International Journal of Geosciences, 2013, 4, 1-7
http://dx.doi.org/10.4236/ijg.2013.45B001 Published Online September 2013 (http://www.scirp.org/journal/ijg)
Experimental Study of Invasion and Biofouling of
Freshwater Mussel Limnoperna fortunei
Mengzhen Xu1, Zhaoyin Wang1, Cheng Chieh Lin2, Baozhu Pan3, Na Zhao1
1State Key Laboratory of Hydroscience and Engineering, Tsinghua University, Beijing, China
2Program in Civil and Hydraulic Engineering, Feng Chia University, Taiwan, China
3Changjiang River Scientific Research Institute, Wuhan, China
Received May 2013
ABSTRACT
Golden mussel Limnoperna fortunei (Dunker 1857) is a filter-collector species of fresh water mussel originating from
southern China. In the water transfer tunnels from the East River to Shenzhen and Hong Kong, golden mussels attach to
the walls of pipelines and gates, causing serious biofouling, increased flow resistance, and resulted in corrosion of the
tunnel wall. Golden mussel has very high environmental adaptability and may colonize habitats with low dissolved
oxygen and a wide range of trophic levels. The colonization process of the species on solid surface was studied in the
Xizhijiang River, a tributary of the East River and the main water resource of Shenzh en from March 2010 to April 2011.
The results showed that the golden mussel completed three generations and reproduced six cohor ts per year in the tropic
zone. Water temperature was the controlling factor for the growth rate and maturity of each cohort. Based on the results,
an ecological method for controlling the invasion of golden mussels in water transfer tunnels was proposed.
Keywords: Bi ofouling of Water Transfer Tunnel s ; Golden Mus s e l (Limnoperna fortunei); Reproduction; Invasion
Control
1. Introduction
Inter-basin water transfer projects have been widely used
to ease uneven distribution and shortage of water re-
source in China. However, biological invasion and bio-
fouling caused by golden mussel (Limnoperna fortunei)
has become a challenge to these projects. The golden
mussel is a native freshwater species of southeast China.
It shares invasive biological and ecological features such
as size, growth speed, and colonization on hard substrata
by strong byssuses with the North American invasive
pest zebra mussel [1]. Recently, the golden mussel has
been observed in waters in Beijing [2 ]. Nevertheless, it
has also invaded in aquatic ecosystems and hydraulic
structures in South America and other Asian countries
([3,4]). Golden mussel invasion in a new habitat causes
ecological imbalance owing to changes in fishes’ feeding
habits and macroinvertebrate composition [5].
Golden mussel invasion and biofouling in water trans-
fer tunnels, pipelines, and pumps have caused damages in
water transfer projects. The biofouling density of golden
mussel reaches as high as 50,000 ind./m2. The thickness
of golden mussel clusters may exceed 10 cm on pipe walls,
valves, gates, and other structures, thus results in concrete
wall corrosion and high flow resistance [2]. Golden mus-
sels consume a significant amount of dissolved oxygen
due to their respiration and metabolism, causing water
quality degradation in water transfer tunnels [3]. Clog-
ging of pipes by high density of golden mussels caused
shutdown of a hydra ulic power plant in J a pa n [6]. Invas ion
and biofouling of golden mussels also damaged several
most i mpor ta nt nuc lea r po wer pla nts in So uth Amer ica [3].
Controlling of golden mussel biofouling has been stu-
died for decades. Various chemical and physical meas-
ures of getting rid of golden mussels such as coating pipe
walls, poisoning with pesticide, spraying with hot water,
trapping with filters, ultr aviolet irradiation, washing with
high velocity flow artificial removal with scrapers have
been employed [7]. However, such measures can them-
selves contribute to water pollution or high cost or dam-
age of structures for practical application. It is found that
prevention of golden mussel invasion is the only effec-
tive and environmental-friendly way to control the bio-
fouling [8]. Therefore, it is essential to study golden
mussels’ invasion pattern and intensive invasion season,
which are related to their reproduction characters for
controlling their invasion and biofouling in this paper.
2. Study Methods
2.1. Experimental Arrangement
The experiment was conducted at the Xizhijiang River, a

M. Z. XU ET AL.
2
secondary tributary of Pearl River in South China from
March 2010 to April 2011. The Xizhijiang River is a
water resource of the water transfer projects in Shenzhen
and Hongkong, China, which suffered serious golden
mussel invasion. The annual precipitation in the river
basin varies from 1,500 to 2,400 mm, and the air temper-
ature varies in 5˚C - 35˚C with an annual average of
21.5˚C.
Figure 1 shows the experimental materials. Three
cages were fixed on one bamboo, which is popular habi-
tat material for golden mussels [9] , and the bamboo was
planted into the river bed. The water depth of the three
cages was 1 m, 2.5 m, and 4 m, respectively. Three regu-
lar bricks (6 cm × 24 cm × 0.8 cm) were fixed in each
cage. The cages were used for protecting the colonized
golden mussels from being preyed by fish [10]. Th e cag-
es, the bamboo, and the bricks in the cages provided new
habitats for golden mussels to colonize.
In the experiment, a total of 42 cages on 14 bamboos
were fixed in the river water, of which 9 bamboos with
27 cages were planted into the river bed on March 13,
2010, and the other 5 bamboos with 15 cages were suc-
cessively planted into the river bed in July, August, Sep-
tember, October, and November in 2010. The new habi-
tats were colonized by golden mussels, as expected. One
month after the start of the experiment, the first bamboo
with cages was taken out and colonies of golden mussels
on the bricks were analyzed. From then on, the bamboos
with cages were taken out successively at one month
Figure 1. Experimental materials-bamboos with cages and
bricks were planted into the river bed.
interval over 9 months. Therefore, the results with colo-
nization time from 1 month to 9 months were obtained.
Additionally, data from the other 5 bamboos that were
given 5 months of colonization time but with different
start seasons were obtained for comparison. The density,
shell length, and distribution of golden mussels were
measured. During the experiment, density of golden
mussel larvae in the river water was measured once a
week. Water temperature (WT) and dissolved oxygen
(DO) levels were measured at the experiment site.
2.2. Data Analysis
For different colonization times, the colonized golden
mussels at the water depth of 2.5 m were used for cohort
decomposition using modal progression analysis in Fi-
SAT II (FAO-ICLARM Fish Stock Assessment Tools,
Version 1.2.2), a software program that decomposes the
length-frequency distributions into their unimodal com-
ponents by using Bhattacharya’s method [11]. The gol-
den mussels were grouped according to their shell length
into 0 - 1 mm, 1 - 2 mm, 2 - 3 mm, and so on up to 24 -
25 mm groups. The length-frequency data of all groups
were analyzed by means of modal progression analysis
(MPA) in FiSAT II, in which frequency is the number of
golden mussels found in each shell length group. Age
was assigned to each component according to the modal
progression analysis method [12], assuming that each
progression of the component represents a cohort. In this
manner, all of the cohorts were distinguished.
3. Results and Discussion
3.1. Colonization Density of Golden Mussels
Table 1 lists the experimental results. The maximum
shell length Lmax increased s lowly in the first two months,
and very fast in the third, fourth, and fifth months, and
slowly again from then on. The minimum shell length
Lmin was 0.28 mm, which was regarded as the smallest
mussel that can attach to solid surfaces with byssuses.
Figure 2 shows the densities of the golden mussel co-
lonized at water depths of 1 m, 2.5 m, and 4 m as a func-
tion of colonization time. The golden mussel density was
different for different water depths and colonization time.
The density was high at water depths of 1 m to 2.5 m,
and slightly lower at water depth of 4 m. The dashed
curve represents the average density of golden mussels at
different water depths. The density was nearly zero in the
first two months and increased sharply from the third
month, resulting in three peaks of density in May (third
month), July (fifth month), and October (eighth month).
In general, the golden mussel prefers a surface with
algae (Frances 2006). In our experiment, a layer of algae
was detected on the brick surfaces after two months.
150
bed surface
150
water surface
9010
50 100
80

M. Z. XU ET AL.
3
Table 1. Results of the new habitat colonization experiment.
No. Start time End time
△
t (month) Colonization on bricks at water depth of 2.5 m
D (ind./m2) Lmax (mm) Lmin (mm) Lave (mm)
1# 2010/3/13 2010/4/14 1 12 2.27 2.27 2.27
2# 2010/3/13 2010/5/14 2 12 3.23 3.23 3.23
3# 2010/3/13 2010/6/17 3 6620 11.46 0.28 3.46
4# 2010/3/13 2010/7/15 4 17593 13.25 0.78 5.22
5# 2010/3/13 2010/8/15 5 35370 16.32 1.29 7.38
6# 2010/3/13 2010/9/15 6 35069 17.74 1.79 8.49
7# 2010/3/13 2010/10/12 7 17824 20.77 1.11 9.97
8# 2010/3/13 2010/11/16 8 32593 21.96 0.99 9.37
9# 2010/3/13 2010/12/14 9 23264 22.22 0.76 10.90
10# 2010/8/15 2011/1/14 5 15677 18.1 0.75 4.93
11# 2010/9/15 2011/2/15 5 1736 13.42 0.76 4.05
12# 2010/10/12 2011/3/14 5 2523 14.12 1.04 4.53
13# 2010/11/16 2011/4/14 5 4444 14.56 1.17 4.15
Note: △t—colonization time ; D—the colonized density of golden mussels; Lmax, Lmin and Lave—the maxim um, minimum and average shel l lengths, respectively.
Figure 2. Density of golden mussels as a function of coloni-
zation time.
Therefore, very few golden mussels colonized the new
habitats in the first two months. Once golden mussels
colonized on the bricks, the colonization density varied
along the water depth due to variation in water tempera-
ture and dissolved oxygen concentration [13]. The gol-
den mussel larvae prefer water temperature in the range
of 16-28 resulting in high colonization density of golden
mussels in such temperatures; the minimum DO con cen-
tration for survival of golden mussels is 1.0 mg/L [14].
Figure 3 shows the representative DO concentration
and water temperature along water depth in different
seasons in the experiment. The DO concentration was
higher than the minimum threshold 1.0 mg/L at all sam-
pled water depths and seasons. Therefore, DO was not
the critical parameter that affected the vertical distribu-
tion of gold en mussels. As show in Figure 3(b), the most
suitable water temperature appeared at water depth of 1
m in the spring (May 22). Most golden mussel larvae
colonized to this water depth. Correspondingly, the colo-
nization density was the highest at the water depth of 1 m
in the spring. Afterwards, the water temperature in-
creased and even exceeded 28˚C at the water depth of 1
m from June to October, and the most suitable water
temperature occurred at the water depth of 2.5 m. Con-
sequently, the colonization density became the highest at
the water depth of 2.5 m from June to October. In Octo-
ber and December, water temperature at water depth of 1
m fell to the range of 16˚C - 28˚C, and the colonization
density became the highest at the wat e r depth again.
As shown in Figure 4, colonization density was also
found to be closely related to the orientation of the habi-
tat surface. The colonization density on the downward-
facing surfaces was 2 - 8 times higher than that on the
upward-facing surfaces owing to precipitation of silt and
clay on the latter. When we measured the amount of clay
and silt precipitates on the brick surfaces, we found that,
the amount on the upward-facing surfaces was six times
of that on the downward-facing surfaces. It is also re-
ported that the colonization densities of the golden mus-
sel on vertical surfaces and face-down surfaces were ob-
viously higher than that on face-up surfaces [13]. Clay
and silt precipitates interfere with filter -feeding, breath-
ing, and attachment of golden mussels.

M. Z. XU ET AL.
4
(a)
(b)
Figure 3. Dissolved oxygen and water temperature in dif-
ferent seasons. (a) Vertic al distribution of dissolved oxygen;
(b) Vertical distribution of water temperature.
Figure 4. Different colonization densities of golden mussels
on upward- and downward-facing surfaces.
3.2. Community Composition
Figure 5 shows an example of modal progression analy-
sis of the golden mussels colonized at the water depth of
2.5 m. The histogram is the frequency (the number of
golden mussels) as a function of shell length, and the
curves represent the decomposition of the cohorts. Three
cohorts (A, B, C) were decomposed.
As shown in Figure 5(a), for the colonization time
from March to July the average shell length and number
A
B
C
(a)
A
B
C
(b)
Figure 5. Cohort decomposition of golden mussels colonized
on new habitats. (a) March to July 2010; (b) March to Au-
gust 2010.
of golden mussels were 9.64 mm and 133 in cohort A. In
other words, cohort A colonized the habitat in the first
month and grew up to an average size of 9.64 mm in 4
months. The total number of the mussels was 133. For
the same colonization time, the average shell length and
number of mussels in cohorts B and C were 5.55 mm and
1137, and 1.87 mm and 460, respectively. For coloniza-
tion time from March to August (Figure 5(b)), the three
cohorts had developed further. The average shell length
and number of mussels in cohorts A, B, and C became
12.36 mm and 568, 9.83 mm and 550, and 4.77 mm and
1912, respectively. The larger numbers for cohort C were
due to more golden mussels recrui t e d in August .
Table 2 lists the results of all of the cohort decomposi-
tion for the 13 samples. Six cohorts—A, B, C, D, E, and
F were decomposed. Golden mussels maturate for re-
production when the shell length reaches 6 - 8 mm [9].
Therefore, the relations between the 6 cohorts can be
inferred according to their average shell lengths. Cohorts
A and B were the parental generation, and they colonized
the habitats in March-April and May-June, respectively.
The average shell length of cohort A was about 8 mm
longer than the average shell length of cohort C. The
average shell length of cohort B was about 10 mm longer
than the average shell length of cohort D. The length
differences between the generations were coincidentally
equal to the sexual maturity size of the species 6-8 mm.
Therefore, cohorts C and D were the offspring genera-
tions of A and B. Cohorts C and D colonized the habitats
in June to July and August to September, respectively.
For the same reason, cohorts E and F were the offspring
generations of C and D, respectively. From these results,
it is concluded that the species completed three genera-
tions in one year, and each generation experiences two
0
1
2
3
4
5
0.0 2.0 4.0 6.0 8.0
Water depth (m)
DO (mg/L)
22-May 10-Sep
12-Oct 18-Dec
0
1
2
3
4
5
15 18 21 24 27 30
Water depth (m)
Water tempreture (℃)
22-May 10-Sep
12-Oct 18-Dec
0
5000
10000
15000
20000
25000
30000
35000
40000
678955555
Density (ind/m
2
)
Colonization time (month)
Upwards
Downwards

M. Z. XU ET AL.
5
Table 2. Average shell length of cohorts.
No. Colonization time Average shell length of each cohort with standard deviation (mm)
A B C D E F
1# 2010.03-2010.04 2.27 ± 0.2
2# 2010.03-2010.05 3.32 ± 0.22
3# 2010.03-2010.06 6.83 ± 1.57 2.85 ± 1.47
4# 2010.03-2010.07 9.64 ± 1.7 5.55 ± 2.14 1.87 ± 0.63
5# 2010.03-2010.08 12.36 ± 0.91 8.93 ± 1.76 4.77 ± 1.99
6# 2010.03-2010.09 16.53 ± 0.45 11.88 ± 2.45 6.43 ± 0.77 1.86 ± 0.4
7# 2010.03-2010.10 18.2 ± 1.52 12.98 ± 0.88 8.3 ± 0.66 3.47 ± 0.77
8# 2010.03-2010.11 17.86 ± 0.8 14.46 ± 1.34 11.84 ± 0.8 7.02 ± 1.09 2.34 ± 0.9
9# 2010.03-2010.12 20.11 ± 1.38 16.56 ± 1.55 13.56 ± 2.5 8.35 ± 1.84 4.45 ± 1.4
10# 2010.08-2011.01 10.23 ± 2.52 5.75 ± 1.71 1.45 ± 0.89
11# 2010.09-2011.02 6.5 ± 2.14 1.57 ± 0.82
12# 2010.10-2011.03 7.66 ± 2.33 1.92 ± 1.46
13# 2010.11-2011.04 7.72 ± 1.27 3.29 ± 1.22
reproductive periods.
3.3. Larvae Density
Figure 6(a) shows the larvae density of the golden mus-
sel and water temperature from February to December
2010. Six peaks in larvae density, indicated as a, b, c, d, e,
and f, occurred in early March, middle April, late May,
late July, late August, and late September, respectively.
The larvae density was very low in February-March,
when the water temperature was below 20˚C. The six
peaks of the larvae density a-f supplied the larvae of the
cohorts A-F, respectively. For instance, the larvae densi-
ty peak “a” colonized the habitat in one month and de-
veloped into cohort A. Taking th e average shell length of
the cohorts in Table 2 into account, it is inferred that the
larvae density peak “a” gave rise to mussels of length of
2 mm in one month, 6 mm in three months, and 12 mm
in 5 months.
Figure 6(b) shows the larvae density of the golden
mussel and accumulated temperature from February to
December.
The accumulated temperature is given by:
1
(5 C)
n
i
i
TT
=
= −
∑∑

(1 )
where Ti is the daily average water temperature on the
ith-day, n is the number of days for sexual maturity of
the species. The accumulated water temperature was
1700˚Cday for cohort A (or a) to reproduce the cohort C
(or c) and for cohort C to reproduce cohort E (or e);
whereas the accumulated water temperature was 2100˚C
day for cohort B (or b) to reproduce the cohor t D (or d),
and for cohort D to rep roduce cohort F (or f).
3.4. Controlling Strategy
Invasion of the golden mussel into the water transfer
tunnels in South China occurred due to spreading of lar-
vae with water flow, which was similar to the golden
mussel invasion pattern in Sou th America [15], J apan [6],
and Korea [16]. Therefore, trapping the larvae during the
reproduction periods is one of the most efficient strate-
gies for controlling golden mussel invasion.
Previous investigation of larvae density in the whole
Xizhijiang River basin indicates that the density de-
creased to very low level at the downstream of a big re-
servoir, in which the flow velocity was near zero; and no
golden mussels survived at the sand bed of the reservoir
[9]. Therefore, a very effective approach for trapping
larvae can be the use of reservoirs with suitable size and
sand or silt bed at the upstream of water intake can effec-
tively cut off larvae supply. Our previous experiment
found that if the flow velocity in the reservoir is less than
0.01 m/s, the length of the reservoir is longer than 30 m,
and the bottom is covered with fine sediment almost all
larvae can be trapped and settled at the bed of the reser-
voir [9]. As stated in Section 3.1, clay and silt precipi-
tates in the reservoir interfere with filter -feeding, breath-
ing, and attachment of golden mussels settled at the re-
servoir bed. Carps, which are strong predators of golden
mussels [9], are recommended to be kept in the reservoir
to restrain golden mussel density.

M. Z. XU ET AL.
6
(a)
(b)
Figure 6. Larvae density of golden mussel and water temperature. (a) Larvae density and water temperature from February
to December; (b) Larvae density and accumulated temperature from February to December.
Intensive controlling measures should be performed
especially in the peak reproduction seasons of golden
mussel. In Hong Kong, which is close to our study area,
golden mussel has two reproduction seasons: one is from
June to July when water temperature is about 27˚C -
28˚C, and the other is from January to February when the
water temperature is about 16˚C - 17 ˚C [17]. According
to our experimental results, the golden mussel reproduces
frequently with six spawning peaks, resulting in six co-
horts, and completes three generations in each year. The
peak reproductio n seasons are very long , covering March,
May to June, July to September, and October to Novem-
ber. Therefore, controlling measures should be per-
formed continuously, at least cover the peak reproduction
period March to November.
Nevertheless, appropr iate depth of water intake from
the resource river can also reduce golden mussel invasion
in water transfer projects. As stated in section 3.1, the
vertical distribution of golden mussel density in river
water was affected by the distribution of water tempera-
ture along the water depth. Therefore, the depth of water
intake should be adjusted to avoid high golden mussel
density according to water temperature. For instance, it is
suggested to withdraw surface water (H = 1 m) in March
to August, because the golden mussel density was low at
this water depth; whereas it is suggested to withdraw
water from deeper layer (H = 4 m) in September to No-
vember because of lower golden mussel density at such
depth.
4. Conclusions
Golden mussels invade water transfer tunnels and lead to
bio-fouling, increased flow resistance, and tunnel wall
corrosion. Our findings ind icate that golden mussel colo-
nizes new habitats by spreading of larvae via flowing
water during reproduction seasons. The golden mussel
reproduces during March to November with six spawn-
ing peaks and results in six cohorts, and completes three
generations in each year. Its reproductive activity is in-
fluenced by water temperature and the completion of
generations is affected by the accumulated water temper-
ature above 5˚C.
Colonization of golden mussels often forms dense
clusters on the habitat surfaces. The colonization density
is affected by the orientation of the habitat surfaces ow-
ing to precipitation of silt and clay on the surfaces. Pre-
cipitation of silt and clay hampers filter-feeding, breath-
ing, and attachment to habitat and, therefore, greatly af-
fects the colonization density of golden mussel. Fur-
thermore, the vertical d istribution of colonization density
is mainly affected by the distribution of water tempera-
ture alon g the water depth.
Construction of reservoirs at the upstream of water in-
take of a water transfer project can effectively reduce
golden mussel larvae and is useful for controlling golden
15
17
19
21
23
25
27
29
31
33
1
10
100
1000
10000
1-Feb3-Mar2-Apr2-May1-Jun1-Jul31-Jul30-Aug 29-Sep29-Oct 28-Nov 28-Dec
Water tempreture(℃)
Larvae density（inds/m
3
)
2010
Larvae DensityTempreture
ab
cdef
0
500
1000
1500
2000
2500
1
10
100
1000
10000
1-Feb 3-Mar2-Apr 2-May 1-Jun1-Jul31-Jul 30-Aug 29-Sep29-Oct 28-Nov 28-Dec
Accumulated tempreture（℃*day）
Larvae density（inds/m3)
2010
Larvae Densitya-c b-d c-e d-f
ab
cdef

M. Z. XU ET AL.
7
mussel invasion in water transfer tunnels. Additionally,
appropriate depth of water intake from the resource river
can also reduce golden mussel invasion in water transfer
projects.
5. Acknowledgements
This study was supported by the Ministry of Water Re-
sources of China (200901078), Natural Science Founda-
tion of China (41071001), and Tsinghua University
(2009THZ0223 4) .
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