Variations of fish composition and diversity related to environmental variables in shallow lakes in the Yangtze River basin
Aquat. Living Resour.
Variations of fish composition and diversity related to environmental variables in shallow lakes in the Yangtze River basin
Lin Cheng 0 1 2
Sovan Lek 0
Géraldine Loot 0
Sithan Lek-Ang 0
Zhongjie Li 2
0 UMR 5174 EDB, CNRS-University Paul Sabatier , 118 route de Narbonne, 31062 Toulouse Cedex , France
1 Graduate School of the Chinese Academy of Sciences , 100039 Beijing , China
2 State Key Laboratory of Freshwater Ecology and Biotechnology, Institute of Hydrobiology, The Chinese Academy of Sciences , 430072 Wuhan , China
- Variations in fish communities of shallow lakes in the Yangtze basins were investigated from September 2007 to September 2009. Six lakes were chosen for comparative study of species composition and diversity in relation to environmental variations. Lake heterogeneity was described with environmental physico-chemical variables, using principal component analysis. Sixteen families, composed of 75 species of fish were found in the studied lakes, Cyprinidae being the dominant group. Fish species were divided by habitat preference and trophic guild: benthopelagic and herbivorous fish were the most common guilds in all lakes. Species diversity and richness were significantly higher in spring, while the evenness, expressed by equitability of Simpson's index, was not significantly different among seasons. Species richness and diversity were significantly higher in vegetated lakes (e.g. Liangzihu Lake) than in nonvegetated lakes (e.g. Biandantang Lake), with the largest area (Liangzihu Lake) harbouring the largest species richness and the greatest diversity. The relationship between environmental variables and fish assemblage were analysed using canonical correspondence analysis (CCA). The dominant gradients describing species composition and abundance among the sampling sites were: total phosphorus, total nitrogen, chlorophyll a, transparency and water depth. Our study led to the following conclusions: 1) the water quality was better - i.e. high transparency, low total phosphorus (TP) and total nitrogen (TN) and chlorophyll a- in vegetated lakes than in unvegetated lakes; 2) vegetated lakes had higher fish diversity than unvegetated lakes; 3) fish relative abundance (CPUE: number of fish per fishing pass) was significantly related to water chemical parameters. Consequently, the details of the findings are useful and relevant for developing suitable conservation strategies to sustain the integrity of fish communities in these lakes.
Freshwater fish / Environmental variables / Lake / Fish diversity / CCA / PCA / Yangtze River basin
As habitat degradation continues to accelerate on a global
scale, maintenance of species richness and biodiversity has
become a central issue of conservation biology
(Jones et al. 2004;
Lenihan and Peterson 1998; Rouget et al. 2003; Sinclair et al.
. Inland aquatic systems are crucial for the conservation
of local and global biodiversity (Moss 2000). In fact, there is
a great diversity in the form and function of these aquatic
systems, presenting a wide range of habitats
(Allan and Flecker
1993; Moss 2000; Williams et al. 2004)
. This is particularly
the case for the fish fauna of shallow waters
. Indeed, shallow lakes vary considerably in
species richness, supporting considerably more species,
including more unique and more scarce species than other type
of water body (i.e. rivers, streams and ditches) at a regional
(Williams et al. 2004)
; lakes, therefore, make the
greatest contribution to sustaining biodiversity. Unfortunately, most
shallow lakes occur in lowland areas, often with high human
population densities. As a consequence, their environmental
value is being dramatically affected, as demonstrated by
(De Meester and Declerck 2005; Xie and Chen
1999; Fang et al. 2008)
Conditions in the lakes seem to be a major factor
affecting diversity. It is widely accepted that environmental
variation plays an important role in the organization of lacustrine
(Jackson and Harvey 1989; Tejerina-Garro
et al. 1998; Amarasinghe and Welcomme 2002)
most important factors in determining species composition
differ between water bodies, ranging from physical habitat, like
lake morphology, to water chemistry
(Rahel 1984; Jackson
and Harvey 1989; Tejerina-Garro et al. 1998; Amarasinghe
and Welcomme 2002; Zhao et al. 2006; Petry et al. 2003;
Teixeira-de Mello et al. 2009)
. Comparisons of fish
community structure in lakes, conducted at the regional scale, have
highlighted predictable links between community structure
and environmental variations. The investigation of how
environmental factors (physico-chemical and biotic) determine the
structure of natural assemblages has benefited greatly from the
“natural experiments” of comparative studies
1978; Werner et al. 1978)
. This method can fairly quickly
generate and test hypotheses, assess mechanisms, and produce
acceptable explanations for community-level problems under a
wide variety of conditions
(Tonn and Magnuson 1982)
The Yangtze River, also called the Chang Jiang meaning
“long river”, flows for 6300 km from the Tibetan Mountains to
the East China Sea. Its catchment covers 1/5 of the land area of
China. The Yangtze River basin accounts for 40% of China’s
freshwater resources, more than 70% of fishery production,
and 40% of the China’s GDP
(Wong 2007; He et al. 2010)
In addition to its social and economic importance, the Yangtze
River basin is a centre of immense biological wealth. However,
human activities have profoundly degraded the ecosystem of
lakes in the Yangtze basin, with consequences such as water
quality degradation, threats to biodiversity and algal
(Xie and Chen 1999; Fang et al 2008)
. As lakes are seen
as highly productive water bodies, more effort is put into
increasing fishery production and less into the conservation of
fish biodiversity. Few studies of fish communities were found
on this area.
Xie et al. (2001)
compared small fish community
differences between zones with or without submersed
macrophytes and found fish communities in submersed macrophytes
zones to have significantly higher diversity, density and fish
Ye et al. (2007)
examined the spatial and seasonal
variations of the fish community relative to two key
environmental factors: the macrophyte complex and the water depth.
In fact, studies on species richness patterns in these lakes are
rare and have been largely ignored by monitoring and
Our aim in the present study was to analyse factors
explaining patterns of fish assemblages in different lakes within
the Yangtze River basin, based on environmental variations.
Two objectives were thus addressed: 1) to describe the fish
assemblages present in these lakes by examining the abundance
and occurrence of the fish species composition, the assemblage
patterns and the influence of environmental variables on the
assemblages; 2) to test hypotheses of possible links between
changes in the fish community and those in the environment.
Consequently, we hope the details of our findings will be
useful and relevant to developing suitable conservation strategies
to sustain the integrity of fish communities in the area.
2 Material and methods
2.1 Study area
The six lakes (Fig. 1) chosen for our study are located in
the central zone of the Yangtze River basin in Hubei, China.
This area is in the temperate zone and has a great number of
lakes (Hubei alone has over 1300 lakes), therefore providing
the necessary conditions to study the insular biogeography of
fish assemblages and well suited for comparative studies in
In selecting the lakes for this study, we picked those with
different macrophyte coverage, surface area and maximum
depth, but similar fishery activities. The six lakes chosen are
relatively close together and exposed to the same species pool,
but differ in surface area and macrophyte coverage. All of the
lakes are suburban lakes except Tangxunhu Lake. The lakes
present different macrophyte cover: Liangzihu Lake (LZH)
and Niushanhu Lake (NSH) have a high coverage (dominated
by Vallisneria natans (Lour.) Hara and Hydrilla verticillata)
throughout all the year; Luhu Lake (LUH) and Wuhu Lake
(WUH) were also covered by macrophytes, but these only
appeared in spring and summer. Potamogeton crispus Linn.
was the dominant macrophyte species in Luhu Lake, but this
species senesced and died after summer. The water depth in
Wuhu Lake increased sharply (by nearly one metre) in
summer, which led to macrophyte death. Biandantang Lake (BDT)
and Tangxunhu Lake (TXH) were devoid of macrophytes. The
details of lake characteristics and locations are summarized in
2.2 Data collection
Habitat descriptions and fish sampling were conducted
from September 2007 to September 2009. Within-lake habitat
measurements and fish samples were taken at four to twelve
locations in each lake, depending on the total surface of the lake
and the macrophyte coverage. Sampling started from a
random point and then proceeded at evenly distributed intervals
along the margins of the water body. We collected data
seasonally from each lake. Fish samplings were sometimes repeated
the following year because the sampling tools were often
destroyed by crabs (Eriocheir sinensis) aquaculture in the study
lakes. However, only the samplings from intact sampling tools
and one dataset for each season in each lake were selected,
meaning that about 100 datasets were used in this research.
Eleven environmental characteristics were considered in
this research Physical parameters included water body area
(AREA), water temperature (WT), water depth (WD),
transparency - measured by Secchi depth (TRA) -, and distance to
the bank (DIS). Chemical parameters included total
phosphorus (TP), total nitrogen (TN), chemical oxygen demand (COD)
and dissolved oxygen (DO). In addition, chlorophyll a (CHI)
and coverage by macrophytes (CRM) per square metre at each
sampling station were also taken into account in this study.
Fish sampling and measurements
To capture fish, one multimesh gill-net and one trap-net
were set together at each site. The multimesh gill-net method
followed that used by
. The total length of
each net was 20 m using mesh sizes between 5 and 55 mm
knot to knot. The mesh sizes followed a geometric series, with
a ratio of about 1.25 between mesh sizes, and were assembled
in the following order: 43, 19.5, 6.25, 10, 55, 8, 12.5, 24, 15.5,
5, 35 and 30 mm. Using randomly selected mesh sizes, the nets
were set in the water at 6:00∼7:00 p.m. and were hauled out
at 6 : 00 ∼ 7 : 00 a.m. the following day. Fish collected were
immediately identified to the species level, counted, weighed
to the nearest gram, measured (total length, to the nearest mm)
and then classified into habitat and trophic guild. The habitat
and trophic guild of each species was based on our previous
research and on FishBase data
(Froese and Pauly 2010)
relative abundance of each species at each sampling was
expressed in terms of catchper-unit effort (CPUE, mean number
of individuals per fishing pass). In order to record all the fish
species, for the analysis of fish species composition, we also
investigated the fish species from fishery catches.
Lake environmental variations were analysed using
principal component analysis (PCA) with the physico-chemical data.
Geometrically, PCA is a rigid rotation of the original data
matrix, and can be defined as a projection of samples onto a new
set of axes. The maximum variance is projected or “extracted”
along the first axis, the maximum variation uncorrelated with
axis-1 is projected on the second axis, the maximum
variation uncorrelated with the first and second axis is projected on
the third axis, and so on. PCA is now used routinely by
(Townsend et al. 1997; Grossman et al. 1998; Lamouroux
et al. 1999; Brosse et al. 1999, 2001)
to simplify large data sets
while reducing information loss, and to assess intercorrelation
among variables of interest
(Grossman et al. 1991)
Three diversity indices were chosen for diversity
comparisons among different lakes and different seasons, these were:
species richness, Simpson’s inverse index and equitability of
Simpson index. Simpson’s inverse index was calculated by
the formula: D = 1/Σ pi2, where pi is the proportional
abundance of species i. Equitability of Simpson inde was
calculated by dividing Simpson’s inverse index by S, where S is the
species richness. This index allows species richness to be
removed, in order to consider only the distribution of the
different species; it is, thus, used to express the evenness of a
community. The differences of diversity among the four seasons
and the six lakes were tested by Kruskal-Wallis and multiple
comparison tests. The Kruskal-Wallis test is a nonparametric
(distribution free) test used to compare three or more groups of
sample data and the multiple comparison tests are a group of
tests made following a one or two-factor ANOVA or a
The relationships of fish community structure with
environmental factors were analysed using canonical
correspondence analysis (CCA). CCA is a direct gradient analysis
method that concomitantly analyses the species and
environmental data and produces two types of site score. Weighted
average site scores were used in this study. For detail on the
merits and shortcomings of CCA in relation to other ordination
. The species
occurring in only a single lake were omitted from the
analysis. Before the analysis started, data on relative species
abundances were log-transformed (ln(x + 1)) to reduce the weight
of a few dominant species. The environmental variables were
log10(x + 1)-transformed to approximate normal distributions,
but normal distribution was not achieved for all of the
variables. A Monte Carlo permutation test (999 permutations) was
performed to test the significance of the relationships between
environmental variables and species composition among study
sites (i.e. the significance of the sum of all eigenvalues). The
Monte Carlo method is for testing whether a set of data is
consistent with a null hypothesis. It is appropriate for a
situation where the theoretical distribution of the test statistic is
unknown. The test procedure is to use Monte Carlo methods to
generate 999 further data sets of the same size as the true data,
under the conditions defined by the null hypothesis. The value
for the test statistic is calculated for each of these data sets
and the distribution of these values is examined. If the value
of the test statistic for the actual data is similar to the values
obtained from the artificial data sets, then the null hypothesis
is accepted, whereas if it is more extreme than the observed
value, the hypothesis is rejected.
PCA and CCA ordinations were carried out in the R
software using the “ade4” package
(Dray and Dufour 2007)
3.1 Environmental variations in the studied lakes
Physico-chemical parameters of the six studied lakes were
analysed by PCA (Fig. 2) to identify the differences /
similarities between the lakes. The first two principal components
described 42% and 17% of the total hydrological variation,
respectively. The six lakes were mostly differentiated by the
first PCA axis. Hydrological variables had high loadings on the
first component, such as: chlorophyll a (0.923), total
phosphorus (0.886), total nitrogen (0.886) and transparency (–0.806).
Thus, the first component represents a contrast between the
lakes presenting eutrophication and those with macrophyte
The comparison between the six lakes is shown (Fig. 2a).
Tangxunhu Lake (TXH) was very different from other lakes,
as this is an urban lake with high total phosphorus, total
nitrogen, chlorophyll a and chemical oxygen demand. The other
five lakes are suburban lakes. Among these, Liangzihu Lake
(LZH) and Niushanhu Lake (NSH) were found to be close
to each other and far from Tangxunhu Lake. The macrophyte
coverage of these two lakes was relative higher than for the
other lakes, and they had lower TP, TN, COD, chlorophyll a
and higher transparency (Table 1). The remaining three lakes
were located in the intermediate position between Tangxunhu
Lake and Liangzihu Lake + Niushanhu Lake (Fig. 1a). Their
similarities were the absence of macrophytes, in Biandantang
Lake (BDT), or temporary macrophyte coverage in spring and
summer (as at the end of summer, most of the macrophytes
had senesced and died), in Luhu Lake (LUH) and Wuhu Lake
3.2 Fish composition of shallow lakes in the Yangtze
All together, 75 species of fish were found across the six
lakes. A maximum number of 64 species was found in
Liangzihu Lake, while a minimum of only 37 species was found in
Biandantang Lake. The fish identified belonged to 16
families, of which Cyprinidae had the greatest species richness; 43
species from this family being found. The largest number of
species from this family was found in Liangzihu Lake. The
members of other families had relatively low species richness,
ranging from 1 to 5 species per family. In total, 17 791
individual fish were caught during the sampling period. Six species
represented less than 0.5% of the total catches. The six most
abundant species were Toxabramis swinhonis, Hemiculter
leucisculus, Rhodeus ocellatus, Pseudorasbora parva,
Rhinogobius giurinus and Squalidus nitens, which accounted for 74% of
fish collected. Eight species were found to have more than 25%
occurrence frequency: H. leucisculus, T. swinhonis, P. parva,
R. giurinus, S. nitens, Carassius carassius, Culterichthys
erythropterus, and R. ocellatus. H. leucisculus was the most
frequent of all fish species found (Table 2).
Divided by habitat, benthopelagic fish was the most
common type in all the lakes. It is worth mentioning that 31 species
of benthopelagic fish were found in Niushanhu Lake, which
represents 54% of the total number of species in this lake
(Table 3). Only five species of pelagic fish were found in
the lakes. The number of demersal fish species varied from
8 to 18 in the studied lakes. A larger number of demersal fish
species were found in macrophyte lakes, such as Liangzihu
Lake, Niushanhu Lake and Luhu Lake.
Based on their trophic level, we divided all captured fish
into three groups: carnivorous, herbivorous and benthivorous.
A total of 16 carnivorous fish species were identified, 15 of
which were found in Liangzihu Lake, the highest number of
BDT LUH LZH
carnivorous fish species among all the lakes (Table 3).
Pseudobagrus nitidus only appeared in Liangzihu Lake, while
Leiocassis longirostris, as an aquaculture species, was only found
in Niushanhu Lake. Herbivorous fish were the most frequent in
all the lakes. A total of 33 herbivorous fish species were
identified. Niushanhu Lake had the largest number of herbivorous
fish species - up to 28 species - while only 15 were found in
Wuhu Lake and Tangxunhu Lake. In total, 26 species of
benthivorous fish species were found, the largest number of which
were found in Liangzihu Lake (23 benthivorous species).
3.3 Spatial and temporal changes of fish
3.3.1 Seasonal variability
The seasonal changes in fish diversity are shown (Fig. 3).
Species richness (Fig. 3a) and diversity (inverse Simpson
index, Fig. 3b) were significantly different among the four
seasons, but the evenness (expressed by equitability of Simpson
index, Fig. 3c) showed no significant difference among
seasons (Kruskal-Wallis test). Subsequent multiple comparison
tests showed that species richness was significantly higher in
spring than in autumn, and that there were no significant
differences between other seasons (Fig. 3a). Species diversity was
also significantly higher in spring than in autumn and summer
3.3.2 Fish diversity comparisons among lakes
Fish species richness, diversity and evenness comparisons
were found significantly different among the six lakes (Fig. 4).
Biandantang Lake had the lowest value both in species
richness and diversity (Fig. 4a, 4b). When examined with
multiple comparison tests, species richness was found significantly
lower in Biandantang Lake than in Luhu Lake and
Liangzihu Lake; the difference of species richness was also
significant between Liangzihu Lake and Niushanhu Lake (Fig. 4a).
Fish diversity was higher in Liangzihu and Luhu lakes than in
the other lakes. Biandantang Lake and Tangxunhu Lake had
the lowest value of diversity, which was significantly
different from Luhu Lake and Liangzihu Lake (Fig. 4b). Niushanhu
Lake had the highest value of evenness, which was
significantly higher than in Tangxunhu Lake (Fig. 4c).
3.3.3 Environment and fish community
The most important variables (Table 4, Fig. 5) describing
the species composition among the study sites were
chlorophyll a TP, and TN for axis 1 (X-axis); and transparency,
water depth and area for axis 2 (Y-axis). In ecological terms,
axis 1 showed gradients in species composition related to
water chemistry variables Species composition varied along a
gradient from small, eutrophic lakes (high TN and TP) to
larger lakes with low TN, TP and chlorophyll a; axis 1
described mainly the relationships of spatial and temporal
variation with the assemblage composition. The first two CCA axes
Fig. 3. Seasonal changes in the fish community. (A) Species richness
comparisons among the four seasons; (B) Simpson’s Inverse Index
comparisons among the four seasons; (C) Equitability of Simpson
index comparisons among the four seasons. SPR: spring; SUM:
summer; AUM: autumn; WIN: winter.
had eigenvalues of 0.19 and 0.13, explaining 28% and 19% of
variation in the relationship between species composition and
environmental factors, respectively. The overall Monte Carlo
randomization test showed a significant result for the sum of
all eigenvalues (999 permutations, p < 0.01).
Based on the CCA scores, three clusters of fish were
identified (Fig. 5). Cluster I (upper part of Fig. 5) was positively
related to TN, TP, chlorophyll a and COD, dominated by
Neosalanx taihuensis, Hemibarbus maculatus, Cyprinus
carpio, Culter dabry and Xenocypris davidi; Cluster II (left part
of Fig. 5) was mostly related to limnological characteristics
parameters (e.g. lake surface area and water depth) dominated
by Squalidus nitens, Squalidus argentatus, Abbottina rivularis
and Acheilognathus chankaensis; Cluster III (low-right part of
Fig. 5) was positively related to transparency, macrophyte
coverage, negatively related to TN, TP, chlorophyll a and COD,
and dominated by Rhodeus fangi, Hypseleotris swinhonis and
Rhinogobius giurinus. Cluster III can be subdivided into two
groups (groups a and b). Cluster III was made up of the most
frequent species in the lakes, and concentrated in the
middleleft part of Fig. 5; Cluster III was positively related to
macrophyte coverage and transparency.
4 Discussion and conclusion
As environments are modified by anthropogenic pressures,
the species composition of fish communities has been found
, i.e. decreasing biodiversity in lakes
(Xie and Chen 1999; Fang et al. 2008)
, and increases in
omnivorous fish as river system habitats deteriorate
Roth et al. 1996; Schleiger 2000)
. Studies of fish communities
that focus on the relationships between fish and their habitats
are of particular importance because of their value in
quantifying the effects of habitat supply limits, which by some means
control the size and dynamics of fish communities
and Brown 1974; Amarasinghe and Welcomme 2001; Zhao
et al. 2006)
. Among ecosystems that support a higher
biodiversity, wetlands occupy only about 1% of the earth’s surface, but
4.1 Fish community structure and patterns in shallow
lakes of the Yangtze basin
Fish species richness was relatively high in this area
(varying from 40 to 64 species in our study lakes) compared with
lakes in other regions (average of 35 species) in China
et al. 2006)
and lakes in other temperate regions, i.e. 25 species
in the temperate regions of Europe and Asia; 67 species in the
temperate region of South America
(Teixeira-de Mello et al.
2009; Tonn and Magnuson 1982; Persson et al. 1992; Fischer
and Eckmann 1997)
, and the dominant fish in our studied lakes
The dominance of herbivorous fish in the study lakes can
be explained by equilibrium-based mechanisms. It is well
known that equilibrium-based mechanisms are inseparably
linked to niche structure in communities
(Tonn and Magnuson
. In saturated communities, species richness is proposed
to be a function of the resource availability, tolerable niche
overlap and minimum niche size along a resource gradient
(Menge and Sutherland 1976; Connell 1978)
. If the
tolerable niche overlap and minimum niche size are relatively
(Roughgarden 1974; Werner 1977)
, species richness
should depend mainly on habitat complexity. Similarly, more
productive habitats allow greater dietary specialization and
should support more species
fish species such as Parabramis pekinensis, Xenocypris
argentea and Ctenopharyngodon idellus feed mainly on plants or
plant debris, while other herbivorous fish such as Aristichthys
nobilis, Hemibarbus maculatus and Hemibarbus labeo feed
mainly on plankton. As a result, it does not matter whether
macrophytes or phytoplankton are the main primary producer
in these lakes, in both cases there is a rich food source for
herbivorous fish and they could thus become the dominant fish
species in all the investigated lakes.
In this research, we found fish community structures and
diversity were co-affected by limnological characteristics (lake
area, water depth), macrophyte coverage and some
physicochemical parameters. Different fish species were related to
different environmental parameters.
Grouped by CCA scores, three clusters of fish were
identified as being highly related to different environmental
variables. The first of these clusters was dominated by Hemibarbus
maculatus, Carassius auratus and Culter dabry. This group of
fish was positively related to TN, TP and chlorophyll a. Most
of these species are tolerant of environmental changes,
presenting high growth rates in the presence of high levels of
algae or in eutrophic lakes. As these tolerant species have a high
resistance to environmental stress, it is assumed that the
raising of their population density could be used as an indicator to
reflect any worsening of environmental conditions
Fausch et al. 1984; Roth et al. 1996; Schleiger 2000)
The second group of fish was dominated by Squalidus
nitens, Squalidus argentatus, Abbottina rivularis and
Acheilognathus chankaensis, and was related to limnological
characteristics parameters (i.e. lake area and water depth)
High density (CPUE) of these fish would be found in shallow
habitats and large lakes. Furthermore, most of the species in
this cluster were common and widespread in the studied lakes.
The third group of fish species seems to be sensitive to
environmental changes (e.g. Rhodeus fangi). They were
negatively related to TN, TP and chlorophyll a, and positively
related to transparency and macrophyte coverage. The cluster
can be subdivided into two groups (Fig. 5). One group is made
up of the most frequent species in the lakes, concentrated in
the central part of Figure 5, such as Toxabramis swinhonis and
Hemiculter leucisculus. This group is less sensitive to the
environment compared with the other group of fish in this
cluster. The second group brings together sensitive species from
the lakes. Most of these species are only found in clear waters
with high macrophyte coverage. This group of fish, dominated
by Rhodeus fangi, Hypseleotris and Odontobutis obscurus, can
probably be used as indicators in these lakes.
All the lakes are exposed to the same species pool and
are relatively close to each other (Fig. 1), but the patterns
of fish community were found to be different among them.
Fish species richness, diversity and evenness were relatively
higher in vegetated lakes (i.e. Liangzihu Lake, Luhu Lake
and Niushanhu Lake) than in unvegetated lakes (i.e.
Biandantang Lake and Tangxunhu Lake). Based on our
investigations, the environments of the lakes were shown to be
significantly different. The relationships between the environment
and fish communities are valuable for lake management.
Several decades ago, all of the shallow lakes in the Yangtze basins
were highly covered by macrophytes
(Xie and Chen 1999;
Fang et al. 2008)
. However, due to water pollution and
intensive fishery activities, some lakes are now found to be
completely devoid of macrophytes
(Xie and Chen 1999; Fang et al.
We have shown that as the coverage of macrophytes
decreases, the water conditions deteriorate (i.e. increasing TN,
TP and COD, decreasing transparency).
Some previous studies have shown a significant effect of
macrophytes on fish communities in Yangtze basin shallow
lakes. They found that this type of habitat along with
structural complexity of macrophytes harboured high fish diversity
(Xie et al. 2002; Ye et al. 2006)
. In this study, the fish
community was also linked to macrophytes. Our study confirmed
the following conclusions: 1) the water quality was better (i.e.
high transparency, low TN, TP and chlorophyll a) in vegetated
lakes than in unvegetated lakes; 2) vegetated lakes had higher
fish diversity than unvegetated lakes; 3) relative fish abundance
(CPUE) was significantly related to the macrophyte coverage.
We suggest that a good method for restoring water quality
would be to replant macrophytes in these lakes. The choice
of macrophyte species used for restoration should be based on
their biological characteristics. Species with a long life-span
(e.g. Vallisneria natans (Lour.) Hara and Hydrilla verticillata)
would be most suitable, while species that senesce and die at
the end of the summer, like Potamogeton crispus Linn. (which
widely distributed in Luhu Lake) should be introduced into
these lakes with caution.
It is worth mentioning that two non-native species
(Cirrhinus molitorella and Leiocassis longirostris) were found
during our samplings. Cirrhinus molitorella (only found in Luhu
Lake) had probably escaped from nearby fish ponds and does
not pose a serious problem as an invasive species at present
because it is a sub-tropical species that requires high water
temperatures to survive in winter. However, the ecological risk
of Leiocassis longirostris (escaped from cages in Niushanhu
Lake) is under estimated. We strongly suggest that more
caution should be taken with the ecological security when
introducing an alien species to lakes and nearby water bodies.
Acknowledgements. We express our thanks to Prof Xie Songguang,
Liu Jiashou, Brian R. Murphy, and Dr. Zhang Tanglin, Ye Shaowen,
Cândida Shinn, Clément Tisseuil for their kind help and suggestions
in research design and manuscript writing. Mr. Li Wei, Lin Mingli,
Zhang Chaowen, Guo Chuanbo, Zhu Fengyue, Tang Jianfeng and
Miss. Liu Xin, Zhang Lihong and Du Xue for their assistance in
the field sampling and sample measurements. Special thanks go to
Loïc Tudesque for helping to draw the map. We appreciate the
helpful comments of the two anonymous referees and the editor on the
early version of this paper. This research was financially supported
by grant No. 30830025 and 30900182 from the National Natural
Science Foundation of China, and No. 200903048-04 from the R & D
project of Ministry of Agriculture of China.
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