Journal of Environmental Management 333 (2023) 117446

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Journal of Environmental Management

journal homepage: www.elsevier.com/locate/jenvman

Research article

Diversity and distribution variation of urban spontaneous vegetation with

distinct frequencies along river corridors in a fast-growing city
Xiaopeng Li a, *, Sining Zhang a, Rui Huang a, Li Feng b, Sihui Xu a, Baichuan Liu a
a
School of Architecture, Southwest Jiaotong University, Chengdu, 611756, Sichuan, China
b
Chengdu Park City Construction and Development Research Institute, Chengdu, 610031, Sichuan, China

A R T I C L E I N F O A B S T R A C T

Keywords: River corridors are vital to urban ecology, regulating climate and providing habitats for animals. Spontaneous
Biodiversity conservation plants naturally colonize various spaces therein, where they play important roles. Previous studies have explored
Habitat heterogeneity many factors driving spontaneous plant diversity at the city scale and in specific habitats. However, we lack a
Landscape design and management
holistic understanding of the diversity and distribution of variation of spontaneous vegetation that directly re­
Rare species
flects the effects of urbanization. We conducted a field study of 1250 sample plots along river corridors in
Richness and diversity pattern
Urban spontaneous vegetation Chengdu, a fast-growing city in China, and the spontaneous plants in fifteen microhabitat types were surveyed.
Diversity and distribution patterns were examined for species with distinct dispersal abilities. There was a far
greater richness of occasional species compared to dominant and common species, but occasional species were
markedly less abundant. Vacant lots and natural riverbanks harbored the most species, but revetment walls and
lawns also supported a considerable number of species, especially in the city center. Species diversity among
different urban areas was strongly related to microhabitat conditions. The proportions of dominant, common,
and occasional species varied among communities. In communities on vacant lots of less urbanized areas, the
richness of dominant species was greater, whereas in green spaces created by planted vegetation, occasional
species were more diverse. Green space microenvironments are hospitable to some rare species. Urbanization
and microhabitats have co-effects and thus ultimately determine diversity and distribution patterns. Such pat­
terns, if linked to ecological and ornamental value, can provide a new perspective and nature-based solutions to
urban rewilding and landscape design.

1. Introduction defined as an area characterized by a high degree of self-regulation in

In the past decades, natural bio-resources have declined at a startling
speed. Habitat loss, ecosystem imbalance, environmental change, and
pesticide use during urbanization construction are the prominent causal
factors (Fuller et al., 1995; Fahrig, 2003; Aronson et al., 2014; Ali et al.,
2021; Guo et al., 2021). Into the present day, severe homogenization has
been prevailing in most cities as a consequence of the overuse of culti­
vated exotic flora in urban green spaces (McKinney, 2006; Groffman
et al., 2014). The expansion of urban sprawl is still accelerating. It is
predicted that 68% of the world’s population will live in urban areas by
2050 (United Nations, 2018). The trend in urbanization is unstoppable,
and thus, rewilding in cities and preserving spontaneous vegetation are
considered a critical nature-based solution approved and advocated by
progressively more scholars (Sikorska et al., 2021). Urban wilderness is
ecosystem processes (Kowarik, 2021). Spontaneous plants are essential
components of urban ecosystems as local biodiversity refuges and food
for animals (Robinson and Lundholm, 2012; Deng and Jim, 2017),
pollen sources for pollinators (Larson et al., 2014; Yang et al., 2019), and
regulators of microclimates, surface soil, and water (Del Tredici, 2010;
Seiter and Future Green Studio, 2016). Additionally, they are crucial
urban biological resources themselves, containing abundant species
with various types of flowers, leaves, fruits, and biomass and a partic­
ularly higher proportion of native species compared to cultivars (Li
et al., 2019a). Integrating spontaneous vegetation into urban landscapes
can promote urban rewilding and biodiversity conservation and further
promote the development of eco-friendly cities (Omar et al., 2018; Qian
et al., 2020; Hu et al., 2022). The transformation of attitudes regarding
the perception of “weeds” to “spontaneous plants/vegetation” in China

* Corresponding author.
E-mail addresses: penguinlee26@126.com (X. Li), ZSNing@swjtu.edu.cn (S. Zhang), rain@swjtu.edu.cn (R. Huang), 328268238@qq.com (L. Feng), 774785359@
qq.com (S. Xu), 413595264@qq.com (B. Liu).

https://doi.org/10.1016/j.jenvman.2023.117446
Received 6 September 2022; Received in revised form 28 December 2022; Accepted 1 February 2023
Available online 7 February 2023
0301-4797/© 2023 Elsevier Ltd. All rights reserved.
X. Li et al.
was just beginning in the early 2010s, but this perspective is now widely
accepted and has been studied by scholars at a surprising speed in the
fields of both urban ecology and landscape architecture (Pan et al.,
2022). Many aspects of spontaneous plants remain to be assessed, not
only in terms of academic research but also in terms of practices
applicable to local use and management.
Urban development processes create diverse habitat types (e.g.,
vacant lots, sidewalks, parks, and city walls), which can support spon­
taneous vegetation that is unique to urban sites. Urban plant assem­
blages are shaped by a series of filters, from interspecific interactions to
human-modified landscapes (Cervelli et al., 2013; Bonthoux et al., 2019;
Huang et al., 2019; Li et al., 2019a; Hu et al., 2021). Previous studies
have mostly investigated factors influencing the diversity of urban
spontaneous plants, from the city scale to the microhabitat scale. These
studies indicate that multiple abiotic and biotic factors jointly determine
the diversity and species composition of spontaneous plant commu­
nities, but the major underlying factors differ among distinct habitats. At
the geographic scale, species composition is more affected by climatic
factors (Hu et al., 2022). At the whole city scale, urbanization level, land
use types, and topographical factors explain much of the variation in
species composition (Qian et al., 2020; Hobohm et al., 2021a). The
species diversity of spontaneous plants is positively correlated with soil
depth and negatively correlated with the degree of trampling (Cervelli
et al., 2013). On vacant lots, soil and anthropogenic substrates, such as
concrete, pebbles, sand, and rubble, have more influence in determining
plant species richness, composition, and diversity. The amount of trash
in a lot has also been shown to be a significant variable (Godefroid et al.,
2007; Anderson and Minor, 2018). In an urban park, canopy density,
intensity of disturbance, water conditions, and planted vegetation type
are all critical factors (Li et al., 2019 a; and b). Within urban tree bases,
the biological characteristics of species and the characteristics of tree
planting pools both have an impact on the distribution of spontaneous
plants (Omar et al., 2018). The level of urbanization represented by the
percentage of urban developed area or the distance from the city center
can be primary predictors of species richness and abundance patterns.
The rate and duration of urbanization also impact the extinction rate of
urban plants (Veselkin et al., 2017; Hu et al., 2022). However, despite
our accumulated knowledge about the diversity of spontaneous plants
and the influence of site environmental conditions, the distribution
patterns and assemblages of different species in urban areas remain
scarcely explored. We lack a holistic understanding of how spontaneous
plants are distributed in a city, which could reflect the effects of ur­
banization processes on them.
River corridors support a suite of key ecosystem services that are
important to citizens, including water circulation, flood prevention,
microclimate regulation, recreation, and refuges for flora and fauna
(Ward et al., 2001; Williams et al., 2004; Lerner and Holt, 2012). The
vegetation that grows in the riparian zone forms a riparian corridor and
acts as a buffer between terrestrial and aquatic ecosystems. Various
functions are enhanced by riparian vegetation, such as reducing the risk
of bank erosion and providing shade, shelter, and organic matter for
aquatic organisms (Nobis et al., 2017). Meanwhile, urban construction
along rivers has created diverse microhabitats, providing an excellent
natural experiment for the study of spontaneous vegetation patterns
responding to urban environment. In most studies, urban spontaneous
vegetation is primarily considered as a distinct component that is
examined only through overall species richness; the diversity of species
with different dispersal and reproductive abilities is rarely explored.
Studies indicate that spontaneous plants in urban areas are dominated
by a few widespread species, while a large number of species are
narrowly distributed (e.g., Cervelli et al., 2013; Qian et al., 2020).
Common and rare species are certainly of differing conservation sta­
tuses. Understanding how species with different local levels of preva­
lence contribute to overall patterns of diversity is crucial in conservation
(Gaston et al., 2008; Lennon et al., 2011). Furthermore, species distri­
bution patterns are increasingly important in optimizing restoration
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Journal of Environmental Management 333 (2023) 117446
(Ancillotto et al., 2022; Yang et al., 2022). Integrating data on plant and
habitat distribution patterns has been justified to be useful in optimizing
planting regimes and setting restoration targets (Nobis et al., 2017;
Elliott et al., 2022). Spontaneous plants can find suitable niches in
almost any available gaps and even in inhospitable sites (Cervelli et al.,
2013; Bonthoux et al., 2019). Combining species diversity and distri­
bution patterns with various microhabitats created by urbanization
processes may thus contribute to a better understanding of urbanization
effects on spontaneous plants. Species with extremely limited areas are
expected to have outstanding value in nature conservation (Mouillot
et al., 2013; Hobohm, 2014; Huang et al., 2019), and rare species
contribute substantially to community trait space (Zhang et al., 2022).
Many negative effects of urbanization on plant assemblages have been
reported, especially affecting those sensitive and rare species (Ruas
et al., 2022). Creatively designed landscapes integrating the survival of
species have the potential to alleviate species extinction at a global scale
(Hobohm, 2021a). To achieve this target, a scientific foundation estab­
lishing how urban spontaneous plants with distinct dispersal abilities
can contribute to local plant diversity and responses to urbanization and
microhabitats is particularly essential.
To address the above issue, we conducted a substantial field study to
disentangle the diversity and distribution patterns and the variation
among spontaneous plant species with different frequencies (dominant,
common, and occasional species) under the interactions between ur­
banization development and microhabitats along city river corridors in
Chengdu, a fast-growing city in China and the first proposed “Park City”
as outlined by President Xi Jinping, indicating the objective of it
becoming a more natural, green, ecological, and livable city. Specif­
ically, three hypotheses were tested: (A) urbanization expands the gap
between richness and abundance among distinct species frequency types
(i.e., dominant, common, and occasional species); (B) within commu­
nities, the proportion of different species types vary in their response to
urbanization process; (C) species with distinct dispersal abilities colo­
nize various habitats differently, in a way that is dependent on the
developing features of urbanization. Our results address each of the
hypotheses, allowing us assess the following specific predictions. (1)
There is greater species richness among occasional species, though they
are not more abundant than dominant or common species, and their
richness is lower in highly developed areas. (2) At the community level,
the diversity of dominant species is higher in the city center where oc­
casional species contribute little to diversity, while the diversity of
common species is stable regardless of location. (3) Artificial habitats
support less diversity than natural habitats.
2. Methods
2.1. Study area and selection of sample transects
This study was conducted in Chengdu, Sichuan, China
(102◦ 54′ –104◦ 53′ E, 30◦ 05′ –31◦ 26′ N), which is located in Southwest
China and the west of the Sichuan Basin and is characterized by flat
terrain, a crisscrossing river network, and species richness. It has a
subtropical monsoon humid climate, with an annual average tempera­
ture of 16 ◦ C and an annual rainfall total of about 1000 mm. The city
covers an area of 14,335 km2, and the permanent population was 21.192
million by the end of 2021. There are 70 rivers within the outer ring road
of Chengdu, spanning a total length of about 413 km (Li and Xu, 2018).
Since ancient times, the water resource of Chengdu has inspired its title
“Land of Abundance.” In recent years, urbanization construction has
fragmented the water network; therefore, the existing river corridors
play a prominent role in ecological services. Chengdu is a fast-growing
city in China, and it only recently became a megacity in 2021 (China
National Bureau of statistics, 2021).
The study area is within the Ring Expressway, Chengdu’s main urban
area, spanning a total area of 540 km2. Three river corridors running
through the main urban area were selected as the sample transects,
X. Li et al.
including six important water systems (Fig. 1(a)). The total length of the
three sample transects is about 99.2 km. The scope of the studied river
corridors was defined as follows: for the river channel without any
formal green space, the scope was inside the municipal roads or areas
not more than 5 m from both sides of the river; for rivers surrounded by
green space, the municipal road was used as the research boundary.
2.2. Sampling design
Sampling was conducted using the random line transect method
combined with the typical sample plot method. A sample point was set at
an average interval of about 200 m for each sampling transect according
to the accessibility and abundance of spontaneous plants. The inacces­
sible sample points were not included in the scope of the investigation. A
Global Positioning System (GPS) device was used to locate all the points
for sampling. According to the growing conditions of spontaneous
vegetation, one or more 3 m × 2 m plots were selected near each sam­
pling point. The number of plots increased correspondingly in the sec­
tions with more spontaneous plants. Additionally, to avoid omitting
species, when new species and new microhabitats appeared between
two sample points, they were also recorded. Once a plot was selected,
two quadrats, each with an area of 1 m × 1 m, were set to record
spontaneous plants. The field study was conducted twice, in spring and
summer, respectively. In total, 1257 plots including 625 plots in spring
and 632 plots in summer, were surveyed. They are not strictly over­
lapped because of the varing plant assemblages. To measure the effect of
habitat, plots were classified into three types according to their location
and distance from the river corridors, namely riverfront (FR), revetment
(R), and waterside (W). Each habitat was further classified into different
microhabitat types according to the microenvironment and planted
vegetation, i.e., vacant lot (V), gravel vacant lot (VG), natural riverbank
(R–N), semi-natural revetment (R–SN), revetment wall (R–W), pave­
ment (P), and green space (FRG) (including arbor (A), brushwood (B),
flowerbed (F), lawn (L1), non-gramineous lawn (L2), and multiple layer
communities (M) (Fig. 1(a)). Of these microhabitat types, vacant lot and
natural riverbank are more spontaneous, while revetment wall, pave­
ment, and planted vegetation in green space are more artificial.
To assess the variation in the distribution of spontaneous vegetation
Fig. 1. (a) Study area, sampling transects and the number of plots, and the 15 microhabitat types; (b) the seven zones with different degrees of urbanization.
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Journal of Environmental Management 333 (2023) 117446
in response to urbanization, all plots were divided into one of seven
zones and then into one of three categories based on the degree of ur­
banization, i.e. low, medium, and high (Fig. 1(b)). The zones and cate­
gories within each zone are summarized below.
Zone 1, Low-1 (308 plots), is located in the northwest of the city,
with the lowest degree of urbanization, characterized by semi-rural
countryside with a lower density of residential areas, large areas of
factory district, some ecological area, some farmlands, and undeveloped
lands. Most revetments are concrete and cement, with intermittent
natural riverbanks covered by plants.
Zone 2, Low-2 (260 plots), is in the north of the city, with a lower
density of residents, commercial land, and large patches of green space.
Over half of river transects are natural and semi-natural revetments
mixed with bare soil and bricks, and the others are walls.
Zone 3, Low-3 (94 plots), is located in the northeast of the city, with a
relatively lower density of developed area. Areas adjacent to Dongfeng
canal are covered by agricultural production green space and green
buffer. Nearly half of the revetments are natural riverbanks, while the
other half are walls.
Zone 4, Medium-1 (148 plots), is located in the west of the city,
which is relatively dense in buildings, with small patches of peripheral
ecological area. Around its river corridor are distributed some urban
parks and a longer green belt that are moderately maintained. Natural
riverbank is more common in this zone.
Zone 5, High-1 (164 plots), is located in the city center, with a greater
density of buildings and a municipal road network. Intensively main­
tained green space and a green belt as well as commercial buildings are
around both sides of the river corridor. Most of the revetments are walls.
Zone 6, High-2 (139 plots), is located in the south of the city, char­
acterized by a higher degree of urbanization, with a high density of
residential and commercial lands. The river corridor is surrounded by
intensively maintained green space, and most of the revetments are
walls.
Zone 7, Medium-2 (144 plots), is located in the southeast of the city,
characterized by a moderate density of residential buildings mixed with
industrial areas, medium-sized patches of ecological area, and a few
large parks with intense or extensive maintenance. Revetment walls are
common, but some segments have natural riverbanks.
X. Li et al.
2.3. Data collection
The fieldwork was conducted during the growing season from March
to October 2021. The spring survey was carried out in March and April,
while the summer survey was in late August, September, and October.
All the collected data on spontaneous plants within each 1 m × 1 m
quadrat included species name, abundance (number of individuals),
coverage, and height. Information about the plant family, life form, and
origin of each species was retrieved from the Flora of China (Flora
Committee of Chinese Academy of Sciences, 2013).
2.4. Statistical analyses
(a) To assess α diversity, the species richness and community di­
versity of spontaneous plants were estimated by using the Patrick
index and the Shannon-Wiener index. These values were calcu­
lated in R (v. 4.0.4) (R Core Team, 2021) using the vegan and
spaa packages (Zhang, 2016; Oksanen et al., 2017).
Patrick richness index: R = S, Shannon-Wiener index (Magurran,
∑ at the community level, the diversity of dominant species was actually
2004): H = - Pi(ln Pi).
Here, S is the number of species, and Pi is the relative abundance of
the ith species.
(b) To assess β diversity, the beta diversity of pairs of plots was
evaluated using the Sørensen (1948) index.
S’ = (b + c)/(2 a + b + c)
where a is the number of species shared between two plots, and b and c
are the numbers of unique species occurring in each plot.
(c) The k-means clustering method was performed to classify all
species as dominant, common, or occasional species according to their
frequency. Duncan’s new multiple range test was performed to compare
the diversity and dissimilarity of spontaneous plants between each
microhabitat and zone. In all figures, groups labeled with different
lowercase letters were significantly different at a p < 0.05 threshold. To
clearly show the results, groups with intermediate levels of variation
were not marked. Species composition analyses are presented using al­
luvial plots and chord diagrams, which were generated using the
‘ggplot2,’ ‘ggalluvial,’ ‘statnet,’ and ‘circlize’ packages in R. The pack­
age ‘spatstat’ was used to analyze species and diversity distribution
patterns as well as plot the maps. Non-metric multidimensional scaling
(NMDS) analysis was conducted to assess the community composition
variation among zones and habitats using the ‘vegan’ and ‘yyplot’ R
packages. Unless otherwise indicated, all analyses were conducted in R
(v. 4.0.4).
3. Results
3.1. Species composition, richness, abundance, and distribution patterns
A total of 336 species belonging to 245 genera and 85 families were
identified. Of them, 42 and 35 species were from the families Asteraceae
and Gramineae, respectively, 121 species (36.01%) were annual or
biennial herbs, and 100 species (29.76%) were perennial herbs. These
336 species were clustered into dominant species (frequency >20%; 12
species, 9 native), occasional species (frequency <6.2%; 287 species,
213 native), and common species (frequency between 6.2% and 20%; 37
species, 31 native). About 26% of occasional species were exotic. In
terms of plant life forms, most dominant and common species were
herbs; arbor, fern, and vine also each included a few dominant and
common species. Half of the dominant species were from the families
Asteraceae and Gramineae, and the remaining half were from other
families, but with fewer than eight species per family. Different species
types were relatively evenly distributed across different zones and
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Journal of Environmental Management 333 (2023) 117446
habitats, except for native occasional species, which were more often
accumulated in zones Low-1 and Low-2 and riverfront habitat (Fig. 2).
These results confirmed hypothesis (A).
Distribution patterns revealed that all types of spontaneous plants
were more abundant in Low-1, Low-2, and Medium-1 zones. It was
apparent that the number of individuals of most species in High-1, High-
2, and Medium-2 zones were less numerous, especially in spring. The
abundance of different species types was approximately the same in
most zones, confirming that although occasional species were far more
diverse, they were markedly less abundant. The most frequent native
and exotic species were found almost throughout the whole city, and
native species were normally highly abundant. In contrast, species
appearing only once mostly colonized Low-1 and Low-2 zones, with
small populations. Species with large coverages also appeared more
often in the Low-1, Low-2, and Medium-1 zones (Fig. 3).
3.2. Spatial variation in community diversity and dissimilarity among
different urbanization zones
The results confirmed hypothesis (B), but contrary to the prediction,
not significantly higher in the city center where occasional species can
exhibit even higher diversity in summer. While common species are
polarizing, they tend to be remarkably higher both in the less and more
developed city zones (Low-1 and High-1). This suggests niche differen­
tiation and complementarity between species with different adapta­
tions. Even so, β diversity (community dissimilarity) showed a clearly
tendency to be higher in less developed zones (Fig. 4(a)). Indeed, the
richer species in somewhat rural areas contributed substantially to the
differences among communities.
3.3. The effects of microhabitats and their differential influence in
combination with urban zones
Overall, as assumed, microhabitats explained much of the richness
and diversity of different species assemblages. The natural microhabitats
riverfront vacant lot (FR-V), waterside vacant lot (W–V), and natural
riverbank (R–N) harbored more species. The 12 dominant species
appeared in almost all microhabitats. Exceeding expectations, the arti­
ficial microhabitat revetment wall (R–W) supported 139 species, only 8
species fewer than R–N (Table 1). Lawns (FRG-L1) harbored 35 common
and 96 occasional species. At the community level, species richness,
diversity, and dissimilarity significantly varied among microhabitat
types, and these variables fluctuated with the season. Common and
occasional species were richer in the natural microhabitat gravel vacant
lot (FR-VG and W-VG) and the artificial microhabitat under arbor areas
within riverfront greenspace (FRG-A) (Fig. 4(b)).
Furthermore, in relating microhabitats with urban zones, some
notable patterns were revealed. Common and occasional species were
the most numerous in vacant lots and natural riverbanks of the less
developed zones, while the walls and lawns in the city center and highly
developed zones supported the most occasional species (Fig. 5(a) and
Table S1). At the community level, there were remarkable differences in
both species richness and the diversity of different species assemblages
in distinct microhabitats across zones. In some cases, the species richness
of dominant and common species was far greater than that of occasional
species, for example, in riverfront pavement microhabitats in Medium-1
and High-1 zones, and vacant lot microhabitats in zone Low-1. Never­
theless, in microhabitats created by planted trees, brushwood, and lawns
in zone Low-2 and flowerbeds in zone High-1, occasional species in
communities were notably more diverse (Fig. 5(b) and Fig. S1). Non-
metric multidimensional scaling (NMDS) analysis of data from spring
(Fig. 6) and summer (Fig. S2) showed similarities of plots in terms of
community assemblage among different zones and habitats. Some zones
had large areas of overlap, and habitat types differed in species
composition between common and occasional species. Dominant species
X. Li et al.
Fig. 2. Species composition among different species types, habitat types, and zones. Species type: S1-native dominant species, S2-exotic dominant species, S3-native
common species, S4-exotic common species, S5-native occasional species, S6-exotic occasional species; Habitat type: H1-Riverfront (FR), H2-Revetment (R), H3-
Waterside (W); Zone: Z1-Low-1, Z2-Low-2, Z3-Low-3, Z4-Medium-1, Z5-High-1, Z6-High-2, Z7-Medium-2. The same below.
were highly homogenized across habitats, but some assemblages were
differentiated among specific zones.
4. Discussion
4.1. Response to hypothesis (A): overall richness and distribution pattern
of different spontaneous plant assemblages
Despite their low abundance, occasional species were found to have
far greater species richness, and their abundance was generally lower in
highly developed areas. Overall, unexpectedly, the 336 spontaneous
species found along Chengdu river corridors greatly exceeded the
number in most cities, except for a record of 386 species observed in
Kunming (Gao et al., 2021), another city in Southwest China. The large
fraction of occasional species (accounting for 80% of all the 336 species
in this study) has promoted not only inter-urban but also intra-urban
heterogeneity. One investigation documented 1211 spontaneous
plants in 59 cities in China, and found that at the regional scale, native
and occasional species constitute a high proportion of all species (Hu
et al., 2022). However, an unanticipated difference in abundance among
dominant, common, and occasional species was identified. Even though
the number of occasional species (287) was far greater than the number
of dominant and common species (12 and 37, respectively), their total
abundances were approximately the same (Fig. 3), which reflects the
severe homogenization associated with dominant species. Occupying
the vacant niches left by the declining species, some species, though few
in number, can indeed increase in population size and distribution
5
Journal of Environmental Management 333 (2023) 117446
(Sánchez-Bayo and Wyckhuys, 2019). This phenomenon is more con­
spicuous in more urbanized areas, where occasional species are affected
more severely than in less developed areas.
In terms of the variation in distribution, the most frequently
observed native and exotic species are less affected by urbanization,
with a wide distribution over all sites. In contrast, species that appeared
only once and had greater coverage were more abundant in areas with
less urbanization degrees (Fig. 3). Of course, dominant and common
species in the present study are generally more adapted to the urban
environment. Moreover, many of them are often considered “weeds”
with extensive distributions globally. In contrast, occasional species and
some common species are more often endemic and geographically spe­
cific. For example, Impatiens rhombifolia is naturally distributed in
Chengdu and is endemic to Sichuan Province. Fagopyrum dibotrys is
listed in the “National Key Preserved Wild Plants” (National Forestry
and Grassland Administration, 2021). The low reproductive capacity
and low dispersal ability as well as environmental sensitivity of occa­
sional species generally make them less abundant and more spatially
scarce (Lowe and McPeek, 2014; Vellend, 2016). Notably, urban envi­
ronmental conditions could be a key factor for specific species. For
example, Torilis scabra is rare in urban sites but widely grows in suburbs
and mountain areas across Chengdu as large local populations. Thus,
this species may be accidently introduced into urban areas by human
activities.
X. Li et al. Journal of Environmental Management 333 (2023) 117446

Fig. 3. Distribution patterns of distinct species groups. The 4 most frequent native species: Youngia japonica and Artemisia argyi in spring; Broussonetia papyrifera and
Digitaria sanguinalis in summer. The 6 most frequent exotic species: Trifolium repens, Erigeron sumatrensis and Erigeron canadensis in spring; Alternanthera philoxeroides,
Bidens pilosa and Symphyotrichum subulatum in summer. Species appeared only once: including 58 species in spring and 67 species in summer; Species appeared with
larger coverage (sum ≥100% in two quadrats): 55 species including A. argyi and Ulmus pumila etc.

4.2. Response to hypotheses (B) and (C): co-effects of degree of
urbanization and microhabitat on richness and diversity pattern of
spontaneous plants
Variation in diversity at the community level among different plant
assemblages was not directly associated with the observed urbanization
process. It is widely acknowledged that higher urban management in­
tensity reduces biodiversity (Chollet et al., 2018). However, if urbani­
zation is linked to microhabitats, interesting results can be revealed.
Studies have shown that microenvironmental variables at the local
scale, like soil depth, soil moisture, solar access, and planted vegetation,
have substantial influence on composition and diversity of spontaneous
plants (Godefroid et al., 2007; Fagot et al., 2011; Cervelli et al., 2013;
Omar et al., 2018; Li et al., 2019a). Within the data, it was clear that
peripheral urban sites with a low density of buildings nearby predomi­
nantly had a higher richness of occasional species, mostly associated
with their colonizing of vacant lots (Fig. 5). These sites were less
disturbed than the intensively developed city center, and they are
considerable habitats for biodiversity conservation (Anderson and
Minor, 2018). However, unexpectedly, the community richness and
diversity of occasional species in rural areas within vacant lots were far
lower than those in the areas where more parks are located along the
river transects with diverse planted vegetation (Figs. 4 and 5). It is
thought that rising temperatures influence resource availability, and
increased CO2 concentrations are predicted to affect the water and
nutrient requirements of plants (Duarte, 2007). The heat-island effect of
urban areas can result in higher proportions and abundances of ther­
mophilous species, which generally have strong reproductive and
dispersal abilities (Ruas et al., 2022). Producing large quantities of small
seeds (e.g., Youngia japonica and Artemisia argyi) also confers a selective
advantage in colonizing disturbed sites (Rees and Westoby, 1997).
Therefore, the microclimate of the city center and more harsh wasteland
soil conditions (Shuster et al., 2014; Sharma et al., 2015) in the urban
periphery may increase the dominance of common species in the com­
munity. Conversely, the microenvironment of green space is favorable
for some rare species. Compared to a park in Beijing where sunlight
strongly promoted the diversity of spontaneous plants (Li et al., 2019a),
Chengdu harbors more species that prefer shade and humid environ­
ments. In microhabitats under trees and shrubs, more shade-preferring
occasional species occupy their proper niches. This finding indicates
the necessity and feasibility of creating microhabitats for specific
spontaneous plants in green spaces (Kühn, 2006; Li et al., 2019b).
Moreover, the influence of specific urban conditions on novel wilderness
sites can promote unpredictable community assemblages with novel
mixtures of native and exotic species (Kowarik, 2011). This was
consistent with the results for common and occasional species, as shown
in Fig. 5(b) and Fig. S1. As dispersal and reproductive abilities are
correlated with seed size and competitive advantages can change with
the environmental conditions (Rees and Westoby, 1997; Levine and
Rees, 2002), the proportions of dominant, common, and occasional
species in communities varied substantially. Further study on the spe­
cific influence of environmental variables on species diversity and

6
X. Li et al. Journal of Environmental Management 333 (2023) 117446

Fig. 4. Species richness, diversity and dissimilarity at the community level among: (a) zones and (b) microhabitats. Note: groups labeled with different lowercase
letters are significantly different according to Duncan’s multiple range text (p < 0.05).

community assembly in southwestern regions of China is merited.

Table 1
Thus, the proportion of different species types varied in their
Numbers of species of different types in each microhabitat.
response to urbanization, in support of hypothesis (B), such that at the
Microhabitat Plot Dominant Common Occasional Total community level, species richness and diversity in artificial habitats
number species species species species
created by planted vegetation can be significantly higher than that in
FRG-A 27 12 25 55 92 natural habitats. In terms of total richness, natural habitats indeed
FRG-B 21 12 23 50 85
support more species, but the artificial habitats also harbored remark­
FRG-F 40 12 33 71 116
FRG-L1 138 12 35 96 143
able numbers of species. This was attributed to habitat heterogeneity
FRG-L2 104 12 34 86 132 along the river corridors, which demonstrated that for spontaneous
FRG-M 48 11 27 60 98 plants, urbanization development should not be evaluated as a
FRG-V 61 12 33 75 120 straightforward negative factor. Distinct microhabitats support different
FR-P 38 12 24 41 77
species and community assemblages, and the habitat composition varies
FR-V 307 12 37 187 236
FR-VG 11 10 24 24 58 among different zones in Chengdu. In the highly urbanized areas, arti­
R–N 93 12 37 98 147 ficial lawns occupy a large proportion and coverage. In this study, the
R–SN 42 11 31 54 96 number of species observed in lawn microhabitats therein was the
R–W 165 12 34 93 139
highest among all zones (Fig. 5(a) and Table S1). In contrast, lawn mi­
W–V 154 12 35 113 160
W-VG 8 9 24 19 52
crohabitats have been found to support low diversity by many scholars.
However, if maintained and managed according to an appropriate
regime, this type of microhabitat can harbor spontaneous plants with

7
X. Li et al.
Fig. 5. (a) Plot and species number of different species types in all microhabitats in each zone. (b) Co-effects of zone and microhabitat on species richness and
diversity at the community level in spring.
high richness and plant assemblages that differ from those of vacant lots.
Owing to human intervention, lawns select species that are tiny, tough,
and can survive mowing (Li et al., 2019a); thus, innovative rewilding of
lawns is a key objective (Ignatieva and Hedblom, 2018). Similarly, the
number of species found in revetment wall microhabitats (139) was not
remarkably lower than that in natural riverbanks (147) because of the
large proportion of walls among all the river transects leading to more
colonizing space (Lomolino, 2000) (Table 1). In addition, the capacities
of the microhabitats to support diversity are dependent on various fac­
tors. For example, in the city center with a longer history of develop­
ment, the revetment wall has established a moister and more stable
environment; thus, the richness and diversity of spontaneous plants
were not significantly lower than those of the same habitat in less
8
Journal of Environmental Management 333 (2023) 117446
developed zones. This is consistent with previous studies indicating that
urban walls can be reliable refuges for spontaneous plants (Huang et al.,
2019; Chen, 2020). Nonetheless, at the community level, the richness
and diversity of spontaneous plants in lawn and revetment wall micro­
habitats were exceedingly lower than those in other microhabitats,
especially in spring (Fig. 4(b)). This finding still underlines the impor­
tance of preservation and construction of habitats in more natural ways.
By contrast, although gravel vacant lots were few in number, they
effectively supported biodiversity. This can be attributed to the fact that
gravel habitats near rivers are effective for biodiversity restoration,
producing heterogeneity and creating ideal conditions for diverse spe­
cies (Jepson and Blythe, 2021).
Moreover, it was confirmed that variables associated with the
X. Li et al.
Fig. 6. Non-metric multidimensional scaling (NMDS) analysis of different species types in different zones and habitats in spring.
landscape composition subgroup were more influential on community
structure within the riparian zone. The adjacent land cover and land­
scape pattern of a river is a pivotal factor, and fine-scale adjacency ef­
fects are stronger (Fernandes et al., 2011; Chen et al., 2017). For
example, in this study, the lower species abundance and diversity of
Low-3 and Medium-2 zones, which also included more vacant lots, could
be attributed to a wide swath of hard revetment in parallel with cement
roads that are disturbed more frequently by people. The smaller width of
the vacant lot patches between the river corridor and roads is also
influential. Such landscape patterns reduce the habitat area and frag­
ment potential habitats of spontaneous vegetation. The species richness
and diversity patterns of spontaneous plants in this study are deeply
shaped by landscape composition at a broad scale and
9
Journal of Environmental Management 333 (2023) 117446
microenvironment at a fine scale.
4.3. Strategies for conservation and landscape design of spontaneous
vegetation along river corridors
Nature-based solutions that take advantage of naturally occurring
processes to maximize the provisioning of ecosystem services have been
increasingly recognized (Sikorska et al., 2021). A series of spontaneous
vegetation distribution maps are essential in future landscape design to
promote the growth of natural areas. Thus, these maps can be powerful
tools for the conservation and optimization of spontaneous vegetation in
cities. The distribution pattern of occasional species (particularly the
rare species that appeared only once) can be used to identify sites with
X. Li et al.
key microhabitats and species to be protected, as well as their com­
panion species. For example, the species Veronica polita and Maesa
japonica appeared only once each in the city center, and they colonized
lawns, which could be protected at these sites and elsewhere (Fig. 3).
The species appearing only once are mostly scarce in urban sites across
China, such as Delphinium anthriscifolium, Sagina japonica, Cynanchum
auriculatum, and Eragrostis ferruginea. Many of these species specifically
have high ornamental value, with eye-catching flowers. Although each
occasional species exhibits an extremely restricted and narrow distri­
bution, they overall contribute to the highly heterogeneous biodiversity
across Chengdu. One strategy to mitigate homogenization that was
previously neglected in urban greening is the protection and propaga­
tion of occasional spontaneous plants. Furthermore, rare species have
interspecific relationships with common species in communities. For
example, they share pollinators whose food resources are primarily
provided by common plants. Consequently, the long-term survival of
rare species very likely depends on the more common species. Hence
conservation management should also consider the associated common
species (Gibson et al., 2006). Many detailed interspecific interactions
merit further assessment.
Exceptional green spaces that combine novel wilderness with design
interventions are simultaneously effective in supporting ecosystem ser­
vices and attracting visitors (Kowarik, 2018, 2021). Integrating multiple
patterns can be used to develop tailored planting schemes and target
ecosystem conditions that are specific to a given site, hence maximizing
biodiversity and economic outcomes (Elliott et al., 2022). Specifically,
in this study, when multiple microhabitats overlapped, we found that
combinations of waterside habitat and natural riverbank were more
effective in supporting biodiversity than either microhabitat alone.
Linear landscape elements (e.g., drainage ditches), fragile zones, and
buffer zones along rivers need to be distinguished and improved (Donath
et al., 2003; Wynhoff et al., 2011). Furthermore, designing and main­
taining habitat mosaics with different structures in wilderness areas is
important (Planchuelo et al., 2019; Kowarik, 2021). A new heteroge­
neous anthropogenic habitat in a landscape can promote biodiversity
(Hobohm, 2021b; Li et al., 2019a). For example, in this study, species
colonizing areas of planted vegetation can be primarily clustered into
four different ecotypes: tiny lawn-adapted species, shade-loving species
found under arbor or brushwood areas, cultivar -competitive species,
and generalists. These specific species types are of critical value in the
planning and design of community structures. Notable ecologist Aldo
Leopold stated that “no tract of land is too small for the wilderness idea”
(Diemer et al., 2003). As such, it is clear that the protection and
enhancement of spontaneous plants in small-sized microhabitats like
vacant lots and pavement cracks will be meaningful (Kowarik, 2011,
2021; Bonthoux et al., 2019; Li et al., 2019a and b). Moreover, although
great areas of overlapping state space can be observed in the NMDS
plots, there is still divergency in community assemblages between each
zone and each microhabitat (Fig. 6 and Fig. S2). A more rational allo­
cation of various microhabitats to increase heterogeneity could be
achieved by intentionally propagating assemblages with higher speci­
ficity. For example, the water-loving species Echinochloa caudata and
Epilobium hirsutum have high ecological and ornamental value, but were
very rare. Breeding these species and intentionally sowing their seeds in
a similar microhabitat would be helpful to expand their distributions.
Last but not the least, maintenance practices have been proven to be a
key driver of species diversity; hence, intentional policy innovation and
well-designed maintenance interventions that balance various objec­
tives and metrics are critical (Rupprecht et al., 2015; Phillips and
Lindquist, 2021). For example, spontaneous plants that are invasive or
undesirable can be removed (Del Tredici, 2014).
5. Conclusion
This study focused on the variation in diversity and distribution of
spontaneous plants occurring with different frequencies along river
10
Journal of Environmental Management 333 (2023) 117446
corridors running through distinct zones of Chengdu, China. Our find­
ings corroborate the broad hypothesis that interactions between urban
development and microhabitat composition have shaped novel species
richness, diversity, and distribution patterns, as well as stochastic
community assemblages. Specifically, urbanization expands the gap
between richness and abundance among dominant and occasional spe­
cies, and at the community level the proportion of different species types
vary in their response to urbanization process. Spontaneous plants
colonize various habitats differently, and artificial microhabitats can
also support diverse species. Some key problems were revealed, such as
the severe homogenization caused by dominant and common species.
Meanwhile, some optimistic findings were preliminarily demonstrated,
e.g., the considerable value of planted vegetation in green space,
revetment walls, and lawns in providing refuge to some occasional
species, particularly in highly urbanized areas. Together, these patterns
can inform robust and quantitative plans for spontaneous vegetation
conservation and urban rewilding that are adaptable to microhabitat
conditions and, therefore, can effectively improve the biodiversity of
urban ecosystems, not only along river corridors (Ancillotto et al., 2019;
Huang et al., 2021).
Credit author statement
Xiaopeng Li: Conceptualization, Methodology, Software, Investiga­
tion, Data curation, Writing- Original draft preparation. Sining Zhang:
Visualization, Writing- Reviewing and Editing. Li Feng: Supervision,
Validation. Rui Huang: Supervision, Validation. Sihui Xu: Software,
Investigation. Baichuan Liu: Software, Investigation.
Funding
This work was supported by the National Natural Science Foundation
of China (NSFC) (Grant No. 52108065), the Fundamental Research
Funds for the Central Universities, China (Grant No. 2682021CX095),
Park City Project of Chengdu Landscape Architecture Planning and
Design Institute (Grant No. R114620H01038) and Natural Science
Foundation of Sichuan Province (Grant No. 2022NSFSC1108).
Declaration of competing interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influence
the work reported in this paper.
Data availability
Data will be made available on request.
Acknowledgements
The authors would like to acknowledge Norbert Kühn and Ingo
Kowarik (Technische Universität Berlin) for their advices in landscape
design incorporating of spontaneous vegetation. Thanks to the valuable
comments of all the reviewers.
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://doi.
org/10.1016/j.jenvman.2023.117446.
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