Science of the Total Environment 710 (2020) 136242

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Short communication

Extinction of one of the world's largest freshwater fishes: Lessons for

conserving the endangered Yangtze fauna
Hui Zhang a,⁎, Ivan Jarić b,c, David L. Roberts d, Yongfeng He a,⁎, Hao Du a, Jinming Wu a,
Chengyou Wang a, Qiwei Wei a,⁎
a
Key Laboratory of Freshwater Biodiversity Conservation, Ministry of Agriculture and Rural Affairs of P. R. China, Yangtze River Fisheries Research Institute, Chinese Academy of Fishery Sciences,
Wuhan, Hubei Province 430223, PR China
b
Biology Centre of the Czech Academy of Sciences, Institute of Hydrobiology, České Budějovice 37005, Czech Republic
c
University of South Bohemia, Faculty of Science, Department of Ecosystem Biology, České Budějovice 37005, Czech Republic
d
Durrell Institute of Conservation and Ecology, School of Anthropology and Conservation, University of Kent, Canterbury, Kent CT2 7NR, UK

H I G H L I G H T S G R A P H I C A L A B S T R A C T

• The giant Chinese paddlefish (up to 7 m

in length) was found to be extinct.
• The timing of extinction was estimated
to be by 2005–2010.
• The paddlefish became functionally ex-
tinct by 1993, prior to extinction
• Conservation efforts on endangered
Yangtze fishes are urgently needed.

a r t i c l e i n f o a b s t r a c t

Article history: The mega river ecosystem of the Yangtze River was once home to diverse aquatic megafauna but is increas-
Received 30 September 2019 ingly affected by various anthropogenic stressors that have resulted in continuous loss of biodiversity, such
Received in revised form 16 December 2019 as the probable extinction of Yangtze River Dolphin. The Chinese paddlefish, Psephurus gladius, was one of
Accepted 18 December 2019
only two extant members of a relict lineage that was most diverse and widespread 34–75 million years ago.
Available online 23 December 2019
It is also one of the largest freshwater fish species, reaching up to 7 m in length. The Chinese paddlefish was
Editor: Sergi Sabater once common in the Yangtze River, with c.25 t being harvested per annum during the 1970s. Populations
have, however, declined drastically since the late 1970s as a result of overfishing and habitat fragmentation.
Keywords: Here, a basin-wide capture survey during 2017–2018 found 332 fish species, but did not find a single spec-
Chinese paddlefish imen of Chinese paddlefish. Furthermore, 140 historically reported fish species have not been found and
Endangered fish most of them are considered highly endangered. Based on 210 sightings of Chinese paddlefish during the
Sighting record period 1981–2003, we estimated the timing of extinction to be by 2005, and no later than by 2010. In addi-
Functional extinction tion, the paddlefish probably became functionally extinct (i.e. it was unable to reproduce) by 1993, before it
Extinction debt
went extinct. It is likely that the lack of reproduction was among the major causes of extinction. As no indi-
Yangtze River
viduals exist in captivity, and no living tissues are conserved for potential resurrection, the fish should be

⁎ Corresponding authors at: Yangtze River Fisheries Research Institute, Chinese Academy of Fishery Sciences, No.8, 1st Wudayuan Road, Donghu Hi-tech Development Zone, Wuhan
430223, Hubei Province, PR China.
E-mail addresses: zhanghui@yfi.ac.cn (H. Zhang), heyf@yfi.ac.cn (Y. He), weiqw@yfi.ac.cn (Q. Wei).

https://doi.org/10.1016/j.scitotenv.2019.136242
0048-9697/© 2019 Elsevier B.V. All rights reserved.
2 H. Zhang et al. / Science of the Total Environment 710 (2020) 136242

considered extinct according to the IUCN Red List criteria. The delayed extinction of Chinese paddlefish re-
sulted from multiple threats, suggesting that optimizing conservation efforts on endangered Yangtze fauna
is urgently needed.
© 2019 Elsevier B.V. All rights reserved.

1. Introduction (Tenualosa reevesii, Data Deficient, DD) are thought to be functionally

The Yangtze River is the third longest (N6300 km) mega river eco-
system in the world. The river crosses four climatic zones from the
west to the east of China, with a basin area of 1.8 × 106 km2 and an
elevation difference of 5400 m. The diversity of climate, hydrology and
habitat create a rich aquatic biodiversity. It is thought that there are
N400 fish species in the basin, of which 45% (n = 178) are endemic
(Ye et al., 2011). The Yangtze River is also home to several species that
are globally recognized for their unique evolutionary and biological
characteristics, such as aquatic mammals Baiji or Yangtze River Dolphin
(Lipotes vexillifer, listed as Critically Endangered/CR species on the IUCN
Red List of Threatened Species) and the Yangtze finless porpoise
(Neophocaena asiaeorientalis, Endangered, EN), reptiles such as the
Chinese alligator (Alligator sinensis, CR), as well as an amphibian, the
Chinese giant salamander (Andrias davidianus, CR; MEP-PRC and CAS,
2015; IUCN, 2019).
The Yangtze River Basin has undergone intensive economic develop-
ment since the 1950s, which has supported three megalopolis clusters
(N40 cities alongside the river), and is now producing 40% of the GDP
of China and supporting a third of the Chinese population (0.4 billion;
Yang et al., 2007, 2009, 2011). However, intensive development led to
severe declines in the region's biodiversity (Vörösmarty et al., 2010;
Qiu, 2012; Liu et al., 2019). Currently, N70 aquatic animals are listed as
internationally and/or nationally protected species (MEP-PRC and CAS,
2015; IUCN, 2019). Several species, such as Baiji and Reeves shad
extinct, while Chinese paddlefish (Psephurus gladius) and Chinese stur-
geon (Acipenser sinensis) are listed as CR species (MEP-PRC and CAS,
2015; IUCN, 2019). Habitat loss, fragmentation, and deterioration due
to damming, water flow regulation, lake reclamation, water pollution,
navigation, urbanization, and aquaculture development, as well as pop-
ulation declines due to overfishing, represent the major threatening
factors (Yang et al., 2007, 2009, 2011). For instance, the Yangtze River
is currently the busiest river globally in terms of inland navigation. Nev-
ertheless, the river is currently under a new development plan (the
“Yangtze River Economic Belt”) to further promote the economic devel-
opment of China, which will further increase human pressures on this
unique river system. If no effective protection measures are taken, the
Yangtze River aquatic ecosystems will be in danger of collapse.
The Chinese paddlefish (P. gladius) was one of the two recently
extant species of paddlefishes (the other being the American paddlefish,
Polyodon spathula; Vulnerable/VU; Bemis et al., 1997), and has been
recorded to attain a length of up to 7 m, making it one of the largest
freshwater fishes in the world (Xie, 2003). Historically, the Chinese pad-
dlefish was distributed throughout many large rivers that flow into the
West Pacific (Fig. 1A). Since 1950, it was commonly seen only in the
Yangtze River (Fig. 1B; YARSG, 1988), where it was commercially
exploited, with an estimated 25 t harvested annually during the 1970s
(Wei et al., 1997; Phelps et al., 2016). Since the late 1970s populations
have gone through a drastic decline due to overfishing and habitat frag-
mentation, especially after the construction of the first and lowermost

B Maximum: 7 m in record

Fig. 1. Chinese paddlefish (Psephurus gladius) and its distribution range, (A) Contraction of the distribution range over the past hundred years; historic distribution included Liao, Hai,
Yellow, Huai, Yangtze and Qiantang rivers, and coastal areas; however, since the 1950s it was commonly sighted only in the Yangtze River and its estuary (in pink; YARSG, 1988),
(B) A specimen found below the Gezhouba Dam in 1993.
 H. Zhang et al. / Science of the Total Environment 710 (2020) 136242
dam (Gezhouba Dam) on the Yangtze main stem in 1981 (Wei et al.,
1997; Phelps et al., 2016). The dam separated the fish into two isolated
populations and blocked the migration route, preventing adult fish from
moving to the upper reaches of the river to spawn.
With the drastic decline, the fish has been listed as one of the top-
level protected animals in China since 1989. During its most recent
threat assessment by the IUCN in 2009 (Wei, 2010), the Chinese paddle-
fish was re-evaluated as Critically Endangered (CR, Possibly Extinct),
having been previously listed as CR in 1996. The species continued to
decline following the construction of the Gezhouba Dam, with occa-
sional sightings until the last known record in 2003 (Zhang et al.,
2009, 2016). The species continued survival is increasingly questioned.
Here, we evaluated the status of Chinese paddlefish based on a large-
scale basin-wide capture survey in the Yangtze River basin during
2017 and 2018, and an assessment based on sighting records from
1981 onwards (Collen et al., 2010; Boakes et al., 2015). The aim of this
study is to assess the population status of Chinese paddlefish, and
whether the conservation measures are timely and efficient, and there-
fore likely to support future conservation and rehabilitation of other
endangered aquatic species.
2. Methods
2.1. Study area
The Yangtze is the longest river in Asia with the main stem crossing
11 provinces in China (Fig. 2). Most of the areas within the basin have a
subtropical monsoon climate. The basin has 437 tributaries with the
catchment area of N1000 km2, and 27 lakes of N100 km2 (Yu and Lu,
B C
500 m
Fig. 2. Study area of the Yangtze River basin, (A) The Yangtze River basin and 65 stationary monitoring sites, (B) Historic spawning reach of Chinese paddlefish, image made on June 28,
2011 (Google Earth), (C) The Gezhouba Dam, the first and lowermost dam on the Yangtze main stem, built in 1981 with no fish passages, image made on February 23, 2018 (Google Earth).
3
2005). In 1981, the Gezhouba Dam (Fig. 2C, located c. 1700 km from
the river estuary and without a fish passage) was built on the Yangtze
main stem and is the lowermost dam on the main stem. The dam
blocked the migration of all migratory fishes including that of Chinese
paddlefish. As the spawning area of Chinese paddlefish was located in
the upper Yangtze (Fig. 2B, c. 2740 km from the estuary), adult fish
are prevented from swimming upstream to spawn, while juveniles can-
not reach downstream feeding areas.
2.2. Field survey
In 2017–2018, a basin-wide capture survey was conducted to inves-
tigate the present status of Chinese paddlefish, specifically focusing on
the area of the Yangtze main stem, important tributaries (Yalong
River, Heng River, Min River, Tuo River, Chishui River, Jialing River,
Wu River, Han River), and two large lakes (Dongting Lake and Poyang
Lake; Fig. 2). In 2017, the whole basin was divided into a 20 × 20 km
sampling grid, and the aim of our survey was to cover all of the grid
cells within the survey area. The survey was conducted four times
(once in each season) in all of the areas where the weather conditions
permitted field work. Sampling methods included active and passive
sampling. Active sampling comprised multiple fish sampling methods
whenever possible, such as gill nets, ground cages, set nets, electro fish-
ing, etc. As the survey area comprises variety of aquatic habitats, the
length of gillnets varied among sampling localities to better fit the
local conditions. In general, a triple gillnet with 30–50 m long, 2–4 m
high and 2–3 cm mesh size was used. The fishing gear was set up at
dusk and taken in the early morning of the following day, with the
usual sampling time of about 10 h. Passive sampling was represented
500 m
4 H. Zhang et al. / Science of the Total Environment 710 (2020) 136242
by investigation of the traditional fisheries markets in the major cities
and towns within the study area. The traditional fisheries markets are
those which sell fresh, wild-caught fish, and are usually located near riv-
ers or lakes. All the methods were set to provide full coverage of fish
species in the studied area. Each survey lasted about 1–2 months. In
2018, the survey was mainly conducted in 65 stationary monitoring
sites (Fig. 2A), as well in the areas in their vicinity. Two surveys were
conducted, in spring and autumn. For each survey in each site, catches
from 30 or more fishing boats were checked, or the survey was alterna-
tively continued for 15 days when the fishing boats were rare.
2.3. Sighting collection
We collected all available literature (scientific articles, books,
reports, and news articles) related to Chinese paddlefish from various
databases (including China Knowledge Resource Integrated Database,
Chongqing VIP Database, and Web of Science). In total, 80 publicly avail-
able documents were assessed and sightings were gathered (mainly
originating from catch, bycatch, sightings of dead specimens, etc.). We
also collected historic unpublished data from the Yangtze River Fisher-
ies Research Institute, Chinese Academy of Fishery Sciences, Wuhan,
which has the longest record in studying sturgeons and paddlefish in
the Yangtze River.
2.4. Extinction inference
Extinction was inferred by the Optimal Linear Estimation (OLE)
method (Roberts and Solow, 2003; Solow, 2005). This method assumes
that the joint distribution of the most recent sightings follows the form
of the Weibull extreme value distribution, and by estimating the shape
parameter of this distribution it infers the time of extinction by the
optimal linear estimation (Solow, 2005). While there are various
methods designed to infer extinction based on sightings (Boakes et al.,
2015), we chose to use OLE because it has been widely used and consid-
ered to be the most robust approach within this group of methods, espe-
cially in case of declining populations (Rivadeneira et al., 2009;
Clements et al., 2013, 2014). OLE assumes presence of a continuous
sighting effort throughout the studied period, but it makes no assump-
tion regarding the actual sighting rate, which makes it more flexible
compared to other methods (Boakes et al., 2015). It was also shown to
perform fairly well under various types and magnitudes of sighting
effort and sighting probability (Rivadeneira et al., 2009). Furthermore,
although it was not originally developed to account for varying reliabil-
ity of sightings (Roberts et al., 2010), application of the weighted resam-
pling (BBJ) method (Brook et al., 2019) allows it to effectively
incorporate observational reliability into extinction estimates. It has
been also advocated as a suitable tool for extinction risk analysis for
Critically Endangered - Possibly Extinct species CR (PE) within the
IUCN Red List (Collen et al., 2010). As such, it was considered to be the
optimal method for the present study. Since there was no information
on the exact dates of each sighting, multiple sightings within a year
were pooled and considered as a single sighting. OLE was used to esti-
mate the time of extinction and the upper bound of one-sided 95% con-
fidence interval.
2.5. Functional extinction inference
Functional extinction, which represents time-delayed deterministic
extinction caused by a permanent lack of reproduction or recruitment,
was estimated based on the method by Jarić et al. (2016), using the
sighting time and age estimate of each observed specimen. We also
explored the effect of impeded migration route by the Gezhouba Dam
(completed in 1981) on the functional extinction of this species. The es-
timation of functional extinction is based on a simple population
dynamics model and the analysis of a likelihood of a temporal gap
with no confirmed birthyears of observed individuals near the end of
the sighting record, based on their age estimates (Jarić et al., 2016).
The necessary input data are sighting records with the time of sighting
and the age estimate of each individual. Analyses of the method perfor-
mance and sensitivity indicated that it performs well even with limited
data available, except in scenarios with very low natural mortality and
small number of sightings, where the method tends to be more conser-
vative (Jarić et al., 2016). The method was developed specifically for
data-deficient situations, where the population of studied species and
its demography are known only through rare sightings (Jarić et al.,
2016), which makes it suitable for the present study.
For sightings with body size reported but not the age, we used the
von Bertalanffy growth curve (VBGF) equation from Ma et al. (1996)
to estimate the age. The VBGF equations for males and females are:
h i
Male : Lt ¼ 368:5  1−e−0:0901ðtþ1:66Þ
h i
Female : Lt ¼ 436:3  1−e−0:0724ðtþ2:29Þ
where Lt is the total body length, t is the age, and the theoretical maxi-
mum length of females and males are 436.3 and 368.5 cm, respectively.
If sex was not determined, we used the average age estimated based on
male and female VBGF. Perturbation analysis, which assessed the effect
of imprecise age estimation of fish on the functional extinction infer-
ence, was conducted in line with Jarić et al. (2016), by randomly chang-
ing each estimated age by ±3 years.
2.6. Species longevity estimates
There are no available data on Chinese paddlefish longevity. Based
on the available data on the age at maturity of females (6–15; Mims
et al., 1993; 6–16; Ma et al., 1996), natural mortality estimated in the
present study (0.119–0.132, see below), and the methods that establish
relationship between the age at maturity and longevity (Jarić and Gačić,
2012), as well as between natural mortality and longevity (Hoenig,
1983), paddlefish lifespan was estimated to range from 29 to 38 years.
However, such estimates are likely to be gross underestimations of spe-
cies longevity in an undisturbed population, since the data was based on
heavily exploited and in other ways impacted population.
3. Results
3.1. On-site surveys
During our basin-wide capture survey 332 fish species were identi-
fied (Supplementary Table 1), including 274 inland, 32 estuarine and
26 marine species, with 293 species reported previously in the Yangtze
Basin. Based on the data on fish diversity in the same area from Fu et al.
(2003; 361 species) and Ye et al. (2011; 416 species) species recorded
during our survey represented between 70.4% and 81.2% of the total
fish fauna. The remaining 39 species (11.8%) have never been previously
reported in the area. On the other hand, there were 140 species previ-
ously reported to occur in the Yangtze basin, which were not recorded
during the survey, most of them highly endangered species. Around
69% (99/144) species listed on the IUCN Red List were recorded by our
survey. In addition, we recorded 25 exotic species among 332 identified
species, out of which 11 were never observed in the basin before. The
high number of the observed species, and especially the number of spe-
cies observed in the area for the first time, implies that our sampling
effort was sufficient to cover most of the species present in the area.
However, we did not find a single specimen of Chinese paddlefish.
3.2. Sighting record collected
A total of 210 sightings during 1981–2003 (Fig. 3, Supplemental
Table 2) were collected, of which 45 sightings have detailed information
 H. Zhang et al. / Science of the Total Environment 710 (2020) 136242
(body length, age, sighting location, etc.; Supplemental Table 3). Most of
the sightings (200, 95.2%) occurred before 1995, with a peak in sighting
frequency around 1985. Although the commercial capture on Chinese
paddlefish was prohibited since 1983, the bycatch from intensive
fishery (gill nets, setlines, etc.) below the dam did not decrease. Approx-
imately 22.4% (47) sightings were found in the upper Yangtze, while
most of the sightings (75.7%, 159) were found below the Gezhouba
Dam (Fig. 3C). A total of 45 sightings have detailed biological informa-
tion (Supplemental Table 3), with the total length and body weight
ranging from 47 to 363 cm and 0.4 to 200 kg, respectively, and with
10 female and 25 male individuals, and 10 with sex unknown.
3.3. Inference of extinction and functional extinction
The OLE method suggested that the Chinese paddlefish became
extinct by 2005, with an estimated upper bound of a 95% confidence in-
terval for extinction time in 2010. To explore the potential impact of
Gezhouba Dam (built in 1981; Zhang et al., 2017) on the functional
extinction of the species (Jarić et al., 2016), 45 sightings were divided
into upstream (above the dam: includes Yibin, Luzhou, and Chongqing
reach) and downstream section (below the dam), as well as assessed
as a single dataset (i.e., both above and below the dam). Analysis con-
firmed functional extinction of the species for the upstream section
(i.e., null hypothesis that functional extinction has not occurred was
rejected, p = 0.002), but not for the downstream section (p = 0.119).
The upper bound of the confidence interval for the functional extinction
of the upstream population was in 1993. Perturbation analysis (by ran-
domly changing each estimated age by ±3 years) indicated that the
results are robust - functional extinction was supported 98% of time.
This indicates that the lack of reproduction was likely one of the major
causes of extinction in the upstream section. Functional extinction by
1993 (the upper end of the confidence interval) is also supported by a
long period since that year, which almost reaches the estimated lifespan
of this species (29–38 years) and would thus make continued survival
with no reproduction highly unlikely. The method also provided
estimates of natural mortality for the species. The annual natural mor-
tality of Chinese paddlefish for the upstream and downstream popula-
tion was estimated to be 0.119 +/− 0.002 and 0.132 +/− 0.020 SD,
respectively.
4. Discussion
Although the accuracy of the methods applied here may be affected
by various environmental factors (Clements et al., 2013, 2014),
sightings represent a useful source of information to infer the extinction
and the time of extinction when the taxon is extremely rare and has a
Fig. 3. Sightings of Chinese paddlefish Psephurus gladius in the Yangtze River from 1981 onwards, (A) All sightings and sightings with biological information by year, (B) Sightings in each
river reach by year: Yibin reach (river km/rkm 2741.7–2622.2), Luzhou reach (rkm 2622.2–2511.2), Chongqing reach (rkm 2511.2–1813.2), and reach below the Gezhouba Dam (rkm
1678.0–0), (C) Total sightings in each reach.
5
long life-span (Roberts and Solow, 2003; Solow, 2005; Boakes et al.,
2015; Jarić et al., 2016). Here, a total of 210 sightings from 1981 to
2003, including 45 with detailed information (e.g. body length, age,
sighting location), were collected from available literature as well as un-
published data. While other survey data do exist, notably that of Turvey
et al. (2010), they were not included as they are based on oral, indirect
and potentially unreliable reports by fishermen, and thus represent a
different type of information that requires a different set of methods
that account for uncertainty.
Both historic and present field surveys provided validation of the
inference of extinction (Fig. 4). Eight hydroacoustic (zigzag mobile sur-
vey by a 199 kHz split-beam echo sounder) and trial capture surveys
(by drift nets and setlines) during 2006–2013 in the historic spawning
grounds of the species (covering an ~400 km reach in the upper Yangtze
main stem) did not result in a single specimen (Zhang et al., 2009, 2016).
Based on the series of bulletins that include information on annual mon-
itoring activities of rare and endemic aquatic animals, fishery resources,
environment, and water quality, issued by the Ministry of Environmental
Protection of the People's Republic of China during 1997–2017, it can be
concluded that the Chinese paddlefish is one of a very few fish species
which has never been observed for N15 years (Supplementary Table 1).
Functional extinction analysis indicated that the lack of reproduction
was likely one of the major causes of extinction in the upstream section,
which was probably driven by habitat fragmentation that blocked
upstream spawning migrations, and the resulting reduced spawning
probability (i.e., Allee effects). As a large piscivorous fish and an apex
predator, it is likely that its population was never large, and was further
reduced by over-exploitation during 1970s and dam construction, thus
making breeding individuals rare and successful reproduction unlikely.
Functional extinction, as a persistent reproductive failure, can be driven
by various factors and mechanisms, including the lack of key habitats,
genetic erosion and inbreeding, phenological shifts, diseases, sex-ratio
bias and population fragmentation (Jarić et al., 2016). Which of these
factors contributed to functional extinction of Chinese paddlefish in
the upstream Yangtze section, and to what extent, is yet to be fully
resolved. However, functional extinction was not confirmed for the
downstream section, which is probably to be expected since the down-
stream section remained open for spawning migrations of this anadro-
mous species. There, bycatch and habitat degradation likely played a
key role in extinction.
We found that Chinese paddlefish is likely to have become extinct
before the most recent IUCN Red List assessment in 2009, based on
extensive surveys and a statistical assessment of its sighting record. Cur-
rently, no individuals of the Chinese paddlefish exist in captivity, and no
living tissues have been conserved for the potential resurrection of the
species. Although artificial gynogenesis (Zou et al., 2011a, 2011b) and
6 H. Zhang et al. / Science of the Total Environment 710 (2020) 136242
Carcass found below the dam
221 BC - 1271 AD,
Acipenseriformes commonly seen in
existed 200 Yangtze, Yellow River, etc.
Million Years
Before Present 1949, P. R. China
(MYBP) established
1970s, annual
harvest ~25 t in
34-75 MYBP, fossil Yangtze River
records of the four 1271-1840 AD,
Number (ind.)
exnct paddlefish gradually disappeared
species in Yellow River
Fig. 4. Timeline of key events related to the decline and extinction of the Chinese paddlefish (Psephurus gladius).
surrogate broodstock technologies (Takeuchi et al., 2004) are at dis-
posal, ready to be used for rescuing the species, success of such efforts
depends upon finding live individuals (Phelps et al., 2016). Based on
the weight of the evidence, the species can be declared as extinct with
high certainty. The Acipenseriformes, which include sturgeons and
paddlefishes, have existed since the Lower Jurassic, 200 million years
ago, and occupy a special place in fish evolution (Bemis et al., 1997).
Among them, six paddlefishes (Polyodontidae) are known, with four
only known from the fossil record (Phelps et al., 2016). Given that the
Chinese paddlefish was one of the two extant species of paddlefishes,
loss of such unique and charismatic megafauna representative of fresh-
water ecosystems is a reprehensible and an irreparable loss. In case of an
unlikely event of sighting and capture of a Chinese paddlefish, such
individual would have to be carefully cultivated in controlled environ-
ment, with surrogate broodstock technologies applied to restore the
species by germ cells obtained from the captured individual (Takeuchi
et al., 2004). In addition, the ongoing search and rescue efforts for
other possibly extinct species, such as Baiji and Reeves shad, should be
intensified and maintained as long as possible, following established
criteria for declaring species extinct (Akçakaya et al., 2017).
There are multiple lessons to be learned from the extinction of Chi-
nese paddlefish. Firstly, there is a need for a new comprehensive survey
of the whole Yangtze River basin, which should be thereafter conducted
on a regular basis (for instance, at a five-year interval). The first compre-
hensive survey in the Yangtze basin was conducted during 1973–1975.
Our present survey was the second comprehensive survey of the basin,
with a gap of about 41 years between the two surveys. The 2017–2018
survey failed to provide any records of 140 fish species, but due to a lack
of regular and consistent surveys it is not possible to determine the fate
of these fishes, nor to implement timely and effective protection mea-
sures. Secondly, fish communities can manifest lagged responses to
multiple human threats (Kuussaari et al., 2009). This study indicated
that the critical time to rescue Chinese paddlefish was before 1993
(i.e., before its functional extinction), or at least before 2005 (the esti-
mated year of extinction). Unfortunately, all substantial rescue efforts
(including the hydroacoustic and trial capture surveys in historic
spawning grounds, the research on artificial gynogenesis techniques,
etc.) were conducted after 2006 (Zhang et al., 2009, 2017; Zou et al.,
2011a, 2011b), when it was already too late to prevent the extinction.
As there are many species in the Yangtze River basin that are considered
to be on the edge of extinction, it is critical to estimate their extinction
risk using various suitable methods, including the use of extinction
likelihood estimators based on sighting events (Boakes et al., 2015)
and population viability analysis (Brook et al., 2000) Extinction risk as-
sessments for all endangered Yangtze species should be conducted as
1993, funconal
exncon
2005,
exncon
1981, Gezhouba 2017-2018, not
Dam on Yangtze 2003, Three Gorges found in a basin-wide
main stem built Projects iniated capture survey
2000
1981-2002, 2006-2013, not found
bycatch below 2003, last live in historic spawning
Gezhouba Dam specimen observed reach (~400 km)
30
20
10
0
1980 1985 1990 1995 2000 2005
Year
soon as possible, to allow the development of appropriate conservation
measures and their prioritization, and thus avoid potential further
extinctions. While for some species the window of conservation oppor-
tunity may have already closed, for those that still persist it is important
to seize what opportunity remains. Thirdly, when the investments are
limited, conservation priorities should be focused on species that have
not been found for many years, with no natural production, or with
rapidly declining populations due to critical habitat loss. For example,
the long spiky-head carp (Luciobrama microcephalus) and Sichuan
shoveljaw fish (Onychostoma angustistomata) have not been observed
for many years, even though neither of them have been so far evaluated
within the IUCN Red List (MEP-PRC and CAS, 2015; IUCN, 2019). In
addition, some species that are confirmed to be present are known to
be rapidly declining with no natural reproduction observed for years,
such as Chinese sturgeon, Yangtze sturgeon (A. sinensis), Chinese sucker
(Myxocyprinus asiaticus, NE, not evaluated by IUCN), Sichuan taimen
(Hucho bleekeri, CR) and largemouth bronze gudgeon (Coreius
guichenoti, NE; MEP-PRC and CAS, 2015; IUCN, 2019). Prioritizing con-
servation efforts on those species with the greatest extinction risk is
particularly urgent.
In the Yangtze basin, there are N70 aquatic animals listed as interna-
tionally and/or nationally protected species, while some other species
are known to be highly endangered but not yet listed as such. To protect
these species, firstly appropriate habitat conservation measures should
be implemented and extended to the whole Yangtze basin. From this
point of view, ongoing in situ, ex situ, and socio-economic measures,
such as establishment of aquatic protected areas, stock enhancements
for rare and endemic species, extinction risk assessments, increasing
the number of species listed as protected, implementing action plans
for key species, habitat restoration, and strengthening laws and law en-
forcement, seem to have limited effects in the diverse and highly
affected river ecosystems within the Yangtze basin. To be truly effective,
these measures will have to be greatly intensified in the future. How-
ever, corresponding measures should have to be specifically designed
for each species based on its biological and ecological characteristics.
Sustainable development and biodiversity conservation will have to
be widely recognized as priorities for the whole Yangtze River basin,
to prevent and mitigate ongoing irreversible changes in the region.
Supplementary data to this article can be found online at https://doi.
org/10.1016/j.scitotenv.2019.136242.
CRediT authorship contribution statement
Hui Zhang: Conceptualization, Data curation, Writing - original
draft, Funding acquisition. Ivan Jarić: Formal analysis, Methodology,
 H. Zhang et al. / Science of the Total Environment 710 (2020) 136242
Writing - original draft, Writing - review & editing, Funding acquisition.
David L. Roberts: Formal analysis, Methodology, Writing - review &
editing. Yongfeng He: Data curation, Formal analysis, Methodology.
Hao Du: Data curation. Jinming Wu: Data curation. Chengyou Wang:
Data curation. Qiwei Wei: Conceptualization, Data curation, Funding
acquisition.
Declaration of competing interest
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influ-
ence the work reported in this paper.
Acknowledgments
This study was supported by the National Key Research & Develop-
ment Program of China (2018YFD0900801), the Project of Yangtze Fish-
eries Resources and Environment Investigation from the MARA, P. R.
China, and the China Three Gorges Corporation (No. 0799564). IJ was
supported by the J. E. Purkyně Fellowship of the Czech Academy of Sci-
ences, and the Alexander von Humboldt Foundation.
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