics and physiological adaptations to the climate
or microenvironment along the altitude slope. Temperate taxa show a
wider altitude range-size than tropical taxa, as they may have
experienced higher variability in environmental factors during their
evolutionary and geographic history (Oommen \& Shanker, 2005; Wang, Tang
\& Fang, 2007). Therefore, information from various regions is required
to determine the link between ecological and physiological properties
and biogeographic affinity.

Another hypothesis made using the ERR is that species diversity
decreases as altitude increases. However, the results of previous
studies have been inconsistent (Feng, 2016; Zhou et al., 2019). For ERR,
a multimodal trend other than unimodal decreases has been observed
worldwide, and thus further surveys are required. Species with tropical
affinities can migrate their habitats along warm climate zones (Bergamin
et al., 2020; Feeley, Rehm \& Machovina, 2012). This suggests that
biogeographic affinity is capable of differentiated adaptation to
environmental factors including the altitude of different taxa. Species
with different biogeographic affinities may exhibit varying patterns of
abundance with altitude, which may explain the differences in support
for the abundance-altitude hypothesis of the ERR (Feng et al., 2016;
Zhou et al., 2019).

In this study, (1) the ERR was applied to Seorak Mountain (1708 m),
which is relatively lower than the world's highest mountains (e.g.,
Himalayas at 8848 m or Andes at 6961 m); (2) we determined the plant
distribution pattern according to the boundary effect and plant
geographical affinity; and (3) information regarding the species
range-size distribution patterns on the Seorak Mountain area was
obtained.

Materials and Methods

2.1 Study area

The Seorak Mountain (128\selectlanguage{ngerman}°18\selectlanguage{english}'N, 38deg05\selectlanguage{english}'E) is in eastern Korea and covers
an area of approximately 398 km\textsuperscript{2} (Figure 1a). The main
peak of Seorak Mountain is Daecheongbong (1708 m), and it is the second
highest mountain in Korea. The climate of the region is temperate, with
a mean annual temperature of 3.05degC and mean annual precipitation of
1537.39 mm (Kim, Lim \& Yun, 2019). Its temperate forests
comprise\emph{Pinus densiflora} or \emph{Abies holophylla} in the
lowlands and\emph{Betula ermanii} , \emph{Pinus koraiensis} ,
\emph{Quercus mongolica} , and \emph{Abies nephrolepis} in the
highlands. There are also dwarf tree species near the peak and in the
highlands, including\emph{Pinus pumila} , \emph{Taxus caespitosa} , and
\emph{Thuja koraiensis} , as well as arctic-alpine plants, such as
\emph{Arctous ruber} , \emph{Crataegus komarovii} , and \emph{Vaccinium
uliginosum} . To evaluate the relationship between elevation range-size
and species richness of vascular plants along the elevation gradients,
the elevation range (500--1708 m) of our study area was divided into 13
elevation bands (100 m bands, Figure 1b).

2.2 Data collection

A list of 238 plant taxa including varieties and subspecies belonging to
163 genera and 70 families was compiled for Mount Seorak based on field
data collected during vegetation surveys in this region from 2016 to
2020 at the most favorable season for plant flowering (i.e., when most
plants could be identified). For this field survey, the study area was
divided into 13 elevation bands, each with 100 m line transects
established on the mountain along elevational gradients using the
Misiryeong, Hangyeryeong, Danmokryeong, and Osaek trails centered on
Daecheongbong, which is the highest peak.

The lengths of the Misiryeong, Hangyeryeong, Danmokryeong, and Osaek
trails are approximately 15.4, 9.5, 15.5, and 10.3 km, respectively. The
four line transects were divided into 13 elevational bands at 100 m
intervals from 500 to 1708 m, and 228 plots were randomly investigated
in every elevation band of the line transects. A 400
m\textsuperscript{2} survey zone within each transect was set to conduct
plant surveys at controlled points. For surveys within each plot, the
cover-abundance scale and plant species were recorded using a previously
described vegetation survey method (Braun-Blanquet, 2013). The location
of each plot was recorded using Garmin Montana 64s GPS equipment
(Garmin, Olathe, KS, USA).

2.3 Plant life-forms and taxonomy

Following the Raunkiaer system (Raunkiaer, 1934), each species was
classified as tree, shrub, liana, herb, pteridophyte, or woody species
(including trees, shrubs, and lianas) based on the species descriptions
in the illustrated plant books by Lee (2003) and KNA (2008, 2010).
Species were classified as common species, rare, or endemic to Korea
(Lee, 2003; KNA, 2008; KNA, 2010; KFS, 2010a; KFS, 2010b). In addition,
rare plants were classified into different groups based on the red list
in IUCN 2020. The rare and red list species found in Mountain Seorak
have phytogeographic affinities (e.g., \emph{Pinus pumila}
,\emph{Leontopodium leiolepis} , \emph{Arctous ruber} ,
and\emph{Thalictrum coreanum} ).

2.4 Species richness

Species richness was defined as the total number of species in randomly
selected plots within the 100 m elevation bands, referred to as gamma
diversity. That is, a species was defined as being present in every 100
m band between its upper and lower elevational limits (Bhattarai \&
Vetaas, 2006; Feng et al., 2016; Stevens, 1992; Vetaas \& Grytnes, 2002;
Zhou et al., 2019). We calculated the species richness for the
distribution patterns of the total plant species; each life-form; and
each rare, endemic, and red list species (IUCN 2020).

2.5 Elevation range-size

To compare the elevation patterns of the different components using the
same standard, Steven's method was used to estimate the elevation range
of each species (Stevens, 1989). We identified the minimum and maximum
elevation for the distribution of each plant species in every 100 m
elevation band. A range of 100 m was provided for species found only on
a single plot and was included in the analysis.

We used our own field observations based on Steven's method and
generalized additive models to explore the range of diversity (Feng et
al., 2016; Zhou et al., 2019). The elevation range-size for each species
was estimated using the distribution patterns between the minimum and
maximum elevations, and a cubic smooth spline was used to evaluate the
significance of specific trends in the elevation range-size and species
richness (Feng et al., 2016; Hastie \& Tibshirani, 1990; Zhou et al.,
2019). These analyses were carried out using R 3.6.3 (R Core Team,
2020).

Results

3.1 Patterns of species richness along the elevation gradient

A total of 238 plant taxa, 163 genera, and 70 families were reported to
be growing from elevations of 500 m up to the tree line on the Seorak
Mountain (above 1500 m). The total species richness exhibited a
positively multimodal pattern along the elevation gradient, with a
pronounced mid-peak at 1008.4 m above sea level (a.s.l.). At this peak,
41 taxa were identified in each band (Figure 2a). After rare and endemic
species were excluded, the results were similar to those of the
pronounced mid-peak at 1004.1 m a.s.l; 39 taxa were observed at the peak
(Figure 2b). For the different life-form groups, there were decreases in
tree species at 927.3 m, shrub species at 932.7 m, and climber species
at 937.4 m (Figure 2c, d, f). In contrast, the multimodal pattern
exhibited decreased herb species at 1435.5 m, pteridophytes at 1104.1 m,
and woody species (including trees, shrubs, and lianas) at 1004.8 m
(Figure 2e, g, h).

3.2 Mean elevation range-size for the distribution of life-form groups

The mean elevation range size for total species richness exhibited a
pronounced downward trend after the mid-peak at 1291.7 m (Figure 3a).
The mean elevation range-size of excluding rare, endemic, and red list
species showed a downward trend at 1292.7 m in the elevation regions
(Figure 3b), as observed for total species richness. Similarly, trees,
shrubs, lianas, and woody groups exhibited a sharp downward trend after
the mid-peak. Pteridophytes exhibited a gentle downward trend at 1061.7
m, but no trend was detected for herbs (Figure 3).

3.3 Elevation patterns of the rare, endemic, and red list species (IUCN)

Rare, endemic, and red list species with phytogeographic affinities
intermittently appeared on the Seorak Mountain from 500 m; in contrast
to the elevation patterns of the life-forms or total species, the rare
species increased continuously from 1613.3 m (Figure 4a), and endemic
species exhibited a positive unimodal pattern at 1430 m (Figure 4b). Red
list species showed similar elevation patterns as rare species (Figure
4c). Furthermore, rare and endemic species were almost absent from the
Seorak Mountain below 800 m (Figure 4). The mean revolution range was
slightly adjusted to the range-size compared to species richness and was
similar to the species richness elevation pattern.

Discussion

The results of the empirical data collected were similar to those found
in previous studies for various taxa (e.g., birds, land snails, and
fish), as they were observed to decrease in abundance along the
elevation slopes (Carvajal-Quintero et al., 2015; Liew, Schilthuizen \&
Lakim, 2010; Pan et al., 2016). Although Seorak Mountain is a low
mountain with an altitude of 1708 m, its treeline begins at 1500 m,
above which characteristic dwarf and arctic-alpine plants appear (Kim,
Lim \& Yun, 2019).

Mount Seorak is home to a temperate climate zone forest. In tropical and
subtropical mountains, unimodal patterns of organisms are common, and
are more likely to appear because of peak diversity below the mid-point
of elevation (Cirimwami et al., 2019; Feng et al., 2015; Guo et al.,
2013; Rahbek, 2005; Zhou et al., 2019). Mountains always exhibit a
larger altitude range and longer climatic slopes, and thus they
generally have unimodal patterns. In this study, we conducted surveys in
multiple elevation bands to minimize the possibility of bias due to
uneven sampling. Field surveys were conducted using the same sampling
intensity via a phytosociology-based line transect survey method.

The maximum species abundance of vascular plants on Seorak Mountain was
found to have a multimodal pattern, with a mid-peak at 1008.4 m a.s.l.,
along the elevation gradients (Figure 2a). By excluding rare and endemic
species, the abundance patterns were found to decrease above 1004.1 m
a.s.l. (Figure 2b). Although the different life-forms groups and
vegetation species composition were similar, the total species showed a
slightly adjusted range-size pattern depending on the elevation. The
abundance of each component varies greatly along the elevation gradients
(Figure 3). The height of plants, such as trees and shrubs, showed a
high level of diversity at lower elevations, and the herbaceous and
pteridophytes patterns were combined with those of woody plants (i.e.,
trees, shrubs, and lianas), significantly reducing the maximum species
abundance (Figure 2a). The significant decrease in the proportion of
woody plants in relatively low mountain areas, including Mount Seorak,
reflects their physiological adaptations. The unimodal pattern of these
woody plants suggests that a strong boundary effect can result in a
pattern of decreasing species density (Colwell, Rahbek \& Gotelli, 2005;
Feng et al., 2016; Grytnes \& Vetaas, 2002).

Furthermore, the presence or absence of rare and endemic species
(maximum peaks at 1613.3 and 1430 m, respectively; Figure 4) greatly
influences the altitude patterns of the total species abundance
(elevation 900 to 1100 m; Figures 2a, b). For species abundance, when
rare and endemic species were excluded, the rule of decreasing species
abundance from approximately 1000 m above sea level and with higher
altitudes was confirmed. This is likely because of the shorter growing
seasons, lower temperatures, lower mass circulation, and treeline-like
environments (i.e., physical environments such as hard rock formations
and physiological constraints due to extreme climatic conditions). The
region around the Daecheongbong Peak of Mount Seorak has a strong wind
and rocky terrain; this will benefit species with a small distribution
range such as rare or endemic species (Kim et al., 2017; Kim, Lim \&
Yun, 2019).

In the Himalayas (Bhattarai \& Vetaas, 2006; Feng et al., 2016; Vetaas
\& Grytnes, 2002), Andes (Cuesta et al., 2017; Hutter, Guayasamin \&
Wiens, 2013), and various African mountain ranges (Cirimwami et al.,
2019; Zhou et al., 2019), the species abundance of rare and endemic
species continuously increased at higher altitudes, a pattern similar to
that of most mountains. In the Himalayan Mountains of Nepal, endemic
species increase with elevation gradients to an altitude of 4200 m
(Vetaas \& Grytnes, 2002), and rare and endemic species in Korea's
Seorak (maximum peak 1708 m) increased to 1613.3 and 1430 m,
respectively (Figure 4). Because of the physiological adaptations of the
plants, the range-size of endemic species or rare species may have been
reduced differently compared to that of general species, particularly in
the lowlands. Therefore, endemic and rare species may peak at altitudes
that are higher than those for the total species.

Tropical or endemic species exhibited a small elevation range-size (Zhou
et al., 2019). Similarly, on Mount Seorak, most rare or endemic species
remaining after the interglaciation showed a small elevation range-size
(Figure 4). As altitude increases, the range-size of species in the
assembly is explained as the result of individuals having to withstand
extreme climatic conditions at higher altitudes (Feng et al., 2016;
Gaston, 1996; Gaston \& Chown, 1999; Morin \& Lechowicz, 2011). Thus,
even if the overall elevation range is small, a species pool has a
similar shape (ecosystem), and an adjusted range-size can be predicted
for each species.

This elevation pattern also affects the mean elevation range of the
total species (Figure 2c, d). The mean elevation range for total species
was skewed along the elevation slope at 1291.7 m (Figure 2c), and the
mean elevation range excluding rare and endemic plants was found to have
the same elevation gradients at 1292.7 m (Figure 2d). Strong support for
the range-elevation relationship predicted by the ERR was observed in
herbs, rare, endemic, and red list species (Figure 3e; Figure 4a--c).
According to the boundary effect, the decreasing trend in the average
altitude range at high altitudes may be affected by environmental or
climatic conditions (Bhattarai \& Vetaas 2006; Feng et al., 2016; Vetaas
\& Grytnes, 2002). Considering that this study showed support for the
ERR as an increasing trend in the elevation relationship of the
range-size of herbs, rare, endemic, and red list species, the boundary
effect did not appear to have a strong effect. The proportion of endemic
and rare species increasing along the elevation gradient can affect the
relationship between the mean elevation range and elevation of species
assemblages (Pottier et al., 2013; Vetaas \& Grytnes, 2002). On Mount
Seorak, rare and endemic species were distributed continuously and
appeared at high elevation gradients (species included \emph{Adenophora
grandiflora} , \emph{Weigela subsessilis} , \emph{Lonicera subsessilis}
,\emph{Viola diamantiaca} , \emph{Syringa wolfii} , \emph{Rodgersia
podophylla} , \emph{Smilacina bicolor} , \emph{Patrinia saniculifolia} ,
and \emph{Pinus pumila} , which were from different life-form groups).

Conclusions

A field survey of plant diversity from the lowland forests of Mount
Seorak to its maximum peak was conducted, and the plants identified were
divided into their different components to test the ERR. The altitude
range of the herbs, rare, endemic, and red list species was
significantly higher than that of woody plants. Particularly, the rare,
endemic, and red list species can withstand extreme climatic conditions
through physiological adaptations, as their ranges reached the highest
elevation, and therefore can be applied to the ERR. However, the hard
boundary effect in this region consistently supported different
life-forms (i.e., trees, shrubs, lianas, pteridophytes, and woody
species). Overall, the ERR was inconsistent between plants of different
life-forms in the same region.

Data accessibility statement

All data for analysis are available at Dryad
(https://doi.org/10.5061/dryad.d51c5b02x).

Competing interest statement

The authors declare that they have no competing interests.

Author contributions section

Ji-Dong Kim: Conceptualization (lead); Formal analysis (equal);
Investigation (equal); Methodology (equal); Project administration
(lead); Software (equal); Writing\selectlanguage{english}-original draft (equal). Mi-Hyun Lee:
Data curation (equal); Formal analysis (equal); Software (equal);
Visualization (lead); Writing\selectlanguage{english}-original draft (equal). Ju-Hyeon Song:
Investigation (equal); Writing\selectlanguage{english}-review \& editing (equal). Seong-Yeob
Byeon: Formal analysis (equal); Investigation (equal). Jeong-Eun Lee:
Data curation (equal); Investigation (equal). Ho-Jin Kim: Investigation
(equal); Writing\selectlanguage{english}-review \& editing (equal). Seung-Beom Chae:
Investigation (equal); Visualization (equal). Chung-Weon Yun: Funding
acquisition (lead); Project administration (equal); Resources (equal);
Supervision (lead).

Acknowledgments

We thank the students of the Forest Ecology Laboratory at Kongju
National University for their assistance with field surveys.

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Figure Legends

\textbf{FIGURE 1} (a) Location of the Seorak Mountain (peak 1708 m) in
Korea; (b) 100 m vertical elevation bands of the Seorak Mountain. The
survey route was: \selectlanguage{english} Misiryeong, \selectlanguage{english} Hangyeryeong, \selectlanguage{english} Danmokryeong, and \selectlanguage{english}
Osaek.

\textbf{FIGURE 2} Elevation patterns of species richness of vascular
plants on the Seorak Mountain. Total species richness (a); species
richness excluding rare, endemic, and red list species (b); and species
richness of trees (c); shrubs (d); herbs (e); lianas (f); pteridophytes
(g); and woody species (h).

\textbf{FIGURE 3} Mean elevation range-size of different of life-form
groups along the elevation gradient of the Seorak Mountain: (a) total
plants; (b) total plants excluding rare, endemic, and red list species;
(c) trees; (d) shrubs; (e) herbs; (f) lianas; (g) pteridophytes; and (h)
woody species.

\textbf{FIGURE 4} Elevation patterns in species richness and mean
elevation range of (a) rare species, (b) endemic species, and (c) red
list (IUCN) species on the Seorak Mountain.

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