Journal of Hydrology 587 (2020) 124938

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Journal of Hydrology
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Research papers

Impacts of a large river-to-lake water diversion project on lacustrine T

phytoplankton communities
Jiangyu Daia, , Shiqiang Wua, , Xiufeng Wua, , Xueyan Lvb, Bellie Sivakumarc, Fangfang Wanga,
⁎ ⁎ ⁎

Yu Zhanga, Qianqian Yanga, Ang Gaoa, Yuhang Zhaoa, Lei Yua, Senlin Zhua,
⁎

a
State Key Laboratory of Hydrology-Water Resources and Hydraulic Engineering, Nanjing Hydraulic Research Institute, Nanjing 210029, China
b
Jiangsu Environment Monitoring Center, Nanjing 210029, China
c
Department of Civil Engineering, Indian Institute of Technology Bombay, Powai, Mumbai, Maharashtra 400076, India

ARTICLE INFO ABSTRACT

Keywords: Allochthonous inputs of species and nutrients and hydrodynamic disturbance induced by water diversion pro-
Water diversion jects are two critical factors of ecohydrological effects in eutrophic lakes. However, identification and quanti-
Allochthonous input fication of potential contributions of allochthonous inputs and physicochemical habitat shifts to variations in
Physicochemical habitat phytoplankton communities remain challenging. The present study addresses this issue with a study of the Water
Phytoplankton
Diversion from the Yangtze River to Lake Taihu in China. To explore the effects and contributions of seasonal
Lake Taihu
water diversion activities on lacustrine phytoplankton communities, the comparative analysis was conducted to
compare the biotic and abiotic variables between the water diversion and non-diversion periods in different
seasons. The results showed that seasonal water diversion activities, in addition to significantly reducing organic
pollutant concentrations, definitely increased the average concentrations of nitrate and phosphorus in the
Gonghu Bay. Compared with the results in the Gonghu Bay on the non-diversion days, phytoplankton diversities
increased and the community compositions were altered, with the Bacillariophyta species and non-Microcystis
Cyanophyta species dominating in the Gonghu Bay on the water diversion days in different seasons. The venn
diagram analysis showed that the highest potential contribution of the allochthonous species to the increase in
phytoplankton diversity in the Gonghu Bay on the water diversion days was about 15.8%. The physicochemical
habitat disturbance induced by the water diversion contributed about 12% to 31.3% of the phytoplankton di-
versity and 23.3% to 31.3% of the phytoplankton community variations in the Gonghu Bay. The allochthonous
phytoplankton species may contribute directly to the lacustrine phytoplankton community variations. However,
due to the high loads of nutrients from inflow rivers, positive effects of water diversion shaping the phyto-
plankton communities were always short-term. Pollutant control and multi-objective operation, considering
flood control, water supply, and water environment improvement, are indispensable for the long-term man-
agement of water diversion projects.

1. Introduction countermeasures, including mechanical salvage, phytoremediation, and

Lakes not only provide water resources and eco-services for the
human society, but also reflect regional ecological health of the aquatic
ecosystems (Scheffer, 1997). Due to the influences of anthropogenic
activities (Ali et al., 2019) and climate change, the limnetic eu-
trophication process accelerates. This brings about a series of ecological
problems, such as cyanobacterial blooms, vegetation degradation, and
diversity reduction, and threatens the water supply function and eco-
logical balance of lakes (Paerl et al., 2016). To cope with the harmful
cyanobacterial blooms and restore lake ecosystems, many
fish predation (Cooke et al., 2016; Jeppesen et al., 2007; Qin, 2009),
and various engineering measures, such as water diversion and ecolo-
gical dredging (Hu et al., 2008; Lürling and Faassen, 2012), have and
continue to be attempted. Among these measures, water diversion from
rivers to lakes has received significant interest from scientists and lake
basin administrations, because of the rapidity and effectiveness with
which it relieves the cyanobacterial blooms. It has also become a
popular and quick measure to improve water quality and to prevent or
mitigate the ecological deterioration of many lakes, including Lake
Green in USA (Oglebsy, 1969), Lake Veluwe in the Netherlands

⁎
Corresponding authors.
E-mail addresses: jydai@nhri.cn (J. Dai), sqwu@nhri.cn (S. Wu), xfwu@nhri.cn (X. Wu), slzhu@nhri.cn (S. Zhu).

https://doi.org/10.1016/j.jhydrol.2020.124938

Available online 11 April 2020

0022-1694/ © 2020 Elsevier B.V. All rights reserved.
J. Dai, et al.
(Jagtman et al., 1992), Lake Moses in USA (Welch et al., 1992), Lake
Tega in Japan (Amano et al., 2010), Lake Xuanwu in China (Song et al.,
2018), and Lake Xihu in China (Zhang et al., 2018), among others.
Due to the ecological and drinking water crisis induced by harmful
cyanobacterial blooms in large eutrophic lakes in China, such as Lake
Taihu, Lake Hongze, Lake Chaohu, and Lake Dianchi, water diversion
projects have become a very commonly used engineering measure for
alleviating water crisis in such lakes. The ecohydrological effects of
water diversion projects on large eutrophic lakes have also been at-
tracting significant attention in recent years (Dai et al., 2016; Dai et al.,
2018; Hu et al., 2008; Huang et al., 2015, 2016; Li et al., 2011, 2013;
Liu et al., 2014; Yu et al., 2018). Much of the research in this area has
focused on the hydrodynamic and environmental effects, such as the
effects on water level (Hu et al., 2008), water age (Huang et al., 2016;
Li et al., 2011; Zhang et al., 2016), flow field (Hu et al., 2008; Li et al.,
2013), water quality (Dai et al., 2016; Hu et al., 2010; Liu et al., 2014),
and ecological health based on water quality (Zhai et al., 2010). Only a
few studies have addressed the issues related to the biological effects of
water diversion in large eutrophic lakes (Huang et al., 2015; Lin et al.,
2017). As an abundant and active biota in lakes, phytoplankton are
significantly sensitive to the lake conditions, such as the trophic level
and the hydrodynamic disturbance (Schwalb et al., 2013; Yang et al.,
2016). Conversely, the phytoplankton community structure can also
reflect the environmental changes and can be considered as a suitable
index to represent the evolution of aquatic ecosystems induced by ex-
ternal disturbance (Yang et al., 2017), such as the disturbance by water
diversion.
The ecohydrological effects of water diversion on lakes are mainly
related to three main factors: hydrodynamic disturbance of the inflow
discharge, allochthonous nutrient input, and allochthonous biological
species. The inflow discharges could shorten the lake residence time (Li
et al., 2011, 2013) and alter the lake currents, which can influence the
transportation, transformation, and distribution of nutrients and bio-
logical communities of some small lakes (Oglebsy, 1969; Jagtman et al.,
1992; Welch et al., 1992; Hosper, 1998; Amano et al., 2010). However,
for large eutrophic lakes, the residence time is generally very long and
only a limited inflow discharge of any water diversion cannot sig-
nificantly shorten the lake residence time. For instance, for Lake Taihu,
the residence time is almost always longer than 200 days, and an inflow
discharge lower than 150 m3/s does not have much influence on the
lake residence time (Li et al., 2011). Since the upstream rivers that
divert water to lakes generally have great nutrient loads, the al-
lochthonous nutrients from the upstream rivers will play a more im-
portant role in structuring phytoplankton communities in lakes, by
considering the rapid in situ growth of the phytoplankton in eutrophic
waters (Swarbrick et al., 2019). Moreover, allochthonous phyto-
plankton species from rivers could also directly affect the phyto-
plankton communities in lakes. In China, there are often concerns re-
garding the ecohydrological effects of high allochthonous nutrient
inputs induced by water diversion projects. However, to what extent
the allochthonous inputs from water diversion projects contribute to
variations in the phytoplankton communities remains unknown.
Lake Taihu is the third largest shallow eutrophic lake in China, and
is located in the lower reaches of the Yangtze River. To cope with the
frequent cyanobacterial blooms of Lake Taihu and meet the water de-
mands of the Taihu Basin, Water Diversion from the Yangtze River to
Lake Taihu (WDYT) was initiated in 2002 and has been operated reg-
ularly from 2007 (Qin et al., 2019). The Taihu Basin Authority (TBA)
has been responsible for the regulation of the WDYT project and has
formulated the dispatching rule according to the average real-time
water level of Lake Taihu. Normally, when the average real-time water
level of Lake Taihu is lower than the dispatching control water level,
the Wangyu River will input allochthonous water from the Yangtze
River, and then divert it into Lake Taihu through the Wangyu River
after the water quality of the Wangyu River reaches or precedes the
third level of surface water quality standards in China. Many past
2
Journal of Hydrology 587 (2020) 124938
studies have demonstrated that the seasonal water diversion activities
of the WDYT could improve the hydro-environment of Lake Taihu but
also increase the loading of nutrients, such as the nitrate and total
phosphorus in some regions (Dai et al., 2018; Hu et al., 2008, 2010).
However, only a few studies have addressed the influence of the phy-
sicochemical habitat variations induced by the seasonal water diversion
activities on phytoplankton communities in the water-receiving regions
of Lake Taihu. This provides the motivation for the present study.
In this study, we consider the WDYT project as an example to in-
vestigate the phytoplankton communities and physicochemical para-
meters in the water-receiving regions of Lake Taihu and to compare the
biotic and abiotic variables between the water diversion and non-di-
version periods in different seasons. For the latter, we use the sourcing
and multivariate statistical analysis. Based on these, we examine three
issues: (1) responses of the lacustrine phytoplankton communities and
physicochemical habitat to short-term water diversion in different
seasons; (2) relationships between the physicochemical habitat shifts
induced by water diversion and phytoplankton communities; and (3)
contributions of the allochthonous phytoplankton species and physi-
cochemical habitat disturbance to lacustrine phytoplankton commu-
nities during the water diversion periods. The outcomes of this study
could help elucidate the positive and negative effects of water diversion
on the cyanobacterial blooms in the large eutrophic Lake Taihu. They
could, consequently, provide useful hints to understand the short-term
ecohydrological effects of water diversion on large eutrophic lakes.
2. Materials and methods
2.1. Study area and sampling sites
A schematic diagram of the methodology used in this study is shown
in Fig. 1. Lake Taihu is the geographic and hysteresis storage center of
water resources for the Taihu Basin. The Wangyu River is one of the
main water diversion channels of the WDYT and directly connects the
Yangtze River and Lake Taihu (Fig. 2). The Changshu and Wangting
Hydro-junctions are the two main operating sections regulating the
importation of the Yangtze River water and the inflow discharges into
the Gonghu Bay from the Wangyu River, respectively. Gonghu Bay is
located in the northeastern part of Lake Taihu, which covers an area of
150 km2 and has the multi-year average water depth of 1.8 m (Zhong
et al., 2012) (Fig. 2). The southwestern part of the Gonghu Bay connects
the lake center and the Meiliang Bay, which is the largest and hyper-
eutrophic bay of Lake Taihu. Since 2005, large quantities of cyano-
bacterial blooms have begun to cover most water areas of the Gonghu
Bay, with a significant increase in 2007 (Qin et al., 2010). In recent
years, with the improvement of water quality in the Gonghu Bay,
contributed by the integrated management of the whole basin, Gonghu
Bay has become one of the main drinking water sources for Wuxi City
and Suzhou City in the Taihu Basin.
For the data considered in this study, the sampling sites were lo-
cated in the Wangyu River (sites R1–R3), Gonghu Bay (sites A1–E1,
A2–E2 and A3–E3) and the lake center (sites L1–L3) of Lake Taihu
(Fig. 2). The sites R1–R3 were located in the upstream of the Wangting
Hydro-junction, to investigate the inflow water characteristics. The
layout of the sites in the Gonghu Bay intended to investigate the gra-
dients of the physicochemical and biological parameters from the
Wangyu River mouth to the lake center during the water diversion
periods. Since the lake center is adjacent and open to the Gonghu Bay,
there are exchanges of hydraulics, nutrients and phytoplankton species
between the Gonghu Bay and the lake center. The lake center is less
influenced by water diversion than the Gonghu Bay. The intention of
laying the sampling sites in the lake center is to elucidate the potential
contribution to the phytoplankton community variations in the Gonghu
Bay. The Wangyu River and the lake center were the two adjacent re-
gions to the Gonghu Bay. To elucidate the potential contribution of the
Wangyu River, we should also know the potential contribution of the
J. Dai, et al. Journal of Hydrology 587 (2020) 124938

Fig. 1. The flow chart of methodologies used in this study.

lake center. In lake regions, the sampling sites were located with the
same distance of 2 km between the two adjacent sites.
2.2. Sample collection and measurement of abiotic variables
In the middle and lower reaches of the Yangtze River, January,
April, August, and November are always taken as the representative
months for winter, spring, summer, and autumn, respectively.
According to the dispatching operation of the WDYT in 2013, 2014, and
2015, August in 2013 and January and November in both 2014 and
2015 were all in the water diversion periods. Other months were all in
the non-diversion periods. A 2 L of surface water sample (50 cm under
water surface) from each sampling site was collected on the middle day
of each selected month during 2013–2015 (Table 1). Of this 2 L water
sample, 1 L was fixed using 15 ml 1% mass concentration of Lugol’s
iodine fixing reagent for identification of the phytoplankton commu-
nity. The water samples collected for each site were stored in the sterile
plastic bottle and transported to the laboratory for analysis within 24 h.
The sampling time began at 9:00 on each day.
The data of inflow discharges of the Wangyu River and the water

Fig. 2. Locations of the studied water regions and sampling sites in Lake Taihu. (Sites R1-R3 were in the Wangyu River; sites A1-A3, B1-B3, C1-C3, D1-D3, E1-E3
were in the Gonghu Bay; and sites L1-L3 were in the lake center of Lake Taihu.)

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J. Dai, et al. Journal of Hydrology 587 (2020) 124938

Table 1
The weather and hydrologic parameters in the investigated regions on the sampling days from 2013 to 2015.
Sampling day Wind direction* Wind speed Rain† Lake water Cumulative inflow amount Duration days of one continuous Average inflow discharge during
(m/s) level (m) (108 m3) diversion (d) duration days (m3/s)

2013-01-10 E 3–4 No 3.3 / 10 days after /

2013-04-15 S 3–4 No 3.1 / / /
2013-08-18 SE 5–7 No 3.1 2.12 22 111.5
2013-11-20 E 4–6 No 3.2 / 10 days before /
2014-01-14 NW 2–3 No 3.0 3.32 45 85.4
2014-04-23 SE 5–7 No 3.2 / / /
2014-08-19 SE 7–8 No 3.7 / / /
2014-11-21 SW 2–3 No 3.2 0.97 27 41.6
2015-01-17 N 3–4 No 3.1 4.70 74 73.5
2015-04-21 SE 2–3 No 3.4 / / /
2015-08-19 SE 3–4 No 3.6 / / /
2015-11-24 E 1–2 No 3.5 0.66 13 58.8

* E denotes east, S denotes south, N denotes north, SE denotes southeast, NW denotes northwest, SW denotes southwest.
†
No represents no rain; / represents no water transfer.

level of Lake Taihu during the sampling days were collected from the
Annual Reports of the WDYT Project in 2013, 2014, and 2015 (TBA,
2013, 2014, 2015). Wind direction and speed in each sampling site
were measured in situ using the hand-held anemometer NK 4500
(Kestrel & NK, Beijing, China). Water temperature, pH, dissolved
oxygen (DO), and turbidity were all measured in situ by the Portable
Multi-parameters Detection HQ30d (HACH, Shanghai, China). The total
nitrogen (TN), total phosphorus (TP), ammonia (NH3-N), nitrate (NO3-
N), soluble reactive phosphorus (SRP), permanganate index (CODMn),
dissolved silicate (SiO3-Si), and chlorophyll a (Chl-a) were all de-
termined in the laboratory according to the method used by Jin and Tu
(1990).
and 2015) were analyzed by the Analysis of Similarities (ANOSIM)
method (Fanini and Lowry, 2016) and plotted using the non-metric
multidimensional scaling (NMDS) method with the PRIMER-E software
(Quest Research Limited, Auckland, New Zealand). For the NMDS
analysis, the matrix data constructed using the genera cell numbers of
all samples were first log10(x + 1) transformed and then reassembled
using the Bray-Curtis similarity index. Redundancy analysis (RDA) was
carried out to reveal the correlations and contributions of the in-
vestigated physicochemical parameters to lacustrine phytoplankton
communities by using the CANOCO 5 software (Lai, 2013). The sig-
nificance (p < 0.05) of the calculated results was verified with an
unrestricted Monte Carlo permutation.

2.3. Identification of phytoplankton communities 3. Results

The 1 L Lugol’s iodine reagent fixed water sample for each site was
transferred into a separating funnel. After stewing for 24 h, the 50 ml
precipitated bottom sample was collected to store in the sterile glass
bottle and kept in a dark place until the beginning of the phytoplankton
identification (Jin and Tu, 1990). A 0.1 ml of the concentrated sample
was injected into the algae counting box and put in the optical micro-
scope Axiovert 200 (CARL ZEISS, Jena, Germany) to uniformly capture
100 random views. These random views were used to identify the
phytoplankton species and to count the cell numbers with the help of
the T300 Algae Smart Identification software (Shineso, Hangzhou,
China). This procedure was repeated three times for each sample. The
identification of the phytoplankton species was done according to the
method used by Hu and Wei (2006).
2.4. Statistical analyses
The phytoplankton diversity indexes of the species number and the
Shannon-Wiener index (H′) for each sample were calculated using the
PRIMER-E software (Quest Research Limited, Auckland, New Zealand).
The differences in the physicochemical parameters and phytoplankton
diversity indexes in the Gonghu Bay between the water diversion and
non-diversion periods were compared with the One-way ANOVA
method using the SPSS 16.0 statistic software (IBM, Armonk, USA). The
Venn diagram, plotted using the VennPainter software (Lin et al.,
2016), was adopted to determine the potential contribution of al-
lochthonous phytoplankton species of the Wangyu River to that of the
Gonghu Bay. The gradients in the average relative proportions of the
phytoplankton community composition from the Wangyu River (sites
A1–A3) to the lake center (sites L1–L3) were plotted using the Sigma-
plot 12.5 software (Systat Software Inc., London, UK).
The similarities between the phytoplankton communities in the
Gonghu Bay among the same seasons in the three years (2013, 2014,
3.1. Physicochemical parameters
During the sampling days, there was no rain in the sampling regions
and most of the wind speeds on the lake surface were lower than 4 m/s.
The cumulative inflow during the water diversion period in winter until
the sampling day was higher than the annual average water storage of
the Gonghu Bay (about 2.70 × 108 m3), while that in the summer and
autumn diversion periods until the sampling day were lower (Table 1).
The average inflow (111.5 m3/s) during the summer water diversion
period was higher than that during the winter (85.4 and 73.5 m3/s) and
fall (41.6 and 58.8 m3/s) water diversion periods.
Significant differences in the concentrations of pH, SiO3-Si, TN,
NO3-N, TP, and CODMn were found between the diversion period and
non-diversion period in each season, respectively (One-way ANOVA,
p ≤ 0.05; Fig. 3). Since there were significant differences in the phy-
sicochemical parameters between the Wangyu River and the center of
Lake Taihu, water diversion activities in different seasons significantly
decreased the values of pH and CODMn and increased the contents of
SiO3-Si, TN, NO3-N and TP.
3.2. Phytoplankton diversities and overlaps of genera among three regions
on water diversion days
The average diversities in the Wangyu River and the Gonghu Bay on
different sampling days were, respectively, all significantly higher than
those in the lake center (Fig. 4; One-way ANOVA, p < 0.05). For the
Gonghu Bay in the same season of different years, the two diversity
indexes (i.e. phytoplankton species number and the Shannon-Wiener
index) on the water diversion days were significantly higher than those
on the non-diversion days, except for the diversities in January (Fig. 4a
and 4b; One-way ANOVA, p > 0.05). Whether between the diversion
days or between the non-diversion days, there was also no significant

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J. Dai, et al. Journal of Hydrology 587 (2020) 124938

Fig. 3. Variations in physicochemical parameter values of the three regions on sampling days in different seasons from 2013 to 2015. (The single asterisk represents
significant difference (One-way ANOVA, p ≤ 0.05) between the mean values of parameters in the Gonghu Bay on the sampling days in two different years; the double
asterisk means highly significant difference (p ≤ 0.01)).

divergence in both the two diversity indexes in this study (Fig. 4; One-
way ANOVA, p > 0.05).
The unique genera only found in the Gonghu Bay accounted for
17.4% (Fig. 5a), 12% (Fig. 5b), 23.1% (Fig. 5c), 15.8% (Fig. 5d) and
31.3% (Fig. 5e) of the total phytoplankton diversity in the Gonghu Bay
on the water diversion day in January of 2014, January of 2015, August
of 2013, November of 2014, and November of 2015, respectively.
Moreover, the common genera only found between the Wangyu River
and Gonghu Bay on the water diversion days showed that the al-
lochthonous genera from the Wangyu River potentially accounted for
0%, 8.3%, 7.7%, 15.8%, and 0% of the total phytoplankton diversity in
the Gonghu Bay on the above-mentioned water diversion days,
respectively.
The genera only found in the Gonghu Bay on the water diversion
days in different seasons were affiliated with the Cyanophyta,
Chlorophyta, Bacillariophyta, Chrysophyta, and Xanthophyta.
However, all of the relative proportions in the cell abundances of these
genera were lower than 1.0% (Table 2). None of these was the domi-
nant genera in the Gonghu Bay during the water diversion days. There
were several genera, such as Westella spp., Phacus spp., Arthrospira spp.,
Golenkinia spp., Neidium spp., Ankistrodsemus spp., and Synura spp.,
overlapped only between the Wangyu River and the Gonghu Bay on the
water diversion days in different seasons.
In January and November, the relative proportions of Aulacoseira

Fig. 4. Comparisons in the number of phytoplankton species (a) and Shannon-Wiener index (b) in the studied regions among the sampling days in the same season
from 2013 to 2015. (Jan, Apr, Aug, and Nov are the abbreviations of January, April, August, and November, respectively. The single asterisk represents significant
difference (One-way ANOVA, p ≤ 0.05); the double asterisk means highly significant difference (p ≤ 0.01)).

5
J. Dai, et al.
Fig. 5. The Venn diagrams showing numbers of the identical and common genera among different regions on the water diversion and non-diversion days in January
((a) and (b), August (c) and November ((d) and (e)), respectively. (R, G, and L represent the regions of the Wangyu River, Gonghu Bay, and the central lake,
respectively.)
granulate, Cyclotella spp., and Synedra spp. in the Wangyu River were
evidently higher than those in the Gonghu Bay on the water diversion
days and non-diversion days, respectively (Table 2). In August, the
Arthrospira spp. (46.7%) and Planktothrix spp. (40.2%) were the domi-
nant Bacillariophyta genera in the Wangyu River, which were sig-
nificantly higher in relative proportions than those in the Gonghu Bay
on the water diversion days and non-diversion days, respectively
(Table 2).
3.3. Phytoplankton abundances and community compositions
The spatial variations in the total phytoplankton cell abundances
and community compositions on each sampling day are shown in Fig. 6.
There were eight phytoplankton phyla recovered on the sampling days
in different seasons, including the Cyanophyta, Chlorophyta, Bacillar-
iophyta, Cryptophyta, Euglenophyta, Pyrrophyta, Chrysophyta, and the
6
Journal of Hydrology 587 (2020) 124938
Xanthophyta.
In January, the average values of the total phytoplankton cell
abundances in the Gonghu Bay on the non-diversion day (Fig. 6a) were
significantly higher than those in the Gonghu Bay on the water diver-
sion days (Fig. 6b and c; One-way ANOVA, p < 0.05). The Bacillar-
iophyta dominated in the Gonghu Bay on the water diversion days
(Fig. 6b and c), whereas the Cyanophyta was more abundant on the
non-diversion day (Fig. 6a).
In April, the Cyanophyta predominated in the Gonghu Bay and the
lake center on all the sampling days (Fig. 6d, 6e, and 6f). In August, the
average phytoplankton cell abundances in the Gonghu Bay on the water
diversion day in 2013 (Fig. 6g) and on the non-diversion day in 2015
(Fig. 6i) were significantly lower than those on the non-diversion day in
2014 (Fig. 6h; One-way ANOVA, p < 0.05). On both the water di-
version and the non-diversion days, the Cyanophyta predominated in
all the three sampling regions (Fig. 6g, h, and i).
J. Dai, et al. Journal of Hydrology 587 (2020) 124938

Table 2
The name and affiliations of genera and their average relative proportions in the investigated regions on the water diversion days of different seasons.
Water diversion days Group (x)* Genus (a, b, c)† Phylum

2014-01-14 GB (4) Westella spp. (0, 2.0%, 0); Tetrachlorella alternans (0, 0.3%, 0) Selenastrum spp. (0, 0.08%, 0); Chodatella spp. (0, 0.2%, 0) Chlorophyta
WR-GB (0) / /
WR-GB-O (10) Microcystis spp. (24.8%, 25.3%, 20.3%) Cyanophyta
Chlorella spp. (5.8%, 5.3%, 3.0%); Crucigenia spp. (2.8%, 2.7%, 0.05%); Chlamydomonas spp. (0.7%, 0.06%, 2.0%) Chlorophyta
Navicula spp. (2.3%, 0.6%, 0.8%); Synedra spp. (0.7%, 2.2%, 2.2%); Aulacoseira granulate (25.5%, 24.0%, 21.9%) Bacillariophyta
Cyclotella spp. (32.9%, 27.9%, 7.0%)
Aphanothece spp. (3.9%, 3.3%, 12.9%) Cryptophyta
Mallomonas spp. (0.7%, 0.3%, 0.3%) Chrysophyta
2015-01-17 GB (3) Eudorina spp. (0, 0.3%, 0); Tetrachlorella Korshikov (0, 0.4%, 0) Chlorophyta
Gyrosigma spp. (0, 0.6%, 0) Bacillariophyta
WR-GB (2) Westella spp. (5.7%, 2.0%, 0) Chlorophyta
Phacus spp. (0.4%, 1.0%, 0) Euglenophyta
WR-GB-O (8) Chlorella spp. (10.4%, 2.1%, 3.0%); Ankistrodsemus spp. (0.4%, 0.7%, 0.6%); Schroederia spp. (1.6%, 0.3%, 0.1%) Chlorophyta
Navicula spp. (0.8%, 0.3%, 0.8%); Synedra spp. (3.6%, 3.8%, 2.2%); Cyclotella spp. (70.4%, 56.2%, 7.0%) Bacillariophyta
Aphanothece spp. (4.7%, 1.5%, 12.9%) Cryptophyta
Synura spp. (0.7%, 0.7%, 0.7%) Chrysophyta
2013-08-18 GB (9) Merismopedia spp. (0, 11.0%, 0); Raphidiopsis curvata (0, 0.02%, 0) Cyanophyta
Westella spp. (0, 3.3%, 0); Cocconeis spp. (0, 0.02%, 0); Ankistrodsemus spp. (0, 0.03%, 0); Acrochaetium arcuatum (0, Chlorophyta
0.01%, 0); Actinastrum spp. (0, 0.3%, 0)
Fragilaria spp. (0, 0.02%, 0) Bacillariophyta
Dinobryon spp. (0, 0.03%, 0) Chrysophyta
WR-GB (3) Arthrospira spp. (46.7%, 4.4%, 0) Cyanophyta
Golenkinia spp. (0.2%, 0.02%, 0) Chlorophyta
Neidium spp. (0.2%, 0.05%, 0) Bacillariophyta
WR-GB-O (7) Pseudanabaena spp. (3.3%, 4.6%, 1.9%); Oscillatoria spp. (40.2%, 9.2%, 4.0%) Cyanophyta
Pediastrum spp. (1.9%, 0.7%, 2.6%) Chlorophyta
Synedra spp. (1.2%, 0.3%, 0.1%); Aulacoseira granulate (0.7%, 3.1%, 1.0%); Cyclotella spp. (3.3%, 1.5%, 0.4%) Bacillariophyta
Aphanothece spp. (0.2%, 0.4%, 0.01%) Cryptophyta
2014-11-21 GB (3) Tetraedron spp. (0, 0.2%, 0); Schroederia spp. (0, 0.1%, 0) Chlorophyta
Tribonema minus (0, 0.6%, 0) Xanthophyta
WR-GB (3) Ankistrodsemus spp. (2.6%, 0.3%, 0) Chlorophyta
Phacus spp. (0.5%, 0.2%, 0) Euglenophyta
Synura spp. (7.1%, 0.5%, 0) Chrysophyta
WR-GB-O (9) Microcystis spp. (1.0%, 60.2%, 77.1%) Cyanophyta
Chlorella spp. (9.7%, 1.1%, 2.1%); Scenedesmus spp. (4.1%, 2.2%, 5.0%) Chlorophyta
Navicula spp. (1.0%, 0.2%, 0.02%); Synedra spp. (5.1%, 1.7%, 0.7%); Aulacoseira granulate (27.3%, 14.6%, 8.4%); Bacillariophyta
Cyclotella spp. (12.1%, 3.1%, 2.2%)
Aphanothece spp. (13.8%, 6.3%, 3.6%) Cryptophyta
Mallomonas spp. (1.0%, 0.2%, 0.2%) Chrysophyta
2015-11-24 GB (5) Cocconeis spp. (0, 0.1%, 0); Closterium spp. (0, 0.6%, 0) Chlorophyta
Cymbella spp. (0, 0.1%, 0); Coscinodiscus spp. (0, 0.06%, 0) Bacillariophyta
Phacus spp. (0, 0.06%, 0) Euglenophyta
WR-GB (0) / /
WR-GB-O (4) Navicula spp. (6.1%, 1.9%, 0.02%); Synedra spp. (9.7%, 3.8%, 0.7%); Aulacoseira granulate (38.2%, 11.4%, 8.4%); Bacillariophyta
Cyclotella spp. (27.9%, 30.6%, 2.2%)

GB: Only found in Gonghu Bay on the water diversion day.

WR-GB: Only overlapped between Wangyu River and Gonghu Bay on the water diversion day.
WR-GB-O: Overlapped among Wangyu River and other regions.
The dominant genera and their average relative proportions were in bold font.
* The value in the bracket is the number of genera in this group according to the Fig. 5.
†
The a, b and c are the average values for the relative proportions of the genus in the Wangyu River, Gonghu Bay on the water diversion day and in the Gonghu
Bay on the non-diversion day, respectively.

For November, there was no significant divergence in the total c).

phytoplankton cell abundances in the Gonghu Bay among the three
sampling days (Fig. 6j, k, and l; One-way ANOVA, p > 0.05). On the
non-diversion day in 2013, the average relative proportions of the Ba-
cillariophyta in the Gonghu Bay (Fig. 6j) were significantly lower than
those on the two water diversion days in 2014 and 2015 (Fig. 6k and l;
One-way ANOVA, p < 0.05).
Although the Cyanophyta predominated in the water regions stu-
died during both diversion and non-diversion periods in August, the
dominant Cyanophyta genera in the Wangyu River and the Gonghu Bay
on the water diversion day were different from those on the non-di-
version days (Fig. S1). The Planktothrix spp., Merismopedia spp., and
Arthrospira spp. were dominant in the Wangyu River and in most of the
regions in the Gonghu Bay on the water diversion day (Fig. S1a), while
the Microcystis spp. was the absolute dominant genus in the three water
regions studied during the non-diversion days in summer (Fig. S1b and
According to the NMDS and ANOSIM results, there were significant
differences in the phytoplankton community structures of the Gonghu
Bay among the water diversion and non-diversion days in each season
(Fig. 7a, b, and c), except for the comparison between the community
structures in November 2013 and 2014 (Fig. 7d). However, for January
and August, the R values of the ANOSIM between the water diversion
and non-diversion days were relatively higher than those between the
non-diversion days in each season (Fig. 7a and c).
3.4. Correlations between physicochemical habitats and phytoplankton
communities
In January, the values of water temperature, TN, NH3-N, and SRP
were significantly correlated with the variations in the phytoplankton
communities in the Gonghu Bay (Fig. S2a). In August, the pH, TN, water

7
J. Dai, et al. Journal of Hydrology 587 (2020) 124938

Fig. 6. Relative proportions of different phytoplankton phyla and phytoplankton cell abundances along the gradient from the Wangyu River to the central lake on the
sampling days in different seasons. (R and L represent the regions of the Wangyu River and the central lake, respectively. A, B, C, D, and E represent the cross sections
from sites A1-A3 to sites E1-E3. Jan, Apr, Aug, and Nov are the abbreviations of January, April, August, and November, respectively.)

temperature, and silicate were the significant physicochemical para-
meters correlated with the variations in the phytoplankton commu-
nities in the Gonghu Bay (Fig. S2b). As for November, the values of
water temperature, TN, SRP, pH, and NO3-N were significantly corre-
lated with the variations in the phytoplankton communities of Gonghu
Bay among the three sampling days (Fig. S2c).
The physicochemical parameters studied herein explained 39.8%,
37.2%, and 46.7% of the variances in the phytoplankton communities
for the water diversion and non-diversion days in January, August, and
November, respectively (Fig. 8a, 8b and 8c). The physicochemical
parameters sensitive to the water diversion explained 23.3% (Fig. 8a),
24.5% (Fig. 8b), and 31.3% (Fig. 8c) of the variances in the phyto-
plankton communities in January, August, and November, respectively.
4. Discussion
4.1. Responses of phytoplankton diversity to WDYT in different seasons
The lacustrine phytoplankton diversity, as an important ecological
indicator in lakes, is sensitive to water diversion in different seasons
(see Fig. 4). The effects of water diversion on the phytoplankton
diversity in lacustrine ecosystems has been reported in many previous
studies (Fornarelli et al., 2013; Lin et al., 2003; Wang, 2008; Jiang,
2013). Most of these studies have observed an increase in the diversity
in water-receiving aquatic ecosystems (Lin et al., 2003; Wang, 2008;
Jiang, 2013). Our study has also revealed that the phytoplankton di-
versity on the water diversion days was higher than that on the non-
diversion days in August and November (see Fig. 4). However,
Fornarelli et al. (2013) reported significant positive correlations be-
tween the diversity indexes (Shannon-Weaver index and evenness) of
phytoplankton and the retention time in a reservoir influenced by inter-
basin water transfers. This finding suggests that the increase in diversity
induced by water diversion might be reasonable in the context of ap-
propriate retention time.
The allochthonous species and physicochemical habitat variation
are often regarded as the two critical contributors to variations in
phytoplankton communities in freshwater ecosystems (Dai et al., 2018;
Paerl et al., 2016; Yang et al., 2017). The phytoplankton species only
found in the Gonghu Bay on the water diversion days were the main
contributors to the increase in diversity induced by water diversion in
different seasons (see Fig. 4 and Table 2). This result revealed that the
variations in physicochemical habitat of the Gonghu Bay contributed

8
J. Dai, et al. Journal of Hydrology 587 (2020) 124938

Fig. 7. The non-metric multi-dimensional scaling analysis (NMDS) and analysis of similarities (ANOSIM) among the phytoplankton communities in the Gonghu Bay
on the water diversion days in January (a), April (b), August (c), and November (d), respectively. (Jan, Apr, Aug, and Nov are the abbreviations of January, April,
August, and November, respectively.)

more to the phytoplankton diversity increase on the water diversion
days. The lacustrine habitat disturbed by the water diversion was
mainly reflected in the variations in the hydrodynamic and physico-
chemical conditions. Since WDYT has been found to mainly influence
the water age and circulate flow field in the Gonghu Bay under limited
inflow discharges (Li et al., 2011; Lv, 2013), the horizontal and vertical
water exchange in the Gonghu Bay may be the dominant factor shaping
the phytoplankton communities during the water diversion periods.
This was supported by the observation that most of the phytoplankton
genera only found in the Gonghu Bay on the water diversion days were
affiliated with the Chlorophyta and Bacillariophyta (see Table 2).
Compared with the Cyanophyta species, most of the Chlorophyta and
Bacillariophyta species have faster growth rates and are more adaptive
to the turbulent flowing environment (Istvánovics and Honti, 2012).
Moreover, the decrease in organic pollutants and increase in active
silicate and nutrients during the water diversion periods can also pro-
vide beneficial physicochemical conditions for some of the Chlorophyta
and Bacillariophyta species (Rao et al., 2018; Xu et al., 2017; Zhang
et al., 2018). However, the phytoplankton species only found in the
Gonghu Bay were not dominant (i.e. most of them with relative pro-
portions lower than 1%) during the water diversion days in different
seasons (see Table 2), which revealed that these species might be rare
ones originally survived in Lake Taihu. When the environmental con-
ditions vary during the water diversion periods, the cell abundances of
these rare species will increase to the detectable level for microscopic
counting. Additionally, in January, there was no pronounced difference
in diversity among the three sampling days (see Fig. 4), which was
ascribed to the fact that the sampling day on the non-diversion period
was 10 days after the previous water diversion period (TBA, 2013), and
the influence of the previous water diversion on diversity may have
remained to some extent.
4.2. Impacts of WDYT on lacustrine phytoplankton abundances and
communities
The efficacy of water diversion in reducing the phytoplankton cell

Fig. 8. Variations in the phytoplankton

communities of the Gonghu Bay explained
by water diversion-sensitive physicochem-
ical parameters (W) and other physico-
chemical parameters (O) between water di-
version and non-diversion days in January,
August, and November. (W|O means the
variation portion explained by only water
diversion-sensitive physicochemical para-
meters; O|W means the variation portion
explained by only other non-sensitive phy-
sicochemical parameters; W ∩ O means the
common variation portion explained by
water diversion sensitive parameters and
other parameters.)

9
J. Dai, et al.
abundance and primary productivity has been reported for some lakes,
such as Lake Green (Oglebsy, 1969) and Lake Moses (Welch et al.,
1992) in the state of Washington, USA, and Lake Nanhu in Changchun
city, China (Wang, 2008). The lower phytoplankton cell abundance,
especially of the Microcystis spp., in the turbulent flowing water en-
vironment of rivers is the main reason (Huisman et al., 2004; Li et al.,
2013). The water mixing of the Wangyu River and the Gonghu Bay
directly diluted the phytoplankton cell abundance in the Gonghu Bay.
In this study, the planktonic cell abundances in the Wangyu River were
always lower than those in the lake regions in different seasons (see
Fig. 6). On the other hand, it is also important to note that water di-
version enhances the turbulent diffusivity of water-receiving regions,
which could promote the vertical mixing of planktonic cells (Huisman
et al., 2004). Since the buoyant Microcystis spp. is not well adaptable to
the turbulent mixing environment (Paerl et al., 2011; Reynolds et al.,
1994), the proliferation of the Microcystis spp. could be precluded to
reduce the phytoplankton cell abundance in the water-receiving lake
region.
The phytoplankton community composition is another index re-
presenting the planktonic response to biotic and abiotic disturbances
(Ko et al., 2017; Yang et al., 2017). Due to the influences of climate
warming and anthropogenic activities, in the eutrophic Lake Taihu, the
Microcystis spp. predominated the northern regions for over ten months
per year (Ma et al., 2016). The stronger adaptation to water tempera-
ture and sufficient nutrients in Lake Taihu create opportunities for
forming frequent Microcystis blooms almost throughout the year (Qin
et al., 2018). In this study, water diversions in different seasons were all
found to be capable of decreasing the relative proportions of the Mi-
crocystis spp. and increasing the proportions of the other phyla (e.g.
Bacillariophyta and Chlorophyta) in most regions of the Gonghu Bay
(see Fig. 6 and Fig. S1). This phenomenon, i.e. shaping the phyto-
plankton communities of lakes by water diversion, has also been re-
ported for other eutrophic lakes, such as Lake Veluwe in the Nether-
lands (Jagtman et al., 1992), Lake Nanhu in China (Wang, 2008), and
Lake Tega in Japan (Amano et al., 2010). However, in the case of Lake
Taihu and under conditions of only limited inflow discharges (i.e.
average 90 m3/s during water diversion periods in the last 10 years)
(TBA, 2015), this ecological effect was just limited to the Gonghu Bay
and the lake center was marginally influenced by the water diversion.
However, on the protection of the drinking water source in the Gonghu
Bay, the ecological effect has been found to be positive (Qin et al.,
2010).
Since many phytoplankton species have only a short life-cycle
period, i.e. several days, and are sensitive to physicochemical condi-
tions (e.g. the Microcystis spp., Scenedesmus spp., and Cyclotella spp.)
(Reynolds, 2006), variations in the physicochemical habitat are the
critical impetus to drive the succession of the phytoplankton commu-
nities in freshwater lakes (Li et al., 2019). In the present study, the
physicochemical parameters studied in the Gonghu Bay explained
about 40% of the variations in the phytoplankton communities among
the water diversion and non-diversion days in each season (see Fig. 8).
The variations in the phytoplankton communities in the Gonghu Bay
were explained about 23.3%–to 31.3% by the physicochemical habitat
shifts induced by water diversion in different seasons (see Fig. 8). As for
November and January, the dominant Bacillariophyta in the Gonghu
Bay can be partially explained by the increase in active silicate con-
centrations on the water diversion days (Zhang et al., 2018). For Au-
gust, however, the increase in the other Cyanophyta species abun-
dances, except for the Microcystis spp., can be mainly attributed to the
input of the allochthonous species from the Wangyu River. Most of the
variations (i.e. about 60%) in the phytoplankton communities between
the water diversion days and the non-diversion days cannot be ex-
plained by the variations in the physicochemical parameters studied
(see Fig. 8). Therefore, the allochthonous species as input from the
Wangyu River should be another reason for the variations in the phy-
toplankton communities induced by water diversion.
10
Journal of Hydrology 587 (2020) 124938
In this study, the average relative proportions of the Chlorella spp.,
Aulacoseira granulate, Cyclotella spp., Planktothrix spp., Aphanothece
spp., and others in the Wangyu River of the five water diversion days
were significantly higher than those in the Gonghu Bay on both the
water diversion and non-diversion days (see Table 2). Further, because
the average relative proportions of these genera in the Gonghu Bay on
the water diversion days were also higher than those in the Gonghu Bay
on the non-diversion days (see Table 2), the allochthonous species from
the Wangyu River definitely contributed to the increase of these genera,
at least to some extent. However, due to the only limited sampling time
of each water diversion period, quantitative assessment on the con-
tributions of the allochthonous species to the variations in the phyto-
plankton community compositions in the Gonghu Bay on water diver-
sion days requires further exploration.
For the water diversion day in summer, the dominant groups of
phytoplankton genera in the Gonghu Bay were Merismopedia spp. and
Planktothrix spp., while Microcystis spp. predominated in the non-di-
version period (Table 2). It is worth noting that Merismopedia spp. be-
longs to the phytoplankton functional group L0 (Reynolds et al., 2002),
which is commonly found in the epilimnion of mesotrophic lakes. The
common habitat of the Planktothrix spp. is also the shallow lake with
frequent hydrodynamic disturbance. Moreover, it can be seen that
diatoms predominated in the Gonghu Bay on the water diversion days
in autumn and winter (see Fig. 6). Among the diatom genera, Aulaco-
seira granulate and Cyclotella spp. were the dominant groups and belong
to phytoplankton functional groups P and A, respectively (Reynolds
et al., 2002). In general, Aulacoseira granulate is mostly found in shallow
eutrophic lakes and has good adaptability to hydrodynamic dis-
turbance. Cyclotella spp., on the other hand, is commonly found in well-
mixed clear water lakes and has good tolerance to lake habitats with
nutrient limitation. These genera dominant in the Gonghu Bay during
the water diversion period imply the potential impact of water diver-
sion on the habitat conditions of Gonghu Bay, which might be more
conducive to the health of the lake aquatic ecosystem.
4.3. Implications for water diversion management in the Taihu Basin
During the past three decades, over forty water diversion projects
for national and regional water resource allocation have been con-
structed or under construction in China. As a typical water diversion
project for regional water supply, the WDYT also provides the ad-
vantage for alleviating cyanobacterial blooms in the large eutrophic
Lake Taihu (Hu et al., 2008; Li et al., 2011; Qin, 2009). However, the
allochthonous nutrient input from the Yangtze River and the tributaries
of the Wangyu River (Dai et al., 2018) brings negative impacts on the
water-receiving lake regions. The government has made investments on
hydraulic engineering construction and ecological restoration to im-
prove the water quality of the Wangyu River. In our study, the con-
centrations of NO3-N and TP in the Wangyu River were found to be
significantly higher than those in the Gonghu Bay on the water diver-
sion days (see Fig. 3). Since our monitoring data revealed that the
concentrations of NO3-N and TP in the western tributaries of the
Wangyu River were significantly higher than those in the Yangtze River
and Lake Taihu, the construction projects controlling water pollution of
the west bank of the Wangyu River could offer benefits in improving the
water quality of the Wangyu River. However, if the government wants
to enhance the use of the WDYT through importing water from the
Yangtze River by other water diversion channels, such as the Xinmeng
River and the Xingou River, it would be very important to first in-
vestigate how to reduce the nitrogen and phosphorus concentrations in
these inflow channels. In the middle and lower reaches of the Yangtze
River, due to the anthropogenic influences, the nutrient concentrations
in most of the rivers are higher than that in lakes (Wang et al., 2019).
Thus, to reduce the ecological risks of water diversion projects, con-
trolling the pollutant afflux from tributaries and increasing the self-
purification capacity and time of the inflow river channels are two
J. Dai, et al.
critical measures. Furthermore, long-term monitoring needs to be
continuously carried out to assess the ecological effects of water di-
version projects in China.
For the WDYT, according to the dispatching control water level,
water diversion activities are always run in the fall and winter of each
year, supplying water for the basin (TBA, 2013, 2014, 2015). During
summer and spring, due to the flood control requirement of the Taihu
Basin, water diversion activities are rare and always taken as the
emergency measure to deal with the Microcystis blooms aggregated in
the drinking water sources in the Gonghu Bay (TBA, 2013). In our
study, in the fall of 2015, the effect of decreasing the relative proportion
of the Cyanophyta in the Gonghu Bay was observed after 13 days of
diversion with a 58.8 m3/s average inflow (see Table 1). Although the
effect of decreasing Cyanophyta proportions in the winter of 2014 and
2015 was also evident, allochthonous nutrient inputs from the Wangyu
River also increased with the higher inflow during the winter of 2014
and 2015. Before resolving the problems associated with the high nu-
trient loads in the Wangyu River, controlling the inflow of the Wangyu
River can decrease the nutrient loads into Lake Taihu. However, how to
coordinate the multiple objectives, including the ecological restoration
of Lake Taihu, water supply of the Taihu Basin, and flood control, needs
further exploration. Appropriate diversion discharges, diversion dura-
tion times, and diversion seasons should be evaluated to enhance the
positive ecological effects and reduce the negative impacts.
5. Conclusions
The following conclusions are drawn from this study.
(1) The Water Diversion from the Yangtze River to Lake Taihu (WDYT)
reduced the organic pollutant concentrations in the water-receiving
Gonghu Bay in different seasons, but the nitrate and phosphorus
levels in the inflow to the Wangyu River were significantly higher
than those in Lake Taihu. Allochthonous nutrients from the Yangtze
River and the western tributaries of the Wangyu River should be cut
down with the help of water pollution control projects.
(2) Water diversion activities can profoundly increase the phyto-
plankton diversity and alter the community composition of the
Gonghu Bay in different seasons. In fall and winter, the
Bacillariophyta, instead of the Cyanophyta, dominated in the water-
receiving Gonghu Bay, while the relative proportions of the non-
Microcystis Cyanophyta species significantly increased in summer.
(3) Allochthonous input of the phytoplankton species and the physi-
cochemical habitat disturbance were identified to be the two major
reasons for the variations in phytoplankton communities in the
water-receiving regions. The physicochemical habitat disturbance
induced by water diversion potentially contributed 23.3% to 31.3%
of the community variations. The highest potential contribution of
the allochthonous species to the increase in phytoplankton diversity
in the Gonghu Bay was about 15.8%, while the physicochemical
habitat disturbance contributed about 12.0%–31.3% of the phyto-
plankton diversity in the Gonghu Bay.
(4) Before resolving the widespread problem of high loads of al-
lochthonous nutrients in river networks in the Taihu Basin, the
WDYT can only be taken as an urgent and temporary measure to
alleviate cyanobacterial blooms in Lake Taihu. However, due to the
limitation in this field study, how to conduct the ecological op-
eration of the WDYT in different seasons remains unknown.
Furtherly, the extensive investigations in other similar lakes and the
harmonization among the multiple objectives including flood con-
trol, water supply, and water environment improvement, should be
conducted for better management of water diversion projects in
China.
11
Journal of Hydrology 587 (2020) 124938
CRediT authorship contribution statement
Jiangyu Dai: Conceptualization, Investigation, Formal analysis,
Writing - original draft. Shiqiang Wu: Supervision, Project adminis-
tration, Funding acquisition. Xiufeng Wu: Project administration,
Funding acquisition. Xueyan Lv: Investigation, Resources. Bellie
Sivakumar: Writing - review & editing. Fangfang Wang: Investigation.
Yu Zhang: Formal analysis. Qianqian Yang: Investigation, Resources.
Ang Gao: Investigation. Yuhang Zhao: Investigation. Lei Yu:
Investigation. Senlin Zhu: Writing - review & editing.
Declaration of Competing Interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influ-
ence the work reported in this paper.
Acknowledgements
We are grateful to the Hydrology and Water Resources Investigation
Bureau of Wuxi for their help in the field survey and sample collection.
This work was jointly funded by the National Key R&D Program of
China (2018YFC0407200), the Projects of the National Natural Science
Foundation of China (51679146), the Young Elite Scientists
Sponsorship Program by CAST (2017QNRC001), and the Special
Research Fund of the Nanjing Hydraulic Research Institute (Y118001;
Y120010). We thank the editor and three reviewers for their con-
structive comments and useful suggestions on this manuscript.
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://
doi.org/10.1016/j.jhydrol.2020.124938.
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