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Box 1. Tutorial
This paper contains five sections. The “Introduction” briefly describes the background of the review.
Section2 is about river damming impact on fish physical habitat, including river connectivity, hydrolog-
ical regime, sediment regime and morphology, water temperature regime, and dissolved gas. Section3
is about river damming impact on key fish species across different continents, including salmonids,
Chinese carps, sturgeon, eel and lamprey, and other typical fish species. Section4 focuses on major
conservation measures, which include fish passage facilities, artificial breeding and release, reservoir
ecological operation, and habitat compensation in tributaries, for fish in dammed rivers. The last section
highlights future research perspectives on impact assessments and mitigations, effect of climate and
land cover changes, and long-term systematic observations.
Abstract River damming has brought great benefits to flood mitigation, energy and food production, and
will continue to play a significant role in global energy supply, particularly in Asia, Africa, and South America.
However, dams have extensively altered global river dynamics, including riverine connectivity, hydrological,
thermal, sediment and solute regimes, and the channel morphology. These alterations have detrimental effects
on the quality and quantity of fish habitat and associated impacts on aquatic life. Indeed, dams have been
implicated in the decline of numerous fishes, emphasizing the need for effective conservation measures.
Here, we present a global synthesis of critical issues concerning the impacts of river damming on physical
fish habitats, with a particular focus on key fish species across continents. We also consider current fish
conservation measures and their applicability in different contexts. Finally, we identify future research needs.
The information presented herein will help support sustainable dam operation under the constraints of future
climate change and human needs.
Plain Language Summary River damming yields great social-economic benefits, but also causes
significant eco-environmental impacts, particularly on fish. Dams block fish migration routes, alter hydrological
and water temperature regimes, and modify channel morphology. These changes impact fish physical habitats
and associated communities. Here, we synthesize the impacts of river damming on fish physical habitats, and
review potential conservation measures that could be used to off-set or mitigate the impacts of dams on fish
habitats, populations and communities.
CHEN ETAL.
© 2023. The Authors.
This is an open access article under
the terms of the Creative Commons
Attribution-NonCommercial-NoDerivs
License, which permits use and
distribution in any medium, provided the
original work is properly cited, the use is
non-commercial and no modifications or
adaptations are made.
River Damming Impacts on Fish Habitat and Associated
Conservation Measures
Qiuwen Chen1,2,3 , Qinyuan Li3 , Yuqing Lin1,2,3, Jianyun Zhang1, Jun Xia4,
Jinren Ni5 , Steven J. Cooke6, Jim Best7 , Shufeng He3, Tao Feng3, Yuchen Chen3, Daniele Tonina8 ,
Rohan Benjankar9 , Sebastian Birk10 , Ayan Santos Fleischmann11 , Hanlu Yan3, and Lei Tang12
1Yangtze Institute for Conservation and Green Development, Nanjing Hydraulic Research Institute, Nanjing, China, 2State
Key Laboratory of Hydrology-Water Resources and Hydraulic Engineering, Nanjing Hydraulic Research Institute, Nanjing,
China, 3Center for Eco-Environment Research, Nanjing Hydraulic Research Institute, Nanjing, China, 4State Key Laboratory
of Water Resources and Hydropower Engineering Sciences, Wuhan University, Wuhan, China, 5The Key Laboratory of Water
and Sediment Sciences, Ministry of Education, Peking University, Beijing, China, 6Department of Biology and Institute of
Environmental and Interdisciplinary Science, Carleton University, Ottawa, ON, Canada, 7Departments of Earth Science and
Environmental Change, Geography and GIS and Mechanical Science and Engineering, and Ven Te Chow Hydrosystems
Laboratory, University of Illinois at Urbana-Champaign, Urbana, IL, USA, 8Center for Ecohydraulics Research, University of
Idaho, Boise, ID, USA, 9Civil Engineering Department, Southern Illinois University Edwardsville, Edwardsville, IL, USA,
10Faculty of Biology, Aquatic Ecology, and Centre for Water and Environmental Research, University of Duisburg-Essen,
Essen, Germany, 11Mamirauá Institute for Sustainable Development, Tefé, Brazil, 12College of Water Resources, North China
University of Water Resources and Electric Power, Zhengzhou, China
Key Points:
• Dam construction alters river
hydro-geomorphological conditions
and hence influences fish habitat
quality and quantity
• Knowledge of river
hydrogeomorphology and reservoir
properties can inform which
conservation measures may benefit
fish conservation
• Long-term monitoring is needed
to understand causal effects and
synergies with climate change
Supporting Information:
Supporting Information may be found in
the online version of this article.
Correspondence to:
Q. Chen and J. Zhang,
qwchen@nhri.cn;
jyzhang@nhri.cn
Citation:
Chen, Q., Li, Q., Lin, Y., Zhang, J., Xia,
J., Ni, J., etal. (2023). River damming
impacts on fish habitat and associated
conservation measures. Reviews of
Geophysics, 61, e2023RG000819. https://
doi.org/10.1029/2023RG000819
Received 23 JUN 2023
Accepted 29 NOV 2023
Author Contributions:
Conceptualization: Qiuwen Chen
Data curation: Shufeng He
Funding acquisition: Qiuwen Chen
Project Administration: Yuqing Lin
Supervision: Qiuwen Chen, Jianyun
Zhang
Validation: Yuqing Lin
Visualization: Qinyuan Li, Yuchen Chen,
Hanlu Yan
Writing – original draft: Qiuwen Chen,
Qinyuan Li, Yuqing Lin, Shufeng He, Tao
Feng, Yuchen Chen, Lei Tang
10.1029/2023RG000819
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Box 2. Glossary
ATT: accumulated temperature threshold for fish gonad maturity.
ATU: accumulated thermal units.
CI: connectivity index. For each river basin, CI is quantified for each fish species by combining its
occurrence range with a high-resolution hydrography and the locations of the dams (Barbarossa
etal.,2020).
CSI: connectivity state index. CSI of river reaches is determined with four dimensions, including
longitudinal connectivity between up- and downstream, lateral connectivity to floodplain and ripar-
ian areas, vertical connectivity to groundwater and atmosphere, and temporal connectivity based on
seasonality of flows (Grill etal.,2019).
CTT: critical temperature threshold for fish spawning.
FMCCs: four major Chinese carps.
GBD: gas bubble disease.
IHA: indicators of hydrologic alteration.
LHPs: large hydropower plants.
MPUE: mass per unit effort.
NbS: nature-based solutions.
NFR: natural flow regime.
RVA: the range of variability approach.
SHPs: small hydropower plants.
TDG: total dissolved gas.
TGR: Three Gorges Reservoir.
1. Introduction
Since the first masonry dam was constructed around 3000 BCE on the Wadi Rajil at Jawa, Jordan, river damming
has made significant positive contributions to the supply of water and energy, flood management, irrigation, and
navigation worldwide, especially in the last century (Best, 2019). The first global boom of dam construction
occurred after the Second World War, peaking in the 1960s and 1970s, mainly in Western Europe and North
America (Figures1a and1b; Lehner etal.,2011). Following increasing concerns about the social and ecological
impacts caused by river damming, the trend of global dam construction slowed in the 1990s (Moran etal.,2018).
However, to meet the rapidly growing demands for energy and water for socio-economic development, there has
been a second boom in dam construction, mainly in developing countries and emerging economies in Asia, Africa
and South America (Figures1c and1d), and particularly in large river basins such as the Amazon, Congo, and
Mekong (Winemiller etal.,2016; Zarfl etal.,2015). To date, approximately 3,700 major dams are either planned
or under construction, and the number is predicted to increase further (Zarfl etal.,2015), although removal of
aged dams has become an issue for engineering security and possible restoration of damaged river ecosystem in
some countries (Habel etal.,2020; O'Connor etal.,2015). According to the International Commission on Large
Dams (ICOLD, https://www.icold-cigb.org/), the number of registered large dams (dam height ≥15m, or 5–15m
with reservoir capacity ≥0.03km
3) worldwide reached 58,713 by April 2020. Dams can be categorized accord-
ing to their height or the associated reservoir regulation capacity. Large dams usually convert the upstream lotic
rivers into lentic reservoirs, and have an annual to seasonal regulation capacity that can significantly modify the
hydrological regime downstream; small dams (dam height <15m) and run-of-river dams typically have a small
reservoir, or even no reservoir, and are associated with weekly to sub-daily regulation capacities that impose rela-
tively low impacts on the hydrological regime downstream (D. Anderson etal.,2015; Timpe & Kaplan,2017).
River damming disrupts free flows, and hence results in habitat fragmentation (Grill etal.,2015,2019). Globally,
63% of the world's large rivers (>1,000km) are no longer free flowing (Grill etal.,2019). The construction of
over 3,700 expected hydropower dams is estimated to increase global river habitat fragmentation to 93% in the
future, which has already reached 48% due to the 6,374 existing large dams (Grill etal.,2015; Zarfl etal.,2015).
Writing – review & editing: Qiuwen
Chen, Qinyuan Li, Yuqing Lin, Jianyun
Zhang, Jun Xia, Jinren Ni, Steven J.
Cooke, Jim Best, Shufeng He, Daniele
Tonina, Rohan Benjankar, Sebastian Birk,
Ayan Santos Fleischmann
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The seasonal and interannual dynamics of river flow and water temperature regimes have been greatly dampened
by river dam operation (Poff etal.,2007). As the reservoir operating time increases, sediment is deposited in
reservoirs, while the downstream riverbed tends to experience scour, eventually altering the river morphology
(Kondolf etal.,2014,2018; Schmitt etal.,2019). Moreover, the flood discharge and energy dissipation associated
with high dams can result in supersaturation of total dissolved gas in the downstream waters (Q. Ma etal.,2018;
Weitkamp & Katz,1980).
The alterations of river geophysical conditions caused by dams have significant effects on river ecology, among
which the impact on fish is of great concern because fish contribute to human welfare as key food resources.
Inland fisheries provide the equivalent of all the dietary animal protein for 158 million people globally (McIntyre
etal.,2016), particularly for poor and undernourished populations in food-insecure regions such as the Amazon
(Winemiller etal.,2016) and Mekong (Orr etal.,2012; Ziv etal.,2012) basins, where river fish are a key source
of protein. For example, in the Lower Mekong River basin, fisheries are the primary source of protein for 60
million residents (Ziv etal.,2012). Rivers also provide essential habitats for a diverse array of fishes. Alterations
in river geophysical conditions arising from dams have considerably impaired fish biodiversity and resources
worldwide (Reid etal., 2019; G. H. Su etal.,2021), as dams modify the natural flow, thermal and sediment
regimes, and decrease fish access to spawning and nursery habitats (Freeman etal.,2007). These alterations
cause dramatic reductions in fish productivity and lead to declines in fish populations, threatening global fishery
production and regional food security. The effects of river damming on fish populations have received consider-
able attention (Fullerton etal.,2010), with many studies investigating impacts on globally important commercial
and endangered fish species, such as salmonids (Hilborn,2013), eels (Atkinson etal.,2020), Chinese carp (Q.
Chen etal.,2021), and sturgeon (Z. Huang & Wang,2018).
To mitigate the negative impacts of river damming on fish, a variety of conservation measures, spanning off-setting
to mitigation, have been developed and applied. Reservoir operations have been optimized to mimic natural flow
regimes (NFRs) to satisfy fish living conditions (W. Chen & Olden,2017; Sabo etal.,2017), and fish passage
facilities have been installed at river dams to maintain biological connectivity (Katopodis & Williams,2012;
Noonan etal.,2012). Some small dams and barriers have been removed in tributary channels to rehabilitate and
Figure 1. Global dam construction and distribution. (a) Distribution of global dams. (b) Number of global total dams and number of dams constructed in each decade
from 1900 to 2017. (c) Distribution of global dams in planning and under construction. (d) Number of dams under planning and construction in each continent. Data
of panel (a) and (b) are from Global Reservoir and Dam Database (Lehner etal.,2011). Data of panel (c) and (d) are from Future Hydropower Reservoir and Dams
Database (Zarfl etal.,2015).
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compensate for fish habitats lost in the dammed mainstems (Marques etal.,2018; L. Tang etal.,2021). Artificial
breeding programs have been implemented to restore fishery resources, which to some extent have succeeded
by increasing the target fish populations (Holsman etal.,2012; J. Yang etal.,2013). However, the efficiency
and effectiveness of these conservation measures have not been well investigated, leading to some controversial
perceptions concerning fish conservation measures in dammed rivers. Therefore, assessing the efficiency and
cost-effectiveness of different conservation measures could contribute significantly to identifying and promoting
efficient practices.
The number of studies documenting the impacts of river damming on fish habitats, fisheries resources, regional
food insecurity, and associated conservation measures has rapidly increased in the past decade. This review seeks
to analyze and synthesize state-of-the-art studies concerning the impacts of river damming on fish physical habi-
tats and associated conservation measures. Specifically, we focus on the geophysical aspects of hydrological and
water temperature regimes, dissolved gas supersaturation, sediment dynamics and biogeomorphology in relation
to fish habitats, with emphasis on several key fish species that occur on various continents. In addition, we assess
the cost-effectiveness of existing fish conservation measures. Finally, we formulate future key research directions
for fish conservation in dammed rivers, with the aim of helping foster sustainable hydropower development.
2. River Damming Impacts on Fish Physical Habitat
Dam construction alters river hydro-geophysical characteristics, including river connectivity, hydrological
regime, sediment regime, and morphology, water temperature regime, and dissolved gas, which have essential
impacts on fish physical habitat.
2.1. Impacts on River Connectivity
River longitudinal and lateral connectivity play a vital role in maintaining the structure and functioning of river-
ine ecosystems (Díaz etal.,2020). River systems are hierarchical tree-like networks, whose ecological function
is highly dependent on physical connectivity (Fuller etal.,2016; Grant etal.,2007). The needs of fish for diverse
habitats are strongly dependent on river connectivity and natural mobility (Arthington etal.,2016). Longitudinal
connectivity is essential for fish migration (Figures2a and2b), and lateral connectivity provides fish the access
to spawning and rearing grounds in floodplains, side channels, oxbows and floodplain lakes (Figure2c).
With the increasing number of dams, the longitudinal connectivity of global rivers is significantly under threat.
Currently, about half of global river reaches show diminished longitudinal connectivity, with the Connectivity
State Index (CSI) below 100%; nearly 10% of global river reaches have CSI below 95%, which is the minimum
value for a high level of connectivity (Grill etal.,2019). Large river networks with completely natural connectiv-
ity (CSI=100%) exist only in remote regions of the Arctic, Amazon and Congo basins (Grill etal.,2019). During
the second half of the 20th century, the impairment of river network connectivity became more severe and is now
widespread throughout the entire pan-European continent. There are at least 1.2 million river barriers (an average
density of 0.74 per km) in the 36 European countries (Belletti etal.,2020), and in general more than 50% of river
length is affected (Duarte etal.,2021). The highest barrier densities are found in rivers in central Europe where
river connectivity has been severely altered, whilst the lowest barrier densities are found in the most remote and
less populated alpine areas. Relatively undisturbed rivers exist only in parts of the Balkans, Baltic States, and
Scandinavia in Europe (Belletti etal.,2020). As far as large rivers (catchment area >10,000km
2) are concerned,
the Mediterranean and Western Atlantic regions are those most affected by fragmentation in terms of the number
of basins, while the Black Sea and Caspian Sea regions are the most affected by fragmentation in terms of river
length (Duarte etal.,2021). In the USA, large dams have increased river segmentation by 801% compared to
free-flowing streams in the absence of dams, and 79% of stream length is disconnected from their outlet of oceans
or Great Lakes (Cooper etal.,2017). In South America, more than a hundred hydropower dams have fragmented
the rivers in the Amazon basin. For the eight major river systems in the Andean region of the Amazon headwaters,
the 142 existing or under construction dams have fragmented the tributaries of six major river systems, and the
160 proposed dams could further result in a significant loss of river connectivity in the mainstreams of five major
river systems (E. Anderson etal.,2018; Flecker etal.,2022; Latrubesse etal.,2017,2020; Lees etal.,2016).
In general, the Connectivity Index (CI) of rivers is the lowest in the Europe, United States, South Africa, India
and China, and the completion of the dams currently under construction or planned will further reduce river
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connectivity, particularly in countries such as India and China that are facing a boom in dam construction.
However, commensurate studies are insufficient in these developing countries (Barbarossa etal.,2020).
Dam construction also reduces lateral connectivity of rivers, which decreases the interactions between rivers
and floodplains/wetlands in river basins (Latrubesse etal.,2017; Stoffels etal.,2022), and thereby affects the
productivity of floodplain and wetland ecosystems (Palmer & Ruhi,2019). In the Atreyee River basin in India,
the active floodplain area was reduced by 66.2%, and 48.9% of total pre-dam wetland was completely obliterated,
due to the reduction in lateral connectivity caused by the Mohanpur Dam (Saha etal.,2022). In the wet-dry
tropics of Australia, dam construction reduces the average duration of the lateral connectivity between flood-
plain wetlands and their main river channels by 1% and 2% in the Flinders and Gilbert catchments, respectively
(Karim etal.,2015). In the upper Paraguay River basin in South America, the Manso dam decreased the lateral
connectivity between the Cuiaba River and Pantanal wetlands, one of the largest wetland systems in the world,
which weakened the exchange of sediments and nutrients (Jardim etal.,2020). Regulated releases also impact
on the vertical connectivity between surface and subsurface waters, which alters the exchange of solute, heat and
nutrient between surface and sediment waters (Figure2d) (Sawyer etal.,2009).
It is also necessary to emphasize river fragmentation caused by small dams, due to their large number and wide-
spread distribution, which are usually neglected (E. Anderson etal., 2018; Castello & Macedo, 2016; Fuller
etal.,2016; Rodeles etal.,2017). Couto & Olden(2018) estimate that 82,891 small hydropower plants (SHPs)
are operating or under construction across 150 countries, which is more than the number (58,713 by April 2020)
of large hydropower plants (LHPs) recorded by ICOLD. In addition, there are 181,976 SHPs planned, 10,569 of
which are to be implemented in the coming decades, indicating that the number of SHPs will continue to increase
rapidly. In developed countries, as the hydropower potential of large rivers has been mostly exploited, an increasing
number of SHPs are under planning in some of these nations, such as Austria, to meet energy demands (Wagner
etal., 2015). In water-rich countries in Asia, Africa, Latin America and southeastern Europe, the potential of
SHPs has gained particular interest from policymakers and more SHPs will be constructed (Harlan etal.,2021).
SHPs are conventionally considered as a type of clean energy resource with less environmental impact than LHPs
(Dursun & Gokcol,2011; Nautiyal etal.,2011). However, SHPs are usually set in high-gradient alpine streams
and a river basin usually contains a large number of SHPs, which lead to cumulative, and hence more severe,
impacts on river fragmentation than LHPs (Timpe & Kaplan,2017). For instance, the average loss of connectivity
Figure 2. River connectivity and the impacts of dams. (a) Migration and spawning grounds of fish in free-flowing river.
(b) Migration and spawning grounds of fish in dammed rivers, (c) Spawning and rearing grounds of fish in floodplain. (d)
Hyporheic exchange and redd habitats.
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due to SHPs is much higher than that due to LHPs in Brazil (Couto etal.,2021). Great concerns have been raised
about the large expansion of SHPs in rivers feeding the Pantanal wetlands, as these SHPs have affected the lateral
connectivity between rivers and wetlands (Figueiredo etal.,2021).
River fragmentation has significant impacts on freshwater fish (Barbarossa etal.,2020). Dams act as physical
barriers in spawning or foraging routes and limit the expansion of fish populations (Figure2b). Widely distributed
dams have impeded fish migration and generated population isolation, leading to declines in fish populations and
ultimately to local or total extinction, especially for migratory fish species (Duponchelle etal.,2021; Rodeles
etal.,2017; van Puijenbroek etal.,2018). Over the past 50 years, freshwater migratory fishes have declined by
96% globally, becoming the vertebrate group suffering the most severe decline (Deinet etal.,2020). Studies
concerning the impacts of river fragmentation on migratory fish have mainly analyzed the life history of certain
migratory species, measured their habitat characteristics at different life stages, and assessed the potential impacts
of damming on their habitats (Goodwin etal.,2014; Liermann etal.,2012; Wofford etal.,2005). River frag-
mentation also leads to fish populations becoming isolated and genetically fragmented, exposing them to severe
effects of genetic drift and inbreeding (Brinker etal.,2018; Cheng etal.,2015). Moreover, river fragmentation
changes the structure of riverine food webs, reducing the taxonomic diversity of fish (Freedman etal.,2014).
The beta diversity of native fishes in pools and non-native fishes in riffles decreased with an increase in the ratio
between the length of the longest non-fragmented sections in the river network and the total length of the river
network (Díaz etal.,2020). The decreased lateral connectivity due to dam construction causes loss of access
for some fish species to their spawning and rearing grounds in floodplain and wetlands (Figure2c) (O'Mara
etal.,2021). In the Amazon River, decreases in river-floodplain interactions due to dam construction have caused
significant reduction in catch-per-unit-effort and shifts in the functional composition of fisheries in the Madeira
River floodplain (C. C. Arantes etal.,2021). In addition to fragmentation effects of large dams, widespread small
dams also impose significant, particularly cumulative, impacts on fish distribution and diversity (Consuegra
etal.,2021). Studies show that two-thirds of 191 migratory fish species in Brazil would be affected by river
fragmentation due to the increasing number of SHPs, which is greater than the connectivity loss caused by LHPs
(Couto etal.,2021).
The impacts of river fragmentation on fish diversity and distributions at larger scales (e.g., continental and global
scales) may differ from that at local scales (e.g., basin and sub-basin scales), as fish beta diversity shows signifi-
cant differences among river networks with similar degrees of connectivity (Díaz etal.,2020). In recent years, the
effects of dam-induced river fragmentation on fish, and the mechanisms by which fish diversity responds to river
fragmentation at large scales, have become new areas of interest. Grill etal.(2015) assessed the effects of river
fragmentation on fish habitat at high spatial resolution from sub-basin to watershed scales. Given current progress
in river connectivity studies from watershed to continental scales (Belletti etal.,2020; Duarte etal.,2021; Grill
etal.,2019), it is anticipated that research concerning the impacts of river fragmentation on fish diversity and
distributions will quickly expand to global scales.
Most current studies concerning river fragmentation adopt physical connectivity indices (e.g., CSI, CI); however,
the hydrological and hydraulic disconnections of dammed rivers should receive sufficient attention as they
impose significant impacts on fish habitats. In addition, the impacts of cascade dams on river connectivity and
their cumulative effects on fish are typically much greater than a single dam, and thus deserve more investigation.
At present, indices of physical connectivity are mainly based on geo-spatial data, which may be insufficient in
resolution or possess inadequate records for small dams. As such, the impacts of small dams on river connectivity
worldwide are seriously underestimated, and demand urgent attention.
2.2. Alterations to River Hydrological Regimes
Hydrological regime, including variables such as discharge, flow velocity, water depth, and peak flow, plays an
essential role in riverine bio-habitats and ecosystems (Y. Chen etal.,2016). Fish spawning, rearing and wintering
are strongly related to hydrological conditions. For fish species spawning drifting eggs, continuous stimulation
of flow velocity is required, and thus flooding processes can provide favorable conditions for their reproduction
(Young etal.,2011). With sufficient flow velocity, drifting eggs are not susceptible to sinking, which improves
their survival rate (George etal.,2017). River flow can mediate the dispersal of fish eggs, extending their survival
range and promoting the stability of fish communities (Castello & Macedo,2016). The annual rise and fall of
river depth induces lateral exchange of materials between the river channel and floodplain, which extends fish
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physical habitat and increases baits for them (Castello & Macedo,2016; Paillex etal.,2013). In cold regions,
fish require water bodies with sufficient depth that have warm water in the deeper layers for overwintering (Cott
etal.,2008).
River hydrological regime depends mainly on climatic conditions and possesses distinct features in different
regions (Fiseha etal.,2014). In tropical regions, most rivers have a large annual runoff and their discharge
possesses strong interannual fluctuations with little seasonality (J. P.Syvitski et al.,2014); some rivers have
well-defined high and low discharges in correspondence to a unimodal rainy period, such as the Ganges River in
south Asia and the Purus River in the Amazon basin; some rivers have two peak flows per year in correspond-
ence to bimodal rainy periods, such as the Magdalena River in South America and the Congo River in Africa
(Latrubesse etal.,2005; J.P.Syvitski etal.,2014). In subtropical and temperate regions, rivers usually have a
relatively smaller annual runoff than those in tropical regions, and their discharges have distinct seasonality that
is influenced by seasonal rainfall and snowmelt, such as the Yangtze River and Yellow River in eastern Asia (W.
Yang etal.,2020). In cold regions, rivers are characterized by low flow in winter due to ice cover and high flow
in spring due to ice-jam floods (Peters etal.,2014; Prowse,2001), such as the Mackenzie River in the North
America and Lena River in Russia. In some rivers, peak flows are dominated by snowmelt in spring (from March
to May), and these rivers are mainly distributed in southern North America, eastern Europe and westernmost
Asia; peak flows may appear near June due to relatively late snowmelt (Hansford etal.,2020), and these rivers
are located mainly in regions of low altitude and mid-high latitude (Q. Liu etal.,2022). Peak flows can also be
brief and intense as determined by monsoon rains, and these rivers are mainly distributed in eastern and southern
Asia, and eastern Australia (Dettinger & Diaz,2000). In some rivers, there are no significant peak flows due to
stable rainfall or mixed climates, such as the St. Lawrence River in central-eastern North America and rivers in
southern Finland (Haines etal.,1988).
Dam construction may significantly alter river hydrological regimes (Timpe & Kaplan,2017). The conversion
from lotic river to lentic reservoir upstream of a large dam fully modifies the original hydrological regime.
Dams directly decrease river flow velocity (Y. Yang etal.,2017). Stevaux etal.(2009) reported that the annual
mean flow velocity of the Paraná River in South America was 0.88m/s in the pre-dam period, and decreased
to 0.56m/s after dam construction. Reservoirs impound water in wet seasons and release it in dry seasons, thus
decreasing the seasonal variability of discharges in rivers (Figure3a). The discharge of the Mekong River in the
dry season is now 63% higher than that in the pre-dam period, while the discharge in the wet season has declined
by 22% (Chong etal.,2021). Although run-of-river reservoirs do not alter the seasonal pattern of discharge
(Figure3b), they can significantly increase the variability of diel or daily discharge through hydropeaking oper-
ations (R. M. Almeida etal.,2020). River damming decreases the number and duration of peak flows, and alters
the frequency of water level variability (Timpe & Kaplan,2017). After the construction of the Gezhouba Dam
and Three Gorges Dam, the number of flow pulses in the downstream Yangtze River decreased by 22%, and their
maximum duration decreased from 16days to 4–6days (Y. Wang etal.,2016). River damming decreases the
maximum discharge and increases the minimum discharge of river, resulting in reduced water level fluctuation
zone (Poff etal.,2007). River damming may also reduce the extent of active floodplain, the period and duration
of flooding, as well as the exchange of materials between main channel and floodplain (Jardim etal.,2020; Moi
etal.,2020). Dam construction in the Balonne River in Australia has resulted in a 23% loss of active floodplain
area and decreased the availability of nutrients from the floodplain (Thoms,2003). To quantify alterations to river
hydrological regime incurred by dam construction, Richter etal.(1996, 1998) have developed the Indicators of
Hydrologic Alteration (IHA) method (Table1), which is widely used to evaluate alterations to the hydrological
regime of dammed rivers. Based on the IHA, the range of variability approach (RVA) has been developed to
evaluate the degree of alteration within specific ranges (Richter etal.,1997).
Alterations to hydrological regime caused by river damming could detrimentally impact the spawning, migration
and feeding behavior of fish (Mims & Olden,2012). A decrease in flow velocity caused by river damming has
been found to reduce the stimulation for fish spawning, resulting in declines of fish reproduction (Figures3c
and3d). In dammed rivers, fish gamma diversity, which indicates the regional or total diversity of fish, is nega-
tively correlated to magnitude in flow velocity (Jarzyna & Jetz,2016; McGarvey & Terra,2016; Timpe &
Kaplan,2017). For fish that fertilize in vitro, the decelerated flow of dammed rivers can reduce the success
rate of fertilization (Campton,2004). Decreases in flow velocity may lead to sinking of drifting eggs or failure
to reach their destination for successful hatching (George etal.,2017). For fish species spawning sticky eggs,
artificial hydropeaking incurred by reservoir operation can cause unsuitable conditions for their reproduction
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(Vilizzi,2012). The reduced seasonal variability of discharge in dammed rivers can decrease floodplain produc-
tivity, which leads to poor growth and low survival of the fry, resulting in reduction of fish populations (Ficke
etal.,2007; Reinfelds etal.,2013; Terra etal.,2010). Meanwhile, the degradation of plants in floodplain areas
reduces the matrix for sticky eggs to attach, leading to a decrease in the survival of the eggs (Perna etal.,2012).
The decrease in high and low flow pulses and their duration due to river damming can eliminate the hydrolog-
ical cues for fish migration, which reduces foraging opportunities and increase the risk of fish stranding (Bao
etal.,2022; Reid etal.,2019). Rapid reductions of river water surface elevation due to dam operation may result
in stranding of fish species (E. Bell etal.,2008; Irvine etal.,2015). Such fish stranding has been reported during
flow reductions downstream of dams, leading to mortality of salmonids and sturgeon (Johnston etal., 2020;
Nagrodski etal.,2012). Stranding is mainly caused by changes in flow magnitude, known as ramping rates in
dam operations (Le Coarer etal.,2022; Poff etal.,1997). However, the potential for fish stranding also depends
on factors such as fish species and their life stages, stream temperature, and the time of the day, and thus fish
stranding may be site specific (Auer etal.,2022; Benjankar etal.,2023; Glowa etal.,2022). In newly constructed
reservoirs, the decomposition of organic matter can provide more bait, and thus omnivorous fishes can increase
in the short term (Bunn & Arthington,2002; Junk etal.,2013). The increase in water depth and alteration of the
substrate textures in reservoirs reduce the suitable habitat for periphytic algae and benthic macroinvertebrates
(Holt etal.,2015; Taniwaki etal.,2013). This affects the fish feeding on periphytic algae and benthic macroin-
vertebrates, and increases the number of fish preferring phytoplankton and zooplankton, thereby altering the fish
community structure in dammed rivers (W. Zhang etal.,2020). In winter, the rise of water depth in dammed
rivers buffers the drop of water temperature, increasing the chances of fish survival during overwintering (Fuchs
Figure 3. (a) Hydrograph of the Pingshan hydrological station (averaged monthly discharge during 2007–2010, free-flowing)
and the Xiangjiaba hydrological station (replacement of Pingshan hydrological station, averaged monthly discharge during
2016–2020, regulated by a large reservoir) of the upper Yangtze River. (b) Hydrograph of the Ningnan hydrological station
of the Heishui River, a tributary of the upper Yangtze River, before (2015) and after (2019) the removal of the Laomuhe
Dam (a small hydropower dam with no reservoir regulation capacity). (c) Discharge from the Three Gorges Reservoir (TGR)
and fish egg density of four major Chinese carps (FMCCs) measured at the Yidu cross-section of the Yangtze River during
the ecological operation of the TGR in 2016. (d) The number of annual spawned eggs of FMCCs measured at the Yidu
cross-section of the Yangtze River before and after the operation of the TGR. The data of panel (a) and (b) are available from
Hydrological Data of Changjiang River Basin in Annual Hydrological ReportP.R. China, and the data of panel (c) and (d)
are from the authors.
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etal.,2021; Keefer etal.,2008). In summary, alterations to river hydrological regime caused by dam construction
impact significantly on fish habitats, further affecting the population and diversity of fish in dammed rivers.
Existing studies mainly focus on the impact of altered hydrological regime on the spawning aspects of fish
reproductive biology. However, adequate attention should be paid to the impact of altered hydrological regime on
the hatching processes of spawned eggs in dammed rivers, which also plays an essential role in early-stage fish
resources. The reduced water levels during the flood season diminish material exchanges between channels and
floodplain, which reduces feeding field and food abundance of fish. In addition, most available studies investigate
the behavioral response of fish to hydrological alterations, and it is imperative to better understand the physiolog-
ical mechanisms of fish reactions to dam-induced hydrological alterations.
IHA statistics group Regime characteristics Hydrologic parameters Examples of river ecosystem influences
Magnitude of monthly discharge
conditions
Magnitude Mean discharge for each calendar month Habitat availability for fish
Timing Influences water temperature, oxygen
levels, and photosynthesis in water
Magnitude and duration of annual
extreme discharge conditions
Magnitude Annual minima (1-day means; 3-day means;
7-day means; 30-day means; 90-day means)
Balance of competitive, ruderal, and
stress-tolerant fish
Structuring of river ecosystems by abiotic
versus biotic factors
Duration Annual maxima (1-day means; 3-day means;
7-day means; 30-day means; 90-day means)
Structuring of river channel morphology
and physical habitat conditions
Number of zero-flow days Volume of nutrient exchanges between
rivers and floodplains
Duration of stressful conditions (e.g., low
oxygen, concentrated chemicals) in
river ecosystem
Seven-day minimum flow divided by mean flow
for year (base flow)
Duration of high flows for aeration of
spawning beds in river sediments
Timing of annual extreme discharge
conditions
Timing Julian date of each annual 1-day maximum
discharge
Compatibility with life cycles of fish
Predictability/avoid ability of stress for fish
Access to special habitats during
reproduction or to avoid predation
Julian date of each annual 1-day minimum
discharge
Spawning cues of migratory fish
Evolution of life history strategies and
behavioral mechanisms
Frequency and duration of high/low flow
pulses
Magnitude Number of high pulses each year Availability of fish habitat in floodplain
Nutrient and organic matter exchange
between rivers and floodplains
Frequency Number of low pulses each year Soil mineral availability
Duration Mean duration of high pulses within each year Water birds enter foraging, resting, and
breeding places
Mean duration of low pulses within each year Bed load transport, channel sediment
structure, and substrate disturbance
duration (high pulses)
Rate/frequency of hydrograph changes Means of all positive differences between
consecutive daily values
Entrapment of fish on islands and
floodplains (rising levels)
Frequency Means of all negative differences between
consecutive daily values
Rate of change Number of flow reversals Desiccation stress on low-mobility stream
edge fish
Table 1
Summary of Hydrologic Indicators and Their Ecological Influences (Richter etal.,1998)
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2.3. Changes to River Sediment Regimes and Morphology
Sediment is an essential component in rivers, and plays a pivotal role in maintaining the ecological status of global
river systems (Chapman & Wang,2001; Förstner etal.,2004,2008; Netzband etal.,2007; Ralph etal.,2009).
River sediment is entrained, transported and deposited, thus shaping river morphology and affecting fish habitat
(Nichols,2009). The balance between sediment supply and transport capacity of a river system is a fundamental
driver for river geomorphology, which not only dictates the aggradational or degradational state of a system,
but also controls channel morphology and substrate textures (Dietrich etal.,1989; Lisle etal.,1993; Pitlick &
Wilcock,2001). River planform can adopt a range types, form straight to sinuous, braided, anabranching or anas-
tomosing shapes (Latrubesse,2008; Leopold & Wolman,1957). These different river planforms induce different
hydrodynamic conditions, which in turn cause different patterns of sediment erosion and deposition, forming a
variety of geomorphic units such as riffles, pools, barforms and bedforms, and flood plains, which thus increase
the diversity of biological habitats, for example, those used for fish spawning and wintering grounds (Chapuis
etal.,2015; Namour etal.,2015). Rivers also transport large quantities of organic matter, providing food sources
for aquatic organisms (Karr,1991). The content of organic matter in sediment is affected by sediment character-
istics, including particle size and density, surface site density, and particle morphology (Y. Wu etal.,2020). In
most rivers, bed sediments show an overall downstream-fining trend (Luo etal.,2012), as illustrated in Figure4a.
Sediment from global rivers delivered to the oceans was estimated to be about 20 Bt/yr, prior to significant dam
interception (Milliman & Syvitski,1992). Dams alter the natural balance of sediment flux in rivers by trapping
sediment in reservoirs, discharging waters often free of sediment downstream (Morris & Fan, 2010). Global
dams decreased about 5Gt/yr of sediment flux to the oceans by the 21st century (Milliman & Syvitski,1992).
Upstream of a dam, the reduction in flow velocity may facilitate sediment deposition in the river channel and
floodplain (Fencl etal.,2015; X. Su etal.,2017; Walter & Merritts,2008). As shown in Figure4b, coarse parti-
cles, such as gravel and coarse sand, are the first to settle, forming a delta at the point where the backwater effect
ends; fine sediment particles enter the reservoir and are transported by turbid density currents or non-stratified
flow, and may be deposited near the dam (Fan & Morris,1998; Garde & Raju,1979). In addition, the peak
sediment load in dammed rivers is separated in time from the maximum water flow (Dang etal.,2010; Topping
etal.,2000), which also promotes the deposition of sediment. In the river downstream of a dam, the reduction
in sediment content usually leads to channel incision, chronic erosion of the bed and banks, and even the loss of
delta plain (Bittencourt etal.,2007; Graf,2006; Magilligan & Nislow,2005; Petts & Gurnell,2005). Intensified
scour and floodplain incision are often observed in sediment-starved rivers, as the flows entrain bed material
equaling to their transport capacity (Csiki & Rhoads,2010). In some cases, dams reduce the number of intense
floods and lead to sedimentation downstream, which raises the river bed (Kotti etal.,2016; Słowik etal.,2018).
After damming, pools may occur more frequently and individual pools are longer in the lower reach than that in
Figure 4. Impact of river damming on river bed sediment. (a) Bed sediment in free-flowing rivers. (b) Bed sediment in dammed rivers.
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the upper reach; in contrast, individual riffles in the upper reach are longer than that in the lower reach (Kobayashi
et al., 2012). Low sediment loads downstream after dam construction also leads to reductions in associated
nutrient transport, and thus affects fish feeding grounds (C. Guo etal., 2020). Moreover, the balance between
inorganic and organic sediment is disrupted by damming, with mineral particles deposited predominately in the
reservoirs, and the increased biological production causing the suspended load of the reservoir outflow to be
largely composed of organic matter from aquatic organisms (Sokolov etal.,2020).
Alterations to sediment regimes due to river damming have potential impacts on fish habitat. The changes to
erosion and deposition patterns, and hence the river morphology, may impede suitability for fish spawning and
overwintering grounds (Kruk & Penczak,2003; McLaughlin etal.,2006). For resident or potamodromous fish,
river damming changes the number and distribution of fish habitats for spawning, feeding and overwintering
downstream, leading to increases in competition for spawning and wintering sites as well as food resources
(Cambray etal.,1997). The changes of mesoscale riverbed morphology, such as riffles and pools in the upstream
and downstream, after river damming can indirectly lead to changes in the diversity and distribution of fish
communities (Calderon & An,2016; Langeani etal.,2005). In addition, different fish species may have a differ-
ent preference for sediment properties. For example, larvae of white sturgeon (Acipenser transmontanus) prefer
substrates of clean gravel and cobbles (Nguyen & Crocker,2006), while larvae of Schizothorax wangchiachii
change their preference for substrate at different life stages (Chai etal., 2019). Typically, lamprey spawn in
riverbeds covered with a mixture of sand, gravel, and pebbles (N. S. Johnson etal.,2015). Fine sediment with
relatively high organic matter is a primary source of food and energy for some species, and can even be an inte-
gral requirement within the lifecycles of species such as lamprey ammocete and psammophilous fish. Sediment
coarsening downstream of a dam would result in less bait for these fish (Maitland,2003). Moreover, complex
habitat structure, such as pore space between coarse sediment, can increase predator-free space and thus reduce
predation efficiency, an effect that is most pronounced at low prey densities (Barrios-O’Neill etal.,2015,2016;
Toscano & Griffen,2013). Therefore, the altered river morphology and sediment gradation due to dam construc-
tion may seriously impact fish habitat. The alteration of suspended sediment content can also cause a variety of
effects on fish habitat in dammed rivers. Some fish species prefer turbid over clear water, presumably benefiting
from a reduced risk of predation and increased opportunity of feeding (Cyrus & Blaber,1987,1992). As reser-
voirs convert flowing water to relatively still water, the upstream water changes from turbid to limpid (C. Guo
etal.,2020), and the increase in water clarity can directly affect the habitat of these fish species. In summary,
the alterations of both suspended and bed load sediment regimes have important implications for the whole life
history of fish in dammed rivers.
Despite extensive studies concerning the impacts of sediment regime alterations induced by river damming on
fish, many challenges remain. The effects of changes in transparency and bed color on fish communities after
river damming have not been explored sufficiently. Quantification of the contributions from direct effects, such
as turbidity changes, and indirect effects such as predation behavior changes, also demand investigation. To date,
most studies have focused on the response of fish to alterations in sediment regimes, but neglected the effects
of alterations in fish activity on river sediment and morphology. It has been reported that spawning mucus of
some fish species can modify substrate characteristics and thus affect sediment transport, which highlights the
significance of spawning as a zoogeomorphic activity (Roberts etal.,2020). The effect of changes in spawning
grounds of these fish species on river morphology after dam construction is a topic that demands further study.
2.4. Alterations in River Water Temperature Regimes
Water temperature is an important and highly sensitive factor in river ecosystems, and possesses distinct and
regular seasonality. The natural rhythm of water temperature affects the phenological functioning of aquatic
species, and the longitudinal variation of water temperature plays an essential role in generating spatial patterns
of species communities (Isaak etal.,2012). Water temperature regime can affect the whole life cycle of fish
(Servili et al.,2020), including migration timing, reproductive performance, embryo health, and growth rate
(Figure5c). For migratory fish, the change of water temperature within a suitable range is one of the key envi-
ronmental cues for migration (Harvey etal.,2020; Rijnsdorp etal.,2009). The Atlantic salmon (Salmo salar)
smolts in the three tributaries of the West River in Vermont, USA, begin migration when the water tempera-
ture rises to 5°C, and reach peak migration when water temperature rises to 8°C (Whalen etal.,1999). Fish
have specific reproduction strategies, and a suitable temperature promotes healthy egg development and ensures
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successful spawning (Kurita etal.,2003; Sieiro etal.,2020). For temperate or tropical fishes that spawn in spring
and summer, warm winters could affect the accumulation of nutrients during egg development, resulting in
smaller eggs (Collingsworth etal.,2017; T. M. Farmer etal.,2015). For most cyprinids in temperate and tropical
climates, temperature acts as a time clue for reproduction events, such as ovulation and oviposition (Pankhurst &
King,2010; N. Wang etal.,2010). In addition, changes in plankton and benthos in response to variations in water
temperature directly affects fish food chains, and thus indirectly regulates the energy intake and growth of fish
(Pörtner & Farrell,2008; Prokešová etal.,2020).
The spatio-temporal variation of river water temperature is influenced by multiple factors, including water
flux, climate, latitude and human activity (Caissie, 2006; Collins, 2009; Markovic etal.,2013; B. W. Webb
etal.,2008). The study on a Cairngorm stream in northeast Scotland (Hannah etal.,2004) shows that in terms
of average energy flux affecting water temperature, the main heat sources are sensible heat (38.7%), bed heat
(37.0%) and friction at stream bed and banks (24.3%), while the main heat losses are latent heat (73.1%) and
longwave radiation (26.9%). Typically, the water temperature of headwater streams is often lower than that of
downstream reaches, because many rivers originate in plateaus or snow-covered mountains with water sourced
from ice melting (Cai etal.,2018; Dugdale etal.,2017; Moors etal.,2011). At the global scale, regional trends
of river water temperature are consistent with air temperature (J. Syvitski etal.,2019). In addition, river water
temperature is increasingly affected by climate change (Dugdale etal.,2017; van Vliet, Franssen, etal.,2013; van
Vliet, Ludwig, & Kabat,2013). The response of river water temperature to climate change varies with latitude.
Increases in water temperatures at low latitudes are generally greater than those in high and middle latitudes.
However, water temperature changes in rivers in high-altitude regions, such as the Qinghai-Tibet Plateau (S. Liu
etal.,2020), are different from this broad latitudinal pattern.
River damming can significantly alter water temperature regimes, as illustrated in Figure5a (D. Cheng etal.,2015;
Jung etal.,2022). The alteration of water temperature regime by reservoirs can be influenced by various factors,
including the shape, storage and depth of the reservoir, the inflow water temperature, the hydraulic residence
time, and the operation modes of the reservoir (Lessard & Hayes,2003; Prats etal.,2010; Wotton,1995). The
Figure 5. (a) Water temperature of the Pingshan hydrological station (averaged monthly water temperature during
2007–2010, free-flowing) and Xiangjiaba hydrological station (replacement of Pingshan hydrological station, averaged
monthly water temperature during 2016–2020, regulated by the large Xiangjiaba Reservoir) of the upper Yangtze River.
(b) Water temperature stratification in free-surface (left) and frozen-surface (right) reservoirs. (c) Water temperature in
downstream of the Xiangjiaba Dam on the upper Yangtze River before and after the impoundment of Xiangjiaba Reservoir,
and the phenological properties of fish (OW: overwintering, MG: migration, SP: spawning, GD: gonadal development,
GR: gonadal recovery). Data are available from T. Li etal.(2021). (d) Impact of altered water temperature regime on fish
spawning (CTT: critical temperature threshold for spawning, ATT: accumulated temperature threshold for gonad maturity).
The data of panel (a) are available from Hydrological Data of Changjiang River Basin in Annual Hydrological ReportP.R.
China, and Figure 5d is adapted from T. Li etal.(2021).
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essence of water temperature alteration by reservoirs is that the impounded water is thermally stratified season-
ally and the outflow is mostly from deep layers. Although the impact of reservoirs on river water temperature
regime is complex, the degree of alteration depends principally on dam height and regional climate characteristics
(Maheu etal.,2016b), as summarized in Table S1 in Supporting InformationS1. Reservoirs in different climatic
regions have different effects on the water temperature regimes of dammed rivers (Pieters & Lawrence,2012; D.
Yang etal.,2005). In tropical, subtropical and temperate regions, large reservoirs usually form thermal stratifica-
tion in the spring, summer and autumn, with high temperatures in the surface layer and low temperatures in the
bottom layers (Figure5b). In cold regions, large reservoirs have thermal stratification in freezing periods, mostly
in winter and early spring, with low temperatures in surface layers but relatively high temperatures in the bottom
layers (Figure5b). The seasonal stratification of reservoir waters can, to some extent, alter the natural rhythm
of water temperature downstream (Figure5a), which are characterized by elevated water temperatures in winter,
decreased water temperatures in spring and summer, and reduced amplitude of maximum water temperatures
(Long etal.,2019; Soleimani etal.,2019). In particular, the largest alterations in downstream water temperature
occur in dry seasons (Maheu etal.,2016a). Because of the surface layer discharge mode and non-stratification,
the effects of small dams on river water temperature are different from that of large dams (Maheu etal.,2016b).
The release of surface water from small dams in spring and summer tends to elevate downstream water tempera-
ture (B. W. Webb etal.,2008), which is opposite to water temperature alterations in the summer period caused by
large dams (Skoglund etal.,2011). To quantify the alteration to water temperature in dammed rivers, a variety of
indicators have been proposed, such as changes in suitable water temperature range of target species, and changes
in maximum and minimum water temperatures (T. Li etal.,2021; Maheu etal.,2016a).
Alteration of water temperature regime by dams has serious impacts on river fish habitat (Ahmad etal.,2021;
Couto etal.,2021; Grill etal.,2019; Kuczynski etal.,2017; Prats etal.,2010), which directly affect the spawning,
migration and growth of fish (Figure5c). Fish reproduction is the most vulnerable and sensitive to water tempera-
ture alterations. Due to the operation of the Xiluodu and Three Gorges dams on the upper Yangtze River in China,
the elevated water temperatures in autumn inhibit the reproduction activities of Chinese sturgeon (Acipenser
sinensis) that usually spawn in autumn, causing the effective breeding quantity to reduce to 0%–4.5% (Z. Huang
& Wang,2018). The warming of water during winter due to reservoir operation in northern Europe and North
America leads to delayed timing of spawning and shortened time of egg incubation for winter-spawning Atlantic
salmon and brown trout (Salmo trutta) (Bohlin etal.,1993; Heggenes etal.,2018; Jonsson & Jonsson,2009). In
addition, a phenological knock-on effect could emerge as a consequence of delays in spawning time (Elliott &
Elliott,2010; Skoglund etal.,2011), resulting in low survival rates of Atlantic salmon and brown trout larvae.
In the upper Yangtze River, the ray-finned fish (Coreius guichenoti) spawns in late spring and early summer
when the critical water temperature exceeds 20°C. The timing of the arrival of this critical temperature is delayed
due to the operation of the Xiluodu Reservoir (T. Li etal.,2021). Meanwhile, the warming of water in winter
accelerated the development of gonads and advances the time of egg maturity. The joint effects of warmed water
in winter and cooled water in early summer thus cause ray-finned fish to miss the historical time window for
spawning, resulting in the sharp decline of its population (T. Li etal.,2021; Z. Yang etal.,2021).
The critical temperature threshold (CTT) for fish spawning and the accumulated temperature threshold (ATT) for
fish gonadal development have been used to quantify the impact of water temperature alteration on fish repro-
duction (Figure5d). The ATT is the minimum accumulated temperature required for gonadal maturity, which is
calculated by summing the temperatures higher than the ontogeny temperature in days during gonadal develop-
ment (Chezik etal.,2014; Honsey etal.,2018). For example, the CTT and ATT thresholds for bronze gudgeon
(Coreius heterodon) reproduction in the upper Yangtze River were about 18.4 and 1,324.9°C·day, respectively.
Operation of large reservoirs results in cooling in spring and summer, and warming in autumn and winter (Long
etal.,2019; Y. Tao etal.,2020), which delays the timing for reaching CTT and advances the timing for reaching
the ATT (T. Li etal.,2021). It has been shown that elevated temperatures of 2–3°C in winter can have a signif-
icant impact on the development of fish gonads (T. M. Farmer etal.,2015; T. Li etal., 2022). When the time
to reach the ATT is advanced, fish tend to consume energy in the yolk in order to maintain early exercise and
strengthen metabolism (Mcqueen & Marshall,2017; Wright etal.,2017). When the time to reach the CTT is
delayed and becomes behind the time for reaching the ATT, temperature-sensitive fish cannot spawn in a timely
manner, resulting in over-maturity of their gonads and thereby a decrease in reproduction (Kennedy etal.,2011;
J. King etal.,1998). At present, most studies use the time to reach the CTT as the indicator for reservoir operation
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management (Y. Tao etal.,2020; Y. Wang etal.,2016); however, re-matching the time for reaching both the CTT
and ATT is key to mitigate the impact of water temperature alterations on fish in dammed rivers.
Despite many studies addressing the effects of river damming on water temperature regimes, there is a great
knowledge gap between alterations in water temperature and their impacts on fish, particularly on the whole
life cycle of fish migration, growth and food chain. The limited number studies that document the impacts of
water temperature alterations on fish physiology, such as sex determination and gonad development, have mostly
been conducted at extreme temperature conditions (C. Li etal.,2014; Ribas etal.,2017) instead of natural
temperature conditions (Dorts etal.,2012). In addition, understandings to the long-term impact of water temper-
ature alterations on fish phylogeny are insufficient. In the future, climate change may increase the frequency of
extreme weather events, which could intensify or offset the impacts of reservoir operations on water temperature.
Therefore, the combined effects of river damming and climate change upon water temperature regimes and fish
communities demand further investigations.
2.5. Dissolved Gas Supersaturation
Total dissolved gas (TDG) supersaturation refers to the physical condition that the sum of dissolved gas partial
pressures in water exceeds the sum of gas partial pressures in the air under local atmospheric pressure (Weitkamp
& Katz, 1980). In natural rivers, TDG supersaturation is often caused by a change of water temperature
(Bouck,1984) and water falling (Rowland & Jensen,1988). Dammed rivers can occur severe TDG supersatura-
tion caused by flood discharge during flooding seasons (Figure6a), resulting in serious impacts on fish down-
stream (Ji etal.,2019; Pulg etal.,2016). For instance, the Chief Joseph Dam released flow in excess of the 110%
TDG compliance criterion (EPA,1987) in the Columbia River, causing the downstream Wells Dam to be unable
to comply with water quality criteria for 125 of 133 compliance-mandated days in 2012 (Witt etal.,2017). During
the flood discharge of the Xiluodu Reservoir in the upper Yangtze River in July 2014, the average TDG saturation
of the downstream water was 135%, with a maximum value of 144%, resulting in massive fish mortality in the
downstream reservoir (Q. Ma etal.,2018; Zeng etal.,2020). The release of TDG is slowed with increases in
water depth and decreases in flow velocity. Thus, in a reservoir cascade system, water with supersaturated TDG
from the upstream reservoir is transported to the downstream reservoir and can cause cumulative effects (Q. Ma
etal.,2018). TDG supersaturation effects are thus particularly prominent in reservoir cascade systems.
The generation of TDG supersaturation is mainly affected by water pressure, water temperature, bubble pressure,
bubble retention time, and aeration concentration. The saturation solubility of gas is positively related to water
pressure (J. Feng etal.,2018). When a large volume of water with high head is released from a dam, the water jet
entrains a large amount of air to form aerated water, and drops deeply into the energy dissipation pool near the
dam (Bertola etal.,2018). Due to high water pressure in the pool, the air carried by the water jet dissolves rapidly,
forming TDG supersaturation in the water (Pulg etal.,2016; H. Xue etal.,2018). The degree of TDG super-
saturation in the stilling pool is positively correlated to the bubble retention time (Lu etal.,2019; Y. Peng, Lin,
etal.,2022). The initial saturation of high dam release has little impact on the generation of TDG supersaturation
in the stilling pool, as the water jet is largely saturated after undergoing the process of full gas mixing during the
downward discharge (S. Xue etal.,2019). The influence of aeration on the stable saturation of the water body
Figure 6. Impact of total dissolved gas (TDG) supersaturation on fish. (a) TDG supersaturation due to reservoir flood
dispatch causes fish gas bubble disease. (b) Relation between TDG saturation and half-lethal time of Ya-fish (Schizothorax
prenanti). The data are available from Y. Wang etal.(2015).
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downstream of the dam is also marginal, since the amount of aeration carried by the water jet into the stilling pool
is usually sufficient (Qu etal.,2011).
TDG supersaturated water gradually releases the dissolved gas during its flow downstream, with the release rate
influenced principally by water depth, flow velocity, wind speed and water temperature (C. Cao etal.,2020; Ou
etal.,2016). The release rate of TDG decreases with increase in water depth and decrease in flow velocity. Obser-
vations downstream of the Three Gorges, Ertan and Zipingpu dams, China, showed that the release rate down-
stream of the Zipingpu Dam is the highest, which corresponds to its lowest water depth (J. Feng etal.,2010).
Compared to downstream, the upstream side of a dam has larger water depths and slower flow velocities, result-
ing in lower release rates, as observed in the Kootenay River, North America (Kamal etal.,2018). Increase in
wind speed enhances gas transfer at the gas-liquid interface, which accelerates the release of TDG in the water
(Chu & Jirka,2003; J. Huang etal.,2016). Elevated water temperatures promote the thermodynamic movement
of gas molecules, which can also increase the release of TDG (S. Liu,2013; Ou etal.,2016). Suspended particles
and water plants provide nucleation and aggregation sites for dissociative gas molecules, enhancing free gas
molecules in the water to concentrate and escape in the form of bubbles, and thus promote the release of TDG (J.
Feng etal.,2012; Y. Yuan, Huang, etal.,2018).
When TDG saturation exceeds the sum of atmospheric pressure and hydrostatic pressure, gases in the dissolved
state in fish tissues and body fluids will precipitate and accumulate to form gas bubbles (Figure6a). Gas bubbles
often appear in the tissue of the head, mouth, fins, and gill arches of fish, or in the capillaries of the gill plates
(Lemarie etal.,2011), resulting in gas bubble disease (GBD). GBD could affect the physiology and behavior of
fish, such as loss of balance and abnormal buoyancy, leading to fish death in severe cases (Bouck,1980; J. Huang
etal.,2021). Several fish abnormalities caused by GBD have been reported, such as fast swimming, rapid breath-
ing, back and forth movement, and mouth breathing with sticky bubbles (C. Feng etal.,2019; Y. Yuan, Wang,
etal.,2018). Bleeding may also appear in tissues of gills, fins, muscles, gonads, and intestinal epithelium (Meyers
etal.,2008). These symptoms may impose potential damage on fish, such as tissue necrosis, abnormal growth
(Geist etal.,2013), decreased immunity (Schisler etal.,2000), reduced swimming ability (Y. Wang etal.,2017),
increased risk of predation and increased buoyancy (Shrimpton etal.,1990a,1990b), as well as changes in phys-
iological characteristics (Yuan etal.,2021).
The impact of TDG supersaturation on fish is determined by TDG saturation, exposure time, age and species of
fish, swimming depth, and behavioral habits. High TDG saturation and long exposure time increases the mortal-
ity of fish. The half-lethal time of Ya-fish (Schizothorax prenanti) at 150% TDG saturation is 8.5 times shorter
than that at 120% TDG saturation (Figure6b). Repeated exposure of fish to TDG supersaturated water can lead
to reduced feeding ability and increased susceptibility to fungal and bacterial infections, thereby further reducing
fish tolerance to TDG supersaturated water and affecting fish survival (Brosnan etal.,2016; Huchzermeyer,2003;
Schisler etal., 2000). Fish tolerance to supersaturated TDG also varies between species and growth stages.
During the incubation phase of eggs, increased TDG saturation causes a gradual decrease in the hatching rate (N.
Li etal.,2019; R. Liang etal.,2013). The tolerance threshold of TDG saturation for the juvenile fish of Ya-fish
(Y. Wang etal.,2015), silver carp (Hypophthalmichthys molitrix) (Deng etal.,2020), and Chinese sucker (Myxo-
cyprinus asiaticus) (L. Cao etal.,2016) is higher than that for the juvenile fish of rock carp (Procypris rabaudi
Tchang) (X. Huang etal.,2010) and grass carp (Ctenopharyngodon idella) (F. Wu etal.,2020). However, when
TDG saturation exceeds 135%, the tolerance threshold of TDG saturation for fish is not significantly different
between species and body sizes (S. Xue etal.,2019).
With an increase in water depth, the saturation solubility of TDG increases, and the relative saturation of TDG
decreases. The release of TDG is also related to turbulence, and thus TDG supersaturation is unevenly distributed
vertically (P.Li etal.,2022). Therefore, the tolerance of fish to TDG supersaturation is improved with greater
water depths, which is known as the effect of depth compensation (Yuan etal.,2020). Some fish species can avoid
TDG supersaturation stress using the effect of depth compensation. For instance, when TDG saturation exceeds
120%, Ya-fish, Chinese sucker and elongate loach (Leptobotia elongata) have the ability to detect and avoid TDG
supersaturated water in horizontal and vertical directions (Deng etal.,2020). Rainbow trout (Oncorhynchus
mykiss) has a significantly higher TDG exposure risk than mountain whitefish (Prosopium williamsoni), but
refuge habitats with sufficient water depth can mitigate exposure risk and GBD. To alleviate the stress of TDG
supersaturation, rainbow trout uses depth compensation, with each 1.0m increase in swimming depth offsetting
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9.7% TDG saturation (Pleizier etal.,2020). Therefore, TDG exposure risk and actual risk depend on the interplay
between species-specific ecology and dam-induced TDG patterns (Algera etal.,2022).
TDG supersaturation only occurs occasionally during the discharge of floodwaters, when it is usually risky and
difficult to conduct field surveys (Bertola etal.,2018; Pulg etal.,2016). Therefore, field observation data of
TDG supersaturation are relatively scarce. Also, limited by monitoring technologies, TDG concentration in the
near-field of large dams is difficult to measure well (Algera etal.,2022). Scale models in laboratories possess
substantial difficulties in simulating the processes involved in generation and release of supersaturated TDG in
the field, due to insufficient consideration of the Reynolds and Weber numbers as well as water pressures (P.Li
etal.,2022). Therefore, it remains challenging to model the mass transfer process accurately, and predict bubble
size as well as size distribution in order to support the development of effective mitigation measures. In addition,
the threshold of TDG saturation for fish injury in different tissues requires further investigation. Moreover, fish
adopt an avoidance behavior when TDG saturation exceeds their tolerance threshold, and the tolerance of fish to
TDG supersaturation is enhanced with an increase in water depth. Previous laboratory studies investigating the
response of fish to TDG only consider different TDG concentrations, lacking investigations on depth compensa-
tion effects. Therefore, it is essential to further study the effect of depth compensation, as well as fish physiolog-
ical and behavioral response, in the assessment of TDG supersaturation impacts on fish.
3. River Damming Impacts on Key Fish Species
The interests in the key fish species impacted by river damming vary between different continents. Herein, we
focus on salmonids, Chinese carps, sturgeon, eel and lamprey, which have been intensively studied.
3.1. Impact on Salmonids
Salmonids (family Salmonidae) are one of the most popular commercial fishes, providing high-quality protein to
people around the world, particularly in Europe and the United States (Phillips & Rab,2001). They mainly include
the genera Salmo and Oncorhynchus, which have received most research concerns (Klemetsen etal.,2003). The
genera Salmo contains two typical anadromous species, which are the Atlantic salmon and brown trout. Atlantic
salmon are mainly distributed along both the east and west coasts of the North Atlantic Ocean, while brown
trout is indigenous to Europe, North Africa and western Asia (MacCrimmon etal.,1970; MacCrimmon and
Gots,1979). The genera Oncorhynchus contains the Pacific salmon (Oncorhynchus tshawytscha, named Chinook
salmon; Oncorhynchus keta, named Chum salmon) and Pacific trout (Oncorhynchus mykiss, including rainbow
trout and steelhead trout subspecies), which mainly inhabit the north Pacific and the coastal rivers of both Amer-
ica and Asia (Figure7). The Chinook salmon and Atlantic salmon have received most research attentions due to
the threats of river dams (Harnish etal.,2014; Hilborn,2013; Potter & Crozier,2000).
Figure 7. Distribution of salmonids around the world and their migration route for spawning.
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The anadromous salmonids migrate to freshwater rivers for spawning and juvenile rearing, and to saltwater
oceans for feeding, growing and maturing (Crozier etal.,2021; Groot,1991). The main life cycle of anadromous
salmonids includes a spawning stage (egg, alevin), juvenile stage (fry, fingerling, parr), smolt stage, and adult
stage. In the autumn, female salmonids excavate nest pits (called redds) in river bed gravels and spawn eggs into
them. The optimal spawning temperature for salmonids is in the range of 6–14°C, and the spawning timing is
especially sensitive to temperature changes. Over the winter, the eggs develop into very small salmonids (alev-
ins). In the spring, the alevins swim out of the redds and become fry, with the fry growing into parr that can
protect themselves from predators. The parr grows in freshwater for 2–3years, and transforms into smolts. In
the early spring, silvery smolts swim to the ocean and spend 1–2years maturing into adults. Atlantic salmon
is a cool-water species, and the downstream migrating smolts must swim to the ocean before river tempera-
ture becomes too warm. River discharge plays an important role once migration is under way, and influences
the onset, duration and termination of smolting (Sykes etal.,2009). During the adult stage, salmonids feed
in the open ocean for a certain period of time, which differs between different species. In the summer, adult
salmonids migrate back to the freshwater, strongly following the migratory route they adopted when leaving the
river as smolts (Rivinoja etal.,2001). Increasing flows can facilitate downstream migration of smolts, and flow
augmentation during the upstream migration period can improve the survival of migrating salmonids (Connor
etal.,2003). The upstream swimming capability of salmonids would be reduced at low (below 10°C) and high
(>20°C) temperatures (Alabaster,1990; Johnsen & Jensen, 1994). Therefore, the connectivity between fresh-
water and ocean, water temperature, and river discharge are key factors affecting the physiological behaviors of
anadromous salmonid across their life cycle (Caudill, etal.,2013).
Dams impede both the downstream and upstream migration of salmonids, resulting in the decline of salmonid
populations around the world (Lawrence etal.,2016; Limburg & Waldman,2009). In North America, dams have
been one of the most important factors blamed for the decline of Chinook salmon populations in the Pacific coast
of Oregon and British Columbia (P.H. Wilson,2003). Dams in the upper Columbia River have caused a decline
in the abundance, survival and population of Chinook salmon (Levin & Tolimieri,2001). In South America,
both the catch rate and body size of introduced salmonids decreased in several dammed large rivers in Chile and
Argentina (Arismendi etal.,2019).
Budy etal.(2002) demonstrated that the survival rates of salmonids in dammed rivers decreased mostly in the
smolt-to-adult life stage, rather than in the spawner-to-smolt life stage. When smolts migrate downstream from
the dammed river to the ocean, most of them pass the dams through the spillway, the juvenile bypass system, or
the hydropower turbines. Physiological or behavioral stresses caused by this passage experience on smolts would
result in direct death or a chronic influence, including injury, trauma, diminished physical abilities, and increased
susceptibility to predation and disease, which may eventually lead to death at a later life stage (Budy etal.,2002).
Compared to rivers with a single dam, rivers with cascade dams make juvenile salmonids experience multiple
dam passages, and these cumulative stresses can result in significantly higher mortality (Molina-Moctezuma
etal.,2021). For instance, in rivers with three or four dams, the decline of fish populations can increase to exceed
30% (Lawrence etal.,2016).
When adult salmonids migrate upstream to their natal spawning and rearing habitat, their instinct and imprint of
the downstream passage experience during their juvenile stage leads them to the bypass channel or the turbines,
which increases mortality or results in the situation that they cannot find the correct route, and delays their
migration until they determine the correct route (Rivinoja etal.,2001). On average, there is a 70% loss of poten-
tial Atlantic salmon spawners during their upstream passage at dams in Sweden (Lundqvist et al., 2008). In
Europe, many rivers of the Baltic Sea have lost the natural juvenile reproduction of Atlantic salmon, because river
damming has blocked or reduced the access of adult salmon to their spawning grounds (Rivinoja etal.,2001).
Over several generations, some of the white-spotted charr (Salvelinus leucomaenis) in Japan in dammed-off areas
no longer migrate to the sea and become resident fish due to habitat fragmentation, which leads to a decrease in
their spawning and populations (Morita etal.,2009). Therefore, river fragmentation caused by dams can lead to
the increasingly direct or delayed mortality of salmonids, and thus may severely reduce their population.
Hydrological alterations caused by dam operations may lead to the decline of salmon populations (E. J. Ward
etal.,2015). During the spawning period, the female and male Chum salmon that remain on their redds would
increase swimming activity, reduce digging activity and leave their redds when flow velocities are above a thresh-
old of 0.8m/s. Therefore, artificial hydro-peaks would lead to a decline in spawning rates (Tiffan etal.,2010).
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A significant reduction in discharge under the dam could lead to the stranding of salmonid fry and increase their
mortality (E. Bell et al.,2008). For example, dam operations result in dewatering of Chinook salmon redds,
causing the mortality of eggs and larval fishes (Harnish etal.,2014; Ostberg & Chase,2022; Young etal.,2011).
Regulated flows, even if they do not dewater the redds, can still impact on redd habitats for embryos survival and
development by altering hyporheic exchange of heat and dissolved oxygen (Figure2d) (Bhattarai et al.,2023;
Martin etal.,2020). The upstream migration of adult salmonids may be disrupted by the seasonal and weak
variation in spillway discharge from the dam (Rivinoja etal.,2001). Therefore, alterations to hydrological regime
caused by dams may result in declines in spawning rate, survival of fish eggs and juveniles, and migration of adult
salmonids, thus decreasing the populations of salmonids.
Dam construction can cause the historical spawning grounds of salmonids to become warmer, which makes the
hatchery juveniles progress their life history at an earlier time and thus swim to areas with relatively cool water
temperatures and lower growth opportunities, leading to a decline in their survival and populations (Connor
etal.,2002). However, rising water temperatures in springtime caused by dam operation can negatively affect
the rate of salmonid egg incubation, such as Chinook salmon (Dusek Jennings & Hendrix,2020). Compared to
the daily mean temperature or threshold temperature, temperature experience (accumulated thermal units, ATU)
has more effects on the onset, duration and termination of downstream migration of Chinook salmon smolts
(Sykes etal.,2009). Stich etal.(2015) found that dams decreased the ATU for smolts, thus delaying the time to
initiate their downstream migration. In addition, dams as barriers along migration routes can delay or prolong
the migration of smolts, which may result in a mismatch with migration-timing adaptations. For instance, water
tempera tures in downstream reaches become warmer in the spring due to dam operations, which delays the down-
stream migration of smolts of Atlantic salmon. Consequently, the delayed smolts may face a situation that the
water temperature further increases to a lethal or near-lethal level for smolts (Marschall etal.,2011). Compared
to fish in free-flowing rivers, upstream-migrating fish in impounded systems could encounter potential thermal
barriers in fishways. For instance, fishways offer opportunities for upstream migration of adult Chinook salmon,
but the unfavorable water temperature gradient in the fishway caused by thermal stratification in reservoirs
presents an obstacle to the upstream migration of individuals (Caudill etal.,2013). Therefore, the altered water
temperature regime caused by dams negatively affects egg hatching, the downstream migration of smolts, and the
upstream migration of adult salmonids in fish passages.
The TDG supersaturation caused by flood discharge of dams has little impact on juvenile and adult salmonids
(Geist etal.,2013; Muir et al., 2001). There is little potential of negative effects of TDG supersaturation on
populations of adult Chinook salmon, although fish tissues are probably damaged by the dissolved gases (E. L.
Johnson etal.,2005,2007). Among the fish that pass through dam spillways, the survival of juvenile salmonids
is the highest (Beeman & Maule,2006; Muir etal.,2001).
Overall, most studies focus on the impact of dam construction on the spawning process and migration of salmo-
nid smolts. However, sufficient attention should also be devoted to investigating the direct impact of altered
flow regimes and water temperatures on the upstream migration of adult salmonids, which can provide specific
evidence to improve conservation measures for adult salmonids during upstream migration. Meanwhile, natural
factors such as climate and oceanic conditions also affect the physiological behavior of salmonids. Dam-induced
and natural factors could interact with each other, resulting in nonlinear interdependence (Goodwell &
Bassiouni,2022; Ye etal.,2015). This complex interaction highlights the necessity to better understand how
co-varying dam-induced and natural factors influence the physiological behavior of salmonids across their whole
life cycle. Engineering mitigation measures have been used to reduce the adverse influence of dams on salmo-
nids. However, it is controversial whether these mitigation efforts are effective if differential mortality rates of
salmonids occur for reasons unrelated to river damming (Rechisky etal.,2013; Welch etal.,2008). In the future,
long-term observations of upstream and downstream salmonid are needed to determine whether dams are the
major cause for the decline of salmonids in impounded rivers.
3.2. Impact on Chinese Carps
The four major Chinese carps (FMCCs) are grass carp (C. idellus), black carp (Mylopharyngodon piceus), silver
carp (H. molitrix) and bighead carp (Aristichthys nobilis). FMCCs are freshwater fish whose adults migrate
upstream to spawning sites during flood seasons and spawn semi-buoyant drifting eggs (Figure8). The fertilized
eggs drift a long distance for development, and then juvenile fish swim into riparian lakes, which serve as nursery
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habitats (Q. Chen etal.,2021; W. Xu etal.,2020; B. Yi etal.,1988). The spawning activity of FMCCs requires a
minimum flow velocity to trigger the release of eggs (Q. Chen etal.,2021), and the drifting eggs require a suitable
flow velocity to keep them suspended (Garcia etal.,2013; M. Li etal.,2013; Q. H. Yang etal.,2014;P.Zhang
etal.,2021). High flow events are the most visible physical phenomena associated with the reproductive success
of FMCCs (Coulter etal.,2018; Embke etal.,2016). Under natural conditions, adult fish usually initiate spawn-
ing activity when the water level continues to rise for 0.5–2days, and reduce or even cease spawning when water
levels begin to recede (B. Yi etal.,1988). In China, the Yangtze River basin provides the principal habitats and
spawning sites for the FMCCs, although they are widely distributed across many parts of China. The FMCCs in
the Yangtze River are the dominant natural germplasm resource, which sustains the gene diversity and maintains
freshwater fish aquaculture (J. Wang, Li, Duan, Chen, etal.,2014). Prior to the construction of the Three Gorges
Dam, there were 30 spawning sites for FMCCs in the mainstream Yangtze River (C. Tang etal.,2022), with the
middle reach of Yangtze River being the main reproduction area. There are 12 spawning sites scattered along the
380km long reach from Yichang to Chenglingji, producing about 43% of the total eggs in the Yangtze River (Y.
Yi etal.,2010).
FMCCs play a vital role in providing high-quality animal protein, ensuring national food security, as well as
promoting rural economic development in China (Ban etal.,2019; D. Li, Prinyawiwatkul etal.,2021). However,
FMCCs are an aggressive invasive species in other countries (D. Li, Prinyawiwatkul, etal.,2021), as shown in
Figure8. The silver carp and bighead carp, jointly known as Asian carp, were first imported to North America
in the 1970s as aquaculture fishes. Due to their rapid growth rates and lack of natural predators, they have estab-
lished dense populations in the Mississippi, Ohio, Missouri and Illinois Rivers, and may pose a threat to the Great
Lakes (Heer etal., 2019; Wittmann etal., 2014; Zhu etal.,2018). In Europe, bighead carp are found to have
been widely recruited in northeastern Italy, demanding serious efforts to limit the spread and establishment of
reproducing populations (Milardi etal.,2017). In Australia, Asian carp impose significant impacts throughout the
Murray-Darling basin and other freshwater systems (Marshall etal.,2018).
River damming can block or delay reproductive migration, affect fish assemblages, impair spawning habitat
conditions, and thus reduce the recruitment of FMCCs. After the impoundment of the Three Gorges Reservoir
(TGR) in 2003, the larval abundance of FMCCs in the middle reach of the Yangtze River declined sharply to
less than 20% of the pre-dam abundance (P.Zhang etal.,2021), and the eggs and larvae in the lower reaches of
the Yangtze River declined to 0.34 billion, accounting for only 13% of the pre-dam amount (M. Li etal.,2016).
The Three Gorges Dam reduces river connectivity, preventing FMCCs migration (P.Zhang etal.,2021). Loss
of connectivity in dammed rivers also restricts the gene flow between fish populations located upstream and
downstream of a dam, affecting population genetic structure. Since the construction of the Gezhouba Dam and
Three Gorges Dam, significant genetic differentiations have appeared in the grass carp populations between the
upper and middle reaches of the Yangtze River, which might stimulate population divergence (Zhao etal.,2011).
Although spawning grounds for FMCCs downstream of the dam remain after impoundment, eight spawning
Figure 8. Native and invading distribution of Chinese carps around the world and their migration route for spawning.
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grounds in the reservoir have been lost due to establishment of the lentic ecosystem (Y. Yi etal.,2010). Reservoir
operations cause direct changes in the annual hydrograph and thus water depth and flow velocity, which affect
the spawning activity of FMCCs (Duan etal.,2009; W. Jiang etal.,2010; Z. Wang etal.,2013). Downstream of
the Three Gorges Dam, the suitable flows required for the reproduction of FMCCs are suggested to start from
12,500m
3/s on the day when a spawning event begins, and gradually increase to 18,600m
3/s at day 4 when
spawning actions reach their peak, and then quickly drop to support hatching and larvae survival (Q. Chen
etal.,2021). However, this flow process is mostly not met during the spawning period of FMCCs.
Water temperature has strong impacts on the reproduction of FMCCs. Laboratory studies and field measurements
have shown that the minimum water temperature threshold for gonad development of FMCCs is 18°C, and the
ideal water temperature range for spawning is between 21 and 24°C (Embke etal.,2016; M. Li etal.,2013; Y. Yi
etal.,2010). There is also a significant correlation between spawning time and the arrival date of the cumulative
temperature needed for gonad development. The initial date of FMCCs spawning in the Yangtze River has been
delayed from early May to middle June since operation of the TGR in 2003. The main reason for this delay is that
the water temperature downstream of the dam drops by 2–4°C from March to May, which is the critical period for
gonad development of FMCCs (J. Wang, Li, Duan, Luo, etal.,2014), and thus postpones the arrival date of their
gonad maturity. In addition, flood release could cause the TDG saturation to exceed 120% in the downstream
reach of the Three Gorges Dam during the spawning periods of FMCCs, resulting in the dispersing larvae being
highly vulnerable to the effect of TDG supersaturation. Dead larvae were found due to GBD during the early
operation stage of the Gezhouba Dam between 1981 and 1984 (L. Liu etal.,1986).
However, as invasive species in some regions such as the North America, a viable strategy to prevent the spread
of FMCCs and reduce their population is to manage the existing dams and barriers to limit their dispersion (Fritts
etal.,2021; Whitledge etal.,2019). In the upper Illinois River, reducing gate openings of the locks and dams
during the late spring and summer could provide opportunities to avoid the upstream migration of bighead carp,
and thus limit their recruitment into the upper section of the river (Lubejko etal.,2017). Rather than attempting
to directly block the migration route through dams, another option would be to encourage adult carp to spawn in
the reaches that have suitable spawning sites but lack adequate hydrological conditions to further support embryo
development. Such action could be implemented by adaptive management of hydraulic engineering structures in
rivers (Coulter etal.,2018; Cupp etal.,2021; Prada etal.,2020).
Previous studies attribute the primary cause of decreases in the population of FMCCs in the Yangtze River to the
construction of dams. However, it has also been argued that intensive fishing contributes greatly to their decline
(D. Chen etal.,2009). However, there are few published data on long-term trends in FMCCs catches, making
it difficult to assess whether dam construction or overfishing plays the major role in the degradation of natural
FMCCs resources. Meanwhile, a series of physical barriers, including dams, behavioral deterrents and electric
dispersal barriers, have been applied to block migratory pathways of FMCCs in the rivers in which they have
become invasive, although in practice the effectiveness of this approach is limited. There remains a knowledge
gap in understanding the associated hydrological mechanisms that affect the spawning behavior and early life
development of FMCCs. Studies have indicated that a warm winter-spring period advances the expansion of
Asian carp by altering thermal characteristics to be more favorable for their growth and lessening the time for
competitive interaction with invasive mussels (Alsip etal.,2020; M. Li etal.,2013). It has been observed that the
surface water temperature of the Great Lakes is warming faster than the global rate (Collingsworth etal.,2017;
O’Reilly etal.,2015), and there is thus a great value to investigate whether climate change will increase the risk
of invasion by Asian carp in this region.
3.3. Impact on Sturgeon
Sturgeons are one of the earliest extant vertebrates, and play an important role in the evolution of fishes and even
all vertebrates (Shen etal.,2020). There are 27 species of sturgeons that belong to the Acipenseridae and Poly-
odontidae families. The family Acipenseridae mainly includes Chinese sturgeon (A. sinensis), Russian sturgeon
(Acipenser gueldenstaedtii), Dabry's sturgeon (Acipenser dabryanus), Atlantic sturgeon (Acipenser oxyrinchus),
ship sturgeon (Acipenser nudiventris), sterlet (Acipenser ruthenus), and starry sturgeon (Acipenser stellatus).
The family Polyodontidae, which is commonly called paddlefish, includes the American paddlefish (Polyodon
spathula) and Chinese paddlefish (Psephurus gladius). The number of wild sturgeons in the world has decreased
significantly in recent years, due to the degradation of natural habitats, dam construction, and overfishing (Billard
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& Lecointre,2000; T. Webb & Meyer,2019). Sturgeons are typically anadromous fish that spend most of their
lives in the ocean and return to freshwater to spawn (Figure9). They lay adhesive eggs in rivers with fast currents
and gravel beds. They are subcooled fishes, which prefer to live in water with relatively low temperature (Billard
& Lecointre,2000; McDowall,1997; Siddique etal.,2016), and are mainly found in Eurasia and North America
(Du etal., 2020). According to the International Union for the Conservation of Nature and Natural Resources
(IUCN) Red List of Threatened Species, Dabry's sturgeon has been considered extinct in the wild, Chinese
paddlefish has been considered completely extinct, and 85% of the extant 25 species, such as Chinese stur-
geon, Russian sturgeon, starry sturgeon and sterlet, are endangered (Brevé etal.,2022; IUCN,2022; Lenhardt
etal.,2006; H. Zhang, Jarić, etal.,2020).
Chinese sturgeon has lived in the Yangtze River for about 140 million years, and is often considered a living
fossil. It is a demersal fish species that usually breeds in river reaches with fast currents and gravel beds (Z.
Huang & Wang,2018). Every year, this migratory fish travels 2,000 miles from the East China Sea to its spawn-
ing grounds in the Yangtze River (D. Cheng etal.,2015). The spawning season of Chinese sturgeon is from
October to November, and spawning occurs at water temperatures between 15.3 and 20.0°C, with the optimal
water temperature for spawning being 18.0–20.0°C (Y. Wang etal.,2020). Atlantic sturgeon is one of the seven
sturgeon species found in North America, and extends from New Brunswick, Canada, to the eastern coast of
Florida, USA. Atlantic sturgeon spawns in the spring, and the water temperature requirement for spawning is
13.3–23.0°C (Hager etal.,2014). It spawns in running waters with rocks, pebbles and other hard objects on the
bed, or in pits and pools under waterfalls (Popov,2017). Russian sturgeon is widely distributed in the Caspian
Sea, the Sea of Azov and the Black Sea, as well as in the rivers flowing into these waters (H. Song etal.,2022).
The water temperature requirement for the spawning of Russian sturgeon is 12–14°C (Elhetawy etal.,2020).
Most of the spawning activity of Russian sturgeon takes place in the sloughs of main channels, with a few indi-
viduals spawning in the high tide zone. Sterlet is distributed in the Black Sea, Caspian Sea, Yenisei River and Ob
River in Russia. The optimal water temperature range for sterlet spawning is 13–16°C (Ponomareva etal.,2020),
and the typical spawning grounds of sterlet include riverbed and roaming beach formed by spring water (Lenhardt
etal.,2006). Juvenile sterlets are often found in groups in shallow water, while individual adults are scattered
in deeper water for feeding. White sturgeon is the largest of the eight sturgeon species found in North America.
White sturgeon inhabits the Pacific Ocean from northern Baja California of Mexico to the Aleutian Islands in
Alaska, and the large rivers flowing into the Pacific Ocean between Monterey, California and Alaska. White
sturgeon exhibits freshwater amphidromy, although evidence suggests that only small proportions of their popu-
lations live in marine environments. White sturgeon is an iteroparous broadcast spawner, spawning in the spring
and early summer when water temperatures are between 7 and 18°C (Counihan & Chapman,2018). Therefore,
water temperature and riverbed substrate are major factors that affect sturgeon life cycles, including spawning
migration and reproductive processes.
Figure 9. Distribution of sturgeon around the world and their migration route for spawning.
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Global dam construction has blocked the migration routes and damaged the spawning habitats of sturgeons,
leading to a significant reduction in their populations. The completion of Gezhouba Dam in 1981 and the Three
Gorges Dam in 2003 on the Yangtze River blocked the migration routes, cut off the spawning grounds and
affected the spawning time, spawning size as well as spawning frequency of the Chinese sturgeon, Dabry's stur-
geon and Chinese paddlefish, leading to significant reductions in their populations (Boscari et al., 2022; H.
Zhang, Jarić, etal.,2020). The Gezhouba Dam shortened the migration distance and reduced the capacity of
new spawning grounds of Chinese sturgeon, resulting in the number of hatched larvae being only 25% of that
before dam construction (Z. Huang,2019; J. Tao etal.,2009). The construction of the Volgograd Dam on the
Volga River, Russia, has destroyed the natural spawning area of the Russian sturgeon, starry sturgeon and Beluga
sturgeon, which caused mass mortality of these species. The Volgograd Dam has reduced the area of Russian stur-
geon spawning grounds by 80% (Secor etal.,2000), and the limited spawning area led to high spawning densities,
resulting in egg mortality rates of up to 60% (Popov,2017). In the Don River, Russia, dams cut off the main path
of Russian sturgeon migrating to their spawning grounds, resulting in a sharp reduction in their spawning areas
that are only located downstream of the dam (Boldyrev,2018). At present, Russian sturgeon with natural repro-
duction only exists in the undammed Ural River in Russia and Kazakhstan. In North America, the Eddyville Dam
on the Hudson River has blocked the migratory routes of shortnose sturgeon (Acipenser brevirostrum), leading
to a significant reduction in its population and hence its inclusion in the Endangered Species Act (T. Webb &
Meyer,2019). In the Great Lakes region, dam induced sedimentation in many tributaries has led to a reduction
in spawning grounds and juvenile habitats of lake sturgeon (Acipenser fulvescens), posing great challenges to
the restoration of their population (C. C. Wilson etal.,2022). In Europe, the construction of the Djerdap I and
Djerdap II dams blocked the migration of sturgeon in the Danube River, which seriously reduced populations of
the ship sturgeon and Atlantic sturgeon (Lenhardt etal.,2006). Therefore, river damming seriously compromises
the connectivity of sturgeon habitats worldwide, leading to significant declines in successful spawning and egg
hatching rates, and thus severely decreasing the population of sturgeons.
Water temperature is a primary factor affecting the migration of sturgeons, and determines their spawning time.
However, reservoir operations can alter water temperature regimes and thus affect the migration and spawning
of sturgeon (Y. Wang etal.,2020; H. Zhang etal.,2019). In the Yangtze River, the patterns of water tempera-
ture have been temporally and spatially altered by reservoir operation, which leads to the degradation of gonad
development and delays the spawning of the Chinese sturgeon (Z. Huang & Wang,2018; H. Zhang etal.,2019).
The TGR and Xiluodu Reservoir in the upper Yangtze River have reduced the effective breeding quantity down
to below 4.5% by elevating the water temperature that inhibits breeding activity during the spawning season. The
cumulative effect of the cascade dams, including Wudongde, Baihetan, Xiluodu, Xiangjiaba, Three Gorges and
Gezhouba, has led to an ongoing decline in the abundance of adult Chinese sturgeon in the Yangtze River and
the sea (Chang etal.,2017; Z. Huang & Wang,2018). In the Sacramento River, USA, the optimal water temper-
ature for Chinook salmon spawning is 12°C, while the optimal water temperature for green sturgeon (Acipenser
medirostris) growth is 19°C. At present, the Keswick Dam regulates the water temperature of the river in winter
to match the suitable spawning temperature for the endangered Chinook salmon, causing the cold water to extend
to the habitat of the green sturgeon, whose growth is thereby greatly affected (Zarri etal.,2019). In the Colum-
bia River, cascade dams have fragmented the spawning habitat of white sturgeon in the mainstream into short
sections connected by long-distance impoundments. Due to differences in dam operation and geographically
local environments, the water temperature in different river reaches varies greatly, which in turn leads to differ-
ences in the spawning temperature and time of white sturgeon in different river sections. In the lower Columbia
River, the spawning of white sturgeon downstream of the Bonneville Dam begins at a water temperature of 8°C,
but spawning in the three furthest downstream dam tailraces begins when water temperature reaches at least
10°C. In addition, spawning occurs earlier downstream than upstream in the Columbia River during the spring
season (Counihan & Chapman,2018; Péril,2004).
Dam construction alters the natural hydrological regimes of rivers, and thus affects the growth and reproduc-
tion of sturgeons. The reduced downstream flow results in limited lateral connectivity to the floodplain, which
adversely affects habitat availability and reproduction of sturgeons (F. He etal.,2021). In the Kootenai River, the
demand discharge during the spawning period of the white sturgeon is 1,416–2,832m
3/s; however, the peak flow
is reduced usually to 250–450m
3/s due to operation of the Libby Dam, which has seriously affected the reproduc-
tion of white sturgeon (Paragamian etal.,2001). River impoundment reduces flow velocity and possibly causes
hypoxia in the transition zone of the reservoir upstream, which exposes the pallid sturgeon (Scaphirhynchus
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albus) to low oxygen conditions and thus results in their high mortality (Guy etal.,2015). However, dedi cated
flow regulations can in some cases provide suitable condition for migration and spawning of sturgeon. For exam-
ple, intermittent flood pulsing through dam operation to achieve key water temperature thresholds may have
better facilitated the upstream migration and spawning of shortnose sturgeon adults (Vine etal.,2019).
Previous studies claim that the barrier effect of dams is the primary cause for the decrease in Chinese stur-
geon populations. However, new spawning grounds for Chinese sturgeon may have formed downstream of the
Gezhouba Dam (P.Zhuang etal.,2016), indicating that blockage by the dam has not completely hindered the
natural reproduction of this species. Therefore, it is essential to investigate the impacts of other factors, such as
altered water temperature regimes, in addition to the blockage of migration routes, on the spawning of Chinese
sturgeon. To date, most studies focus on the impacts of river damming on the migration and spawning of sturgeon,
but it is also important to understand the impact on other life stages, such as foraging and overwintering.
3.4. Impact on Eel and Lamprey
Eels (genus Anguilla) and lamprey (genus Lampetra) are among the most valuable fishery species around the
world (P.R. Almeida etal.,2021; FAO,2015). There are 19 species or subspecies of eels being identified glob-
ally (Righton etal.,2021), of which 17 are scattered throughout the Indo-Pacific and the other two species are
distributed in the Atlantic Ocean (Figure10). At present, several species of eel have been listed as “endangered”
or “critically endangered” in the wild (Jacoby etal.,2015; Vié etal.,2009), including the European eel (Anguilla
anguilla L.), American eel (Anguilla rostrata) and Japanese eel (Anguilla japonica). Lampreys are often misi-
dentified as eels because of their similar appearance; however, lampreys belong to the Order Petromyzontiformes
whilst eels belong to the Order Anguillifomes (Renaud,2011). Most lampreys, such as the sea lamprey (Pet ro-
myzon marinus), European river lamprey (Lampetra fluviatilis), Pacific lamprey (Entosphenus tridentatus), and
Caspian lamprey (Caspiomyzon wagneri), are anadromous parasitic fish. The sea lamprey is a notorious invasive
species in the Laurentian Great Lakes (Figure10), and has devastated the fisheries of whitefish and lake trout
(McDonald & Kolar,2007; Zielinski etal.,2019).
The life stages of eels comprise eggs, leptocephalus (larva), glass eel (post-larva), elver (juvenile), yellow
eel (non-mature adult), and silver eel (migratory adult). At each life stage, eels exhibit distinct morphologies
(Tsukamoto etal.,2011). The larva of eels is notably larger than that of almost all the other fish species, and
their morphology is well equipped for both passive and active swimming in their oceanic migration (Righton
etal.,2021). Eels usually form their sex at the silver stage (Miller & Tsukamoto,2016). Unlike salmonid and
sturgeon, eels have reverse migration and spawning behaviors, as they grow up in freshwater and return to the
ocean for reproduction (Figure10). They typically dwell in seawater during the life stages of eggs, leptocephalus
and glass eel, and in brackish and freshwater during the life stages of elver, yellow eel, and silver eel (Haro,2014).
The spawning of temperate eels, such as European eel and American eel, occurs in the Sargasso Sea (Béguer-Pon
Figure 10. Native and invading distribution of eel and lamprey around the world and their migration route for spawning.
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etal.,2015; Schmidt,1923; Tucker,1959), which has been recently reconfirmed by genetic analyses (Barth
etal.,2020). The eggs and spawning adults of Japanese eel have only been discovered within a restricted area
along the seamount chain of the Pacific West Mariana Ridge (Tsukamoto etal.,2011). In contrast to the distribu-
tion of spawning zones, the habitats of eels during the entire growth stage are flexible, encompassing all saltwater
and freshwater areas, because eels can adapt to diverse environmental conditions and different dietary niches
(Tesch,2003). Lampreys have an anadromous migration habit and a wide habitat range (Moser etal., 2021).
Some lampreys are life-long freshwater dwellers, whilst others are seawater dwellers that migrate to freshwater
to spawn (Renaud,2011). The sea lamprey breeds anadromously in basins of Western Europe and eastern North
America, and swims into the North Atlantic and Western Mediterranean. The Korean lamprey (Eudontomyzon
morii) is a small freshwater species that is mostly distributed in the Yalu River as well as some mountain rivers
in northeast China, North Korea and the Russian Far East (Renaud,2011).
Although eels and lampreys have strong acceleration and high mobility (Tytell & Lauder,2004), they cannot
jump over obstacles and their burst swimming speed is relatively low (Kemp etal., 2011). The barrier effect
caused by dams is thus the most immediate impact on eels and lampreys, impeding their migration between
spawning and rearing grounds. The downstream passage through turbines and dams leads to the high disorienta-
tion and mortality of breeding silver eels, which results in significant declines in spawning rates and populations.
It has been reported that 75% of eels are delayed in their downstream migration, and up to 65% are definitively
halted by dams (Besson etal.,2016). Eels are especially vulnerable to screens and turbines due to their anguill-
iform morphotype (Kerr etal.,2015), and thus hydroelectric facilities in dams can cause sublethal injury and
direct mortality to migrating adult eels (Bruijs & Durif,2009). In rivers with cascade dams, the cumulative effect
may result in the level of overall escape of eels not reaching the required conservation criteria, which is 40% for
silver eel in European countries (Pedersen etal.,2012). Habitat fragmentation caused by dams also impedes the
upstream migration of eels (Jellyman,2022). More than 15,000dams have been constructed in the coastal drain-
ages of the North Atlantic, obstructing direct access to 87% of rivers and streams flowing into the Atlantic, and
hence drastically reducing the inland extension of American eels (Jellyman,2022; Miller & Casselman,2014).
The decline of the European eel (Bevacqua etal.,2015), American eel (Kwak etal.,2019) and Japanese eel (J.
Z. Chen etal.,2014) in turn affects the ontogenetic stage and physiological traits of eels (Righton etal.,2021).
The barrier effect of dams is also the main reason for the decline in the global population of most lampreys, such
as about 80% loss of sea lamprey in the Iberian Peninsula and Caspian lamprey in the Volga River, leading to the
collapse of associated commercial fisheries (Atkinson etal.,2020; Jellyman,2022), and strong decline of Pacific
lamprey in the Columbia and Snake Rivers (Moser & Close,2003).
Eels have specific habitat preferences and requirements, with their habitat selection involving water tempera-
ture, water depth, substrate, salinity, flow velocity, oxygen concentration, vegetation cover, prey availability,
and predation threat (Jellyman,2022; Righton etal.,2021). Water temperature appears to be an important factor
throughout the whole lifespan of eels, as they grow faster and mature earlier in warm waters (Tesch,2003). Water
depth can affect the habitat quality of eels, since small eels tend to favor shallow waters and large eels prefer
deep waters (Jellyman,2022; Jellyman & Arai,2016). In particular, large eels spend the daytime in deep waters
and night time in shallower waters (Righton etal.,2021). Eels choose different types of substrates in different
seasons, preferring soft substrates with abundant organics or silts in the spring, muddy substrates in the summer,
and rubble substrates in the autumn (Tomie etal.,2016). Their substrate preference varies with body size, with
smaller eels preferring coarse substrates (gravels, cobbles, and boulders) and larger eels preferring fine substrates.
These habitat variations are likely due to the combined effects of changes in physical space requirements and prey
preference with the increasing body size of eels (Lloyst etal.,2015).
Alterations of water temperature regime, flow velocity and riverbed substrate due to river damming can have
dramatic effects on the living conditions of eels (Righton etal.,2021). The release of hypolimnetic water from
reservoirs has a cooling effect on water temperature in summer and thus degrades the growth of eels downstream
of dams (Maheu etal.,2016a). The migration speed and route selection of eels are affected by flow veloc-
ity, and thus decreases in flow velocity due to dams may cause failure of their migration (Jansen etal., 2007).
During downstream migration, eels usually swim actively with a speed of 0.3–1.2m/s (Behrmann-Godel &
Eckmann,2003), and those that manage to migrate downstream pass over the dam crest only when the flow
velocity is high (Besson etal., 2016). Artificial fluctuations in water depth due to dam operations inevitably
affect the living environment of eels (Righton etal.,2021). Dams are known to cause substrate-sorting effects
and thus result in local habitat homogeneity, which limits the diverse substrate requirements of eels (Naganna
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& Deka,2018). Dams intercept sediment and reduce downstream substrate availability for the critical life stages
such as nesting and refuge of eels (Černý etal.,2003). The decreased sediment flux can also cause delta recession
(Baisre & Arboleya,2006), which increases the susceptibility of eels to natural disturbances (Day etal.,2007;
O'Connor etal., 2015). The introduction of non-native species due to river damming squeezes the habitat and
even leads to extinction of native eels and lampreys (Marohn etal.,2014). Nonetheless, there are also some
marginal benefits of river damming for eels. Reservoir eutrophication has been reported to increase chirono-
mid populations, which comprises the major food of eels (Aprahamian etal.,2021). The increase in plant and
zooplankton production caused by reservoir eutrophication is considered to provide an additional benefit to the
populations of eels (Bunting etal.,2007). In some cases, the isolation created by dams can be beneficial to eels
by preventing the introduction of contaminants, parasites and diseases into their habitats (Liermann etal.,2012;
Righton etal.,2021).
Limited by monitoring technology, comprehensive and reliable data on the migration routes and spawning
grounds of eels are sparse, which hinders accurate assessments of the impact of river damming on eels. To date,
relevant studies outside of Europe, the Americas and Japan are scarce, and the lack of information across Africa
is a severe barrier to the appropriate assessment of the conservation status of eels on a global scale (Righton
etal.,2021). In addition, when compared to intensive studies concerning the impacts of river damming on eels,
similar research on lampreys needs to be enhanced. Although an individual small dam has much less impact on
eels and lampreys than a large dam, the cumulative impact of many small dams requires adequate attention, given
their widespread and intensive distribution (Lehner etal.,2011). Climate change could alter water temperature
regime and thereby induce impacts on the habitat of eels and lampreys, and is regarded as one of the least under-
stood risks to the species worldwide (Jacoby etal.,2015).
3.5. Impact on Other Fish Species
River damming also impacts on other fish species, which have less socioeconomic and cultural value, such as
giant catfish, tilapia and sardines (Figure11).
Giant catfishes are widely known for their giant bodies and mysterious trails in deep water areas, and they
can grow to more than two m in length (Boulêtreau & Santoul,2016). Giant catfish species mainly include
the Mekong catfish (Pangasianodon gigas, Pangasius krempfi, Pangasius sanitwongsei, and Pangasius mekon-
gensis), Amazonian catfish (Brachyplatystoma rousseauxii and Brachyplatystoma filamentous), and European
catfish (Silurus glanis). Megafaunal species have disproportionate per capita effects on community structure
and ecological processes, and any shift in their abundance is likely to affect food webs and ecosystem functions
(Malhi etal.,2016). Giant catfishes generally spawn in freshwater rivers and live in estuaries with long-distance
migration (Figure11). The Mekong catfish migrates long distances to spawn, spending much of their lives in
the brackish waters of the Mekong Delta and in the South China Sea near Vietnam before returning to spawn in
Figure 11. Distribution of giant catfish, tilapia, and sardines around the world, as well as migration route for spawning of giant catfish.
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the Mekong River in Laos and Thailand (Hogan etal.,2007). Amazonian catfish adopt a basin-wide migratory
life cycle between the Andean piedmont and the Amazon estuary, which makes them possess the longest river
migration that may total up to 12,000km in the case of Brachyplatystoma rousseauxii (Duponchelle etal.,2021).
Amazonian catfish are of high commercial importance to fisheries of the Amazon River basin, forming a major
ecosystem service provided by this river system (Fraser,2018). Due to the impact of dam construction, giant
catfish, and especially long-distance migratory species such as the Mekong and Amazonian catfish, have become
endangered in the wild, threatening the stability of ecosystem structure due to their top-tier status in the food web
(Duponchelle etal.,2021; Fraser,2018; Hermann etal.,2016; Hogan,2011). Dams on the Mekong River have
disrupted the migration and spawning of giant catfish, and the situation may become worse as Laos plans to build
more dams on the mainstream of the Mekong River (Soukhaphon etal.,2021). The migration route and access
to a substantial portion of the spawning grounds of the Amazonian catfish in the Madeira River are blocked by
two dams built a decade ago, which profoundly affects the populations and fisheries of the Amazonian catfish
and further alters the food web of the river ecosystem controlled by this apex predator (Duponchelle etal.,2016;
Fearnside,2013; Fraser,2018). It has also been reported that the body size and trophic level of Amazonian catfish
have declined in the reservoir and downstream reach of the Belo Monte Dam on the Xingu River after large
hydropower development (Keppeler etal.,2022). Dams intercept sediment and nutrients that nourish the Amazon
River and its floodplain, and damage fish physical habitats with a cascade of ecological effects (Fraser,2018).
The disruption of nutrient connectivity is more pronounced by dams in the Andean-Amazonian transition, whose
rivers provide most of the sediment and nutrients to the Amazon (Lees etal.,2016).
Tilapia is the third most-cultured finfish in inland aquaculture, with its production reaching 4.41 million tons in
2020, and accounting for 9% of world inland aquaculture (FAO,2022). Tilapia mainly includes the Nile tilapia
(Oreochromis niloticus) and Mozambique tilapia (Oreochromis mossambicus). The Nile tilapia is indigenous in
the Nile River; however, it is highly invasive elsewhere and has invaded many ecosystems worldwide (Faunce &
Paperno,1999). The Mozambique tilapia is indigenous in South Africa, Zimbabwe, Mozambique, and Eswatini
(Moyo & Rapatsa,2021). Tilapia possess a strong survivability and are physiologically tolerant to a wide range of
salinities, dissolved oxygen content and water temperature, and are characterized by multiple spawning, parental
care, and extreme feeding plasticity (Avella etal.,1993; G. Farmer & Beamish,1969; Vitule etal.,2009). Tilapia
inhabits the middle and bottom layers of water bodies, and can survive in waters of 16–40°C, with the optimal
temperature being 28°C. Tilapia is a warm-water fish, which will hibernate or even die when water temperature
is lower than 15°C. Tilapia feeds on the base of the food web at the bottom layer of a water body, most often on
sediment resources such as nematodes, rotifers, bryozoans and hydrozoans, and is well adapted to surviving and
growing in non-native environments (Peterson etal.,2006). Tilapia is widely cultured in reservoirs using cages in
tropical and subtropical regions (Hishamunda,2007), which can alleviate pressures from water scarcity and also
help tilapia to overwinter safely (Moyo & Rapatsa,2021). However, due to the strong environmental adaptation
ability of tilapia, non-native tilapia that escapes accidently from aquaculture in reservoirs could rapidly become
the dominant population. This increases the risk of invasive species to the original ecosystem of reservoirs, result-
ing in the loss of genetic integrity of native species, damage to the native biodiversity, and further affecting the
stability of the native river ecosystem (Bernery etal.,2022; Canonico etal.,2005; Cucherousset & Olden,2011).
Therefore, it is critical to quantify the impact of cultivated tilapia on the native fish species in reservoirs, and
conduct actions to control their habitat range and prevent their escape into the wild, in order to protect the ecosys-
tem stability of dammed rivers.
River damming also affects non-migratory offshore marine fishes such as sardines (Sardina pilchardus), which
are mainly distributed in the Atlantic and Mediterranean (Figure11). Sardines are a small pelagic fish that typi-
cally feeds on plankton, and plays an important role in global fisheries (FAO,2019). The availability of food such
as phytoplankton and zooplankton depend on the nutrient inflows from rivers, and thus river damming indirectly
affects sardine populations (Biswas & Tortajada, 2012). For example, the autumn flooding of the Nile River
irrigates and fertilizes the floodplain annually, and supplies sufficient nutrients to the Mediterranean Sea. Before
1965, the flood of Nile River delivered about 7×10
3 tons year
−1 of nitrogen and 7–11×10
3 tons year
−1 of phos-
phorous to the Mediterranean coast (Nixon,2003). The nutrients in the Nile floodwater support a massive diatom
bloom and a productive fishery, particularly for sardines (Halim,1960; Halim etal.,1964). However, completion
of the Aswan High Dam in 1965 decreased the fall flood by about 90% (Dorozynski,1975) and reduced the fertil-
ity of the southeastern Mediterranean waters, leading to a sharp decline in the marine fisheries (Milliman,1997;
Mohamed,2019; Rzóska,1976). It has been reported that sardine catches along the Egyptian coast declined
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from 18,000 tons in 1962 to a mere 460 tons in 1968 (Abul-Atta,1978). Since the late 1980s, the total fish
catch, including sardines, of the Nile and its estuaries have recovered back to the level before construction of the
Aswan High Dam (General Authority for Fish Resources Development,2012). The cause for this recovery may
be attributed to the increasing discharge of sewage from urban expansion stimulated by damming in the Nile
(Nixon,2003; A. J. Oczkowski etal.,2009; Richards,1982). However, the long-term fertilizing effects of sewage
require further investigation, as the persistent discharge of highly poor-quality water could damage the coastal
ecosystem and lead to an unsustainable increase, or even decline, of the fishery (A. Oczkowski & Nixon,2008).
In many cases, the time of dam construction coincides with adjustment of fishery policies that aim at increasing
fish catch and consequently results in overfishing. The contribution of overfishing and river damming to the
decline of fish catches demands quantification, in order to investigate and reveal the extent of fishery decline
attributed to dam construction (Biswas & Tortajada,2012). Meanwhile, due to rapid urbanization and indus-
trialization, nutrient supply to downstream river reaches and estuaries has largely increased (A. J. Oczkowski
etal.,2009), leading to rebounds of estuarine fisheries after river damming. It is essential to understand whether
this effect is sustainable or brings new challenges. Most current studies have focused on endangered or major
commercial fish species in order to prevent extinction or sustain fishery economies. However, fish diversity plays
an irreplaceable role in the aquatic food web and its integrity, as well as the stability of river ecosystems. There-
fore, we highlight the necessity for future investigations on the impacts of river damming on more fish species
and fish communities.
4. Fish Conservation Measures in Dammed Rivers
To offset or mitigate the impacts of river damming on fish, a variety of conservation measures, including fish
passage facilities, artificial breeding and release, ecological reservoir operation, and habitat compensation in trib-
utaries, have been proposed and implemented. Each measure has its own advantages and limitations, application
conditions, efficiency, and cost-effectiveness, and thus should be selected according to the specific situations of
a given dammed river. Of course, policies concerning fishery conservation play an essential role, but are outside
the scope of the present review. These policies include, but are not limited to, setting up nature reserves and
germplasm resources reserves, implementing policies and laws for no-catch measures, and navigation restrictions
during fish-sensitive seasons such as the spawning season (S. Huang & He,2019; Maxwell etal.,2020; H. Zhang,
Kang, etal.,2020).
4.1. River Connectivity Restoration and Fish Passage Facilities
Restoring longitudinal and lateral connectivity, such as river-floodplain reconnection and fish passage facilities,
could be the most direct approach to rehabilitate physical habitat and migration routes of fish in dammed rivers.
In recent decades, there have been increasing efforts to restore fish floodplain habitats in riverine rehabilitation
practices, as floodplain channels that are lost or disconnected from the main river have demonstrated visible
impacts on river ecosystems. Establishing the connectivity of floodplain channels to the main channel and restor-
ing lateral connectivity of river can provide essential nursery areas for fish and mitigate the loss of fish diversity
(Stoffers etal.,2022). The restored floodplain channels in the Rhine River, Netherlands, have served as suitable
nursery areas for rheophilic fish species (Stoffers etal.,2021). The restoration of hydrologic linkages between
the main channel and floodplain in the Kissimmee River, USA, has demonstrated positive effects on food web
structure and ecosystem functioning (Jordan & Arrington,2014). In the upper Danube River, a secondary flood-
plain channel has been artificially created following a nature-based construction scheme, which has provided
additional habitats and restored migration routes, thereby making an important contribution to restoring the
population of endangered fishes (Pander etal.,2015). The restoration on a tributary of the upper South Esk River,
Scotland, has reconstructed a natural meandering channel guided by historical maps, which reinstates floodplain
connectivity and habitat for Atlantic salmon and trout (Addy etal.,2016).
Fish passage is potentially an effective engineering approach to reconnect fragmented ecological corridor due
to river damming and restore river longitudinal connectivity. It is the earliest, and also the most widely used,
measure to conserve migratory fish in dammed rivers (Schilt,2007). Fish passage mainly includes fishway, fish
lift, fish collection, and transportation, turbine passage, juvenile bypass, and other engineering transport meas-
ures (Figure12). The earliest fishway can be dated to the mid-18th century in Europe (Clay,1995). In the early
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20th century, field and laboratory experiments on different fishway designs were performed. Denil(1909, 1938)
created the unique Denil fishway to reduce the flow velocity inside the path (Figures12a and 13a). In 1946,
a vertical slot fishway (Figures12b and13b) was constructed on both sides of Hells Gate in the Fraser River,
Canada, to allow salmonids to successfully cross the channel barrier caused by a landslide (Jackson,1950). Monk
etal.(1989) proposed a fishway configuration of pool and weir, in which nearly 100% of shad and almost all
Figure 12. Different types of fish passage facilities. (a) Denil fishway. (b) Vertical-slot fishway. (c) Nature-based bypass system. (d) Fish lift. (e) Fish collection and
transport system. (f) Fish-friendly turbine. Panel (a)–(c) are redrawn from Thorncraft & Harris(2000).
Figure 13. Different types of fishways. (a) Denil fishway for Arctic grayling in the Big Hole River watershed in Montana,
Canada (source: Montana State University, photo by Matt Blank, 2015; https://www.montana.edu/ecohydraulics/research/).
(b) Vertical-slot fishway in the Mosel River in Koblenz, Germany (source: The Federal Waterways Engineering and Research
Institute (BAW); https://www.baw.de/en/die_baw/wasserbau/umwelt/umwelt.html). (c) Pool-weir fishway for migrating pink
and coho salmon to spawning grounds in Anderson Creek, Canada (source: McElhanney Company; https://www.mcelhanney.
com/project/anderson-creek-fishway/). (d) Nature-based bypass fishway beside the Se San River of Cambodia (source: the
Xinhua News Agency; https://new.qq.com/rain/a/20191026A099LC00).
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other species could successfully pass through (Figure13c). These early efforts were mostly oriented at salmon
species, with only a small number of studies aiming at shad. Recent legislation for endangered species in the
United States, Canada, and Europe has re-emphasized the importance of fishways for migratory species other
than salmonids and shad; meanwhile, some successful initiatives have been taken for migratory species in other
parts of the world (Katopodis & Williams,2012).
With the rapid growth of large dams, the applicability of fishway has been facing great challenges due to high
cost, low efficiency and engineering complexity, which promotes the exploration of other types of fish passage.
Fish lift has a design principle similar to an elevator, which can actively move and release fish from the down-
stream reach of the dam to the upstream reservoir (Figure12d). These lifts induce fishes into a hopper that is
raised from the bottom to the upstream side of the dam (Santos etal.,2021). Barry and Kynard(1986) found
that the tailrace lift was more efficient than the early fish lift for American shad. It has been reported that the
combination of an Archimedes screw and fish lift can significantly improve the efficiency of fish lift (McNabb
etal.,2003; Zielinski etal.,2022). In some cases of high dams, where the construction of fishway and fish lift
is unfeasible, a fish collection and transportation system could be an appropriate alternative (Figure12e). Fish
collection and transportation is a special form of fish passage facility, which is mainly adopted in high dams or
in projects where fish need to climb several steps continuously. It attracts fishes into cabins or other boxes, and
then transports the fish over the dam by ships or vehicles (Figure14). In 1981, massive collection and transpor-
tation was implemented as an operational program by the United States Army Corps of Engineers (USACE) to
reduce losses of juvenile salmonids during their seaward migration. Fish collection and transportation systems
can improve the survival rate of fish passing through dams and has proved to be more effective than other fish
passage facilities in some contexts (D. L. Ward etal.,1997).
Traditional turbines are extremely harmful for fish to pass through, thus fish-friendly turbines such as Archi-
medes screw turbines and blunt blade turbines have been gradually used (Figure 12f). Fish-friendly turbines
have slowed rotational speeds and large openings, which can allow safe passage of small objects (Bracken &
Lucas,2013; YoosefDoost & Lubitz,2020). Bypass systems (Figures12c and13d), such as the curved-bar rack
bypass and the horizontal rod rack bypass, can guide downstream-moving fish toward a reasonably safe corridor
around water intakes, and thus effectively reduce the mortality of fish passing through dams (Beck etal.,2020;
Meister etal.,2022). In the eight dams of the lower Snake River and lower Columbia River, USA, most of the fish
entering the powerhouse are diverted to a juvenile bypass system, providing safe and efficient passage for juvenile
salmon to migrate downstream (Faulkner etal.,2019). River-like side channels are constructed in some dammed
rivers, which serve as a bypass for fish migration and even as a supplementary habitat for fish reproduction (L.
Zhang etal.,2023). In the Don River, Russia, a natural-type channel has been constructed to allow sturgeon to
bypass the Konstantinovskiy Dam. Stellate sturgeon (Acipenser stellatus) eggs are found in the bypass channel,
indicating that the bypass channel has been used as a spawning habitat (Pavlov & Skorobogatov,2014).
Overall, fish passage facilities can assist fish to pass through the barrier of dams and mitigate the impact of habitat
fragmentation. The efficiency of fish passages differs significantly between different types of fish passages. On
average, the efficiency of downward passage is slightly higher than that of upward passage; pool-weir, vertical
slot and naturalized fish passages have higher efficiency than Denil fishways and fish lifts (Noonan etal.,2012).
Figure 14. Fish collection and transportation system in Fengman hydropower station in Jilin, China (source: the Xinhua
News Agency; https://www.gov.cn/xinwen/2021-08/31/content_5634534.htm#10). (a) Fish collection from the box of fish lift.
(b) Transportation of collected fish to upstream.
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Fishways are used mostly in low dams, which are usually built in low gradient rivers, to improve river longitudinal
connectivity. The Denil and nature-based fishways are the most efficient fish passages (Baumgartner etal.,2018;
Bunt etal.,2001). These fish passages can remain effective even when significant fluctuations in upstream or
downstream water level occur (Quaranta etal.,2019). The Vianney-Legendre vertical slot fishway in the Riche-
lieu River, Canada, has shown to be successful for passing a variety of fish species, including lake sturgeon
(Marriner etal.,2016). The Denil fishways of the 63 diversion dams in the Big Hole River basin in southwestern
Montana provide opportunities for Arctic grayling (Thymallus arcticus) and other fish for year-round access to
critical habitats (Triano etal.,2022). The nature-based fishway installed at a low dam in Indian Creek, Canada,
has proven effective for multiple fish species to traverse the dam (Steffensen etal.,2013). However, fishways
may have a poor effectiveness for some fish. The effectiveness of a fish weir at Foster Dam in the Santiam River,
USA, is low to moderate for the downward passage of juvenile Chinook salmon, while it is consistently high for
steelhead trout, indicating that fish weirs may not be a suitable solution for all species (J. S. Hughes etal.,2021).
Overall, the passage efficiency of fishway is dependent on the slope, baffle and other characteristics affecting
the flow field inside the pools, the water head drop between the pools, as well as the turbulence levels and flow
velocities (Quaranta etal.,2019).
Fish lifts, fish collection and transportation systems are regarded as the most cost-effective fish passage facil-
ities for high dams. The fish lift of large hydropower dams on the Lima River, Portugal, effectively prevented
fragmentation of potamodromous populations between different reaches (Mameri etal.,2019). Fish collection
and transportation systems have the advantages of high flexibility, no interference to the structural layout of the
dam, suitability to large variations in reservoir water level, and small occupation of space for fish to cross the
dam. In the 1940s, a temporary collection and transportation system successfully transferred thousands of adult
salmonids at the Rock Island Dam on the lower Columbia River. Fish collection and transportation systems have
also been used to transport shad successfully at the Mactaquac Dam on the St. John River, Canada, and Essex
Dam on the Merrimack River, USA (Clay,1995). D. L. Ward etal.(1997) reviewed studies conducted by the
US National Marine Fisheries Service from 1968 to 1989 concerning the efficiency of using trucks and barges
to transport migrating juvenile Chinook salmon from the Snake River around dams to reservoirs in the lower
Snake and Columbia rivers, and suggested that the use of barges to transport juvenile Chinook salmon could
improve their survival rate. However, only 47% of Atlantic salmon succeeded in passing through the fish lift of
the Golfech-Malause hydroelectric complex on the Garonne River, France. In the Gezhouba dam, it has proven
successful to lure the bottom fishes by releasing jet flow during the collecting procedure (Y. Liang etal.,2014).
The main difficulty specific to fish lifts involves fish trapping, as the V-shaped entrance of fish lift may inhibit
salmonids entering the holding pool and cannot guarantee that the entered fishes will not return back to the river
(Croze etal.,2008).
The passage efficiency of different fish species under different hydraulic conditions in passage facilities varies
greatly from case to case (Bunt etal., 2016; Nieminen etal., 2016; Williams & Katopodis,2016). Salmonids
and clupeids have been found to efficiently pass through the vertical slot, pool-weir fishway, and Denil fishway,
with an efficiency of 63%, 45%, and 51%, respectively (Castro etal., 2016; Mallen-Cooper & Stuart,2007;
Noonan etal.,2012). Brown bullhead (Ameiurus nebulosus) and striped bass (Morone saxatilis), which have a
smaller body size than adult carp, prefer nature-based fishways, and have a passage efficiency of up to 70% (Bunt
etal.,2012). Fish lifts are the most effective up-migration measure for lamprey and brown trout, but the difficulty
is to capture small-sized individuals (Castro etal.,2016; Pompeu & Martinez,2007). Tummers etal.(2016) high-
lighted that the physical characteristics of baffles and high turbulence may inhibit lamprey ascending the passage,
and Moser etal.(2019) proposed a novel modification of fishway entrance for Pacific lamprey. The passage
efficiency of fishways is also related to the behavior of fishes (Shahabi etal.,2021), as their swimming direction
in fishways is dependent on their experience with the flow field (Goodwin etal.,2014). Inadequate attractiveness
for fish is recognized to be a major factor limiting the efficiency of fish passages (David etal.,2022). According
to Laine(1995), fish often need to become acquainted with the passage entrances before they start to climb the
passage facilities. Mensinger etal.(2021) suggested that fish may segregate at barriers based on their personality
and sizes, and this could be alleviated by increasing fishway attraction and maximizing passage opportunity,
leading to more exploratory eels passing through successfully. Generally, the functionality of fish passages in
alleviating dam barrier effects on fish is limited, and requires both good design of the facilities and good swim-
ming ability of fishes (Noonan etal.,2012).
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Fish passage has demonstrated to be an effective way in many cases to reconnect fragmented fish habitats in
dammed rivers. However, it is argued that moving fish preferring a lotic environment from downstream reaches
to reservoirs could cause further damage to these fish species, as the lentic environment of reservoirs may form
an ecological trap that makes the transferred fish unable to find their migration path or suitable habitats. Fish-
ways are mostly favorable to the fish species with strong swimming ability, which could potentially change the
structure of fish communities both upstream and downstream of a dam, and hence further impair fish biodiversity
in the dammed river. There are also plenty of fish passages that have not achieved their expected functionality.
Their low effectiveness could be attributed to insufficient considerations of fish swimming ability and hydraulic
characteristics in the design of the facilities. This demands renewed efforts to develop innovative solutions, at
the core of which is the need for engineers and biologists to work together and design passages based on the
preferred hydraulic conditions of multiple fish species. In addition, nature-based solutions and application of
natural materials, instead of concrete and metal, in fish passage constructions are an important aspect. The lack
of long-term monitoring and prompt assessment of the effectiveness of the operational fish passages also restricts
the modification and improvement of such facilities.
4.2. Artificial Breeding and Release
Artificial breeding and release of target fish species in dammed rivers is a viable conservation measure to replen-
ish the populations of endangered fish species in the wild and restore fishery resources (Molony etal.,2003;
Naish etal.,2007; J. Yang etal.,2013), although the evidence base for effectiveness is incomplete (Rytwinski
etal.,2021). Artificial breeding and release can be categorized into two main types: “ecological restoration”
and “resource restoration.” “Ecological restoration” aims to conserve the endangered native fish populations
and prevent extinction by releasing hatchery-reared fish into the original habitats of dammed rivers. “Resource
restoration” aims to recover fishery resources and improve the economic fishery in dammed rivers by artificial
breeding and release (L. Wang,2016).
This measure has been adopted as an essential conservation strategy for more than 20 vulnerable fish species in
China (J. Yang etal.,2013). For instance, Chinese sturgeon are preferentially distributed in the lower reaches of
the Yangtze River; however, their spawning migration route is blocked and spawning grounds are damaged by
hydropower dams on the upper Yangtze River, which reduces the length of natural spawning ground from 600 to
7km and causes their gonads to degenerate (L. Wang & Huang,2020; Xie,2003; Y. Zheng etal.,2022). Since
1984, artificial breeding and release of larvae and juveniles (Figure15) into the natural environment has become
an important approach to conservation efforts of the Chinese sturgeon (Chang etal.,2021; Gao etal.,2009; Qin
etal.,2020; Stone,2008; Wei etal.,2004). From 1983 to 1998, approximately 6 million Chinese sturgeon fry
and juveniles were released into the Yangtze River (H. Wang etal.,2019). About 500 to 1,500 adult sturgeons
were released each year into the spawning ground of the Yichang section of the Yangtze River from 1997 to 2003
(J. Li etal.,2021;P.Zhuang etal.,1997). In addition, a breakthrough in artificial breeding technology for the
two sturgeon species has been made, so that the artificially bred Yangtze sturgeons can be developed to a third
generation in the laboratory (D. Li, Prinyawiwatkul, etal.,2021).
The first stock enhancement program by breeding and release in Brazil was carried out for non-native fishes in the
northwest region, which improved fishery yields significantly (Paiva etal.,1994). Nowadays, stock enhancement
Figure 15. Artificial breeding and releasing of Chinese sturgeon. (a) Artificially bred fry and juveniles of Chinese sturgeon.
(b) Release of artificially bred fry and juveniles of Chinese sturgeon into the Yangtze River. Photos are provided by China
Three Gorges Corporation.
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has become a mandatory management action, which is considered a primary measure to mitigate the impacts of
reservoirs on fish populations and protect ichthyofauna in Brazil (F.P.Arantes etal.,2011; Casimiro etal.,2022).
In China, dam construction in the Yangtze River basin has significantly affected natural reproduction processes
and decreased the fishery resources of FMCCs (Q. Peng et al., 2012). After impoundment of the TGR, the
resources of FMCCs have decreased by more than 90%, compared to that in the 1960s (Y. Yi & Wang,2009).
The development of artificial breeding techniques and large-scale implementations of stock enhancement have
significantly recovered the FMCCs resources in the Yangtze River basin. The annual release of over 10,000kg of
bloodstock of FMCCs has been carried out in the Shishou and Jianli sections of the middle Yangtze River since
2010, which has brought substantial economic, social, and ecological benefits (H. J. Chen,2019). The improved
FMCCs fishery has alleviated limitations depending only on the catches of natural FMCCs resources (H. J.
Chen,2019; J. Yang etal.,2013). In the Lancang River, China, fish breeding and release has been introduced for
almost all the dams, which has proven effective in recovering fish resources (H. Xu & Pittock,2018).
Overall, artificial breeding and release of target fish species is an important measure to recover endangered fish
species, maintain genetic diversity, support population expansion, and further preserve the ecosystem integrity of
dammed rivers (Le Luyer etal.,2017; Leinonen etal.,2020). However, monitoring data shows that the number of
juvenile Chinese sturgeon in the estuary of the Yangtze River has not shown a visible increase (Wei etal.,2004),
as there are only small-scale natural breeding activities in the spawning grounds below the Gezhouba Dam
(P.Zhuang etal.,2016). Thus, artificial breeding and release has not recovered the natural reproductive processes
of wild Chinese sturgeon, but only maintains their populations to some extent and prevents them from extinction.
In the artificial breeding process, the selection and renewal of breeding fishes are sometimes overlooked during
artificial reproduction, leading to parent fishes with inferior genetic characteristics arising from inbreeding. The
genetic introgression of the released populations has the potential to reduce the genetic diversity and impair the
performance of wild populations due to genetic drift (Lin etal.,2022). For instance, artificially bred FMCCs
with inferior genetic characteristics escaping from tributaries and connected lakes, have affected the natural
high-quality germplasm resource of FMCCs in the mainstream Yangtze River, further leading to a decrease in
the quality of the germplasm resource and adaptation ability to the wild environment of the wild FMCCs (H. J.
Chen,2019). Fish stocking activities can introduce exotic diseases and parasites into the water body, which is
potentially harmful to the endangered populations (J. Yang etal.,2013). Therefore, the genetic admixture between
artificially bred fishes and wild fishes could cause genetic contamination and affect the genetic structure as well
as the stability of wild populations, leading to genetic and ecological risks (Abdolhay etal.,2011). In addition,
there is competition between large-scale released hatchery fish and wild recipient fish, squeezing the population
of wild species (J. D. Bell etal.,2008). It is necessary to understand the carrying capacity of the receptor envi-
ronment and the size of the wild population before conducting artificial stocking and release, in order to reduce
the negative impacts and maximize the benefits (Agostinho etal., 2016). In particular, artificial breeding and
release of non-native species can lead to biological invasions, which will result in changes and declines in native
fish diversity (Bernery etal.,2022). For instance, tilapia, which possess a wide environmental adaptive ability,
have become the dominant species in reservoirs and lakes due to artificial enhancement of stocking, threatening
the survival of native fish species (Cucherousset & Olden,2011).
Long-term field monitoring has shown that most artificially bred fishes cannot reproduce naturally in the wild,
but merely maintain the size of the target fish population. Failure to reproduce a second generation naturally
leads to the challenge that the wild population of the fish species cannot increase, and thus the stability of the
wild population is vulnerable in the long-term. Therefore, it is important to investigate the mechanisms of natural
reproduction of the target fish species in dammed rivers, including both artificially bred and wild individuals, for
restoring populations. Moreover, due to the lack of long-term and continuous monitoring data, the quantitative
impacts of artificial breeding and release on river ecosystems remain unclear. It is essential to establish a risk
assessment system on genetic admixture and species invasion to evaluate quantitatively the negative effects of
artificial breeding and release. This could help improve conservation measures to increase the target fish popula-
tion and protect river biodiversity.
4.3. Nature-Based Solutions and Reservoir Ecological Operation
Nature-based Solutions (NbS) have been recognized as an umbrella concept to capture eco-friendly strategies
that mimic nature with broad public acceptance (Cohen-Shacham etal.,2016; Y. A. Song etal., 2019). NbS
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follows the laws of nature to protect and restore degraded ecosystems as a whole, which can be applied in various
domains (Maes & Jacobs,2017; Nesshöver etal.,2017; Raymond etal.,2017). After the first appearance in the
official report from the World Bank(2008), the idea has gained increasing attention in scientific communities,
governmental agencies and non-governmental organizations worldwide (Cohen-Shacham etal.,2016; Griscom
etal.,2017; Krull etal.,2015). NbS is particularly suitable for watershed ecosystem protection and river ecosys-
tem remediation so as to preserve and restore the functions and valuable services that nature provides.
Conventional reservoir operations aim predominantly to maximize social-economic benefits, which may cause
severe damage to the ecological structure and functioning of river ecosystems. By contrast, ecological reser-
voir operations follow the idea of NbS, which aims to balance social-economic benefits and river ecosystem
requirements (X. Xia etal.,2009). Studies on reservoir ecological operation have spanned several decades, since
Schlueter(1971) proposed that reservoir operation schemes should take the habitat diversity of river ecosystems
into account when meeting the social-economic water demand. Many studies have been dedicated to developing
optimization models for reservoir ecological operation (D. A. Hughes & Ziervogel,1998; C. Liu etal., 2019;
Suen & Eheart,2006). The United States and Australia are two countries that have undertaken relatively early
implementations of reservoir ecological operation (Higgins & Brock,1999; A. J. King etal.,2010). In recent
years, China has made great efforts in reservoir ecological operations, which have made visible contributions to
restoring fish resources in dammed rivers (Q. Chen etal.,2021; C. Ma etal.,2020).
Flow regulation can create suitable hydrological conditions for fish to acquire their demand environment at
critical life stages, especially during spawning periods (Yin etal.,2011). The NFR, which shapes river species
community and ecosystem structure and function (Poff,2018), is crucial to riverine biodiversity and healthy
(Bower etal.,2022; Poff etal.,1997). Releasing ecological flow (Maavara etal.,2020; Poff,2018) is an impor-
tant measure of reservoir ecological operation (Y. Miao etal.,2020; X. Xia etal.,2009; Y. Xia etal., 2019). Q.
Chen etal.(2012), D. Chen, Chen, etal.(2016) proposed an operation scheme for the Qingshitan Reservoir in the
Lijiang River, China, to maintain a quasi-NFR in the downstream reach for fish conservation when meeting the
demands of irrigation, cruise navigation and water supply. Creating artificial floods through reservoir operation
has been used to stimulate spawning of drifting eggs in dammed rivers (C. Ma etal.,2020; Zhou etal.,2019). In
the Colorado River, artificial flood experiments have been conducted at the Glen Canyon Dam for many years,
which have successfully restored the endangered humpback chub (Gila cypha) and maintained populations of
other native fish species (Jacobson & Galat,2008; Melis etal.,2015; Yao etal.,2015). The Hume Reservoir in
the Murray River, Australia, rebuilds some small-to-medium floods, which modifies the timing of peaks to trig-
ger the spawning of native golden and silver perch, and increase the duration of floods to extend the recruitment
of species from mid-October to mid-December 2005 (A. J. King etal.,2010). In China, ecological operation of
the TGR has substantially increased the spawning of the FMCCs (Figure16). During the 4-day experiment of
ecological operation of TGR conducted in mid-June 2018, an initial outflow of 11,000m
3/s and a flow increment
in the range of 1,000 to 1,500m
3/(sd) was implemented, which formed a concentrated egg spawning activ-
ity of FMCCs in the Yichang section of the Yangtze River (C. Ma etal., 2020). In Brazil, reservoir operators
have recently proposed a flow regime that could restore flooding for 32 fish breeding sites in the Xingu River
(Moutinho,2023).
Discharge regulation alone is sometimes insufficient to meet the demand of fish reproduction, since fish spawn-
ing also requires suitable water temperature. Controlling the selective withdrawal device to adjust the water
temperature of the outflow from temperature-stratified reservoirs can improve the water temperature rhythm to
some extent for fish reproduction downstream of dammed rivers (Saadatpour etal.,2021). A selective-withdrawal
device was installed at the Shasta Dam in 1997 to manage the water temperature regime in the downstream reach
of the dam to meet the year-round thermal requirements of salmonid while fulfilling the obligation on water deliv-
ery, power generation and flood control. Its use resulted in a significant increase in the salmonid fish population
(Bartholow etal.,2001; Hanna etal.,1999). A water temperature regulation experiment was conducted in the
Xiluodu-Xiangjiaba cascade reservoirs in the Yangtze River in May 2017, which increased outflow temperature
and thereby facilitated fish spawning in the reaches downstream of the Xiangjiaba dam. About 10 million and 100
million fish eggs were monitored in the Yibin and Jiangjin section of the Yangtze River, respectively. In particu-
lar, the annual peak in fish spawning occurred during the period of the experiment (Ren etal.,2020).
Sediment regulation of reservoirs can change the morphology of the downstream reaches, and this potentially
affects the location and quality of fish habitats (W. Wang etal.,2012; H. Zhang, Jarić, etal.,2021). Sediment
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regulation has been implemented at the Xiaolangdi Reservoir in the Yellow River since 2002, resulting in the main
channel of the lower reach of the dam being fully scoured (C. Miao etal.,2016). The regulation also brings abun-
dant nutrients and hence plankton blooms downstream, providing suitable feeding sites for pelagic and epipelagic
fish that congregate in the Yellow River estuary (L. Zheng etal.,2014). Some studies show that sediment flushing
could erode the areas of spawning and feeding grounds of fish, and even lead to fish mortality (Kong etal.,2017).
However, proper design of sediment flushing could control suspended sediment concentration and reduce adverse
effects on fish in the downstream reaches (Cattaneo etal.,2021). In addition, fish can possess a certain resistance
to the negative impacts caused by sediment flushing operations (Grimardias etal.,2017). The three-year field
investigation conducted from 2009 to 2011 concerning the controlled sediment flushing of a small reservoir on
the Adda River in Italy showed that fish resources in the downstream reach were not affected significantly (Espa
etal.,2015). Similarly, fish density showed no significant impairment during controlled sediment flushing oper-
ations conducted in 2016 at the Verbois Reservoir in Switzerland (Cattaneo etal.,2021).
Control of intermittent reservoir discharges is possible to reduce the damage of TDG supersaturation on fish.
Modeling results illustrate that intermittent discharges from the Bala Reservoir could diminish TDG supersatura-
tion and reduce the negative effects on fish in the Zumuzu River, China (J. Feng etal.,2014). Based on numerical
modeling of TDG, Wan etal. (2021) found that a proper discharge scheme for the Xiluodu Reservoir on the
Yangtze River could reduce the level and maximum residence time of TDG in downstream waters, thereby alle-
viating negative impacts on fish. The mixing of tail water and spill discharge can create areas of low TDG level,
providing shelter zones for fish to avoid damage from high TDG levels (Wan etal.,2020).
Reservoir ecological operation has developed from single factors to the coupling of multiple factors of fish phys-
ical habitat. W. He etal.(2020) proposed an operation model for the Sanbanxi Reservoir in the Yuanjiang River,
China, to meet the demand of outflow water temperature and downstream ecological flow. Z. Xu etal.(2017)
proposed an eco-friendly operation scheme considering flow velocity and water temperature demands of target
fish, which significantly facilitated the spawning of this fish species. In South Africa, an experiment has been
conducted at the Clanwilliam Dam on the Olifants River by creating small pulses of high flow and making the
water temperature at the spawning site reach above 19°C, which has resulted in a visible increase of successful
spawning activities of yellowfish (Barbus capensis) downstream of the dam (J. King etal.,1998). In the USA, a
Figure 16. Effect of reservoir ecological operation on fish spawning in dammed river. Adapted from Q. Chen etal.(2021). (a) The proposed ecological operation of
the Three Gorges Reservoir (TGR) for improving the spawning of four major Chinese carps (FMCCs) in the downstream river of the TGR. (b) The changes of spawned
eggs of FMCCs in the downstream river of the TGR before and after the proposed ecological operation. Data are available from China Three Gorges Corporation and
China National Environmental Monitoring Center. There was no measurement of spawned eggs during the TGR ecological operation in 2020 due to COVID-19.
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flow pulse has been released from the Gavins Point Dam, together with the improvement of water temperature, to
promote spawning of pallid sturgeon in the lower Missouri River (Jacobson & Galat,2008).
The effect of ecological operation of a single reservoir is sometimes limited. Joint operations of multiple reser-
voirs, which can be in a cascade along one river or are distributed on multiple rivers within the same watershed,
can better coordinate the social-economic interests and ecological requirements. Dalcin etal.(2022) reported a
methodological framework to guide operations of cascade reservoirs for rebuilding expected flow regimes. The
application to the reach between Porto Primavera Reservoir and the Itaipu Reservoir in the upper Paraná River
basin in Brazil provided suitable conditions for the successful recruitment of migratory fish species. Q. Chen
etal.(2013), D. Chen etal.(2015) proposed an adaptive operation scheme for two cascade reservoirs, Jinping-I
and Jinping-II, in the Yalong River, China, to meet the requirements of daily ecological flow and water tempera-
ture for the conservation of the indigenous fish species Schizothorax chongi in the dewatered river reach between
the two dams. D. Chen, Leon, etal.(2016), D. Chen, Leon, Engle, etal.(2017), and D. Chen, Leon, Hosseini,
etal.(2017) optimized the operation of 10 reservoirs in the Columbia River to maximize total power revenue
and fulfill the ecological flow requirements of fish, providing an important reference to ecological operation
of multiple reservoirs. Z. Jiang etal. (2019) developed a multi-objective optimization model to balance flood
control, power production, and ecological flow requirement of a large-scaled cluster of reservoirs with mixed
types in the Pearl River basin, China. The model results played a significant role in the operations to improve both
social-economic benefits and fish conservation.
Although there are studies and engineering practices of reservoir ecological operation concerning multiple
factors, such as flow and water temperature, the spatiotemporal matching of different factors with regards to
habitat requirements of different fish species and their life cycles remain challenging. In the joint operation of
cascade reservoirs, available studies mostly use water balance methods to link the reservoirs. In future, it is better
to adopt hydrodynamic models, which can simulate the distribution of flow fields, water temperature, and TDG
to achieve refined operation strategies for more effective conservation of fish. In addition, climate change could
bring large uncertainties to the inflow of reservoirs (Y. Wang etal.,2019), which implies that current reservoir
ecological operations based on historical data and deterministic models must be updated to incorporate inflow
uncertainties under future climate changes.
4.4. Habitat Compensation and Dam Removal in Tributaries
The shift from a lotic to a lentic environment of the reservoir after river damming leads to a permanent loss of
habitats for the maturation and spawning of fish species (Antonio etal.,2007; Liermann etal.,2012), and such
loss cannot mostly be remediated through reservoir ecological operations. The situation is more severe in rivers
with cascade dams, for instance the Lancang River, where an upstream dam is located in the backwater zone of
the nearest downstream reservoir, and the lentic section stretches for over one thousand kilometers. Under such
circumstances, relatively unaltered tributaries can provide possible alterative places to conserve indigenous species
of the dammed mainstream by serving as high-value natural surrogates or supplements that can restore some
function of the mainstem ecosystems (Neely etal.,2009; Nunes etal.,2015; Pracheil etal.,2009,2013). The
elements of natural flow fluctuations and the availability of food resources and shelter areas could be main-
tained in these tributaries. Moreover, diverse hydraulic conditions exist in the upper, middle, and lower tributary
reaches, giving the migratory or rheophilic fish species chances to colonize new habitats. In addition, for some
native fish species, their survival and life history requirements are directly related to intact longitudinal pathways,
including the possibility of migration into tributaries for reproduction and rearing (Da Silva etal.,2015). There-
fore, the use of unregulated tributaries to alleviate the adverse impact of impoundments in the mainstream has
recently been forwarded as a key alternative for fish conservation in dammed rivers (Figure17).
Successful spawning of humpback chub is found to be related to the migration of adult individuals from the
highly altered Colorado River to a relatively unregulated tributary, which offers necessary spawning habitat
and hydrological variability (Gorman & Stone,1999). Adults of American paddlefish prefer to migrate up the
unaltered Yellowstone River rather than the regulated Missouri River when they move above the confluence of
the two rivers, probably because the more natural flow pattern in the Yellowstone River tributary provides better
spawning conditions than the Missouri River (Firehammer & Scarnecchia,2006). In South America, recent
findings indicate that at least eight long-distance migratory species utilize alternative spawning and nursery
habitat in four tributaries of the upper Paraná River, after the construction of Porto Primavera Reservoir (Da Silva
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etal.,2015,2019). The Congonhas River in Brazil provides possible reproductive routes and feeding sites for six
important rheophilic and migratory species, Piaractus mesopotamicus, Megaleporinus obtusidens, Prochilodus
lineatus, Salminus brasiliensis, Pinirampus pirinampu, and Pseudoplatystoma corruscans, after the construction
of Capivara Dam. These species are long-distance migrators and represent about 29% of all migratory species
inhabiting the upper Paraná River basin (Garcia etal., 2019). In China, indigenous fish species in the upper
Yangtze River and Lancang River are facing great threats incurred by the cascade dams. Conservations of the
indigenous fish in these severely impounded rivers often focus on their large tributaries, which remain natural or
are lightly dammed and have been recognized as alternative habitats for spawning and larvae development (L.
Tang etal.,2021). W. Cao(2000) and Park etal.(2003) suggested that three tributaries, the Jialing, Chishui and
Tuo rivers, in the upper Yangtze River can be a potential refuge for 22 endemic species, including Dabry's stur-
geon that is listed as a first-class protected animal in China. In the Lancang River, its tributary, the Luosuo River,
serves as an important habitat for the migration and spawning of red mahseer (Tor sinensis), one of the most
famous economic fishes of Yunnan province in southwest China (Hong etal.,2022; Y. Peng, Hong, etal.,2022).
These findings demonstrate that unregulated tributaries can provide habitats required by fish at different devel-
opment stages.
Preservation of free flow is essential for tributaries to serve as alternative habitats of fish in dammed mainstreams.
However, numerous tributaries of large rivers have been developed for individual or cascades of SHPs. In the
United States, there are more than 75,000 reservoirs, and many of them are impounded by small dams (<10m in
height) that are now aged and in disrepair (Ahearn & Dahlgren,2005). In China, more than 45,000 small dams
have been built by the end of 2012 to meet rural electricity demand (Ding etal.,2019; Hennig & Harlan,2018),
and approximately 6,590 small dams are reported to be out of service due to their age and loss of function
Figure 17. Habitat compensation and rehabilitation in dammed river. (a) Laomuhe Dam on the Heishui River, a tributary of upper Yangtze River. (b) The removal
of the Laomuhe Dam. (c) The Heishui River after the removal of the Laomuhe Dam. Photo by Lei Tang. (d) Habitat restoration measures in dammed river, including
ecological spur dike construction, creating diverse habitats with large woody debris, big rocks, pool and riffle. (e) Changes of fish diversity and mass per unit effort in
the Heishui River before and after the removal of Laomuhe Dam. Adapted from S. He etal.(2021).
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(Gao etal.,2018). Removal instead of maintenance of these aged facilities has been growing around the world.
Since the early 1900s, nearly 1,800 small dams have been removed from rivers in the USA (Fox etal.,2022). In
Europe, at least 4,000dams and weirs have been removed since the mid-1990s (Kim & Choi,2019). Removal of
small dams in tributaries can open up spawning and rearing habitats in previously inaccessible regions upstream,
which has shown positive effects on fish diversity of the dammed mainstreams (Barbarossa etal.,2020; Lasne
etal.,2015; Magilligan, Graber, etal.,2016; O'Connor etal.,2015; Y. Peng, Hong, etal.,2022). Foley, Bellmore,
etal.(2017) reported that anadromous fishes, such as salmonids, passed the former dam site within days to
weeks after the removal of the Marmot dam in the Sandy River, USA, a tributary of the Columbia River. Simi-
larly, following the removal of small dams in the Jidu River, a tributary of the Lancang River, seven migratory
species recolonized newly accessible habitats (Hong etal.,2022). In addition to the direct effect of reconnection
and upstream access, dam removal has strong influences on fish physical habitat by changing sediment regime
and channel morphology, which increase the heterogeneity of hydraulic features and create new habitats for fish
species (Hatten etal.,2016; Im etal.,2011; Magilligan, Nislow, etal.,2016). The removal of a small dam in the
Heishui River (Figures17a–17c), a tributary of upper Yangtze River, has increased the percentage of suitable
spawning habitat for Jinshaia sinensis by a factor of four (Figure17e), due to the change in river morphology and
hydrological regimes following the removal (L. Tang etal.,2021).
Although removing small dams to restore the natural conditions of rivers is shown effective to conserve fishes
in dammed rivers, removals of large dams still face great challenges due to social-economic constraints and
intensive as well as long lasting ecological consequences (O'Connor etal.,2015). To restore the access of salmo-
nids to their spawning grounds in the mainstream of Elwha River, two large dams, the Elwha Dam and Glines
Canyon Dam, have been removed, resulting in increased species richness and functional stability of the riverine
ecosystem (Foley, Warrick, etal.,2017; Shaffer etal.,2018; Warrick etal.,2015). Similar achievements have
been made in the removal of the San Clemente Dam on the Carmel River, USA (Smith etal.,2020), which has
gradually improved the spawning grounds of steelhead salmon near the original dam site and in its downstream
reach (Harrison etal.,2018). Despite the visible ecological benefits, removal of large dams often comes at a huge
cost, and long-term consequence of fish community demands further investigation.
After dam removal, naturalized artificial habitat, following the NbS concept, can be created to expand living spaces
of fish (Figure17d). The addition of gravel to rivers, known as gravel augmentation, is an early attempt at artificial
habitat creation, which has proven to increase the available spawning grounds for Atlantic salmon and brown trout
in regulated rivers (Barlaup etal.,2008; Pulg et al.,2008). Three types of artificial habitats (straw bales, straw
tubes and moss tubes) have been implemented to enhance egg production of Galaxias maculatus, an important
fishery species in New Zealand (Hickford & Schiel,2013). In the Youjiang River, a tributary of the Pearl River,
China, artificial habitats made of bamboo and palm slices have been deployed to serve as spawning grounds for
fish that produce sticky eggs and as refuges that improve the survival rates of juvenile fishes (D. Guo etal.,2020).
The installation of large woody debris can also restore degraded river ecosystems due to dam construction through
“rewilding,” which has been proven to significantly improve the abundance of food resources and thereby increase
the population of fishes in the restored reaches (Thompson etal.,2018). In the lower Mulde River, Germany, fish
abundance increased nearly 10-fold eight months after the installation of large wood (Anlanger etal.,2022).
Tributaries cannot offer an identical replacement for the degraded habitats of previously undammed mainstreams,
as the former have lower discharge, longitudinal distance, morphological variability, and habitat complexity than
the latter. Discharge and its variability are essential to provide flow-related cues that initiate fish maturation and
spawning, or create conditions for recruitment of larvae and juveniles. The longitudinal distance is critical for
fish species that require long egg-drifting distances for survival. To date, quantitative methods for determining
suitable tributaries for fish spawning, foraging and refuging have not been established. Future studies should
attempt to coordinate the relations between conservation efforts in the mainstream channel and its tributaries to
help achieve maximum effectiveness. In addition, despite the removal of a large number of dams worldwide over
the past decades, our knowledge concerning these effects is still limited due to the lack of long-term monitoring
data, which is critical to quantify the rate, magnitude and sequence of tributary habitat recovery to dam removal.
4.5. Efficiency Assessment on Conservation Measures
A variety of measures to conserve fish species impacted by river damming are available and each measure can
be effective under specific conditions. However, a challenge is to select appropriate measures based on real
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situations and cost-effectiveness. Table2 summarizes the major conservation measures and their advantages,
disadvantages as well as applicable conditions. We acknowledge that environmental decisions are complex and
require a nuanced understanding of local context and will almost always involve trade-offs.
Fishways are of benefit to fish species with strong swimming ability, but are mainly suitable for dams where
the water head is <60m. Fish lifts are space saving and mainly suitable for concrete gravity dams where the
Conservation measures Advantages Disadvantages Suitability of applications
Fish passage facilities Fishway 1. Promote timely fish passage Only effective for fish species
with strong swimming
abilities
All types of dams with a water
head between upstream and
downstream within 60m
2. Reduce damage to fish bodies
Fish lifts 1. Space-saving 1. Complex mechanical facilities Concrete gravity dams with
a water head between
upstream and downstream
over 60m
2. Higher possibility of failure
2. Easy to arrange in the dams 3. Limited number of passed
fishes
Fish collection and
transportation system
1. Flexible 1. High power consumption All types of dams with a water
head between upstream and
downstream over 60m
2. Adjust the replenishment
flow according to the fish
preference
2. Complex operation and
management
3. Fish mortality during
transportation
Fish-friendly turbine passage Ecological friendly turbines can
allow for the safe passage of
small objects
Physically damage or traumatize
fishes
Low and medium water head
hydropower plants
Juvenile bypass system 1. Long length, with good
ecological landscape function
1. Need more space All types of dams with a water
head between upstream
and downstream within
30m, and rely on tributary
projects
2. Effectively reduce the mortality
rate of fish passing through
the dam
2. High requirements of local
terrain conditions
3. Easy to adjust and expand the
trial run after completion
Artificial breeding and release 1. More convenient operation and
management
1. Longer exploration time All dammed rivers, especially
for conservation of
endangered or economic
fish species
2. Relatively mature technology 2. Reducing survival chance of
wild individuals
3. Protect fish species and increase
fish populations
3. Dilution of genetic diversity in
wild populations
Reservoir ecological operation 1. Restore the structure and
function of river ecosystems;
1. Loss of social and economic
benefits;
All types of dams with large or
medium-sized reservoirs
2. Difficulties in coordinating
departments;
2. Promote the fish spawning 3. Difficult to meet the needs
of different fish species
simultaneously
Habitat compensation in tributaries 1. Provide compensatory
habitat for fish affected by
hydropower in the mainstream;
1. Limited compensation ability
of the tributaries without
sufficient free-flowing length;
The presence of tributaries
with high habitat similarity
to the dammed mainstream
2. Restore the natural connectivity
of the river;
3. Promote changes in
the diversity of river
characteristics;
2. Demolition of small dams
in tributaries causes fish
casualties and adenosis
4. Promote fish reproduction
Table 2
Summary of Advantages, Disadvantages and Applicable Conditions of Fish Conservation Measures
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water head is >60m, but they are mechanically complex and have limited passage capacity. Fish collection and
transportation systems are flexible in space and time, and suitable for dams with water head >60m, but they are
complicated to operate and usually cause a high mortality rate during transportation. Fish-friendly turbines are
designed for low and medium water head hydropower plants, which can decrease the mortality and mechanical
damage to fish (Hogan etal.,2014; Pracheil etal.,2016; Watson etal.,2022). Juvenile bypass systems are mostly
used where the water head is <30m and there is a tributary present. Artificial breeding and release are effective to
protect endangered fish species and restore the resources of economic fish species, but it could affect the survival
and genetic diversity of the wild population. Reservoir ecological operation is an effective non-engineering
method for fish conservation in dammed rivers and is particularly applicable to large and medium-sized reser-
voirs, but it may cause certain loss of social-economic benefits. Habitat compensation in tributaries can be a
potential approach to conserve fish species that permanently lose their original habitats in the impounded main-
stream upstream of high dams, but their effectiveness depends on the ecological status of the tributaries and the
future development of the mainstream.
Quantitative evaluation on the effects of conservation measures is essential to select proper approaches and
improve their efficiencies. Table3 summarizes the major indicators and methods to assess these different conser-
vation measures.
Passage effectiveness and efficiency are the main indicators to evaluate the capability of fish passage facilities
(Bravo-Córdoba etal.,2021). Passage effectiveness is used to describe qualitatively the potential effect of fish
passage facilities on fish proliferation by checking that the passage facility is capable of letting all target species
pass through within the range of environmental conditions observed in nature during the migration period.
Passage efficiency is a quantitative evaluation indicator of fish passage effectiveness, which is defined as the
ratio of the number of species and quantities of fish individuals that migrate upstream through the fish passage
facility to the number of species and quantities of fish individuals that demand to pass the dam in a specific period
(Larinier,2008). The average efficiency of fish passage facilities is 50–60% (Hershey,2021). Evaluation of the
effectiveness of artificial breeding and release depends on the specific objectives of the measure, and generally
focuses on the growth of fry and the contribution to target fish resources as well as the related economic, ecolog-
ical and social benefits (Rytwinski etal.,2021). Ecological restoration aims at conserving endangered species,
and the evaluation mainly focuses on the survival rate and natural reproduction of artificially hatched fry after
Conservation measures Evaluation indicators Description and calculation
Fish passage facilities Fish passage effectiveness The potential effect of fish passage facilities on fish proliferation by checking that
the passage facility is capable of letting all target species pass through within the
range of environmental conditions observed in nature during the migration period.
Calculations are based on the monitoring data of fish species, numbers, sizes, life
stages, and behavior under the specific conditions of the operating fish passage
Fish passage efficiency The ratio of the number of fish species and quantities of fish individuals that migrate
upstream through the fish passage facility to the number of fish species and
quantities of fish individuals that demand to pass the dam in a specific period
Artificial breeding and releasing Survival rate after releasing Mark the fry to be released using marking techniques, and calculate the percentage of
fish containing the number of marks in the fish catch by recapture
Catch rate The percentage of the catch in a certain water body and a certain period of time to the
total resource of the fishing object in the water body during the same period
Reservoir ecological operation Ecological flow replenishment Replenishing the flow through reservoir operations to meet the minimum flow
demand of downstream ecosystems
Spawning volume The spawning volume that is monitored for the spawning grounds during the reservoir
ecological operation
Habitat compensation in tributaries Habitat diversity The overall richness of various types of habitats that accommodate various organisms
Fish diversity Biodiversity of indigenous fish species, including species richness, species
abundance, and phylo genetic diversity
Fish populations The total number of fishes that inhabits a certain area
Table 3
Summary of Evaluation Indicators for Fish Conservation Measures
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release (Lyu etal.,2021). Resource restoration aims at restoring fishery resources and improving the economic
benefits of fisheries, and thus the evaluation has mainly focused on the catch rate. The evaluation indicators for
reservoir ecological operation include replenishment of ecological flow and spawning volume of target fish in the
downstream reach of the dam (J. Li etal.,2019), particularly, the amount of spawning volume during reservoir
operation can well evaluate the effectiveness of ecological dispatch. The primary objective of habitat compen-
sation in tributaries is to protect the biodiversity of indigenous fish species, and the indicators for effectiveness
evaluation include habitat diversity, fish diversity and fish populations.
There are significant differences in the cost-benefits of different fish conservation measures for dammed rivers
(Table4). Fish passage has significant variations in cost-benefits, depending on the types of facilities. The Denil
fishway and fish lift have relatively low cost-benefits, while the juvenile bypass system and fish collection system
have relatively high cost-benefits. The Denil fishway costs approximately USD 124,000 per vertical meter and
has low maintenance as well as operation costs, but their average passage efficiency is only 16% (Noonan
etal.,2012). Fish lifts are expensive to build and operate, and costs roughly USD 2.4 million to install and an
annual maintenance charge of 5% (Noonan etal.,2012), although it has relatively high passage efficiency. Fish
collection systems need to temporarily preserve the transferred fish during long-distance transport, leading to
high mortality loss and operational costs. Juvenile bypass systems have relatively low construction and opera-
tional costs, and the average passage efficiency can reach 70%. However, it demands more space, which limits
its applicability.
The disposable investment of artificial breeding and release ranges from millions to hundreds of millions of USD,
and the cost-benefits depends on the bred fish species and scales of release (H. J. Chen,2019). For instance,
Chinese sturgeon spawns in the autumn, so they need to be incubated and reared indoor using heating devices to
promote their growth, which is expensive to maintain (Wei etal.,2004). Ecological restoration for the endangered
Conservation measures Costs Benefits
Fish passage facilities 1. Denil fishway: approximately US $124,000 per vertical
meter with low maintenance and operation costs
1. Denil fishway cost: average passage efficiency is only 16%
2. Fish lifts: roughly US $2.4 million to install and annual
maintenance charge of 5%
2. Fish lifts: relatively high operation efficiency
3. Fish collection systems: high mortality loss and operational
cost
3. Fish collection systems: high passage efficiency
4. Juvenile bypass system: relatively low construction and
operation cost
4. Juvenile bypass system: average passage efficiency is 70%
Artificial breeding and releasing 1. Range from millions to hundreds of millions of USD for
different scales of releasing and different bred fish species
(H. J. Chen,2019)
1. “Ecological restoration”: conserved the endangered native
fish populations and prevent their extinction. “Resource
restoration”: improved fishery with high economic values
2. A total of 31.55 million endemic fishes, including 24.16
million economic fishes which cost about US $0.48 billion
2. The number of fish species increased by 18 species in the
Yangtze River basin, effectively restoring the fishery
resources of the Yangtze River (Sun & Wang,2020)
Reservoir ecological operation 1. The power generation of cascade reservoirs decreased by
1.76%
1. The ecological flow coordination degree increased by
17.45%, promoting the spawning of the four major carps
(Dai etal.,2022)
2. The Gezhouba Hydropower Station lost 0.15% of its power
generation
2. The suitability of the spawning ground for Chinese
sturgeon increased by 39% (Y. Y. Wang etal.,2013)
3. The loss of power generation benefits is about 2.5% 3. Protect at least 50% of the target fish habitat in the river
(D. Chen etal.,2014)
Habitat compensation in tributaries 1. The Waterworks Dam was removed at a cost of US $0.214
million
1. Two years after the removal of the dam, the number of fish
species at the original site of Waterworks Dam increased
from the previous 11 to 26 species (Catalano etal.,2007)
2. The Marmot Dam was removed at a cost of US $4.86 million 2. The removal of the dam restored nearly seven miles of
river to migratory habitat for steelhead, Chinook salmon
and coho salmon (Xiao,2021)
Table 4
Summary of the Costs and Benefits of Conservation Measures
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native fish species can prevent their extinction in dammed rivers, and the value cannot be simply capitalized.
Resource restoration for fishery improvement in dammed rivers can bring high economic values. From 2005 to
2018, a total of 31.55 million endemic fishes, including 24.16 million economic fishes that cost about US $0.48
billion, have been released into the upper Yangtze River. The release of economic fishes has effectively restored
fishery resources, and the release of rare fishes has increased the number of fish species from 22 in 2006 to 40
in 2018 (Sun & Wang,2020).
It is widely perceived that reservoir ecological operations may sacrifice social-economic interests due to the
dedicated discharge of ecological flow or water head loss for withdrawing temperature stratified water. The
temperature regulation device on the Shasta Dam resulted in an estimated US $63 million loss of hydropower
revenue between 1987 and 1996 (Hallnan etal.,2020). However, many studies have shown that by optimizing
the operation of reservoirs, the ecological benefits can be ensured with a marginal loss, or even an increase of
socio-economic benefits. Under the optimal operation scheme, the Jinping cascade reservoirs in the Yalong River,
China, only sacrifice annually 2.5% hydropower production to conserve more than 50% habitat for Schizothorax
chongi in the dewatered reach (D. Chen etal.,2014). In the upper Yangtze River, under an optimal operation
scheme, the hydropower production loss of the cascade dams is only 1.76%, while the fulfillment of ecological
flow increases 17.45%, which greatly promoted the spawning of FMCCs (Dai etal.,2022). X. Wang etal.(2020)
proposed an optimized operation for the Three Gorges and Gezhouba cascade reservoirs that the total hydro-
power production increased by 250,089.2MW·h and the area of suitable spawning grounds of Chinese sturgeon
increased by 2.16%. Cioffi & Gallerano(2012) optimized hydropower production and fish habitat protection for
the Pieve di Cadore Reservoir in the Piave River, Italy, and the results showed that the area of fish habitat could
be increased with little loss of power generation. W. Chen & Olden(2017) designed a reservoir release scheme,
which could create proper conditions to favor native over non-native fish, while human water demands were
barely sacrificed. In general, reservoir ecological operation is an effective non-engineering conservation measure
with high cost-benefits.
Studies have shown that the cost of removing small weirs (≤3.0m in height) is US $69,000 on average or US
$23,000 per meter height, which is less than 20% of the cost of building a fish passage or less than 12% of the
cost of building a fish ladder (Garcia de Leaniz,2008), and dam removal can significantly increase fish species
richness. The removal of the Waterworks Dam in the Baraboo River, USA, a tributary of the Wisconsin River,
cost approximately US $0.214 million in 1998. Two years after the removal, the number of fish species increased
from 11 to 26 at the original dam site (Catalano etal.,2007). The Marmot Dam in the Sandy River, a tributary of
the Columbia River, was removed at a cost of approximately US $4.86 million. The removal has restored nearly
seven miles of river habitat for migratory fishes of steelhead, Chinook and coho salmons (Xiao,2021). In general,
habitat compensation and removing small dams in tributaries is likely a cost-effective approach to conserve fish
diversity in dammed mainstreams.
5. Future Perspectives
5.1. Strategic Plans for Development and Conservation
Mitigation actions should be taken to minimize the potential impacts on fish during the complete process of
river damming, including planning and operation. Planning of dams should be conducted at a system level to
ensure that decisions are made in a more holistic manner. Prior to dam construction, the intensity of hydro-
power development should be determined at a basin scale, in order to balance river ecosystem conservation and
economic benefits. It is also imperative to develop strategic dam planning, especially at the basin or regional
scale, by performing multi-criteria optimization schemes (Flecker etal.,2022). Adequate investigations should
be conducted to identify the siting of dams so as to minimize impacts on fish spawning, feeding and wintering
grounds within the local hydro-geophysical constraints. Research on the impacts of dam construction on river
ecosystem services, such as water supply, sediment transport, and biodiversity maintenance, should be strength-
ened. The planning of dams on mainstream and tributary rivers must be collaborative, and consider the optimal
combination of high dams, low dams and run-of-river dams, which is important to reduce impacts on the river
ecosystem at a basin scale (Couto and Olden,2018; Couto etal.,2021; Schmitt etal.,2019). For fish that have
lost their habitats in a mainstream that has been dammed, it is valuable to investigate the possibility of restoring
tributaries to provide fish with alternative habitats.
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Although there are plenty of studies concerning native fish conservation in dammed rivers, most focus on a
specific issue (e.g., such as fish passage or habitat restoration), and comprehensive conservation strategies that
consider the habitat requirements of fish species throughout their life history are lacking. Existing conservation
measures that aim at restoring the physical habitats for fish have had limited effectiveness, and there is demand
for more studies to refine these measures and ensure they better achieve their intended goals. Specifically, the
effectiveness and efficiency of the measures need to be assessed quantitatively by using long-term monitoring
data of dammed rivers, so that emerging problems can be identified in a timely manner, and thereby aid improved
designs for more efficient measures. It is of great value to develop a framework to assess systematically the
effectiveness and efficiency of conservation measures for target fish species from the genetic to population,
metapopulation, community, and ecosystem levels. NbS have shown a high potential to conserve fish species in
dammed rivers and deserve more attention in future studies. Moreover, it is important to incorporate more target
fish species when investigating and planning a conservation program. Finally, consideration of hydrokinetic
energy and submerged turbines is valuable, as these can generate electricity without many of the drawbacks of
dams, although the power yield is less.
5.2. Long-Term Systematic Observations
One important aspect of the knowledge gap concerning the impacts of river damming on fish and associated
conservation measures is the lack of long-term monitoring data. Compared to the long-term records of natu-
ral rivers from gauge-stations, similar data for dammed rivers are relatively short-term. Dam construction also
causes inconsistencies between data before and after impoundment. Changes in river morphology are usually
slow, and thus the evolution of channel morphology after river damming, and its impacts on fish physical habitats,
demand longer monitoring periods. Hysteresis exists in the response of river ecosystems to hydro-geophysical
changes, which also requires sufficiently long observations. TDG supersaturation only occurs during the occa-
sional flood discharge of dams, providing few monitoring opportunities for data collection. It is also difficult, if
not impossible, to obtain the near-field data of TDG. Moreover, the changes to the physical habitat and adaptation
of fish species in dammed rivers may interact with each other, and these changes increase the complexity of eval-
uating the impact of dam construction on fish communities and developing conservation measures. Therefore,
there is an urgent need to establish dedicated monitoring networks in dammed rivers to strengthen the collection
of long-term and systematic data to improve our understanding of the impacts of river damming on fish. Future
hydro-geophysical monitoring of large rivers can be revolutionalized by observing from satellites, such as the
SWOT mission. Emerging technology, such as eDNA, otolith microchemistry, and biotelemetry, can be used
to characterize the dynamics of fish communities in dammed and undammed rivers, thereby providing useful
comparisons. Such long-term monitoring data concerning hydro-geophysical conditions and fish communities
will deepen knowledge on the impacts of river damming on fish, which will then provide a fuller scientific basis
for developing conservation strategies.
Available information on the impact of dams on fish is based mainly on statistical analyses of data from field
observations and laboratory experiments concerning the relationships between key hydro-geophysical factors and
various fish-related endpoints. Some studies examine the behavioral responses of target fish species to variations
of key hydro-geophysical factors in the laboratory to establish suitability curves, which are then used to develop
fish habitat models for impact assessment or prediction. These studies have made significant contributions to
evaluating the impacts of river damming and the design of fish conservation measures. In future, the adoption of
genomic, transcriptomic, proteomic, metabolomic and bioinformatic methods can be a viable approach to inves-
tigate the physiological mechanisms of how altered hydro-geophysical conditions affect gonad development, sex
differentiation, gene regulation, and gene expression of target fish species (Natri etal.,2019; Ortega-Recalde
etal.,2020). In addition, neurotoxic effects of river damming on fish behavior, personality and cognition could, in
turn, potentially generate feedback loops that may amplify the effects on fish. Therefore, integrative approaches
that combine field observations with novel technologies (e.g., molecular omics techniques, biotelemetry) are
recommended to bridge the knowledge gap in assessing river damming impacts on fish. This will enable the
development of more reliable models to predict long-term consequences, and thus support the implementation of
effective conservation measures.
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5.3. Complex Effects of Climate Change and Land Cover Changes
Climate change has dramatic impacts on river hydrology and thermal regimes, and thus affects the migration,
reproduction, growth and distribution of fish species in various ways. Regional patterns of warming-induced
changes in surface hydroclimate are complex, with evidence of increases and decreases in the magnitude of
precipitation and runoff, as well as frequency, depending on the local context (Milly etal.,2005). The mean and
high water temperature of global rivers are projected to increase under climate change, with the impact varying
between regions and across seasons. Climate change may thus exacerbate the effect of river damming on fish
habitats. Reservoir operation has disturbed natural hydrological regimes and riverine hydrodynamic conditions,
and climate change could bring additional alterations by affecting the patterns of global precipitation and snow-
melt. Climate warming may increase river water temperatures, prolong stratification and decrease vertical mixing
in reservoirs. Such prolonged stratification intensifies anoxia, which promotes the release of nutrients from sedi-
ment and stimulates eutrophication in reservoir waters, thus affecting fish community structure and food web
dynamics. Meanwhile, increases in water temperature reduce the dissolved oxygen content of water bodies, and
the stratification of reservoirs intensifies the deficiency of dissolved oxygen in the bottom layer, which affects
the survival of benthic fish species. However, reservoir operation can also mitigate impacts of future climatic
variability and climate change on fish habitats, for example, by releasing cold water to offset impacts of warming
temperatures on cold-water fish species (Benjankar etal.,2018). Climate change and reservoir retentions jointly
influence river sediment regime, which affects river morphology and substrate composition, thereby impacting
on fish habitat (J. Li etal.,2021). Future studies concerning the effects of climate change on fish should take into
account changes in watershed soil erosion, sediment flux and nutrient loss, which play an increasingly important
role in river ecosystems, when assessing the impacts of dams on fish (Best etal.,2022; J. Li etal.,2021). Due
to the lack of integrated data concerning climate change, dam regulation and the evolution of fish communities,
the interactive multi-stressor impacts of climate change and river damming on fish demand further investigation.
Climate change and dam construction also affect land use, which can directly or indirectly impact river ecosys-
tems. For example, climate change affects the amount and distribution of vegetation, agriculture and forestry, and
dam construction can promote rapid urbanization along the river. This significantly increases impervious surface
areas that lead to higher flood flows and earlier flood timing, and thus intensifies the impact on fish in dammed
rivers. The development of industrial and residential areas results in increased discharge of industrial wastewater
and domestic sewage, which could change local water temperatures and nutrient levels, and thereby affect fish
migration, spawning and feeding. Therefore, the impacts of river damming on fish could become extraordinarily
complex under the effects of climate and land use change. This highlights the need to consider the full range of
stressors affecting rivers, identify the major factors, and assess both their interactions and timescales of their
effects in future studies (Best & Darby,2020). Advances in numerical modeling and data collection could be
used to develop virtual watersheds, where different climate change forcings can be used to simulate processes in
river-reservoir systems (Benjankar etal.,2018; Tranmer etal.,2020). Preliminary implementations of this meth-
odology have shown its benefits in understanding the impact of dam operations on fish habitat.
Data Availability Statement
Most of data supporting the figures are available via the cited references. Data of Figures1a and1b are avail-
able through Lehner etal.(2011). Data of Figures1c and1d are available through Zarfl etal.(2015). Data of
Figure3a are calculated from Hydrological Data of Changjiang River Basin in Annual Hydrological ReportP.R.
China, including average monthly discharge data of Pingshan station (2007–2010) and Xiangjiaba hydrological
station (2016–2020). Data of Figure 3b are available from Hydrological Data of Changjiang River Basin in
Annual Hydrological ReportP.R. China, including average monthly discharge data of Ningnan hydrological
station in 2015 and 2019. Data of Figure3c are from the authors (Q. Li,2023). Data of Figure3d are available
from Q. Chen etal.(2021). Data of Figure5a are calculated from Hydrological Data of Changjiang River Basin
in Annual Hydrological Report P.R. China, including average monthly water temperature data of Pingshan
station (2007–2010) and Xiangjiaba hydrological station (2016–2020). Data of Figure5c are available through
T. Li etal.(2021). Data of Figure6b are available through Y. Wang etal.(2015). Data of Figure16 are available
through Q. Chen etal.(2021). Data of Figure17e are available through S. He etal.(2021).
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Acknowledgments
This work was supported by National
Key Program of Science and Technol-
ogy (2022YFC3203900), and National
Natural Science Foundation of China
(No. 52121006, 92047303). Dr. Chen Q.
was also supported by the Xplorer Prize.
The authors are grateful to Dr. Honghai
Ma, Dr. Ting Li, Mr. Zhiyuan Zhang, Mr.
Yumeng Tang, Ms. Qi Zhang, Ms. Mo
Chen, Ms. Mengru Wei, Mr. Zaoli Yang,
and Mr. Peisi Yang for their contribution
in material collections. We also greatly
appreciate the valuable comments and
constructive suggestions from the anony-
mous reviewers.
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