Journal of Hydrology 625 (2023) 130132

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Journal of Hydrology
journal homepage: www.elsevier.com/locate/jhydrol

Research papers

Analysis of the suitable ecological flow of benthic animals in the lower

reaches of Xiangjiaba Reservoir in the upper reaches of the Yangtze River
based on the physical habitat model
Dingxin Chen a, b, Zefan Yang b, *, Qinghui Zeng b, *, Weize Wang c, Long Yan b, Pu Zhang b,
Xinyu Li b, Peng Hu b, Hao Wang b
a
Tianjin Univ, State Key Lab Hydraul Engn Simulat & Safety, Tianjin 300350, Peoples R China
b
State Key Laboratory of Simulation and Regulation of Water Cycle in River Basin, China Institute of Water Resources and Hydropower Research, Beijing 100038, China
c
State Key Laboratory of Eco-hydraulics in Northwest Arid Region, Xi’an University of Technology, 710048, China

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

Keywords: Abundant water resources have facilitated the completion and operation of numerous large-scale water con­
Water Environmental Model servation projects. However, the operation of cascade hydropower stations has changed the natural hydrological
Decision-making Model regime of rivers, thereby affecting the integrity and stability of downstream benthic animal habitats. To quan­
Model coupling
titatively evaluate the impact of infrastructure development, we proposed a coupling of hydrodynamic and
Ecology, Hydroelectricity
Sustainability
habitat models, with benthic animals as indicator species (because they are highly sensitive to hydrological
changes). This study established a physical habitat model of benthic animals in the upper reaches of the Yangtze
River, from the Xiangjiaba to Zhutuo sections by coupling the MIKE21 hydrodynamic model with a habitat
model. This model coupled the changes in hydrological conditions and ecological processes to quantitatively
reflect the effects of flow changes on river habitat quality, which can be used to predict the effects of hydrological
changes on water ecosystems and provide fundamental information for ecological flow and scheduling decisions.
Our results showed that by jointly regulating the suitable outflow of the Xiangjiaba Reservoir and the inflow of
the Tuojiang, Minjiang, and Hengjiang rivers (three tributaries with water conservancy projects), the minimum
flow in the non-flood season and the maximum flow in the flood season could be controlled more precisely. This
increases the suitable habitat area for benthic animals as well as improves the stability of their habitat. This study
fills the gap in the physical habitat model of benthic animals in the upper reaches of the Yangtze River as well as
enriches the research ideas of ecological flow regulation. Moreover, this indicates the first application of the
physical habitat model to the joint ecological flow regulation of the main tributaries, thus opening new directions
for the application of the physical habitat model.

1. Introduction
Hydroelectric stations, which convert hydropower to electricity, can
yield many socioeconomic benefits; however, they can also transform
rivers into lake-like reservoirs, change downstream hydrological con­
ditions, and threaten the availability of suitable habitats for various
aquatic organisms (Eloranta et al., 2018). The upstream section of
China’s Yangtze River hosts the richest hydropower resources in the
entire Yangtze River Basin and has four giant cascade hydropower sta­
tions (Wudongde, Baihetan, Xiluodu, and Xiangjiaba) that have already
impounded water for power generation (Dai et al., 2020). The upper
Yangtze River is fed by large tributaries with a basin area of 50,000 km2,
including the Jialing, Minjiang, Yalong, and Wujiang rivers. In addition,
the Yangtze River flows through the Qinghai-Tibet Plateau, Hengduan
Mountains, Yunnan-Guizhou Plateau, Sichuan Basin, and other areas
with complex topographies and geomorphologies (Xu, 2006). Overall,
because of its widespread and connected nature, the ecological integrity
of the water in the upper reaches of the Yangtze River is affected by its
associated water conservancy projects as well as by the hydrological
conditions and topography of its tributaries (Chen et al., 2019). To
further reduce the impact of hydropower projects on the ecological
integrity of habitats in the upper reaches of the Yangtze River, the

* Corresponding author.
E-mail addresses: yangzefan@126.com, qhzeng1990@126.com (Q. Zeng).

https://doi.org/10.1016/j.jhydrol.2023.130132

Available online 11 September 2023

0022-1694/© 2023 Elsevier B.V. All rights reserved.
D. Chen et al.
impacts of these projects on downstream hydrological conditions and
water ecology should be accurately assessed and forecasted.
Habitat area is an ecological indicator that reflects the living space of
benthic animals. Owing to the construction of hydroelectric projects, the
areas encompassing these habitats have begun to shrink, weakening the
continuity and integrity of the surrounding habitat (Takada et al., 2019).
Regarding habitat suitability, benthic animals are the most sensitive to
changes in hydrological factors. The flow demand of benthic animals is
approximately 20% greater than that of fish (Li et al., 2011). The role of
benthic animals in rivers and lake ecological networks has recently
garnered more attention within the context of research (Pešić et al.,
2018）.As the natural food for various fish, the benthic animals can
effectively reflect the changes in the habitat of fish and other species in
the river through the flow and material circulation of the food web，
However, few studies have applied these insights to river ecological flow
decision-making（Sharapova et al., 2019). Therefore, studying the
interrelationship between hydrological parameters and the habitat area
of benthic animals will allow for a quantitative assessment of the effects
of hydrological changes on the ecological integrity of their habitat as
well as enrich our understanding of joint ecological flow decisions in the
mainstream and its associated tributaries.
The water flow required to maintain the ecological integrity of an
aquatic habitat is known as the ecological flow. The maintenance of
ecological flow, as the term suggests, requires the continuous flow of
water as well as a fluctuation in the flow amount across different sea­
sons. The protection of ecological flow is the basis for protecting the
river’s habitat area (Tang et al., 2018; Yang et al., 2019). Calculation
methods for ecological flow can be divided into four categories:
ecological hydrology, ecological hydraulics, habitat simulation, and
overall analysis. The ecological hydrology method is based on measured
hydrological data combined with a specific proportion or percentage
quantile to determine the minimum or appropriate ecological flow.
Commonly used methods include the Tennant, 7Q10, annual distribu­
tion, and flow duration curve (FDC) methods, which combine aquatic
biological data with river flow and judge the characteristic flow of a
river by building a relationship between hydraulic parameters and flow.
Common methods include the wet perimeter and R2CROSS methods,
which combine hydrodynamic models with the life history of specific
aquatic organisms at different stages and quantitatively describe the
relationship between flow and habitat area to determine the ecological
flow of a species at a specific life stage. Furthermore, PHABSIM, Riv­
er2D, and physical habitat models are based on river ecological systems
theory to analyze the ecological flow from different angles by inte­
grating the above methods, such as the BBM, DRFT, and ELOHA
methods. Among various methods for calculating ecological flow,
habitat simulation can simulate the complexity of river ecosystems with
the highest accuracy among all types of methods and has become an
important tool for river management (Armour and Taylor, 1991; Bock­
elmann et al., 2004). Ecological flow is the volume of water required to
maintain the ecological integrity of an aquatic habitat. This mainte­
nance necessitates continuous water flow and fluctuations in the flow
amount across different seasons. Protecting the ecological flow is crucial
for safeguarding river habitats (Tang et al., 2018; Yang et al., 2019).
Methods for calculating ecological flow can be classified into four cat­
egories: ecological hydrology, ecological hydraulics, habitat simulation,
and overall analysis. Ecological hydrology uses measured hydrological
data combined with specific proportions or percentage quantiles to
determine the minimum or appropriate ecological flow. Commonly used
methods include the Tennant, 7Q10, annual distribution, and flow
duration curve (FDC). Ecological hydraulics combines aquatic biological
data with river flow to establish a relationship between hydraulic pa­
rameters and flow and determine the characteristic flow a river should
have. The wet perimeter and R2CROSS methods are the most common.
Habitat simulation methods employ hydrodynamic models and the life
histories of specific aquatic organisms at various stages to quantitatively
describe the relationship between the flow and habitat area, thereby
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Journal of Hydrology 625 (2023) 130132
determining the ecological flow of a species at a particular life stage and
the common methods include the PHABSIM, River2D, and physical
habitat models. Based on river ecological systems theory, the overall
analysis method integrates the aforementioned methods to analyze
ecological flow from various perspectives, such as the BBM, DRFT, and
ELOHA methods. Among these methods, habitat simulation can simu­
late the complexity of river ecosystems with the highest level of accuracy
and has become a critical tool for river management (Armour and
Taylor, 1991; Bockelmann et al., 2004).
Among the various methods of habitat simulation, the PHABSIM
model (Bovee, 1986) was the first fish habitat model and is currently
widely used worldwide. However, these models cannot be combined
with habitat and hydraulic models. The RIVER 2D model, a commonly
used commercial model, can perform detailed hydraulic analyses of
habitat units at the micro-level (Steffler and Waddle, 2002); however, it
cannot target benthic species. In contrast, physical habitat models can
establish ecological flow targets for various target species, maximizing
habitat suitability (Byungwoong Choi et al., 2015). Consequently,
considering various factors, we chose the physical habitat model method
for ecological flow analysis in this study.
The physical habitat model is rooted in an In-stream Flow Incre­
mental Methodology (IFIM). This framework allows the accurate eluci­
dation of habitat area dynamics within the context of a target species
under different ecological flow conditions, primarily by coupling the
hydrodynamic model with the habitat model (Volchek et al., 2019). This
method entails the use of a target species’ habitat preferences (within
the context of flow rate, water depth, substrate, and cover) and then
characterizes the required ecological flow of a target species as the
relationship between flow and the weighted suitable habitat area (Cai
et al., 2010). The accuracy of the physical habitat model lies in its ability
to consider both the physical habitat and ecological flow requirements
of the target species across different timescales (Oyague et al., 2020);
this is the main reason for the widespread use of this analytical approach
in the assessment of the ecological health of rivers (Choi et al., 2018;
Kim et al., 2015; Sedighkia et al., 2021; Wang et al., 2019). However,
most studies on ecological flow target only a single river segment;
therefore, studies considering the joint decision-making of the main­
stream and its associated tributaries are not common, limiting our
ability to integrate the regulation of the mainstream and its tributaries
based on the consideration of ecological factors (Nestler et al., 2019).
The region from the Xiangjiaba Dam to Zhutuo Town in the upper
reaches of the Yangtze River in China consists of the Minjiang, Heng­
jiang, Chishui, and Tuojiang rivers. This region lies in the Yunnan-
Guizhou Plateau, Sichuan Basin, and its combined area. Focusing on a
typical river section from the Xiangjiaba Dam to Zhutuo Town, this
study aimed to elucidate the mechanisms underlying the relationship
between ecological flow and habitat suitability. This was performed to
generate a system that could assess the ecological integrity of water
based on the “quantification of the target species’ demand, habitat
simulation assessment, and ecological flow decision optimization” using
a physical habitat model (Cai et al., 2010). First, to achieve these aims, a
suitability index was applied to analyze the monitoring data of benthic
animals in several sections of the mainstream and tributaries of the
Yangtze River. This was performed to characterize the habitat prefer­
ences of the target species. Second, a two-dimensional hydrodynamic
model was established to simulate the spatiotemporal distributions of
habitat flow velocity and water depth. Finally, the distribution of suit­
able habitats for the target species across the different flow rates was
determined. Based on these findings, the relationship between discharge
and suitable habitat areas was analyzed. Additionally, through the lens
of tributary confluence and topography, we simulated changes in the
suitable habitat area of the target species after the impoundment of
cascade dams across different watershed sections. Finally, we combined
the joint regulation of the ecological flow of the mainstream and trib­
utaries to facilitate ecological flow decision-making.
D. Chen et al.
2. Materials and methods
2.1. Overview of the study area
The study area extends from the Xiangjiaba Dam to Zhutuo in the
mainstream of the Yangtze River, where the Xiangjiaba Dam is the last
hydropower station dam. Xiangjiaba is a humid region with annual
rainfall above 897.1 mm. From 1951 to 2021, the annual runoff near
Xiangjiaba was 147 billion m3, with a multi-year average flow rate of
4661 m3/s. The highest and lowest navigable downstream water level of
Xiangjiaba was 277.25 and 265.8 m, with corresponding flow rates of
12,000 (outflow) and 1200 m3/s (outgoing flow), respectively. The
study area was divided into five river sections: Section 1 was from the
Xiangjiaba Dam to the Hengjiang River; Section 2 was from the Heng­
jiang River to the Minjiang River; Section 3 was from the Minjiang River
to the Tuojiang River; Section 4 was from the Tuojiang River to the
Chishui River; and Section 5 was from the Chishui River to Zhutuo Town
(shown in Fig. 1). Three sampling sites were established in each section
as follows: the downstream side of the Xiangjiaba Dam, Yibin, Lizhuang
Town, Luzhou, and Zhutuo Town. Six sets of data were collected twice,
culminating in a total of 180 sets. (see Fig. 2.).
2.2. Sampling methods and data collection
This study collected and monitored data on benthic animals and the
environmental parameters associated with their preferred habitats
across five river sections before the 2021 flood season (June) and at the
end of the 2021 flood season (October). The collected data were then
quantitatively analyzed to determine the habitat preferences of the
target organisms. The benthic animals were collected using a Surber net.
Six samples were randomly collected within a 50-m radius of the
respective river sections. Although the sampling was random, we
ensured that these samples accurately represented a wide array of
habitats occupied by benthic animals in this region; the collection area
for each sample was 1 m2, whereas the total area representing all the
Fig. 1. Benthic animals samples in the downstream section of Xiangjiaba Dam.
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Journal of Hydrology 625 (2023) 130132
sampling efforts was 6 m2.
Water temperature was recorded at each sampling point. The
maximum difference in water temperatures between the two samples of
the five river sections was only 2.1 ◦ C, and the temperatures between the
two samples were highly consistent. Therefore, the effect of temperature
on the biomass of benthic animals was not considered in this study. The
collected samples were washed at the collection site to remove impu­
rities. After repeated washing, the samples were placed in collection
bottles containing 5 mL formalin. All samples were then taken to the
Institute of Hydrobiology, Chinese Academy of Sciences laboratory for
identification, counting, and weighing (the wet weight of the benthic
animals was converted to dry weight).
In order to eliminate the interference of water quality factors, a 5L
water sampler was used in the study. Three bottles of water were taken
repeatedly at each sampling point, and the water samples were collected
and sealed before being sent to the Beijing Institute of Analytical
Technology for water quality analysis. The analysis results showed that
the water quality status at each sampling point was above Class II, and
the impact of water quality on benthic animal habitats could be ignored.
The flow boundary and topographic data used in the hydrodynamic
model were obtained from the Institute of Water Resources of the China
Institute of Water Resources and Hydropower Research; the survey was
conducted in 2020. Benthic animals and their preferred environmental
parameters (water depth, flow velocity, and substrate) must be moni­
tored simultaneously. For example, the water depth can be determined
using a HY.HSW-1000 digital portable ultrasonic sounder (Weifang
Jinshui Huayu Information Technology Co., Ltd., Weifang, Shandong
province, China), whereas the flow velocity can be determined using the
HOH-L-01 Doppler flow profiler (Beijing Jinshuizhongke Technology,
Beijing, China). Once acquired, these data were used to model the
suitability indices for the target species. These indices were used to
construct hydrodynamic models and determine the ideal ecological flow
for the target organisms.
D. Chen et al.
Fig. 2. Hydrodynamic model calibration and verification results (Lizhuang).
2.3. Physical habitat modeling
The physical habitat model consisted of hydrodynamic and habitat
models. The main premise of this model is that aquatic organisms in
river ecosystems can be characterized based on hydrodynamic factors,
as their habitat preferences are primarily rooted in seeking habitats
within the best hydrological conditions (Shan et al., 2022; Zhang et al.,
2021).
2.3.1. Build the hydrodynamic model
Habitat environmental parameters were simulated by applying the
MIKE21FM module in the Danish DHIMIKE software to build a two-
dimensional hydrodynamic model of the river section below the
Xiangjiaba Dam (Krylenko et al., 2018). This was performed to simulate
the spatiotemporal distribution of environmental conditions, such as
habitat flow velocity and water depth. In this study, the Manning coef­
ficient parameter (Abdi and Saghebian, 2021) of the model was deter­
mined to be 0.2789 using water level data from Lizhuang, which was
obtained in the field in 2020. The average error of the average water
level was 0.276 m, whereas that of the Nash-Sutcliffe efficiency coeffi­
cient was 0.967 m. These results demonstrate the high quality and
reliability of the model.
2.3.2. Build habitat model
In this study, the Habitat Suitability Index (HSI) was used to quantify
the data collected on biomass and environmental parameters to derive
the HSI and suitability levels for different ranges of environmental pa­
rameters. The underlying assumption of the HSI is that species select
sites that best meet their habitat needs, with the highest quality habitats
being the most frequently used sites. Therefore, the HSI is calculated as
follows(Ban et al., 2020):
HSI = mi /m （ 1 ） between the measured target species biomass and the potential impact
where mi represents the total biomass of benthic animals observed
within different environmental parameter ranges, and m represents the
total biomass of benthic animals in the study area.
This study synthesized the habitat needs of different genera and
species of benthic animals and drew a habitat suitability curve based on
the HSI value of the total biomass of benthic animals under changes in
environmental parameters. The exponent of the curve ranged from 0 to
1, with 0 and 1 representing habitat conditions that are completely
unsuitable for the target species and the most suitable ones, respectively.
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Journal of Hydrology 625 (2023) 130132
2.3.3. Physical habitat model coupling
According to the simulation results of the hydrodynamic model, the
water depth (D) and flow velocity (V) of each unit can be derived, and
these values correspond to the habitat suitability index (HSI) on the
suitability curve of the habitat model (Dabin, 2019; Ban, X, 2020). The
comprehensive habitat suitability index (CHSI) was calculated based on
the comprehensive impact of various environmental parameters. The
calculation formula is as follows(Yang et al., 2020):
√̅̅̅̅̅̅̅̅̅̅̅̅̅̅̅̅̅̅̅̅̅̅̅̅̅
CHSI = HSI1 × HSI2 （ 2 ）
where HSI1 is the suitability index corresponding to the water depth,
and HSI2 is the fitness index corresponding to each flow rate.
The distribution of the habitat HSI corresponding to each environ­
mental parameter can be obtained.
Habitat ecological flow decisions are based on the relationship be­
tween flow (Q) and the Weighted Usable Area (WUA), which provides a
quantifiable method for making decisions on water ecological sched­
uling by instantly and visually reflecting changes in the WUA of target
species at different flows in the form of curves and tables. The WUA can
be programmed using the following equation(Jianhua L et al., 2023):
∑
n
WUA = ΔMi × CHSIi （ 3 ）
i=1
Where ΔMi is the area of each grid cell, and CHSIi is the composite
suitability index corresponding to that grid cell.
This process was repeated for each simulated flow, and the rela­
tionship curve between the target Q and the WUA was obtained. Com­
bined with the Q-duration curve (Q-t), a corresponding WUA value was
found from the Q-WUA relationship curve for each Q, and the suitable
habitat area duration curve (WUA-t) was finally obtained. Based on the
assumption of higher species biomass for larger suitable areas, a link
of flow change processes on the target species habitat can be established.
3. Results
3.1. Habitat biological demand analysis
3.1.1. Sampling results of benthic animals
Correlations between the biomass of the target organisms (benthic
animals) and the environmental parameters associated with their
preferred habitats across the five river sections were analyzed. Benthic
animals are diverse, their habits vary greatly, and their responses to
D. Chen et al.
hydrology and other environmental parameters are highly variable
(Shen et al., 2022). According to the analysis of the 60 sample sites in
this study, benthic animals in the upper reaches of the Yangtze River
were divided into four phyla: Platyhelminthes, Annelida, Mollusca, and
Arthropoda. This study collected and recorded 42 genera and species of
benthic animals belonging to four phyla and seven classes.
3.1.2. Determining the suitability index of sensitive environmental
parameters (water depth and velocity)
Using Equation (1), the HSI values of the sensitive environmental
parameters (water depth and flow rate) of the target species were
calculated, and the results are shown in Fig. 3. The correlation analysis
between the total biomass of benthic animals and their associated and
environmental parameters (water depth and flow rate) showed that the
correlation coefficient between the total biomass and flow rate was − 0.
62. This indicates a significant negative correlation between the total
biomass of benthic animals and flow rate. Furthermore, the results
showed that benthic animals distributed within the flow rate range of
0–0.3 m/s occupied a habitat whose suitability index was 0.72, corre­
sponding to high suitability. Those in the flow rate range of 0.3–0.6 m/s
occupied a habitat with a suitability index of 0.2, corresponding to
moderate suitability. When the flow rate exceeded 0.6 m/s, the suit­
ability index was 0.08, corresponding to low suitability. In the water
depth range of 0–3 m, the suitability index was 0.54, corresponding to
high suitability. In the water depth range of 3–9 m, the suitability index
was 0.32, corresponding to moderate suitability. In the water depth
range of 6–9 m, the suitability index was 0.09, indicating low suitability.
At depths greater than 9 m, the suitability index was 0.05, indicating low
suitability. Overall, these results indicate that benthic animals prefer
shallow habitats with low flow rates (Ban et al., 2020). The habitat
comprehensive suitability index, which combines water temperature
and flow velocity, was calculated using formula (2) above.
3.2. Simulation of habitat suitability range
In this study, based on the hydrodynamic module of the MIKE-21
model coupled with the habitat model, three typical flow rates in the
study area (across the dry, normal, and flood seasons) were selected to
simulate the distribution of habitat suitable for benthic animals in the
Xiangjiaba–Zhutuo section of the upper Yangtze river. As shown in
Fig. 4, when the flow rate was 1500 m3/s, the most suitable habitat for
benthic animals was concentrated along the river overbank and islands.
Fig. 3. (a) Relationship between the habit suitability indices of benthic animals and water depth. (b) Relationship between the habitat suitability indices of benthic
animals and flow rate.
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Journal of Hydrology 625 (2023) 130132
With increased flow rate and water depth, floodplains on both banks and
river islands gradually merged, gradually expanding suitable habitats.
Additionally, as the water depths of the riverbanks and banks of the river
islands gradually increased, the original suitable areas for benthic ani­
mals disappeared as the flow rate increased. When the flow reached
8000 m3/s, the habitat area of the benthic animals gradually shifted to
the bank. Finally, when the flow reached 15000 m3/s, the habitat area
gradually disappeared with increasing flow, and this disappearance was
largely because the beaches and floodplains of the river islands were
mostly covered by river water.
3.3. Decision-making of habitat ecological flow
3.3.1. Discharge scenario settings
The simulated discharge range was determined with reference to the
actual discharge range in Xiangjiaba, China. The relationship between
the discharge Q and water level E (Q–E) was plotted according to the
discharge and water levels at the Xiangjiaba hydrological station from
2018 to 2021. The simulated discharge range was set at 500–25000 m3/
s. The flow rate within the range of 500–2000 m3 /s was low; therefore,
we used 500 m3/s as the flow rate step size to set the simulated flow rate
in the hydrodynamic model. Because the flow rate was high at
2000–25000 m3/s, we used 1000 m3/s as the flow rate step size to set the
simulated flow rate in the hydrodynamic model.
3.3.2. Overall analysis of ecological flow
See Fig. 5(a)–(e) show the relationship between the habitat area of
benthic animals and discharge in the following sections of the upper
Yangtze River: Xiangjiaba–Hengjiang River, Hengjiang River–Minjiang
River, Minjiang River–Tuojiang River, Tuojiang River–Chishui River,
and Chishui River–Zhutuo Town. The total habitat area of each section
first increased and then decreased as the discharge from Xiangjiaba
increased. The maximum discharges of the Xiangjiaba–Hengjiang River,
Hengjiang River–Minjiang River, Minjiang River–Tuojiang River, Tuo­
jiang River–Chishui River, and Chishui River–Zhutuo Town were 2000,
4000, 16000, 11000, and 15000 m3/s, respectively, and the corre­
sponding maximum habitat areas of each section were 3117, 19971,
190968, 94214, and 39227 m2, respectively. There were variations in
the discharge values corresponding to extreme values in the respective
river sections. The flow rate corresponding to an extreme value up­
stream of the Xiangjiaba–Minjiang River section was relatively small.
This is because this section flows through the Yunnan-Guizhou Plateau,
D. Chen et al.
Fig. 4. Area of the suitable habitat of benthic animals under different discharge flows of Xiangjiaba.
where the river valley is deep with high mountains on both banks,
culminating in a floodplain and a riverbank with a small area. Simul­
taneously, the extreme flow of the river section from the Minjiang River
to Zhutuo was relatively large. This is because after the Yangtze River
enters the hilly area of southern Sichuan in the Sichuan Basin, the width
of the river section gradually increases, increasing the areas of the
floodplain and riverbank.
Fig. 5(f) shows the relationship between the total habitat area and
discharge in the Xiangjiaba–Zhutuo Town section. As shown in Fig. 5
(a)–(f), when the discharge of the river section was 16000 m2, the total
habitat area of benthic animals was the largest, with a maximum value
of 339362 m2. Therefore, based on the current hydrodynamics of the
tributaries, the most suitable ecological flow for the mainstream was
16000 m3/s. In this study, the measured flow of the mainstream and its
tributaries was processed as a comparative working condition, and the
average habitat area of each river section was 281115 m2 under the
current flow rate recorded in 2021. As a result, the maximum habitat
area increased by 20.7% when the ideal discharge was adopted. How­
ever, at that time, only the most suitable flow rate for benthic animals
from the Tuojiang River to the Chishui River was close to this value.
Therefore, when the Xiangjiaba Reservoir is discharged at the most
suitable flow rate, although the total habitat area of benthic animals
from Xiangjiaba to the Zhutuo River is the largest, comprehensively
considering the suitable benthic area for each river section is difficult.
3.3.3. Analysis of ecological flow balance
To ensure that the decision-making of ecological flow is based on the
simultaneous consideration of all associated river sections (and there­
fore is more accurate), this study proposed controlling the maximum
habitat area ratio of each river section when making decisions within the
context of ecological flow. Three scenarios were considered, as shown in
Fig. 6(a)–(f). Specifically, when the total habitat area of each section is
greater than 90% of the maximum value, the ecological flow of each
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Journal of Hydrology 625 (2023) 130132
mainstream section cannot be considered. When the total habitat area of
each section is greater than 85% of the maximum value, the flow of all
sections of the mainstream can be considered to be in the range of
5000–13000 m3/s. However, when the total habitat area of each section
was greater than 80% of the maximum value, the discharge range of all
sections of the mainstream could be considered to be 3000–23000 m3/s.
The actual annual discharge in the lower reaches of Xiangjiaba was
4661 m3/s; During the flood season, the actual maximum discharge
could reach 25000 m3/s. In the non-flood season, the actual discharge
capacity of Xiangjiaba barely reached 4000 m3/s. Therefore, when the
total habitat area of each section was greater than 83% of the maximum
value, the habitat area of all sections of the mainstream could be
considered stable. Furthermore, the ecological discharge in the non-
flood season was suggested to be 4000 m3/s; additionally, the habitat
area of the mainstream can be guaranteed to be above 297180 m2 (5.7%
more than the natural state).
To ensure the stability of the habitat area in each river section, we
suggest that Xiangjiaba be discharged as early as possible during the
flood season, and the maximum discharge should be kept under 13000
m3/s. During the flood season, the habitat area from Xiangjiaba to
Zhutuo was maintained above 85% of the maximum value, minimizing
the impact of the flood season on the population dynamics of benthic
animals.
4. Discussions
4.1. Joint regulation of ecological flow in mainstream and tributaries
Based on the study of ecological discharge in the Xiangjiaba Reser­
voir, this study assumed that the three tributaries with hydraulic pro­
jects, namely, the Hengjiang, Minjiang, and Tuojiang rivers, were driven
by ecological regulatory mechanisms similar to those of the Xiangjiaba
Reservoir. Joint regulation and control schemes of the mainstream and
D. Chen et al.
Fig. 5. Habitat area in relation to the discharge flow at Xiangjiaba for each studied river section: (a) Xiangjiaba-Hengjiang River, (b) Hengjiang River-Minjiang River,
(c) Minjiang River-Tuojiang River, (d) Tuojiang River-Chishui River, (e) Chishui River-Zhutuo Town, and (f) Xiangjiaba-Zhutuo Town.
tributaries are also discussed. Fig. 7 shows the relationship between
habitat area and mainstream discharge from Xiangjiaba to Zhutuo. The
habitat area of each river section first increased and then decreased as
total discharge increased. Additionally, the maximum discharges cor­
responding to each section were 2000, 4000, 18000, 13000, and 20000
m3/s, respectively. The theoretical maximum habitat area was 344401
m2 under the condition that the flow rate could be regulated freely. (See
Fig. 8.).
The actual annual average discharges in the lower reaches of the
Xiangjiaba, Hengjiang, Minjiang, Tuojiang, and Chishui rivers were
4661, 242, 3017, 521, and 283 m3/s, respectively. The actual annual
flow of the mainstream Xiangjiaba section and each tributary section
does not reach the ideal flow, and in this study, we assumed that the
amount of water in each river section could be freely allocated and that
the maximum discharge could reach the annual average discharge.
When the total habitat area was at its greatest, the discharges in the
lower reaches of the Xiangjiaba, Hengjiang, Minjiang, Tuojiang, and
Chishui Rivers were 4661, 242, 3017, 521, and 283 m3/s, respectively.
Additionally, the total habitat area required to reach the most suitable
habitat area was 87% greater than the ecological flow rate of each
section, corresponding to a maximum area of 319130 m2.
It is important to note Considering the following is important: it is
difficult for the actual discharge capacity to reach its theoretical opti­
mum during the non-flood season; controlling the discharge in the
Chishui River is difficult; and finally, the stability of the habitat area is
considered on a section-by-section basis. In this study, the proposed
scheme for the non-flood flow rate for each section considered the
7
Journal of Hydrology 625 (2023) 130132
conditions above. The proposed non-flood control flow rates for the
downstream sections of the Xiangjiaba, Hengjiang, Minjiang, and Tuo­
jiang rivers were 4000, 200, 2400, and 400 m3/s, respectively, resulting
in the ecological flow rate of each section reaching more than 86% of the
optimum habitat area, corresponding to a habitat area of 313316 m2.
Compared to the area of 297180 m2, when only the discharge flow of
Xiangjiaba was controlled, it was further increased by 5.4%. By ensuring
ecological flow during the non-flood season, the habitat area of each
section could be increased from 83% to 86% of the most suitable habitat,
and the stability of the habitat area of each mainstream section could be
further considered.
During the flood season, to ensure the stability of the habitat area of
each river section, we assumed that the habitat area of each river section
reached the standard of the guaranteed habitat area in the non-flood
season, and we suggest a flow of 86% of the optimum habitat as the
standard. We also suggest that the maximum control flows during the
flood season in the lower reaches of the Xiangjiaba, Hengjiang, Min­
jiang, and Tuojiang River sections should be 10000, 1000, 14000, and
9000 m3/s, respectively. These conditions are intended to maximize
habitat stability across the entire study area.
4.2. Comparison of research results and sensitivity analysis
Few studies have been conducted on applying physical habitat
models for benthic animals in aquatic environments. Among them,
Blettler et al. analyzed the impact of hydrological changes on the living
environment of benthic animals in plain systems with large river floods
D. Chen et al. Journal of Hydrology 625 (2023) 130132

Fig. 6. Schematic diagram of the range of discharge flows corresponding to different percentages of habitat area extremes in each river section in the study area
（Only control the flow towards Jiaba, with each tributary having an average annual flow rate）: (a) Xiangjiaba-Hengjiang River, (b) Hengjiang River-Minjiang
River, (c) Minjiang River-Tuojiang River, (d) Tuojiang River-Chishui River, (e) Chishui River-Zhutuo Town, and (f) Xiangjiaba-Zhutuo Town.

(Blettler, M. C. M., et al. 2016). During floods, the habitat of benthic
animals in the main tributaries is greatly affected. Maria et al studied the
sensitivity of benthic animals to water pressure (Maria et al. 2021),
revealing the reasons why benthic animals are more sensitive to the
influence of water depth and watershed.In a study of ecological flow,
Ban Xuan et al. used the physical habitat of benthic animals to analyze
the suitable ecological flow range for benthic animals in the Jianli sec­
tion of the Yangtze River, ranging from 12,000 to 36000 m3/s. However,
this study only provided the ecological flow range for a single river
section. Compared to the results of this study, this study did not consider
the changes in suitable ecological flow between the upstream and
downstream sections of the river, nor did it consider the joint regulation
of the main river and its tributaries. Therefore, this study conducted a
segmented ecological flow analysis and joint regulation of the main
tributaries in the study area, and the study results have more practical
significance than those of previous studies. Research on the application

8
D. Chen et al.
Fig. 7. Relationship between ecological flow and habitat area: (a) Xiangjiaba-Hengjiang River, (b) Hengjiang River-Minjiang River, (c) Minjiang River-Tuojiang
River, (d) Tuojiang River-Chishui River, and (e) Chishui River-Zhutuo Town.
of physical habitat models for benthic animals in aquatic environments
is limited. However, this study only provided the ecological flow range
for a single river section. In contrast to the results of this study, Ban Xuan
et al. did not consider the changes in suitable ecological flow between
the upstream and downstream of the river section or the joint regulation
of the main river and tributaries. Therefore, this study conducted a
segmented ecological flow analysis and joint regulation of the main
tributaries in the study area, with results that have practical significance
compared to those of previous studies.
In addition to the above studies, Yiwen Z et al analyzed the
comprehensive habitat quality index of urban rivers by quantitative
analysis of zooplankton (Yiwen Z et al. 2022). This study is similar to the
idea of this paper, but the index species used are different. This also
validates the rationality of using benthic animals which are most sen­
sitive to the change of hydrological regime as ecological flow decision-
making.
Although this study has made many breakthroughs compared with
previous studies, it has three overall limitations: 1. There was a slight
lack of selection of environmental factors. Owing to the limitations of
the research area and time period, the impact of environmental factors,
such as temperature and substrate, on habitats was not considered in this
study; 2. This study only selected benthic animals as the target species
for research without considering the impact of environmental factors on
other types of biological habitats, such as fish, zooplankton, phyto­
plankton, and benthic diatoms; 3. In the comprehensive impact analysis
of environmental factors, the analysis method used in this study is still
slightly conventional. In future research, expanding the scope of the
research area, research period, and types of target species is necessary.
Other environmental factors, such as substrate and temperature, and
various biological types, such as fish, zooplankton, phytoplankton, and
benthic diatoms, must be added to the construction of physical habitat
models, and a more comprehensive approach should be adopted to
construct the relationship between environmental factors and various
biological habitats.
9
Journal of Hydrology 625 (2023) 130132
Based on an analysis of the degree of habitat area change (i.e., degree
of habitat area change maximum habitat area / minimum habitat area)
and the actual discharge flow range under the Xiangjiaba Dam from
2018 to 2021, the degree of change was more apparent in the Minjiang
River–Zhutuo Town section compared to that observed in the Xiang­
jiaba–Minjiang River section (Table 1). These trends indicate the
sensitivity of changes in the area of suitable habitat to distance as well as
to the topography and geomorphology of each river section (Durden
et al., 2015). Specifically, the Minjiang River–Zhutuo Town section
mainly flows through the Sichuan Basin, where the riverbed is relatively
wide, and the area of the floodplain and river island beaches is relatively
large, resulting in greater sensitivity to changes in the area of suitable
habitat. In contrast, the Xiangjiaba–Minjiang River section mainly flows
through the Yunnan-Guizhou Plateau, characterized by high mountains
and deep valleys. In this section, the river was V-shaped, and the impact
of flow variation on the habitat area was limited. Therefore, under
similar topographical and geomorphological conditions, the overall
degree of variation in suitable habitat areas tended to decrease with
increasing distance.
5. Conclusions
This study coupled the MIKE21 hydrodynamic and habitat ecology
models to establish a physical habitat model for benthic animals in the
upper reaches of the Yangtze River, specifically from Xiangjiaba to
Zhutuo Town. This study assessed the effects of two environmental pa­
rameters (flow velocity and water depth) on the habitats of benthic
animals. We focused on the river section from Xiangjiaba to Zhutuo
Town to study changes in the range of habitats suitable for benthic an­
imals. We studied these habitat area dynamics under changing flow rates
and suggested recommendations for decision-making regarding collab­
orative ecological discharge of the mainstream and its associated trib­
utaries. This study addresses the gap in the physical habitat model of
benthic animals in the upper reaches of the Yangtze River as well as
D. Chen et al. Journal of Hydrology 625 (2023) 130132

Fig. 8. Schematic diagram of the range of flows corresponding to different percentages of habitat area extremes in each river section of the study area（Simulta­
neously controlling the flow rate towards Jiaba and the flow rate of each tributary）: (a) Xiangjiaba-Hengjiang River, (b) Hengjiang River-Minjiang River, (c)
Minjiang River-Tuojiang River, (d) Tuojiang River-Chishui River, and (e) Chishui River-Zhutuo Town.

indicates the first application of the physical habitat model to the joint
ecological water use scheduling of main and tributaries, thus paving the
way for a new direction in the application of the physical habitat model.
The main conclusions of this study are as follows:
1. Based on 180 sets of data from five sampling sites, which include
Xiangjiaba Dam, Yibin, Lizhuang, Luzhou, and Zhutuo Town, the habitat
suitability index (HSI) was used to quantify the collected biomass and
environmental parameter data. The results showed that the environ­
mental parameters associated with the total biomass were water depth
and flow rate. The total benthic animal biomass was negatively corre­
lated with water depth and flow rate. Benthic animals tend to prefer
habitats with low flow rates and shallow waters. A flow velocity within

10
D. Chen et al.
Table 1
Data of habitat area change degree in each river section.
Actual flow River reach Change degree of Distance from
range (m3/s) habitat area (m2) Xiangjiaba to river
reach (km)
0–25000 Xiangjiaba- 1.325 2.81
Hengjiang River
0–25000 Hengjiang River- 1.233 19.35
Minjiang River
0–25000 Minjiang River- 1.397 96.30
Tuojiang River
0–25000 Tuojiang River- 1.293 194.30
Chishui River
0–25000 Chishui River- 1.283 245.95
Zhutuo Town
Correlation coefficient between the degree of change in the 0.080
area of the habitat and the distance from Xiangjiaba to
river reach
the 0–0.3 m/s range was associated with the highest biomass and cor­
responded to a suitability index of 0.72. Water depths in the 0–3 m range
had a high suitability index (0.54).
2. By coupling the hydrodynamic and habitat models, a physical
habitat model was established, and three typical flows were selected
with the actual flow discharge range to simulate a suitable habitat area
distribution in the Xiangjiaba-Zhutuo section during the dry, normal,
and flood seasons. As the flow and water depth increased, there was a
gradual expansion in the suitable habitat area, and the original suitable
area for benthic animals gradually disappeared with an increase in the
flow. These conditions were accompanied by increased water depth
along the riverbank and river island.
3. The hydraulic distribution from Xiangjiaba to Zhutuo under
various flow rates was simulated using the MIKE21 hydrodynamic
model. Discharge from Xiangjiaba to Zhutuo was obtained by coupling
the habitat suitability index of benthic animals with the relationship
between the total benthic area. According to the results, we suggest that
the discharge flow of the Xiangjiaba Reservoir should be maintained
above 4000 m3/s in normal and dry seasons; however, the maximum
discharge should be kept below 13000 m3/s during the flood season to
minimize the influence of extreme water fluctuations during the flood
and dry seasons on the growth of benthic animals in the lower reaches of
the Xiangjiaba Reservoir.
4. Based on the actual discharge range under the Xiangjiaba Dam
from 2018 to 2021, the sensitivity of the habitat area changes in each
section was analyzed. The sensitivity of changes in suitable habitat areas
was related to distance as well as the topography and geomorphology of
each river section. Specifically, the sensitivity in hilly areas was higher
than that in mountainous areas. However, when the topography was
similar, the sensitivity of suitable habitat areas decreased with
increasing distance.
5. By controlling the ecological discharge flow of each tributary and
further analyzing the joint regulation and control scheme of the main­
stream and associated tributaries, the proposed non-flood control flows
downstream of Xiangjiaba and sections of the Hengjiang, Minjiang, and
Tuojiang rivers were above 4000, 200, 2400, and 400 m3/s, respec­
tively, corresponding to a habitat area of 10000 m2. During the flood
season, we propose that the flows downstream of the Xiangjiaba,
Hengjiang, Minjiang, and Tuojiang rivers should be 10000, 1000,
14000, and 9000 m3/s (or less), respectively. This corresponds to a
habitat greater than 313316 m3. This is 5.4% higher than that in the
scenario in which only the discharge flow of Xiangjiaba is controlled.
The ecological flow of each section reached more than 86% of the
optimal habitat area, considering the stability of the habitat area of the
respective mainstream sections.
11
Journal of Hydrology 625 (2023) 130132
CRediT authorship contribution statement
Dingxin Chen: Writing – review & editing, Conceptualization,
Methodology. Zefan Yang: Supervision. Qinghui Zeng: Investigation,
Data curation. Weize Wang: Methodology. Long Yan: Investigation,
Supervision, Validation. Pu Zhang: Investigation, Supervision, Soft­
ware. Xinyu Li: . Peng Hu: Supervision. Hao Wang: Conceptualization,
Methodology, Supervision.
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.
Acknowledgments
This work was supported by the National Key Research Program of
China (No. 2022YFC3202003), the National Natural Science Foundation
of China [No. U2240202, 52122902, 52009146], and the Independent
Research Project of the State Key Laboratory of Simulation and Regu­
lation of Water Cycle in River Basin (No. SKL2022TS07, SKL2022ZD01).
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