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多棘海盘车对魁蚶摄食量、选择性及昼夜摄食差异的初步研究(英文)



全 文 :Vol. 18 No. 1 Marine Science Bulletin May 2016
Received on Apr. 18, 2016
*Corresponding author: zhaoxingqiu@163.com

A preliminary study on food consumption, preference
and day-night predatory difference of Asterias
amurensis on Scapharca broughtonii

ZHANG Tian-wen, LIU Guang-bin, LIU En-fu, ZHENG Yong-yun, QIU Zhao-xing*
Marine Biology Institute of Shandong Province, Qingdao 266002, China

Abstract: In order to study the food consumption, preference and day-night predatory
difference of Asterias amurensis on Scapharca broughtonii, the experiments were
carried out at the temperatures ranging from 16 ℃ to 20 , salinity of 28.2℃ ± 2.1,
dissolved oxygen of 7.5±0.7 mg/L and pH of 8.0 ± 0.3.The results provided basic data
for promoting steady and healthy development of the multiplication and cultivation of S.
broughtonii. The results of food consumption suggested that the food intake of starfish
increased with the density of S. broughtonii at all conditions. The food consumption of
starfish in different sizes on S. broughtonii in the same size had no significant difference
(p>0.05), but it was significantly different among S. broughtonii in different sizes. The
starfish had the highest feeding rate on the smallest S. broughtonii. Our findings also
verified the positive relation between the weight of food intake of starfish in different
sizes on the density and size of S. broughtonii. The feeding rhythm of starfish on
S. broughtonii in different sizes showed a certain day-night difference, higher at night
than in the day. The maximum food consumption occurred when the size of S.
broughtonii became larger. The results showed that 30 ind/m2 was the suitable sea
bottom sowing density for S. broughtonii. Our data indicated that the ability to avoid
predators and rivals increased as the shell length rose. Based on prey selectivity,
bivalves that have a lower commercial value may be used to clear away A. amurensis.

Keywords: Asterias amurensis; Scapharca broughtonii; food consumption; food
preference; day-night difference
Commonly known as ark shell and baby clam, Scapharca broughtonii, belonging to
arcidae of lamellibranchia, is a large-scale bivalve mollusk living by burial and dwelling.
With delicious taste, rich nutrition, and high economic value, it is always the object caught
by fishermen in Yellow Sea and Bohai Sea. In recent decade, the natural resource of S.
broughtonii shrinks extremely for long-term excessive catch [1]. At present, the natural
fishing and catching yield cannot meet the demand in domestic and foreign market, so it is
70 Marine Science Bulletin Vol. 18
imperative to develop culture and proliferation and repair the resource [2]. Since 2010, the
culture and proliferation of S. broughtonii becomes a hotspot in the development and
utilization of shallow-sea resource after holothurian culture [3]. There are various factors
affecting the effects of S. broughtonii culture and proliferation, but the major influencing
factor after the bottom sowing is the invasion of harmful organism, starfish [4].
Starfish is one of major predators in the biological system of intertidal zone and
subtidal zone [5, 6]. Its ingestion can directly or indirectly affect the distribution and diversity
of benthos [7-10]. The starfish distributing in Shandong coastal area is mainly Amurensis
asterias, belonging to asterias of asteroidean. A.amuresis mainly ingests shellfish,
crustacea, polychaetes, and small fish [12], especially bivalve. Moreover, it competes with
benthos for baits [13, 14]. In the natural environment, A.amuresis has extensive feeding habit,
but it also has selectivity in food organism. Such selective feeding not only brings more
living opportunities, but also plays an important role in adjusting population quantity of food
organism [15]. In shellfish culture area, A.amuresis can accurately position burial-dwelling
shellfish and excavate ingestion according to the smell of shellfish [16, 17]. In 1954, the direct
loss in shellfish culture caused by ingestion of A.amuresis was up to 4 billion Yen [18]. In
2007, A.amuresis bursting out in Qinghai coastal area caused severe loss of
bottom-sowing shellfish in Red Island [19]. Studying the ingestion selectivity of A.amuresis
in different sizes and densities by A.amuresis in different sizes and ingestion difference
between day and night, this works provided basic data to know more about the ecology of
A.amuresis to ingest S. broughtonii. It is expected to effectively avoid the influence on
major harmful organisms, improve the survival rate of S. broughtonii, and to better develop
the proliferation of S. broughtonii by the bottom sowing.

1 Materials and methods

1.1 Experiment polder and management
In Rizhao Institute of Oceanic and Fishery, a pond of sediment substrate was
selected for experiment. The ecological method of polder experiment was used to study
the capacity of A.amuresis to ingest juvenile mollusk of S. broughtonii and ingestion
difference between day and night. The experiment was developed in 2 m × 2 m polder, and
30-mesh bolting-silk was used as enclosure ecosystem, the lower part of which was
embedded into bottom mud for 0.6 m. The whole enclosure ecosystem was set in the pond
with timber pile and bamboo as support. 2 days before the experiment, water was flooded
for 96h experiment, during which the water depth in pond was 1 ± 0.1 m. After the
experiment, the statistics was conducted in the empty pond. In the experiment, water
No. 1 ZHANG Tian-wen et al.: A preliminary study on food consumption, preference and ⋯ 71
temperature was 16 ˚C - 20 ˚C, salinity 28.2 ± 2.0, dissolved oxygen (7.5 ± 0.7) mg/L, and
pH was 8.0 ± 0.3.
1.2 Experimental materials
The juvenile mollusks of S.broughtonii for experiment were offspring seeds by
overwintering culture in 2012 from estuary area between 2 cities in Shanhaitian Tourist
Resort of Rizhao, Shandong. After being transported to experiment field, the juvenile
mollusks were reared for 14 days temporarily with circulating water in the storage pond.
During temporary rearing, mangroves and diatom were fed at 06:00, 12:00, 18:00, and
24:00 every day.
The A.amuresis for experiment was from the sea area in Rizhao, Shandong. After
being transported to experiment field, the A.amuresis was reared for 3 days temporarily
with circulating water in the storage pond. During the temporary rearing, no bait was fed.
1.3 Experiment design
1.3.1 Food consumption of juvenile mollusks of S. broughtonii in different densities and
categories by A.amuresis
Juvenile mollusks of S. broughtonii in four categories with the shell length of 1 ± 0.09 cm
and 1.5 ± 0.13 cm were selected and divided into 1 cm group and 1.5 cm group. With
10 ind/m2, 30 ind/m2 and 50 ind/m2 as the experiment density, the weight of soft part was
measured for S. broughtonii in 4 sizes. A.amuresis with peduncle length of 8 cm - 10 cm
and density of 1 ind/m2 was taken for 96h experiment to analyze its capability to ingest S.
broughtonii in different densities and categories. Four parallels and four blank control
groups without A.amuresis were set for every group of experiment.
1.3.2 Food consumption of juvenile mollusks of S. broughtonii in different categories by
A.amuresis in different categories
A.amuresis with wrist lengths of 5 cm - 7 cm, 8 cm - 10 cm, and 11 cm - 13 cm was
selected as small-scale, middle-scale, and large-scale A.amuresis. The density of S. broughtonii
was 30 ind/m2, and that of S. broughtonii was 1 ind/m2. During 96h experiment, the
difference in capabilities among A.amuresis in 3 sizes to ingest S. broughtonii in 4 sizes
was analyzed. 4 parallels and 4 blank control groups without A.amuresis were set in every
experiment.


72 Marine Science Bulletin Vol. 18

1.3.3 Difference of A.amuresis to ingest juvenile mollusks of S. broughtonii in different
sizes between day and night
In the experiment to analyze the difference of A.amuresis to ingest S. broughtonii in 4
sizes between day and night, the density of S. broughtonii was 30 ind/m2, and peduncle
length and density of A.amuresis were 8 cm - 10cm and 1 ind/m2, respectively. The
experiment started at 8:00am and was observed at 1h, 2h, 8h, 12h, 24h, 30h, 46h, 54h,
60h, 72h, and 96h after the experiment. 4 parallels and blank control groups without
A.amuresis were set for every group of experiment.
1.4 Measurement of experiment indicators
The empty shells of S. broughtonii should be counted and natural fatality of seeds
was recorded in every enclosure after the experiment about A.amuresis in different sizes
ingesting S. broughtonii in different sizes and densities.
Number of empty shells of S. broughtonii in enclosures and that in blank groups were
recorded according selected observation time in the experiment about A.amuresis in
different sizes ingesting S. broughtonii in different sizes and densities.
1.5 Data processing and analysis
The food consumption of S. broughtonii in different sizes by A.amuresis is presented
by the number of empty shell of S. broughtonii to be ingested.
K=E-D
where K refers to the number of empty shells of S. broughtonii to be ingested, E the
number of empty shells of S. broughtonii in experiment enclosure, and D the number of
empty shells of S. broughtonii in blank group without A.amuresis.
The difference in food consumption of S. broughtonii in different sizes by A.amuresis
is represented by the percentage of total empty shells accounting for total empty shells of S.
broughtonii.
Ht=(Et-Dt)/(E-D)
where t refers to observation time, Ht the percentage of S. broughtonii to be ingested
accounting for all S. broughtonii to be ingested in experiment at the observation time, Et
empty shells of S. broughtonii in experiment enclosure (ind) at the observation time, and Dt
the empty shells of S. broughtonii in blank control group without A.amuresis at that time.
No. 1 ZHANG Tian-wen et al.: A preliminary study on food consumption, preference and ⋯ 73
Variance analysis was conducted for experiment data with statistical software
SPSS17.0, and multiple comparison conducted by Duncan method. P>0.05 means that
difference is insignificant, while P<0.05 means that difference is significant. Results were
represented by lowercases. Data with same letters means that the mutual difference is
insignificant (P>0.05). During the experiment, the shell length of S. broughtonii was
denoted by L and the meat weight by W.

2 Experiment results

2.1 Analysis on food consumption of S. broughtonii in different sizes and densities
by A.amuresis
Fig. 1 shows that the food consumption of S. broughtonii in different sizes by
A.amuresis rose with the increase in the density of S. broughtonii. When the density of
S. broughtonii was 30 ind/m2, the food consumption of S. broughtonii by A.amuresis
increased without significant difference from that when the density of S. broughtonii was
10 ind/m2(P>0.05). When the density of S. broughtonii was 50 ind/m2, the food
consumption of S. broughtonii in 4 sizes by A.amuresis increased significantly. In the same
density, the food consumption of S. broughtonii by A.amuresis decreased with the increase
of shell length of S. broughtonii. Food consumption in each density of 2.0 cm group was
the smallest. The food consumption of S. broughtonii in different sizes by A.amuresis was
significantly negative correlated to the size of S. Broughtonii(r=-0.984 5, P<0.01). The
fitting formula between both is K= 35.543e-0.457L(R2=0.979 3).


Fig. 1 The amount of food intake of A.amurensis on different kinds and densities
of S. Broughtonii
(Note: Data with the same superscript are not significantly different(P>0.05))

74 Marine Science Bulletin Vol. 18

Fig. 2 The weight of food intake of A.amurensis on different kinds and density
of S. Broughtonii
(Note: Data with the same superscript are not significantly different(P>0.05))
Fig. 2 shows that the food consumption of S. broughtonii by A.amuresis with soft part
in different densities increased with density increase. The weight of soft part to be ingested
was the maximum when the density of S. broughtonii in different sizes was 50 ind/m2. The
difference in weight of soft part to be ingested was insignificant when the density was
10 ind/m2 and 30 ind/m2. The soft part of S. broughtonii in different sizes ingested by
A.amuresis also tended to increase with the enlargement of S. broughtonii. In 3 densities,
the food consumption of soft part in 1 cm experiment group was the minimum, while that in
2.5 cm experiment group was the maximum. However, there was no significant difference
between 2 cm experiment group and 2.5cm experiment group. The food consumption of
soft part of S. broughtonii by A.amuresis was significantly positive correlated to the size of
S. broughtonii(r=0.987 3 p<0.01). The fitting formula between both is W=1.486 2
L(R2=0.972 8).
2.2 Analysis on food consumption S. broughtonii in different sizes by A.amuresis
in different sizes
Fig. 3 shows that there was no significant difference in food consumption of S. broughtonii
in the same size by A.amuresis in different sizes (P>0.05). However, there was difference
in the food consumption in different sizes of S. broughtonii by A.amuresis in the same size
to some extent. The difference in food consumption between 1cm group and 1.5 cm group
was insignificant. With the enlargement of S. broughtonii in experiment group, the food
consumption gradually decreased. The food consumption by A.amuresis in 3 sizes was the
minimum (5 ind at average). In 4 experiment groups, the food consumption of S.
broughtonii by A.amuresis with peduncle length of 5 cm - 7cm was slightly higher than that
in groups with peduncle length of 8~10cm and 10 cm - 12 cm without significant difference
(P>0.05).

No. 1 ZHANG Tian-wen et al.: A preliminary study on food consumption, preference and ⋯ 75
 
Fig. 3 Feeding intensity of different kinds A.amurensis on different kinds S.broughtonii
(Note: Data with the same superscript are not significantly different (P>0.05))
2.3 Difference in food consumption of S. broughtonii in different sizes by
A.amuresis between day and night
There was difference in food consumption of S. broughtonii in 4 sizes by A.amuresis
between day and night to some extent because the ingestion time was mainly at night
(20:00 - 8:00). The experiment started at 8:00, but S. broughtonii in 4 sizes were not
ingested by A.amuresis in 12 hours (at 20:00 on the first day). When observing at 24h (at
8:00 on the second day), except S. broughtonii in 2.5 cm group, those in 3 other sizes were
all ingested by A.amuresis, and the ingestion ratio in 2 cm group was the largest that is,
14.29%. When observing after 30 hours (at 14:00 on the second day), no S. broughtonii
was ingested by A.amuresis in 3 experiment groups, except 1cm group with a few S. broughtonii
ingested. When observing after 48 hours (at 8:00 on the third day), S. broughtonii in 4 sizes
were ingested by A.amuresis in large amount with the highest ratio among all observation
times. When observing after 54 hours (at 14:00 on the third day), only a few S. broughtonii
in 1 cm group was ingested, but no new empty shell was found in other 3 experiment
groups. When observing after 60 hours (at 20:00 on the third day), except S. broughtonii in
2.5 cm group, a few in other 3 groups were ingested. When observing after 72 hours (at
8:00 on the fourth day), S. broughtonii in 4 experiment groups were all ingested, but the
ingestion ratio was significantly lower than that in 48th hour. When observing after 96 hours
(at 8:00 on fifth day), S. broughtonii in 4 experiment groups were ingested with the lowest
ratio.

3 Discussion

3.1 Analysis on the ingestion of S. broughtonii in different densities and categories
by A.amuresis in different sizes
With wide feeding scope and selectivity, A.amuresis is in the top of biogenesis food

76 Marine Science Bulletin Vol. 18
chain [20] with ingestion objects, including available food organisms in environment [21]. The
selectivity of A.amuresis is dependent on the favor to the food organism, as well as the
number and availability of such food organism in environment. Studies on the predatory
relation and ingestion selectivity of starfish started early in foreign countries, including
relevant studies on the selectivity and functional response of A.amuresis and Asterias
forbesi in the Pacific to the size of scallop and mussel [22-25]. However, there is still no study
on the selectivity of A.amuresis to ingest S. broughtonii.
According to the most suitable recipe model, when food number is up to a certain
density, the predator will refuse to ingest the prey with smaller advantage [26]. This
experiment found that the soft part in S. broughtonii ingested by A.amuresis increased with
the enlargement of S. broughtonii and its density. When the density of bottom sowing was
50ind/m2, the soft part of S. broughtonii in 4 sizes ingested by A.amuresis was the largest.
When the densities of bottom sowing were 10 ind/m2 and 30 ind/m2, there was no
significant difference in food consumption of S. broughtonii in 4 sizes by A.amuresis
(P>0.05).
In the production of S. broughtonii by bottom sowing, the yield and economic benefit
will be bad if the density of bottom sowing is as low as 10 ind/m2 even though there are little
harmful organism. If the density is 50 ind/m2, there will be a large amount of S. broughtonii
ingested by harmful organism, causing unnecessary economic loss. Therefore, the density
of bottom sowing is suggested to be about 30 ind/m2, which can ensure the number of
S. broughtonii per unit area, and avoid too much loss for the ingestion of S. broughtonii by
A.amuresis to obtain better effect of culture and proliferation. This suggestive value is close
to 15 ind/m2 - 30 ind/m2 of the general density for bottom sowing of S. broughtonii.
The food selectivity of ingested in marine ecosystem is dependent on the size of food
particle [27]. Therefore, the ingestion of shellfish by A.amuresis has significant relation with
the size of shellfish. According to Kim Y (1968), et al. [18], A.amuresis had capability to
ingest shellfish in various shell lengths, and the rate to ingest shellfish in appropriate shell
length was the largest. In this experiment, there was no significant difference in the food
consumption of same S. broughtonii by A.amuresis in 3 sizes, but there was difference in
food consumption of S. broughtonii in different sizes. Especially, the food consumption of
S. broughtonii in 1 cm group and 1.5 cm group was the largest, while that in 2.5 cm group
was the smallest. This result is the same as that by Liu Jia (2012) [28] that the food
consumption by A.amuresis in the group with the smallest shell length was the largest.
According to the results in this work, S. broughtonii with shell length of 2.0 cm - 2.5 cm is
the most suitable as the standard for bottom sowing, which is close to the conclusion by
Tang Qisheng et al. (1994) [9]. When developing proliferation by bottom sowing, the larger
No. 1 ZHANG Tian-wen et al.: A preliminary study on food consumption, preference and ⋯ 77
the size of S. broughtonii is, the more effectively it can defend enemy. Therefore, the size
of S. broughtonii released in Japan is 2 cm - 3cm, and even 5 cm - 6cm [29]. However, the
large size will certainly extend the culture time and increase the cost. Moreover, juvenile
mollusks of S. broughtonii may be dead from the anoxia in spring, high temperature in
summer, and other uncontrollable factors in a large amount [9]. Therefore, the size of
S. broughtonii for bottom sowing should be determined with the consideration of burrowing,
predator, cost, and other factors.
3.2 Difference in ingestion of S. broughtonii by A.amuresis between day and night
There is difference in the food consumption of S. broughtonii in different sizes by
A.amuresis between day and night. Related to illumination, this ingestion difference
between day and night widely exists among aquatic animals [30]. Belonging to low-grade
Echinodermata, sterias amurensis is underdeveloped in visual and other sense organs, but
many rhombic neurosensory cells with antennal organs and chemical sensors are spread
in epithelium. A large quantity of sucking discs in sucker of A.amuresis play a role of smell [27].
The visual and smell organ systems of A.amuresis are underdeveloped, but ingestion
difference between day and night forms for the gradual adaptation to biological factors,
including illumination and bait in the long-term evolution [19]. In this work, the food
consumption of S. broughtonii in 4 sizes by A.amuresis was low in the first 24 hours of
experiment mainly because A.amuresis requires a period to adapt the surrounding
environment as other aquatic organisms. From 24th hour to 30th hour (daytime on the
second day) and 48th hour to 60th hour (daytime on the third day), only a few S. broughtonii
in 1 cm was ingested by A.amuresis, but S. broughtonii in other seizes showed no obvious
ingestion. From 30th hour to 48th hour (nighttime on the second day), a large amount of
S. broughtonii in 1 cm and 1.5 cm group was ingested with the percentage of empty shell
increasing from 29.39% and 7.69% to 68.18% and 53.85%, respectively. Moreover,
S. broughtonii in 2 cm and 2.5 cm group were greatly ingested from the 60th hour to the 72th
hour (nighttime on the third day) with percentage increasing from 47.62% and 50% to
85.71% and 100%, respectively. According to results in this work, the food consumption of
S. broughtonii ingested by A.amuresis was large at night on the second day after the
bottom sowing if the seed was in the small size (≤1.5 cm), while that was large at night on
the third day after the bottom sowing if the seed was in the large size (1.5 cm - 2.5 cm).
There is difference in food consumption of S. broughtonii in the same size ingested by
A.amuresis between day and night to some extent. The food consumption at night is larger
than that during the daytime, and the larger the size S. broughtonii is, the later the time with
the largest food consumption will be.

78 Marine Science Bulletin Vol. 18

3.3 Strategy to control A.amuresis in proliferation of S. broughtonii by bottom
sowing
A.amuresis is para-oral predator, which can precisely locate the shellfish by the smell.
According to the search theory, predator will always choose food organisms with high
profitability, and neglect those with low profitability [31]. Tamura thought that the shellfish in
shallow water had stronger adductor muscle than that in deep water [32]. In aspect of
availability of food organism, A.amuresis has much stronger adductor muscle than the
shellfish in shallow water, so more energy is required to open them. Therefore, shellfish
with low value from shallow water can be caught near the proliferation area to be trap food
for the elimination of A.amuresis. In this way, the food consumption of S. broughtonii by
A.amuresis can be reduced to obtain better economic benefit.

Acknowledgements

This paper was supported by the National Sea Welfare Project(201205023); China
Agriculture Research System(CARS-48) and the earmarked fund for Modern Agro-industry
Technology Research System in Shandong Province(SDAIT-14).

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多棘海盘车对魁蚶摄食量、选择性
及昼夜摄食差异的初步研究

张天文,刘广斌,刘恩孚,郑永允,邱兆星*
(山东省海洋生物研究院 山东 青岛 266002)

摘 要:在水温为 16 ~ 20℃、盐度 28.2 ± 2.1、溶解氧为(7.5 ± 0.7)mg/L、pH为 8.0 ± 0.3 的条件下,
研究了 3 种规格多棘海盘车(Asterias amurensis)对 4 种规格魁蚶(Scapharca broughtonii)的摄食量、
摄食选择性和摄食昼夜差异。研究表明:多棘海盘车对不同规格魁蚶摄食数量均随魁蚶底播密度的增加而上
升;不同规格多棘海盘车对同一规格魁蚶摄食数量无显著差异(p>0.05),但对不同规格魁蚶摄食数量存在
一定差异,对于最小壳长组的摄食数量最大;多棘海盘车对不同规格魁蚶软体部摄食量与魁蚶规格呈成显著
正相关;多棘海盘车对不同规格魁蚶摄食数量均存在一定的昼夜差异且夜间的摄食数量大于昼间,魁蚶规格
越大,多棘海盘车对魁蚶最大摄食数量时间出现的越晚。结果表明:魁蚶适宜底播密度在 30 ind/m2 左右,
底播规格越大越能有效防御敌害。根据多棘海盘车对贝类捕食特点,可采用浅水低值贝类作为诱捕生物对多
棘海盘车进行清除。

关键词:多棘海盘车;魁蚶;摄食量;摄食选择性;昼夜差异