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\begin{document}

\title{Synergies between mini-patches of plant communities and steady-stage
alpine meadows on the Qinghai-Tibet Plateau}



\author[1]{li lin}%
\author[2]{Xingliang Xu}%
\author[1]{guangmin cao}%
\author[1]{Fawei Zhang}%
\author[1]{Yikang Li}%
\author[3]{Junjie Huang}%
\affil[1]{Northwest Institute of Plateau Biology Chinese Academy of Sciences}%
\affil[2]{Institute of Geographic Sciences and Natural Resources Research Chinese Academy of Sciences}%
\affil[3]{School of Mechanical and Power Engineering Henan Polytechnic University Jiaozuo China 454000}%


\vspace{-1em}



  
  \date{May 16, 2024}


\begingroup
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\selectlanguage{english}
\begin{abstract}
Mini-patches are considered indicators of an ecosystem's response to
interference, particularly those in alpine meadow ecosystems. Thus,
monitoring the characteristics of mini-patches can elucidate the
organization of an ecosystem's components, the strategies it employs to
survive interference, and the mechanisms whereby it maintains stability.
In this research, we used multivariate statistical analysis methods to
investigate the characteristics of the plant community and the
micro-topography of mini-patches in alpine meadows on the Qinghai-Tibet
Plateau from August 2012 to August 2013. Our findings show that (1)
mini-patches were distributed in alpine meadows with different levels of
degradation and the effects of meteorological characteristics
(accumulated temperature above 0\selectlanguage{ngerman}°C and accumulation of precipitation)
and geographical characteristics (altitude, longitude, and latitude)
contributed less than 20\% to their distribution and characteristics;
(2) alpine meadows maintained aboveground biomass within a certain range
under a relative larger range of grazing intensity, illustrating their
ability to regulate community structure and components under various
intensities of disturbance and showing that alpine degradation could
itself counteract grazing disturbance; and (3) overgrazing is the main
driver of multi-steady stage coexistence in alpine meadows, as the
mini-patches that remain involved in plant community succession
function, and as a source of germplasm in the plant community regime
shift under different grazing intensities damaged alpine meadows.%
\end{abstract}\selectlanguage{ngerman}%



\sloppy


\textbf{Synergies between mini-patches of plant communities and
steady-stage alpine meadows on the Qinghai-Tibet Plateau}

\textbf{Li Lin 1, 2, Xingliang Xu 3*, Guangmin Cao 1, Fawei Zhang 1,
Yikang Li 1, Junjie Huang 4}

1 Northwest Institute of Plateau Biology and Chinese Academy of
Sciences, Xining, China, 810008

2 University of Chinese Academy of Sciences, Beijing, China, 100049.

3 Institute of Geographic Sciences and Natural Resources Research,
Chinese Academy of Sciences, Beijing, China, 100101

4 School of Mechanical and Power Engineering, Henan Polytechnic
University, Jiaozuo, China, 454000

* Correspondence: xuxl@igsnrr.ac.cn.

Founder: Natural Science Foundation of Qinghai (2020-ZJ-720)

\textbf{Abstract:}

Mini-patches are considered indicators of an ecosystem's response to
interference, particularly those in alpine meadow ecosystems. Thus,
monitoring the characteristics of mini-patches can elucidate the
organization of an ecosystem's components, the strategies it employs to
survive interference, and the mechanisms whereby it maintains stability.
In this research, we used multivariate statistical analysis methods to
investigate the characteristics of the plant community and the
micro-topography of mini-patches in alpine meadows on the Qinghai-Tibet
Plateau from August 2012 to August 2013. Our findings show that (1)
mini-patches were distributed in alpine meadows with different levels of
degradation and the effects of meteorological characteristics
(accumulated temperature above 0\selectlanguage{ngerman}°C and accumulation of precipitation)
and geographical characteristics (altitude, longitude, and latitude)
contributed less than 20\% to their distribution and characteristics;
(2) alpine meadows maintained aboveground biomass within a certain range
under a relative larger range of grazing intensity, illustrating their
ability to regulate community structure and components under various
intensities of disturbance and showing that alpine degradation could
itself counteract grazing disturbance; and (3) overgrazing is the main
driver of multi-steady stage coexistence in alpine meadows, as the
mini-patches that remain involved in plant community succession
function, and as a source of germplasm in the plant community regime
shift under different grazing intensities damaged alpine meadows.

\textbf{Keywords:}

Mini-patch, alpine meadow degradation, succession process, overgrazing,
multi-steady stage

\textbf{Introduction}

The alpine meadows that occupy more than 35\% of the total area of the
Qinghai-Tibet Plateau are the main location of alpine livestock
husbandry in the area, particularly because of their use as winter and
spring pastures (Zhang et al., 2003). The health of such alpine meadows
must be measured to enable estimation of the capacity of ecosystem
services, such as water and soil conservation and carbon sequestration
(Yang, 2002). In recent years, these alpine meadows have undergone
succession into multi-steady stages due to disturbances arising from
climate change and excessive human activity (Meng et al., 2014). Given
the unique household contract system implemented on family ranches on
the Qinghai-Tibet Plateau, the landscape has fragmented into multiple
succession stages or steady states, resulting in multi-steady stage
coexistence within each topographical and climate area (Cao et al.,
2007).

Ecosystem succession processes always proceed sequentially, such that
the corresponding plant communities organize into a series of
hierarchical structures or discrete levels (Jørgensen et al., 2016;
Wellemeyer et al., 2019). Livestock grazing is the primary form of
disturbance that affects plant community structure, physical and
chemical soil properties, and the ability of alpine meadows to maintain
ecological and productive stability (Proulx et al., 1998; Yang et al.,
2001). In addition, the effects of livestock grazing on plant-soil
system organization are complex (Nicola et al., 2010). For example,
long-term and high-intensity grazing can affect plant community
structure and composition and even trigger the plant community to regime
shift (Lin et al., 2022). Such grazing can also alter the accumulation
and transformation of nutrients above and below the ground and influence
the micro-climate and micro-geomorphology of mini-patches (Gonzalez et
al., 2010). Most prior studies have observed that the intensity of
livestock grazing and the reaction of a plant community to grazing are
coupled to various conditions (Wedin \& Tilman, 1996; Klinerová \&
Dostal, 2020; Lauren, 2018), which means that the examination of
mini-patches can be used to evaluate plant communities and predict the
trends of plant community succession (Wu et al., 2012).

Compared with other parts of ecosystems, mini-patches are more sensitive
and exhibit greater variations in response to disturbance and can thus
be considered as indicators of how ecosystems respond to disturbance.
Monitoring the characteristics of mini-patches can thus reveal the
organizational components of an ecosystem, the survival strategies used
by an ecosystem's plant community to cope with disturbance, and the
mechanisms whereby an ecosystem maintains its stability. Mini-patches in
alpine meadows can therefore be considered as the basic units of the
plant--soil system at fine spatial scales, with their features isolated
in function but correlated in space. Consequently, the study of
mini-patches in alpine meadows can provide answers to many key
questions. For example, are the same types of mini-patches distributed
across alpine meadow area of different grazing intensities or only in
areas of certain grazing intensities? What factors mainly influence
mini-patch distribution and determine the direction of plant community
succession in alpine meadows?

In this research, we aimed to improve understanding on the stabilization
mechanisms of alpine meadow ecosystems and strengthen theories on
restoration management in multi-steady-state coexistence conditions.
Therefore, we explored and examined the factors affecting alpine meadow
ecosystems under various grazing regimes by studying the characteristics
of plant functional groups and topography in mini-patches and grazing
intensity in typical alpine meadows on the Qinghai-Tibet Plateau.

\textbf{1 Materials and methods}

\textbf{1.1 Study area and sampling method}

We chose 109 sampling sites located in the autonomous prefectures of
Haibei, Yushu, Guoluo, Haixi, and Hainan in China (Table 1). The
meteorological characteristics of accumulated temperature above 0°C and
accumulated precipitation over the growing season (mainly in May to
September) were respectively 659.1 to 1228.4°C and 257.5 to 693.7 mm in
all of the sampling sites. The vegetation type was classified as cold,
wet-rainy tundra and alpine meadow according to the Comprehensive and
Sequential Classification System of Grassland, and as alpine meadow
according to the passive-active degradation process hypothesis (Cao et
al., 2007). However, the sampled alpine meadows were in different steady
stages (Cao et al., 2007), so we classified the corresponding sample
sites according to the characteristics of their plant community, mattic
epipedon, and topography into seven steady-stage types of meadow:
Gramineae--\emph{Kobresia}meadow, Gramineae--\emph{K.humilis} to\emph{K.
pygmaea} transitional meadow, healthy mattic epipedon in \emph{K.
pygmaea} meadow, thickening mattic epipedon in\emph{K. pygmaea}meadow,
cracking mattic epipedon in \emph{K. pygmaea} meadow, eroded mattic
epipedon in \emph{K. pygmaea} meadow, and black-soil beach meadow (Table
1). In view of the fact that the quantitative characteristics of plant
communities are prone to reciprocating changes under different grazing
intensity, so we divided the meadows into five categories, namely
Gramineae--\emph{Kobresia} meadow and Gramineae--\emph{K.}
\emph{humilis} to \emph{K. pygmaea} transitional meadow as ``A'',
healthy mattic epipedon in \emph{K. pygmaea} meadow and thickening
mattic epipedon in \emph{K. pygmaea} meadow as ``B'', cracking mattic
epipedon in \emph{K. pygmaea} meadow as ``C'', eroded mattic epipedon in
\emph{K. pygmaea} meadow as ``D'', and black-soil beach meadow as ``E''.

Table 1. Latitude, longitude, and altitude of sampling sites and their
stability stages\selectlanguage{english}
\begin{longtable}[]{@{}lllll@{}}
\toprule
Code & Longitude & Latitude & Altitude (m) & Steady stage\tabularnewline
\midrule
\endhead
1 & 101\selectlanguage{ngerman}°44\selectlanguage{english}'56\selectlanguage{english}" & 34\selectlanguage{ngerman}°39\selectlanguage{english}'52\selectlanguage{english}" & 3596 & Gramineae-\emph{Kobresia}
meadow\tabularnewline
2 & 100\selectlanguage{ngerman}°51\selectlanguage{english}'57\selectlanguage{english}" & 34\selectlanguage{ngerman}°58\selectlanguage{english}'19\selectlanguage{english}" & 3802 & Gramineae- \emph{Kobresia}
meadow\tabularnewline
3 & 100\selectlanguage{ngerman}°29\selectlanguage{english}'32\selectlanguage{english}" & 34\selectlanguage{ngerman}°21\selectlanguage{english}'29\selectlanguage{english}" & 3948 & Gramineae- \emph{Kobresia}
meadow\tabularnewline
4 & 99\selectlanguage{ngerman}°54\selectlanguage{english}'47\selectlanguage{english}" & 33\selectlanguage{ngerman}°34\selectlanguage{english}'28\selectlanguage{english}" & 4067 & Gramineae- \emph{Kobresia}
meadow\tabularnewline
5 & 95\selectlanguage{ngerman}°29\selectlanguage{english}'24\selectlanguage{english}" & 34\selectlanguage{ngerman}°5\selectlanguage{english}'24\selectlanguage{english}" & 4306 & Gramineae- \emph{Kobresia}
meadow\tabularnewline
6 & 94\selectlanguage{ngerman}°33\selectlanguage{english}'0\selectlanguage{english}" & 34\selectlanguage{ngerman}°30\selectlanguage{english}'0\selectlanguage{english}" & 4316 & Gramineae- \emph{Kobresia}
meadow\tabularnewline
7 & 99\selectlanguage{ngerman}°32\selectlanguage{english}'14.34\selectlanguage{english}" & 37\selectlanguage{ngerman}°4\selectlanguage{english}'2.58\selectlanguage{english}" & 3233 & Gramineae- \emph{Kobresia}
meadow\tabularnewline
8 & 101\selectlanguage{ngerman}°18\selectlanguage{english}'57\selectlanguage{english}" & 37\selectlanguage{ngerman}°36\selectlanguage{english}'47\selectlanguage{english}" & 3200 & Gramineae- \emph{Kobresia}
meadow\tabularnewline
9 & 101\selectlanguage{ngerman}°17\selectlanguage{english}'52.02\selectlanguage{english}" & 37\selectlanguage{ngerman}°37\selectlanguage{english}'13.26\selectlanguage{english}" & 3204 & Gramineae- \emph{Kobresia}
meadow\tabularnewline
10 & 96\selectlanguage{ngerman}º55\selectlanguage{english}'40.56\selectlanguage{english}" & 32\selectlanguage{ngerman}º51\selectlanguage{english}'59.4'' & 3982 & Gramineae- \emph{Kobresia}
meadow\tabularnewline
11 & 100\selectlanguage{ngerman}º12\selectlanguage{english}'47.22\selectlanguage{english}" & 34\selectlanguage{ngerman}º27\selectlanguage{english}'47.1'' & 3740 & Gramineae- \emph{Kobresia}
meadow\tabularnewline
12 & 101\selectlanguage{ngerman}º2\selectlanguage{english}'31.14\selectlanguage{english}" & 37\selectlanguage{ngerman}º35\selectlanguage{english}'24.48'' & 3171 & Gramineae- \emph{Kobresia}
meadow\tabularnewline
13 & 101\selectlanguage{ngerman}°27\selectlanguage{english}'09\selectlanguage{english}" & 35\selectlanguage{ngerman}°4\selectlanguage{english}'56\selectlanguage{english}" & 3725 & Craking mattic epipedon \emph{in K.
pygmaea} meadow\tabularnewline
14 & 100\selectlanguage{ngerman}°51\selectlanguage{english}'53\selectlanguage{english}" & 34\selectlanguage{ngerman}°58\selectlanguage{english}'13\selectlanguage{english}" & 3786 & Thickening mattic epipedon \emph{in
K. pygmaea} meadow\tabularnewline
15 & 100\selectlanguage{ngerman}°34\selectlanguage{english}'19\selectlanguage{english}" & 34\selectlanguage{ngerman}°20\selectlanguage{english}'23\selectlanguage{english}" & 4081 & Gramineae-\emph{K.} \emph{humilis}
to \emph{K. pygmaea} transitional meadow\tabularnewline
16 & 100\selectlanguage{ngerman}°29\selectlanguage{english}'33\selectlanguage{english}" & 34\selectlanguage{ngerman}°21\selectlanguage{english}'57\selectlanguage{english}" & 3955 & Gramineae-\emph{K.} \emph{humilis}
to \emph{K. pygmaea} transitional meadow\tabularnewline
17 & 100\selectlanguage{ngerman}°16\selectlanguage{english}'51\selectlanguage{english}" & 34\selectlanguage{ngerman}°25\selectlanguage{english}'16\selectlanguage{english}" & 3768 & Gramineae-\emph{K.} \emph{humilis}
to \emph{K. pygmaea} transitional meadow\tabularnewline
18 & 100\selectlanguage{ngerman}°24\selectlanguage{english}'21\selectlanguage{english}" & 34\selectlanguage{ngerman}°28\selectlanguage{english}'42\selectlanguage{english}" & 4328 & Gramineae-\emph{K.} \emph{humilis}
to \emph{K. pygmaea} transitional meadow\tabularnewline
19 & 100\selectlanguage{ngerman}°16\selectlanguage{english}'22\selectlanguage{english}" & 34\selectlanguage{ngerman}°23\selectlanguage{english}'37\selectlanguage{english}" & 3820 & Gramineae-\emph{K.} \emph{humilis}
to \emph{K. pygmaea} transitional meadow\tabularnewline
20 & 100\selectlanguage{ngerman}°36\selectlanguage{english}'34\selectlanguage{english}" & 34\selectlanguage{ngerman}°19\selectlanguage{english}'47\selectlanguage{english}" & 4151 & Gramineae-\emph{K.} \emph{humilis}
to \emph{K. pygmaea} transitional meadow\tabularnewline
21 & 100\selectlanguage{ngerman}°5\selectlanguage{english}'6\selectlanguage{english}" & 34\selectlanguage{ngerman}°4\selectlanguage{english}'6\selectlanguage{english}" & 4292 & Thickening mattic epipedon \emph{in K.
pygmaea} meadow\tabularnewline
22 & 100\selectlanguage{ngerman}°5\selectlanguage{english}'28\selectlanguage{english}" & 34\selectlanguage{ngerman}°1\selectlanguage{english}'15\selectlanguage{english}" & 4252 & Thickening mattic epipedon \emph{in
K. pygmaea} meadow\tabularnewline
23 & 99\selectlanguage{ngerman}°42\selectlanguage{english}'16\selectlanguage{english}" & 33\selectlanguage{ngerman}°48\selectlanguage{english}'26\selectlanguage{english}" & 3974 & Thickening mattic epipedon \emph{in
K. pygmaea} meadow\tabularnewline
24 & 100\selectlanguage{ngerman}°55\selectlanguage{english}'9\selectlanguage{english}" & 35\selectlanguage{ngerman}°42\selectlanguage{english}'50\selectlanguage{english}" & 3370 & Craking mattic epipedon \emph{in K.
pygmaea} meadow\tabularnewline
25 & 99\selectlanguage{ngerman}°25\selectlanguage{english}'8\selectlanguage{english}" & 33\selectlanguage{ngerman}°45\selectlanguage{english}'6\selectlanguage{english}" & 4012 & Thickening mattic epipedon \emph{in K.
pygmaea} meadow\tabularnewline
26 & 100\selectlanguage{ngerman}°33\selectlanguage{english}'53\selectlanguage{english}" & 32\selectlanguage{ngerman}°46\selectlanguage{english}'36\selectlanguage{english}" & 4207 & Thickening mattic epipedon \emph{in
K. pygmaea} meadow\tabularnewline
27 & 100\selectlanguage{ngerman}°51\selectlanguage{english}'7\selectlanguage{english}" & 30\selectlanguage{ngerman}°17\selectlanguage{english}'28\selectlanguage{english}" & 4401 & Thickening mattic epipedon \emph{in
K. pygmaea} meadow\tabularnewline
28 & 100\selectlanguage{ngerman}°3\selectlanguage{english}'42.00\selectlanguage{english}" & 33\selectlanguage{ngerman}°25\selectlanguage{english}'51\selectlanguage{english}" & 4235 & Thickening mattic epipedon
\emph{in K. pygmaea} meadow\tabularnewline
29 & 100\selectlanguage{ngerman}°23\selectlanguage{english}'9\selectlanguage{english}" & 33\selectlanguage{ngerman}°16\selectlanguage{english}'5\selectlanguage{english}" & 4111 & Gramineae-\emph{K.} \emph{humilis} to
\emph{K. pygmaea} transitional meadow\tabularnewline
30 & 98\selectlanguage{ngerman}°7\selectlanguage{english}'21\selectlanguage{english}" & 34\selectlanguage{ngerman}°46\selectlanguage{english}'6\selectlanguage{english}" & 4328 & Thickening mattic epipedon \emph{in K.
pygmaea} meadow\tabularnewline
31 & 98\selectlanguage{ngerman}°41\selectlanguage{english}'41\selectlanguage{english}" & 37\selectlanguage{ngerman}°31\selectlanguage{english}'53\selectlanguage{english}" & 3642 & Gramineae-\emph{K.} \emph{humilis}
to \emph{K. pygmaea} transitional meadow\tabularnewline
32 & 97\selectlanguage{ngerman}°10\selectlanguage{english}'06.42\selectlanguage{english}" & 33\selectlanguage{ngerman}°50\selectlanguage{english}'12.3\selectlanguage{english}" & 4425 & Thickening mattic epipedon
\emph{in K. pygmaea} meadow\tabularnewline
33 & 96\selectlanguage{ngerman}°10\selectlanguage{english}'48\selectlanguage{english}" & 33\selectlanguage{ngerman}°10\selectlanguage{english}'48\selectlanguage{english}" & 4417 & Gramineae-\emph{K.} \emph{humilis}
to \emph{K. pygmaea} transitional meadow\tabularnewline
34 & 96\selectlanguage{ngerman}°1\selectlanguage{english}'12\selectlanguage{english}" & 33\selectlanguage{ngerman}°20\selectlanguage{english}'24\selectlanguage{english}" & 4303 & Gramineae-\emph{K.} \emph{humilis} to
\emph{K. pygmaea} transitional meadow\tabularnewline
35 & 95\selectlanguage{ngerman}°25\selectlanguage{english}'12\selectlanguage{english}" & 33\selectlanguage{ngerman}°29\selectlanguage{english}'24\selectlanguage{english}" & 4232 & Gramineae-\emph{K.} \emph{humilis}
to \emph{K. pygmaea} transitional meadow\tabularnewline
36 & 95\selectlanguage{ngerman}°28\selectlanguage{english}'48\selectlanguage{english}" & 34\selectlanguage{ngerman}°3\selectlanguage{english}'36\selectlanguage{english}" & 4219 & Gramineae-\emph{K.} \emph{humilis} to
\emph{K. pygmaea} transitional meadow\tabularnewline
37 & 95\selectlanguage{ngerman}°30\selectlanguage{english}'36\selectlanguage{english}" & 34\selectlanguage{ngerman}°4\selectlanguage{english}'12\selectlanguage{english}" & 4219 & Gramineae-\emph{K.} \emph{humilis} to
\emph{K. pygmaea} transitional meadow\tabularnewline
38 & 95\selectlanguage{ngerman}°30\selectlanguage{english}'36\selectlanguage{english}" & 34\selectlanguage{ngerman}°4\selectlanguage{english}'48\selectlanguage{english}" & 4432 & Gramineae-\emph{K.} \emph{humilis} to
\emph{K. pygmaea} transitional meadow\tabularnewline
39 & 100\selectlanguage{ngerman}°45\selectlanguage{english}'9.72\selectlanguage{english}" & 37\selectlanguage{ngerman}°1\selectlanguage{english}'19.2\selectlanguage{english}" & 3350 & Gramineae-\emph{K.}
\emph{humilis} to \emph{K. pygmaea} transitional meadow\tabularnewline
40 & 100\selectlanguage{ngerman}°45\selectlanguage{english}'10.32\selectlanguage{english}" & 37\selectlanguage{ngerman}°1\selectlanguage{english}'21.96\selectlanguage{english}" & 3336 & Gramineae-\emph{K.}
\emph{humilis} to \emph{K. pygmaea} transitional meadow\tabularnewline
41 & 100\selectlanguage{ngerman}°22\selectlanguage{english}'42.9\selectlanguage{english}" & 37\selectlanguage{ngerman}°54\selectlanguage{english}'18.66\selectlanguage{english}" & 3686 & Thickening mattic epipedon
\emph{in K. pygmaea} meadow\tabularnewline
42 & 100\selectlanguage{ngerman}°22\selectlanguage{english}'32.04\selectlanguage{english}" & 37\selectlanguage{ngerman}°54\selectlanguage{english}'27.84\selectlanguage{english}" & 3691 & Thickening mattic epipedon
\emph{in K. pygmaea} meadow\tabularnewline
43 & 100\selectlanguage{ngerman}°15\selectlanguage{english}'26.76\selectlanguage{english}" & 38\selectlanguage{ngerman}°0\selectlanguage{english}'21.06\selectlanguage{english}" & 3985 & Thickening mattic epipedon
\emph{in K. pygmaea} meadow\tabularnewline
44 & 99\selectlanguage{ngerman}°23\selectlanguage{english}'04.74\selectlanguage{english}" & 38\selectlanguage{ngerman}°35\selectlanguage{english}'43.38\selectlanguage{english}" & 3436 & Gramineae-\emph{K.}
\emph{humilis} to \emph{K. pygmaea} transitional meadow\tabularnewline
45 & 99\selectlanguage{ngerman}°23\selectlanguage{english}'02.7\selectlanguage{english}" & 38\selectlanguage{ngerman}°35\selectlanguage{english}'39.96\selectlanguage{english}" & 3428 & Gramineae-\emph{K.}
\emph{humilis} to \emph{K. pygmaea} transitional meadow\tabularnewline
46 & 100\selectlanguage{ngerman}°53\selectlanguage{english}'54\selectlanguage{english}" & 37\selectlanguage{ngerman}°57\selectlanguage{english}'55\selectlanguage{english}" & 3335 & Gramineae-\emph{K.} \emph{humilis}
to \emph{K. pygmaea} transitional meadow\tabularnewline
47 & 101\selectlanguage{ngerman}°2\selectlanguage{english}'56\selectlanguage{english}" & 37\selectlanguage{ngerman}°52\selectlanguage{english}'18\selectlanguage{english}" & 3527 & Gramineae-\emph{K.} \emph{humilis}
to \emph{K. pygmaea} transitional meadow\tabularnewline
48 & 101\selectlanguage{ngerman}°15\selectlanguage{english}'28\selectlanguage{english}" & 37\selectlanguage{ngerman}°42\selectlanguage{english}'08\selectlanguage{english}" & 3278 & Cracking mattic epipedon \emph{in
K. pygmaea} meadow\tabularnewline
49 & 101\selectlanguage{ngerman}°10\selectlanguage{english}'56\selectlanguage{english}" & 37\selectlanguage{ngerman}°40\selectlanguage{english}'54\selectlanguage{english}" & 3239 & Thickening mattic epipedon \emph{in
K. pygmaea} meadow\tabularnewline
50 & 101\selectlanguage{ngerman}°10\selectlanguage{english}'38.28\selectlanguage{english}" & 37\selectlanguage{ngerman}°40\selectlanguage{english}'9.3\selectlanguage{english}" & 3241 & Thickening mattic epipedon
\emph{in K. pygmaea} meadow\tabularnewline
51 & 95\selectlanguage{ngerman}º59\selectlanguage{english}'45.06\selectlanguage{english}" & 34\selectlanguage{ngerman}º36\selectlanguage{english}'20.58'' & 4250 & Gramineae-\emph{K.}
\emph{humilis} to \emph{K. pygmaea} transitional meadow\tabularnewline
52 & 97\selectlanguage{ngerman}º19\selectlanguage{english}'38.34\selectlanguage{english}" & 33\selectlanguage{ngerman}º22\selectlanguage{english}'31.56'' & 4255 & Cracking mattic epipedon
\emph{in K. pygmaea} meadow\tabularnewline
53 & 97\selectlanguage{ngerman}º19\selectlanguage{english}'47.7\selectlanguage{english}" & 33\selectlanguage{ngerman}º22\selectlanguage{english}'33.9'' & 4247 & Cracking mattic epipedon
\emph{in K. pygmaea} meadow\tabularnewline
54 & 96\selectlanguage{ngerman}º55\selectlanguage{english}'39.12\selectlanguage{english}" & 32\selectlanguage{ngerman}º51\selectlanguage{english}'56.04'' & 3988 & Gramineae-\emph{K.}
\emph{humilis} to \emph{K. pygmaea} transitional meadow\tabularnewline
55 & 96\selectlanguage{ngerman}º41\selectlanguage{english}'41.52\selectlanguage{english}" & 33\selectlanguage{ngerman}º7\selectlanguage{english}'41.82'' & 4370 & Thickening mattic epipedon
\emph{in K. pygmaea} meadow\tabularnewline
56 & 96\selectlanguage{ngerman}º10\selectlanguage{english}'10.38\selectlanguage{english}" & 33\selectlanguage{ngerman}º24\selectlanguage{english}'34.68'' & 4504 & Gramineae-\emph{K.}
\emph{humilis} to \emph{K. pygmaea} transitional meadow\tabularnewline
57 & 95\selectlanguage{ngerman}º8\selectlanguage{english}'13.32\selectlanguage{english}" & 34\selectlanguage{ngerman}º43\selectlanguage{english}'12.36'' & 4507 & Cracking mattic epipedon
\emph{in K. pygmaea} meadow\tabularnewline
58 & 95\selectlanguage{ngerman}º20\selectlanguage{english}'51.18\selectlanguage{english}" & 34\selectlanguage{ngerman}º33\selectlanguage{english}'41.16'' & 4234 & Gramineae-\emph{K.}
\emph{humilis} to \emph{K. pygmaea} transitional meadow\tabularnewline
59 & 95\selectlanguage{ngerman}º20\selectlanguage{english}'51.36\selectlanguage{english}" & 34\selectlanguage{ngerman}º33\selectlanguage{english}'10.32'' & 4229 & Thickening mattic epipedon
\emph{in K. pygmaea} meadow\tabularnewline
60 & 94\selectlanguage{ngerman}º49\selectlanguage{english}'48.42\selectlanguage{english}" & 34\selectlanguage{ngerman}º53\selectlanguage{english}'21.06'' & 4531 & Cracking mattic epipedon
\emph{in K. pygmaea} meadow\tabularnewline
61 & 100\selectlanguage{ngerman}º8\selectlanguage{english}'11.1\selectlanguage{english}" & 34\selectlanguage{ngerman}º22\selectlanguage{english}'32.94'' & 3934 & Gramineae-\emph{K.}
\emph{humilis} to \emph{K. pygmaea} transitional meadow\tabularnewline
62 & 99\selectlanguage{ngerman}º48\selectlanguage{english}'11.82\selectlanguage{english}" & 33\selectlanguage{ngerman}º55\selectlanguage{english}'11.58'' & 4185 & Cracking mattic epipedon
\emph{in K. pygmaea} meadow\tabularnewline
63 & 99\selectlanguage{ngerman}º14\selectlanguage{english}'59.94\selectlanguage{english}" & 33\selectlanguage{ngerman}º22\selectlanguage{english}'22.5'' & 4320 & Cracking mattic epipedon
\emph{in K. pygmaea} meadow\tabularnewline
64 & 99\selectlanguage{ngerman}º54\selectlanguage{english}'16.2\selectlanguage{english}" & 33\selectlanguage{ngerman}º34\selectlanguage{english}'16.44'' & 4082 & Thickening mattic epipedon
\emph{in K. pygmaea} meadow\tabularnewline
65 & 101\selectlanguage{ngerman}°16\selectlanguage{english}'32\selectlanguage{english}" & 35\selectlanguage{ngerman}°17\selectlanguage{english}'52\selectlanguage{english}" & 3580 & Eroded mattic epipedon \emph{in K.
pygmaea} meadow\tabularnewline
66 & 101\selectlanguage{ngerman}°16\selectlanguage{english}'17\selectlanguage{english}" & 35\selectlanguage{ngerman}°17\selectlanguage{english}'32\selectlanguage{english}" & 3580 & Eroded mattic epipedon \emph{in K.
pygmaea} meadow\tabularnewline
67 & 101\selectlanguage{ngerman}°19\selectlanguage{english}'31\selectlanguage{english}" & 35\selectlanguage{ngerman}°16\selectlanguage{english}'59\selectlanguage{english}" & 3635 & Black-soil beach
meadow\tabularnewline
68 & 101\selectlanguage{ngerman}°44\selectlanguage{english}'08\selectlanguage{english}" & 34\selectlanguage{ngerman}°39\selectlanguage{english}'06\selectlanguage{english}" & 3593 & Black-soil beach
meadow\tabularnewline
69 & 100\selectlanguage{ngerman}°52\selectlanguage{english}'02\selectlanguage{english}" & 34\selectlanguage{ngerman}°58\selectlanguage{english}'27\selectlanguage{english}" & 3800 & Eroded mattic epipedon \emph{in K.
pygmaea} meadow\tabularnewline
70 & 100\selectlanguage{ngerman}°34\selectlanguage{english}'19\selectlanguage{english}" & 34\selectlanguage{ngerman}°20\selectlanguage{english}'24\selectlanguage{english}" & 4084 & Eroded mattic epipedon \emph{in K.
pygmaea} meadow\tabularnewline
71 & 100\selectlanguage{ngerman}°29\selectlanguage{english}'47\selectlanguage{english}" & 34\selectlanguage{ngerman}°21\selectlanguage{english}'9\selectlanguage{english}" & 3957 & Black-soil beach
meadow\tabularnewline
72 & 100\selectlanguage{ngerman}°16\selectlanguage{english}'8\selectlanguage{english}" & 34\selectlanguage{ngerman}°25\selectlanguage{english}'57\selectlanguage{english}" & 3772 & Black-soil beach
meadow\tabularnewline
73 & 100\selectlanguage{ngerman}°8\selectlanguage{english}'9\selectlanguage{english}" & 34\selectlanguage{ngerman}°9\selectlanguage{english}'2\selectlanguage{english}" & 4089 & Eroded mattic epipedon \emph{in K.
pygmaea} meadow\tabularnewline
74 & 100\selectlanguage{ngerman}°8\selectlanguage{english}'9\selectlanguage{english}" & 34\selectlanguage{ngerman}°9\selectlanguage{english}'19\selectlanguage{english}" & 4091 & Eroded mattic epipedon \emph{in K.
pygmaea} meadow\tabularnewline
75 & 100\selectlanguage{ngerman}°36\selectlanguage{english}'27\selectlanguage{english}" & 34\selectlanguage{ngerman}°19\selectlanguage{english}'42\selectlanguage{english}" & 4145 & Black-soil beach
meadow\tabularnewline
76 & 100\selectlanguage{ngerman}°5\selectlanguage{english}'07\selectlanguage{english}" & 34\selectlanguage{ngerman}°4\selectlanguage{english}'55\selectlanguage{english}" & 4212 & Eroded mattic epipedon \emph{in K.
pygmaea} meadow\tabularnewline
77 & 99\selectlanguage{ngerman}°48\selectlanguage{english}'35\selectlanguage{english}" & 33\selectlanguage{ngerman}°55\selectlanguage{english}'20\selectlanguage{english}" & 4189 & Black-soil beach
meadow\tabularnewline
78 & 99\selectlanguage{ngerman}°48\selectlanguage{english}'38\selectlanguage{english}" & 33\selectlanguage{ngerman}°55\selectlanguage{english}'26\selectlanguage{english}" & 4203 & Eroded mattic epipedon \emph{in K.
pygmaea} meadow\tabularnewline
79 & 99\selectlanguage{ngerman}°42\selectlanguage{english}'10\selectlanguage{english}" & 33\selectlanguage{ngerman}°48\selectlanguage{english}'06\selectlanguage{english}" & 3956 & Eroded mattic epipedon \emph{in K.
pygmaea} meadow\tabularnewline
80 & 100\selectlanguage{ngerman}°13\selectlanguage{english}'07\selectlanguage{english}" & 33\selectlanguage{ngerman}°28\selectlanguage{english}'23\selectlanguage{english}" & 4299 & Eroded mattic epipedon \emph{in K.
pygmaea} meadow\tabularnewline
81 & 100\selectlanguage{ngerman}°12\selectlanguage{english}'59\selectlanguage{english}" & 33\selectlanguage{ngerman}°23\selectlanguage{english}'28\selectlanguage{english}" & 4288 & Eroded mattic epipedon \emph{in K.
pygmaea} meadow\tabularnewline
82 & 99\selectlanguage{ngerman}°23\selectlanguage{english}'23\selectlanguage{english}" & 33\selectlanguage{ngerman}°34\selectlanguage{english}'23\selectlanguage{english}" & 4077 & Black-soil beach
meadow\tabularnewline
83 & 99\selectlanguage{ngerman}°54\selectlanguage{english}'31\selectlanguage{english}" & 33\selectlanguage{ngerman}°34\selectlanguage{english}'6\selectlanguage{english}" & 4078 & Eroded mattic epipedon \emph{in K.
pygmaea} meadow\tabularnewline
84 & 99\selectlanguage{ngerman}°25\selectlanguage{english}'34\selectlanguage{english}" & 33\selectlanguage{ngerman}°45\selectlanguage{english}'21\selectlanguage{english}" & 4009 & Black-soil beach
meadow\tabularnewline
85 & 99\selectlanguage{ngerman}°10\selectlanguage{english}'5\selectlanguage{english}" & 34\selectlanguage{ngerman}°18\selectlanguage{english}'16\selectlanguage{english}" & 4219 & Eroded mattic epipedon \emph{in K.
pygmaea} meadow\tabularnewline
86 & 100\selectlanguage{ngerman}°33\selectlanguage{english}'50\selectlanguage{english}" & 32\selectlanguage{ngerman}°46\selectlanguage{english}'35\selectlanguage{english}" & 4232 & Black-soil beach
meadow\tabularnewline
87 & 100\selectlanguage{ngerman}°33\selectlanguage{english}'42\selectlanguage{english}" & 32\selectlanguage{ngerman}°46\selectlanguage{english}'32\selectlanguage{english}" & 4257 & Eroded mattic epipedon \emph{in K.
pygmaea} meadow\tabularnewline
88 & 100\selectlanguage{ngerman}°29\selectlanguage{english}'51\selectlanguage{english}" & 33\selectlanguage{ngerman}°23\selectlanguage{english}'25\selectlanguage{english}" & 3912 & Black-soil beach
meadow\tabularnewline
89 & 100\selectlanguage{ngerman}°29\selectlanguage{english}'36.6\selectlanguage{english}" & 33\selectlanguage{ngerman}°23\selectlanguage{english}'10\selectlanguage{english}" & 3900 & Eroded mattic epipedon \emph{in
K. pygmaea} meadow\tabularnewline
90 & 100\selectlanguage{ngerman}°51\selectlanguage{english}'07\selectlanguage{english}" & 33\selectlanguage{ngerman}°17\selectlanguage{english}'28\selectlanguage{english}" & 4401 & Black-soil beach
meadow\tabularnewline
91 & 100\selectlanguage{ngerman}°51\selectlanguage{english}'7\selectlanguage{english}" & 33\selectlanguage{ngerman}°17\selectlanguage{english}'31\selectlanguage{english}" & 4420 & Eroded mattic epipedon \emph{in K.
pygmaea} meadow\tabularnewline
92 & 98\selectlanguage{ngerman}°55\selectlanguage{english}'57\selectlanguage{english}" & 34\selectlanguage{ngerman}°56\selectlanguage{english}'29\selectlanguage{english}" & 4435 & Eroded mattic epipedon \emph{in K.
pygmaea} meadow\tabularnewline
93 & 98\selectlanguage{ngerman}°0\selectlanguage{english}'40\selectlanguage{english}" & 34\selectlanguage{ngerman}°37\selectlanguage{english}'18\selectlanguage{english}" & 4239 & Eroded mattic epipedon \emph{in K.
pygmaea} meadow\tabularnewline
94 & 98\selectlanguage{ngerman}°0\selectlanguage{english}'56\selectlanguage{english}" & 34\selectlanguage{ngerman}°37\selectlanguage{english}'26\selectlanguage{english}" & 4131 & Eroded mattic epipedon \emph{in K.
pygmaea} meadow\tabularnewline
95 & 98\selectlanguage{ngerman}°4\selectlanguage{english}'20\selectlanguage{english}" & 35\selectlanguage{ngerman}°0\selectlanguage{english}'48\selectlanguage{english}" & 4237 & Black-soil beach meadow\tabularnewline
96 & 97\selectlanguage{ngerman}°59\selectlanguage{english}'6\selectlanguage{english}" & 34\selectlanguage{ngerman}°31\selectlanguage{english}'35.4\selectlanguage{english}" & 4330 & Eroded mattic epipedon \emph{in K.
pygmaea} meadow\tabularnewline
97 & 97\selectlanguage{ngerman}°48\selectlanguage{english}'46.02\selectlanguage{english}" & 34\selectlanguage{ngerman}°13\selectlanguage{english}'42.6\selectlanguage{english}" & 4674 & Black-soil beach
meadow\tabularnewline
98 & 95\selectlanguage{ngerman}°30\selectlanguage{english}'36\selectlanguage{english}" & 34\selectlanguage{ngerman}°4\selectlanguage{english}'48\selectlanguage{english}" & 4428 & Black-soil beach
meadow\tabularnewline
99 & 97\selectlanguage{ngerman}º59\selectlanguage{english}'48.12\selectlanguage{english}" & 34\selectlanguage{ngerman}º36\selectlanguage{english}'20.16'' & 4255 & Eroded mattic epipedon
\emph{in K. pygmaea} meadow\tabularnewline
100 & 96\selectlanguage{ngerman}º41\selectlanguage{english}'42.78\selectlanguage{english}" & 33\selectlanguage{ngerman}º7\selectlanguage{english}'41.28'' & 4373 & Eroded mattic epipedon
\emph{in K. pygmaea} meadow\tabularnewline
101 & 96\selectlanguage{ngerman}º10\selectlanguage{english}'10.68\selectlanguage{english}" & 33\selectlanguage{ngerman}º24\selectlanguage{english}'35.04'' & 4503 & Eroded mattic epipedon
\emph{in K. pygmaea} meadow\tabularnewline
102 & 95\selectlanguage{ngerman}º8\selectlanguage{english}'13.32\selectlanguage{english}" & 34\selectlanguage{ngerman}º43\selectlanguage{english}'12.36'' & 4507 & Eroded mattic epipedon
\emph{in K. pygmaea} meadow\tabularnewline
103 & 94\selectlanguage{ngerman}º49\selectlanguage{english}'47.76\selectlanguage{english}" & 34\selectlanguage{ngerman}º51\selectlanguage{english}'18.6'' & 4518 & Eroded mattic epipedon
\emph{in K. pygmaea} meadow\tabularnewline
104 & 100\selectlanguage{ngerman}º29\selectlanguage{english}'34.02\selectlanguage{english}" & 34\selectlanguage{ngerman}º21\selectlanguage{english}'58.44'' & 3968 & Black-soil beach
meadow\tabularnewline
105 & 99\selectlanguage{ngerman}º14\selectlanguage{english}'8.64\selectlanguage{english}" & 33\selectlanguage{ngerman}º22\selectlanguage{english}'20.28'' & 4077 & Eroded mattic epipedon
\emph{in K. pygmaea} meadow\tabularnewline
106 & 99\selectlanguage{ngerman}º54\selectlanguage{english}'15.6\selectlanguage{english}" & 33\selectlanguage{ngerman}º34\selectlanguage{english}'18'' & 4082 & Black-soil beach
meadow\tabularnewline
107 & 99\selectlanguage{ngerman}º12\selectlanguage{english}'9.84\selectlanguage{english}" & 33\selectlanguage{ngerman}º26\selectlanguage{english}'8.7'' & 4321 & Eroded mattic epipedon \emph{in
K. pygmaea} meadow\tabularnewline
108 & 99\selectlanguage{ngerman}º12\selectlanguage{english}'7.44\selectlanguage{english}" & 33\selectlanguage{ngerman}º26\selectlanguage{english}'7.98'' & 4318 & Black-soil beach
meadow\tabularnewline
109 & 101\selectlanguage{ngerman}º1\selectlanguage{english}'11.22\selectlanguage{english}" & 37\selectlanguage{ngerman}º37\selectlanguage{english}'6.36'' & 3142 & Black-soil beach
meadow\tabularnewline
\bottomrule
\end{longtable}

Sampling was performed in August in 2012-2013 along 100-m transects
within every sampling site that each contained eight evenly distributed
50 cm \selectlanguage{ngerman}× 50 cm quadrats. The plant community was divided into five
functional groups: Gramineae, Cyperaceae, forbs, Leguminosae, and
shrubs. In each quadrat, we recorded the coverage and dry aboveground
biomass of plant functional groups, the thickness of the mattic
epipedon, and the slope aspect, position, and angle. We also noted the
number of livestock grazing at each sampling site (Lin, 2017).

\textbf{1.2 Sample collection and preparation}

The plant samples were placed in an envelope after collection and
subsequently baked at 60--80°C for 48 to 72 h to a constant weight,
which was then recorded.

An unstructured questionnaire was used to determine the area of the
pasture, the type and quantity of grazing livestock, and the grazing
time. According to the conversion ratio of 4 sheep equal to 1 yak, the
stocking capacity of the pasture was calculated (Xu, 1994), and the
stocking capacity was divided into 5 grades. The average grazing
intensities per grazing season were 0.00 sheep/ha for more than 1 year,
less than 3.65 sheep/ha, 3.65 to 7.50 sheep/ha, 8.25 to 13.00 sheep/ha,
and more than 13.00 sheep/ha. In most of the time when the alpine meadow
degradation into Black-soil beach meadow would become to no grazing in
the pasture, so we investigated the possible grazing intensity used in
the past as the average grazing intensity of the pasture.

\textbf{1.3 Data analysis}

The plant functional group characteristics of mini-patches were
determined by calculating the average coverage, dry aboveground biomass,
and importance value of each plant functional group in each mini-patch
(Lin, 2017). The data collected from every mini-patch were also used to
calculate the niche breadths of plant functional groups based on the
Levins index (Lin, 2017), and mean and standard error on the biomass
aboveground, the importance and the niche breadth were calculated by
performing a one-way analysis of variance in SPSS Statistics Software
(19.0). The probability of each mini-patch reaching a certain
multi-steady stage was calculated as the ratio of the environmental
characteristics in a single quadrat to the total in all quadrats. The
coupling between the plant functional groups of all mini-patch and each
environmental factor was calculated via a redundancy analysis (RDA) in
R3.6.1 software.

\textbf{2 Results}

\textbf{2.1 Characteristics of plant functional groups in mini-patches}

The aboveground primary production of alpine meadows depended more on
their steady stage than on their geographical location or on climate
factors. The aboveground primary production was significantly higher
under zero grazing and light grazing intensity than under a heavy
grazing intensity. The aboveground biomass also differed by grazing
intensity. The aboveground biomass of Gramineae, Cyperaceae, and
Leguminosae functional groups significantly decreased as grazing
intensity increased, while the aboveground biomass of forbs, which were
dominant or subdominant in most steady stages, remained relatively
stable across grazing intensities. Nonetheless, the total productivity
of alpine meadows remained stable across a large range of grazing
intensities and a large spatial scale (Fig. 1).\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image1/image1}

\end{center}
\end{figure}

Fig. 1. Aboveground biomass of plant functional groups (PFGs) in
mini-patches at various steady stages

Note: A, B, C, D, and E represent Gramineae--\emph{Kobresia} meadow and
Gramineae--\emph{K.} \emph{humilis} to \emph{K. pygmaea} transitional
meadow; healthy and thickening mattic epipedon in \emph{K.
pygmaea}meadow; cracking mattic epipedon in \emph{K. pygmaea} meadow;
eroded mattic epipedon in \emph{K. pygmaea} meadow and black-soil beach
meadow, respectively; a, b, c, d represent significance of each biomass
of PFG among different steady stages.

\par\null

The importance of plant functional groups in mini-patches could reflect
their ability of resource obtaining positions in the plant community.
The importence of Gramineae, Cyperaceae and Leguminosae gradually
decreased, that of forbs gradually increased, and shrub kept in a
relative stabilization under increasing grazing intensity. The trends in
plant functional groups differed and could be divided into three
successional periods by estimating the importance in each mini-patch. In
the first period, which was under zero or light grazing intensity,
Gramineae dominated and the structure of the alpine meadows was that of
a grazing climax community. The second period was a transition period in
which Cyperaceae was dominance. The third stage was the black-soil beach
stage in which Gramineae and Cyperaceae were minor groups and forbs
became the dominant group in the plant community, and during this
process the Leguminosae had the analogous reaction to Gramineae (Fig.
2).\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image2/image2}

\end{center}
\end{figure}

Fig. 2. Importance of plant functional groups in mini-patches at various
steady stages

Note: A, B, C, D, and E represent Gramineae--\emph{Kobresia} meadow and
Gramineae--\emph{K.} \emph{humilis} to \emph{K. pygmaea} transitional
meadow; healthy and thickening mattic epipedon in \emph{K.
pygmaea}meadow; cracking mattic epipedon in \emph{K. pygmaea} meadow;
eroded mattic epipedon in \emph{K. pygmaea} meadow and black-soil beach
meadow, respectively; a, b, c, d represent significance among different
steady stages.

\par\null

The niche breadths of Gramineae, Cyperaceae, Leguminosae, and forbs
decreased as grazing intensity increased. This succession process could
also be divided into three periods. The first period was represented by
``A'' and ``B'' meadow in which the niche breadths of Gramineae,
Cyperaceae, Leguminosae, and forbs were highest among all of the steady
stages. The second period was the first regime shift point in which the
niche breadth of Gramineae was superseded by the niche breadth of
Cyperaceae (see the ``C'' and ``D'' steady stages). The third stage was
the ``E'' stage in which the niche breadths of Gramineae, Cyperaceae,
Leguminosae, and forbs all greatly decreased. In this process we also
found the shrub seemed to increase with grazing intensity increasing,
that was not because the steady stage underwent to the shrub meadows,
just because the shrub which always used as the summer pasture had more
severe degree degeneration than the alpine meadow used as the winter
pasture (Fig. 3).

Overall, our findings showed that the steady stage of alpine meadows
maintained the highest plant community primary production relatively
constantly under no grazing intensity and under light grazing intensity.
When alpine meadows were disturbed by a high grazing intensity, their
total yield remained stable but the coefficient of variation in yields
increased, which may indicate that change the structure of the plant
community changed under these conditions to maintain a steady total
biomass.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image3/image3}

\end{center}
\end{figure}

Fig 3. Aboveground niche breadth of functional groups in multi-steady
stages

Note: A, B, C, D, and E represent Gramineae--\emph{Kobresia} meadow and
Gramineae--\emph{K.} \emph{humilis} to \emph{K. pygmaea} transitional
meadow; healthy and thickening mattic epipedon in \emph{K.
pygmaea}meadow; cracking mattic epipedon in \emph{K. pygmaea} meadow;
eroded mattic epipedon in \emph{K. pygmaea} meadow and black-soil beach
meadow, respectively; a, b, c, d represent significance among different
steady stages.

\textbf{2.2 Effects of geographical location and climatic factors on the
formation characteristics of mini-patches in alpine meadow}

The RDA revealed that the independent variation of latitude, longitude,
altitude, accumulated temperature above 0 , and accumulation of
precipitation explained 13.29\%, 8.07\%, 9.05\%, 6.44\%, and 2.44\% of
the variation in the biomass distribution in mini-patches of Gramineae,
Cyperaceae, Leguminosae, forb and shrubs, respectively. With regard to
independent variation, there were significant correlations (\emph{P}
\textless{} 0.01) between geographical location (longitude and latitude)
and the characteristics of Gramineae, the accumulation of growing-season
precipitation, and altitude and the characteristics of Cyperaceae, the
accumulation of growing-season precipitation and longitude and the
characteristics of Leguminosae, the accumulation of growing-season
precipitation and altitude and the characteristics of forbs, and
longitude and the characteristics of shrubs (Fig. 4).

As independent variation explained less than 20\% of the total variation
in the plant functional groups in mini-patches, environmental factors
(latitude, longitude, altitude, accumulated temperature above 0\selectlanguage{ngerman}°C, and
accumulation of precipitation in growing season) were considered to have
no power to explain the variation in the plant functional group
distributions (Fig. 4).

The results also revealed that there were no significant correlations
between plant community characteristics in mini-patches and geographical
location, rainfall, and accumulated temperature characteristics. This
suggests that geographical location and climatic characteristics were
not the primary drivers of differences in plant community organization
and structure in our study area (Fig. 4).\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image4/image4}

\end{center}
\end{figure}\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image5/image5}

\end{center}
\end{figure}\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image6/image6}

\end{center}
\end{figure}\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image7/image7}

\end{center}
\end{figure}\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image8/image8}

\end{center}
\end{figure}

Fig 4. Correlations between plant functional groups and micro-topography
and climate

Note: A, F, and K represent the biomass, coverage, and importance value
of Gramineae, respectively; B, G, and L represent the biomass, coverage,
and importance value of Cyperaceae, respectively; C, H, and M represent
the biomass, coverage, and importance value of Leguminosae,
respectively; D, I, and N represent the biomass, coverage, and
importance value of forbs, respectively; E, J, and O represent the
biomass, coverage, and importance values of shrubs, respectively; and P,
Q, R, S, and T represent the accumulated temperature above 0\selectlanguage{ngerman}°C, the
accumulation of precipitation, and latitude, longitude, and altitude,
respectively.

\textbf{2.3 Relationships between micro-topographic factors and steady
stages}

The characteristics of mini-patches investigated in our research show
that 11.1\% could be classified to Gramineae--\emph{K. humilis} and
Gramineae--\emph{K. humilis} to \emph{K. pygmaea}transitional
mini-patches, 16.3\% could be classified to healthy and thickening
mattic epipedon in \emph{K. pygmaea} meadow, 36.2\% could be classified
to cracking mattic epipedon in \emph{K. pygmaea} meadow, 16.6\% could be
classified to eroded mattic epipedon in \emph{K. pygmaea} meadow, and
19.8\% could be classified to black-soil beach meadow (Table 2).

Most of the mini-patches were located on sunny or semi-sunny slopes with
angles of less than 10° or in valleys between hills. Livestock grazing
intensities affected mini-patches differently and the \emph{K.
pygmaea}mini-patches were more degraded if they were located in the
middle of slopes than if they were located elsewhere. Herders typically
use pastures located lower on slopes or in valleys as reserve forage
production areas because their superior climate conditions facilitate
the growth of forage for use in cases of natural disasters or other
emergencies. Therefore, more than 90 \% of the Gramineae and
Gramineae--\emph{K. humilis} mini-patches occurred in such areas (Table
2).

Our findings indicate that as grazing intensity increased, total plant
community coverage decreased. In addition, the total plant community
coverage in eroded mattic epipedon \emph{K. pygmaea} meadows and
black-soil beach meadows were not significantly different but were lower
than that in other types of meadow. In addition, our findings confirm
the generally accepted belief that healthy alpine meadows invariably
maintain a certain thickness of mattic epipedon (typically 4-5 cm)
(Table 2).

Table 2. Micromorphology in alpine meadows under different grazing
intensities\selectlanguage{english}
\begin{longtable}[]{@{}lllllll@{}}
\toprule
& Degree & Proportion of Gramineae--\emph{K.} \emph{humilis} (\%) &
Proportion of Gramineae--\emph{K.} \emph{humilis} and \emph{K. pygmaea}
(\%) & Proportion of \emph{K. pygmaea} (\%) & Proportion of \emph{K.
pygmaea} with eroded mattic epipedon (\%) & Proportion of black-soil
beach (\%)\tabularnewline
\midrule
\endhead
Grazing intensity (sheep / ha) & No grazing for more than 1 year & 39.5
& 0.0 & 0.0 & 0.0 & 0.0\tabularnewline
& \textless{} 3.65 & 60.5 & 0.0 & 32.6 & 0.0 & 0.0\tabularnewline
& 3.65--7.5 & 39.5 & 14.3 & 26.1 & 8.8 & 0.0\tabularnewline
& 8.25--13 & 0.0 & 53.6 & 38.0 & 45.6 & 17.5\tabularnewline
& \textgreater{} 13 & 0.0 & 32.1 & 3.3 & 45.6 & 82.5\tabularnewline
Total plant community coverage (\%) & \textgreater{} 90 & 100.0 & 24.9 &
45.3 & 32.3 & 20.0\tabularnewline
& 70--90 & 0.0 & 16.1 & 18.6 & 21.0 & 17.5\tabularnewline
& 45--70 & 0.0 & 25.0 & 17.5 & 8.1 & 15.0\tabularnewline
& \textless{} 45 & 0.0 & 16.1 & 18.6 & 38.7 & 47.5\tabularnewline
Slope aspect & Sunny & 66.7 & 23.8 & 43.2 & 50.0 & 60.0\tabularnewline
& Semi-sunny & 33.3 & 76.2 & 56.8 & 50.0 & 40.0\tabularnewline
Slope location & Top & 0.0 & 23.8 & 48.1 & 32.0 & 20.0\tabularnewline
& Middle & 33.3 & 76.2 & 36.5 & 56.0 & 60.0\tabularnewline
& Bottom & 66.7 & 0.0 & 15.4 & 12.0 & 20.0\tabularnewline
Slope angle & \textless{} 5\selectlanguage{ngerman}° & 86.8 & 48.4 & 36.4 & 53.5 &
43.2\tabularnewline
& 5\selectlanguage{ngerman}°--10° & 13.2 & 9.7 & 40.9 & 20.9 & 32.4\tabularnewline
& 10\selectlanguage{ngerman}°--35° & 0.0 & 25.8 & 22.7 & 25.6 & 24.3\tabularnewline
& \textgreater{}35\selectlanguage{ngerman}° & 0.0 & 16.1 & 0.0 & 0.0 & 0.0\tabularnewline
Mattic epipedon thickness (cm) & \textless{} 5 & 73.7 & 46.4 & 44.7 &
6.5 & 90.0\tabularnewline
& 5--10 & 26.3 & 53.6 & 38.3 & 23.9 & 0.0\tabularnewline
& 10--13 & 0.0 & 0.0 & 17.0 & 43.5 & 10.0\tabularnewline
& \textgreater{} 13 & 0.0 & 0.0 & 0.0 & 26.1 & 0.0\tabularnewline
\bottomrule
\end{longtable}

\textbf{3 Discussion}

\textbf{3.1 Alpine meadow multi-steady-stage coexistence is a response
and adaptation to grazing disturbance under harsh environmental
conditions and is mainly controlled by grazing intensity}

\textbf{3.1.1 Alpine meadows are characterized by their long grazing
history and extremely fragile environments}

\emph{Kobresia} evolves approximately 240 million years (Shen et al.,
2008; Herzschuh, 2006) and alpine meadow soil began forming on the
Qinghai-Tibet Plateau 4 ka BP as aeolian dust rapidly accumulated in the
Late Holocene and alluvial sediment parent material formed during the
middle Holocene (\textasciitilde{}7--5 ka BP) (Zhang, 2019). Livestock
grazing has been performed for approximately 7,300 years (Simons \&
Sch\selectlanguage{ngerman}ön, 1998\selectlanguage{english};Simons et al., 1994; Georg et al., 2009) and large-scale
livestock husbandry for approximately 3,600 years (Qiang et al., 2016).
Consequently, alpine \emph{Kobresia} meadow has a long history of
livestock grazing, during which both the climate and population
densities have fluctuated (e.g., there was a large population during the
6 AD) (Liao \& Sun, 2003). Thus, alpine meadows on the Qinghai-Tibet
Plateau have the ability to remain stable and recover after repeated
disturbances.

\textbf{3.1.2 Multi-steady stage coexistence is a strategy to resist the
adverse effects of disturbance in a harsh environment}

Many studies have shown that the degradation succession process in
alpine meadows is a typical jump succession process (e.g., Liu et al.,
1999). Variations in livestock grazing intensities, particularly those
resulting from the household contract system, can change the
characteristics (i.e., coverage, height, leave area index) of plant
communities and the strategy or nature of plant community organization
in response to disturbances (Zhang, 1990).

Most alpine meadow plants (e.g.,\emph{K.pygmaea} or \emph{K.}
\emph{humilis} ) propagate asexually (Zhou et al., 1986). In the current
study, it was found that the density of tillers, leaves, and young
shoots increased with a light grazing intensity, this is because the
colonization of the asexual branches begins from the outermost leaf
layer and the spacing between the propagule shortened with increasing
grazing intensity, and the root-soil ratio (R/S) decreases rapidly,
accelerating the imbalance of limiting nutrient in soil supplying and
the plant growth demanding, especially in its asexual branches core area
(Cao et al., 2010), which leads those branches dead and to emerge bare
mini-patches in the centers of\emph{K.} \emph{pygmaea} patches in heavy
grazing intensity \emph{K.pygmaea} or \emph{K.} \emph{humilis} meadows
(Fig. 5)\emph{.}\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image9/image9}

\end{center}
\end{figure}

Fig. 5. Bare area (indicated by the red ellipse) in the propagule of

\emph{K. pygmaea}

As the ratio between aboveground and belowground biomass decreases with
livestock grazing intensity (Cao et al., 2010), grazing can also
stimulate root production, increase soil compactness (Fan et al., 2020),
decrease water infiltration (Dai et al., 2019), increase the density of
dead roots in the core areas of mini-patches, and expand the area of
bare mini-patches (Lin, 2017). Thus, the aforementioned asexual
reproductive strategy of the dominant species can lead to two results
under increasing grazing intensity: increase in bare areas, and the
internodes becoming even shorter as the mattic epipedon further thickens
(Zhu, 2002).

In alpine \emph{Kobresia} meadows, the formation of mattic epipedon is
an adaptive response to livestock trampling and grazing disturbance (Hou
et al., 2004). The thickening of mattic epipedon due to overgrazing can
affect soil nutrient and water availability and thus change the growth
and composition of the plant community. For example, as the niche of
Gramineae-type plants (including Gramineae and Cyperaceae functional
groups) narrows, the resulting empty resource spaces can be occupied by
poisonous or inedible plants that are resistant to trampling and grazing
(Lin, 2017). In this manner, the vegetation could maintain high
productivity during the growing period and reduced the utility of the
pasture for livestock production. This strategy allows a damaged alpine
meadow to recover its functional ecology via a new succession process
and thereby to adapt to overgrazing. Prior research calculated that it
would take 70--80 years for black-soil beach-type grassland to recovery
to\emph{K. pygmaea} meadow in an area with no grazing (Chen, 2016).

An alpine meadow could also maintain relatively stable productivity when
Gramineae had been reduced as the remaining resources were used by
subdominant, poisonous, and inedible species, particularly in heavily
disturbed ecosystems. This leads to different plant communities forming
in the same topographic and climatic environment and ultimately
large-scale multi-stable coexistence (Cao et al., 2007). Therefore, the
degradation of alpine meadows to areas with lower edible grass
productivity is also a self-regulating process that prevents the
occurrence of heavy disturbances.

\emph{K. pygmaea}meadows occur across a wide region of the Qinghai-Tibet
Plateau and can tolerate substantial livestock grazing intensity. This
characteristic was dependent on the steady condition maintained by
Cyperaceae plant functional groups, without which these meadows lost
their utility for livestock grazing. Therefore, \emph{K. pygmaea}meadows
were considered the last line of defense against alpine meadow
degradation and were the key to improving alpine meadow productivity.

\textbf{3.2 Mini-patch patterns in alpine meadows might drive
multi-steady stage coexistence}

The characteristics of different mini-patches in alpine meadow
ecosystems were isolated in space and yet related in function, which
imbued ecosystems with a self-organization ability that enabled them to
maintain overall stability while permitting small-scale variability. The
mini-patches of plant communities in alpine meadows are typically
clustered (Zhang et al., 2003), which results in the formation of
mosaics of patches in which each patch reflects a certain
micro-topography and micro-climate. According to island biogeography
theory, mini-patches can be thought of as ``islands'' and the
surrounding areas can be thought as ``ocean'' (MacArthur \& Wilson,
1967). The stability of this plant mosaic system depends on the species
turnover rate, patch area, and distance between patches (Bueno \& Peres,
2019). While grazing is invariably disturbs to alpine meadow ecosystems,
its effect on their patch mosaics is non-homogeneous, which facilitates
habitat fragmentation (Jessica et al., 2017). The plant mosaic system
therefore resulted in the formation of many types of plant communities
with a range of characteristics at different stages during the
degradation succession process, with these communities serving as the
natural germplasm necessary for the recovery of damaged meadows.
Moreover, the interrelationships and permeability of the mosaic patch
system in alpine meadows can promote small-scale variability while
maintaining system in large-scale stability (Zhang et al., 2003).

The characteristics of alpine \emph{Kobresia} meadows did not appear
continuously in multi-steady stages because of the positive and negative
feedback effects between various ecosystem components. However, the
alpine meadow system is an open system with the ability to self-regulate
in response to external disturbances, which is generally manifested as a
discontinuous effect in the feedback of external disturbances. This
nonlinear superposition effect typically causes a lag around an
equilibrium point or critical value in the response of alpine ecosystems
to disturbances (Liu et al., 1999). In addition, at a certain critical
value in the density of matrix media between mini-patches, infiltration
and transformation occurs that drives the plant community into new
stages (Oborny \& Hubai, 2014). This means that the grazing disturbance
pressure of the grassland ecosystem could show a gradual and continuous
change process, but the point that leads to the regime shift among
different steady stages discontinuous, that is, only when the grazing
pressure reached a critical value, the regime shift should be occurred,
so succession process among different steady stages of alpine meadows
was sudden change, not gradual change.

Alpine meadows have high plant diversity compared to most plant
communities growing in dry and cold regions, and this is also true for
mini-patches. Therefore, the degree of degradation of alpine meadows was
best estimated by determining the frequency or pattern of mini-patches
rather than their overall richness in plant community species.

Human factors have played important role in changing alpine ecosystems
and resources in China over the past 70 years (Yang et al., 2016). The
basic cause of alpine meadow degradation at small spatiotemporal scales
is unsustainable human exploitation, and intense pasture use inevitably
leads to degradation and subsequent reduction in regional economic and
social services (Wang, 2004). Secondary effects such as rodent gnawing
and digging are not initial causes of degradation but arise from the
degradation caused by overgrazing (Zhang \& Fan, 2002). Thus, while
estimating the service function of alpine meadows improves understanding
of mini-patch patterns on such meadows, the substantial problem of
managing the degradation of alpine meadows remains.

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