Journal of Experimental Marine Biology and Ecology 461 (2014) 337343

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Journal of Experimental Marine Biology and Ecology

journal homepage: www.elsevier.com/locate/jembe

Effects of water current on swimming performance, ventilation

frequency, and feeding behavior of young seahorses
(Hippocampus erectus)
Geng Qin, Yanhong Zhang, Liangmin Huang, Qiang Lin ,1
Key Laboratory of Tropical Marine Bio-resources and Ecology, South China Sea Institute of Oceanology, Chinese Academy of Sciences, Guangzhou, Guangdong 510275, PR China
University of Chinese Academy of Sciences, Beijing 100049, PR China

a r t i c l e

i n f o

Article history:
Received 23 May 2014
Received in revised form 5 September 2014
Accepted 6 September 2014
Available online 26 September 2014
Keywords:
Feeding
Hippocampus erectus
Swimming performance
Ventilation
Water current

a b s t r a c t
Seahorses are poor swimmers, and their dispersion is a topic of increasing interest for their conservation. In the
present study, the swimming performance, ventilation, and feeding behavior of the young seahorses Hippocampus
erectus Perry, 1810 under simulated water currents (control-, low-, mid-, and high velocities) were investigated.
Between 25.0% and 33.9% of the seahorses in the experimental groups (3, 10, and 30 days post birth [DPB]) stayed
attaching to the substrate, and the rest swam if there was no water current; however, the attachment rates of the
10 and 30 DPB seahorses increased signicantly in the high velocity treatment (P b 0.05). The ventilation rates decreased signicantly with the increase of seahorse age under the high water current, regardless of whether they
were swimming or attaching (P b 0.05). For all of the groups, the high water current could signicantly decrease
the total feeding frequency (TFF) and effective feeding frequency (EFF), with the swimming seahorses having a
higher feeding frequency than that of the attaching seahorses. The young seahorses in the high water current treatment had lower growth rates and suffered total mortality by the end of 30 day experiment.
2014 Elsevier B.V. All rights reserved.

1. Introduction
Seahorses (Syngnathidae, Hippocampus) are unusual sh because of
their unique male parental care regimen and morphological appearance
(Linton and Soloff, 1964). Seahorses swim with an upright posture, and
they can continue swimming slowly within the coral, seagrass, and
mangrove habitats in temperate and tropical ocean regions (Lourie
et al., 1999; Teske and Beheregaray, 2009). Seahorses are popular for
use in traditional Chinese medicine, and there has been a growing interest in marine aquaria in recent years (Baum and Vincent, 2005; Lin et al.,
2009a, 2012). The over-exploitation of wild seahorse stocks, however,
has led to a decline, and even depletion, in many tropical regions
(Lourie et al., 1999). Therefore, all seahorses have been regulated for
international trade by Appendix II of CITES (Convention for the
International Trade in Endangered Species of Wild Fauna and Flora,
www.cites.org).
For marine sh, swimming ability during their larval stages is essential for their ability to avoid predators, obtain food, and control dispersal
patterns (Fisher, 2005; Leis et al., 2011). The study of the swimming

Corresponding author at: Key Laboratory of Tropical Marine Bio-resources and

Ecology, South China Sea Institute of Oceanology, Chinese Academy of Sciences,
Guangzhou 510301, PR China. Tel.: +86 20 89023161; fax: +86 20 84452611.
E-mail address: linqiangzsu@163.com (Q. Lin).
1
Postal address: No. 164, Xingang West Rd. Guangzhou 510301, PR China.

http://dx.doi.org/10.1016/j.jembe.2014.09.001
0022-0981/ 2014 Elsevier B.V. All rights reserved.

performance of sh larva has often been focused on the connection between survival, habitat behavior, and surrounding environments
(Fisher and Bellwood, 2002). Seahorses are small carnivorous demersal
sh, and they have poor swimming ability in comparison with other sh
(Ashley-Ross, 2002). The dorsal and pectoral ns in seahorses cannot
provide the propulsion necessary for long distance migration, and
their body shapes appear to be incapable of maintaining high velocity
(Michelle, 1993). Seahorses have prehensile tails for grasping holdfasts
tightly (Porter et al., 2013); this grasping ability helps seahorses attach
to substrates, which allows them to more effectively forage for food and
evade predators (Kleiber et al., 2011). Seahorses can also grasp owing
objects, which provides a mechanism of dispersal for the seahorse populations (Luzzatto et al., 2013; Teske et al., 2007). Compared with adult
seahorses, studies of the swimming behavior and abilities of newborn
seahorses are decient because of the difculty of obtaining small
seahorses from the wild.
Ventilatory activity in sh refers to the ow of water ventilated over
the gills, performed by bucco-opercular movements, aiming to ensure
the intake of oxygen and the release of wastes, such as carbon dioxide
(Martins et al., 2012). In fact, there is a robust correlation between oxygen uptake rate and ventilation frequency for most sh, including
seahorses (Adams et al., 2001). When sh display increased ventilation
frequency, it is commonly viewed as a sign of stress, poor health, and/or
high energy consumption (Martins et al., 2012). For seahorses, the gill is
tufted due to its irregular crumpled shape, which appears to be attached

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G. Qin et al. / Journal of Experimental Marine Biology and Ecology 461 (2014) 337343

to stems (Prein and Kunzmann, 1987). Ventilation frequency elevates

signicantly in seahorses either acutely or chronically when exposed
to ammonia at low concentrations; however, it is not as sensitive to
nitrite exposure (Adams et al., 2001).
The feeding process in seahorses is a particularly interesting topic
because of the special capture behavior and special long-snouted head
of seahorses (Felcio et al., 2006; Lin et al., 2009b,c; Roos et al., 2009;
Van Wassenbergh et al., 2009). Head morphology functions to create a
reduced uid deformation zone, minimizing hydrodynamic disturbance
while feeding strikes occur, and permits syngnathid sh to approach
highly sensitive copepod prey undetected (Gemmell et al., 2013). Feeding rate is a primary factor affecting the growth of seahorses, and environmental factors, such as light intensity, prey type, and temperature
can partially affect seahorse feeding frequency (Celino et al., 2012; Lin
et al., 2010; Sheng et al., 2007).
Water current is one of the most important factors affecting the ontogenetic behavior, growth, and population distribution of sh (Merino
et al., 2007). Fish have evolved their corresponding behavior to adapt to
the ocean current (Blake, 1976). The lined seahorses Hippocampus
erectus are widely dispersed in the western Atlantic Ocean, and their
main habitat is the seagrass or coral reef in the shallow sea. The whole
lifespan and the distribution structure of this species are markedly
affected by the western Atlantic Ocean currents (Lourie et al., 1999).
The present study aimed to investigate the swimming performance,
ventilation rate, and feeding behavior of young seahorses H. erectus
exposed to water currents, so as to predict the relationship between
water currents and dispersion, and the feeding and survival strategies
of seahorses.
2. Material and methods
2.1. Experimental seahorses
F4 generation adult seahorses H. erectus were reared in the Shenzhen
Seahorse Center of the South China Sea Institute of Oceanology, Chinese
Academy of Sciences (SCSIO-CAS) (110.04E, 20.54N) with animal
ethics approval for experimentation granted by the Chinese Academy
of Sciences. Seahorses were maintained in re-circulating holding tanks
(90 80 60 cm) each with ve pairs, and seawater was pumped directly from the South China Sea and treated with double sand ltration.
Attachment substrates for seahorses were nylon mesh and plastic
grasses. Temperature, salinity, pH, light intensity, dissolved oxygen
(DO), and photoperiod were maintained at (mean S.D.) 25

0.5 C, 32 1.0, 7.9 0.4, 2000 lx, 6.5 0.5 mg L1, and 16 L: 8D,
respectively. Seahorses were fed three times a day (0900, 1200, and
1600 h) with live Artemia, and feces and uneaten food were siphoned
out daily. The newborn seahorses were moved into re-circulating holding tanks (50 30 40 cm) and fed four times a day (0830, 1230, 1730,
and 2130 h) with enriched Artemia metanauplii.
The experiment was conducted in annular glass tanks (120 80
80 cm3), and the water depth was 70 cm (Fig. 1). Air ow generated
by an air pump (YFSR, Yafeng, Hebei, China) was used to create and
maintain the experimental water current velocities, which were measured by a current meter (LS-600A, Dongxiyi, China). There was no air
ow in the control tanks. The inlet and outlet seawater ow rates of
each tank were kept at 0.30.4 L min1, and the water ow rate in
the tanks was approximately 0.003 m s 1. Three diameters of nylon
meshes (0.9 mm, 1.5 mm, and 5 mm) were used as the substrates for
the seahorses (Martinez-Cardenas and Purser, 2012). A digital video
camera in the center of the annular tanks was used to record the swimming performance, ventilation, and feeding behavior of the seahorses
(Fig. 1).
2.2. Experimental design and measurements
2.2.1. Swimming performance and ventilation frequency under different
water currents
Four treatments of water currents (control: 0 m s1; low: 0.02
0.01 m s1; mid: 0.10 0.02 m s1; high: 0.20 0.02 m s1, which
were among the common velocity ranges of the local seawater currents
in Daya Bay [110.04E, 20.54N]), each with three replicates, were
performed to examine the effects of water velocity on the swimming
performance of the seahorses. Three different ages of juvenile seahorses
(3 days post birth [DPB] [11.8 0.2 mm]; 10 DPB [25.0 3.1 mm], and
30 DPB [36.1 4.4 mm]) were moved into the experimental annular
tanks, each with 20 seahorses of the same age. During the experiment,
the temperature, salinity, light intensity, and photoperiod in the annular
tanks were 25 0.5 C, 32 1.0, 2000 lx, and 16 L: 8D, respectively.
Artemia metanauplii with the density of 0.1 ind mL1 was utilized to
feed the 3 DPB and 10 DPB seahorses, and Artemia 0.6 0.2 cm in
body length was used for the 30 DPB seahorses. All of the seahorses
were fed three times a day (0900, 1600, and 1900 h). The feces and
dead Artemia metanauplii were siphoned out from the tanks at 0000 h
(midnight) daily.
The swimming performance in this experiment was monitored for
4 days in the annular tanks, and they were recorded ve times a day,

Fig. 1. Annular glass tank (top view) used in this experiment for investigating the swimming performance, feeding behavior, ventilation, and survivorship of young seahorses Hippocampus
erectus.

G. Qin et al. / Journal of Experimental Marine Biology and Ecology 461 (2014) 337343

30 min each time, using the digital video camera (09000930, 1100
1130, 14000430, 16001630, and 19001930 h). Because of the limitation of the number of annular glass tanks, this experiment was conducted sequentially for each age group (3, 10, and 30 DPB), and each
treatment was studied for 4 days. All the experimental seahorses were
from different broods born at the same day. Five different swimming
types of seahorses were recorded, as follows: (1) Ruleless swimming
type (RS) which could change its swimming direction and speed at
any moment and swam freely; (2) Contranatant swimming type (CS)
which swam against the water current; (3) Downstream swimming
type (DS) which swam with the water current; (4) Driftage type (DR)
which oated with the water current; and (5) Attachment type (AT)
which grasped the nylon mesh in the water. The experimental
seahorses that had the same swimming behavior were summed up to
calculate the nal ratios of different types.
The ventilation frequency of the seahorses was also recorded
through counting the frequency of operculum movements per minute
(Van Rooij and Videler, 1996). During 4 days of the investigation for
the swimming performance, the ventilation frequency of six swimming
seahorses and six attaching seahorses in each treatment was recorded,
respectively, through the digital video camera at 09000930 and
14000430 h per day, and the comparison of ventilation frequency
between the swimming and attaching seahorses was conducted after
the 4 days of investigation.
2.2.2. Feeding frequency and daily food intake under different water
currents
The same experiment as Section 2.2.1 was conducted to study the
effects of water currents on feeding frequency and daily food intake
of the seahorses of different ages (3, 10, and 30 DPB). Live Artemia
metanauplii and sub-adult Artemia (0.6 0.2 cm) were used as the
prey. The photoperiod and light intensity in the annular tanks were
24 L: 0D and 2000 lx, and the water conditions in the tanks were the
same as those in experiment in Section 2.2.1.
The experimental seahorses, which were randomly selected from
the temporary tanks, were moved into the annular tanks a day prior
to the start of the experiment. During the experiment, the seahorses
were fed once a day (0900 h) with Artemia metanauplii (the 3 and
10 DPB) and with sub-adult Artemia (0.6 0.2 cm) (the 30 DPB), respectively. The total feeding frequency (TFF), which was dened by
the total number of times of attacking action to the diet (such as the
Artemia in this study) by a seahorse in a minute, and effective feeding
frequency (EFF) of the seahorses, which is dened by the number of
times of successful preying the Artemia by a seahorse in a minute,
were recorded ve times a day, for 15 min each time (09300945,
11301145, 14300445, 16301645, and 19301945 h). Diet densities
in the experimental (Dtest) and control tanks (no seahorse) (Dcontrol)
were measured at the end of the experiment. The daily food intake
(DFI) of the seahorses in different water currents was calculated
through a modied formula (Willadino et al., 2012): DFI = (Ccontrol
Ctest) V/n; V, water volume; n, number of seahorses.
2.2.3. Growth and survival rates under different water currents
Under different water currents (control: 0 m s 1; low: 0.02
0.01 m s1; mid: 0.10 0.02 m s1; high: 0.20 0.02 m s1) with
the same culture conditions as those in experiment in Section 2.2.1,
the 3 DPB seahorses were cultured in each tank for 30 days to examine
the effects of water currents on their growth and survival rates, with
three replicates each with 50 seahorses of the same age. Artemia
metanauplii were used to feed the seahorses at 0900, 1600, and
1900 h every day, and the feces and dead Artemia were siphoned out
from the tanks at 0000 h (midnight) daily. During the experiment, the
number of dead seahorses was recorded daily, and the initial and nal
wet weights and standard body lengths from each replicate were
measured according to the methods of Lourie et al. (1999) and
Job et al. (2002). The specic growth rate (SGR = 100 [ln nal wet

339

weight ln initial wet weight / duration]) of seahorses was calculated

in each treatment.
2.3. Statistical analysis
Statistical analyses were conducted using the software SPSS 19.0
(Statistical Program for Social Sciences 19.0) and Sigma PLOT 10.0
(Version 10.0, Systat Software, Inc). One-way analysis of variance
(ANOVA) was used to assess the differences in swimming performance,
ventilation frequency, feeding frequency and intakes, and the growth
and survival rates of the seahorses among the treatments with a significance level of 0.05. If ANOVA effects were signicant, comparisons
between the different means were made using the post hoc least significant differences (LSD).
3. Results
3.1. Swimming performance under different water currents
The swimming performance of different ages of seahorses (3, 10, and
30 DPB) was signicantly different under the four water current groups.
From 66.1% to 75.0% of the seahorses maintained ruleless swimming
(RS) behavior when they were not experiencing any water current,
and RS behavior still occurred in the low water current group. The
3 DPB seahorses all drifted with the owing water, except for 27.9% of
them attaching to the substrate under the high water current. For 10
and 30 DPB seahorses, attachment and downstream swimming behaviors were the main behavior under the mid and high water currents.
Moreover, 25.6% and 12.1% of the 10 and 30 DPB seahorses, respectively,
drifted under the high water current. Under high water current, only
4.3% of the 30 DPB seahorses could perform contranatant swimming
(CS) behavior. It was obvious that the ratios of attachment rates increased, but the drift rates decreased, among the 3, 10, and 30 DPB
seahorse groups with the increasing water currents (Fig. 2).
3.2. Ventilation frequency under different water currents
The water currents signicantly affected the ventilation frequency of
the young seahorses (one-way ANOVA, F(3, 116) = 10.70, P b 0.05)
(Fig. 3). A positive relationship was found between water current and
ventilation frequency in the 3 DPB seahorses, and ventilation frequency
was 169 8 beats min1 and 160 11 beats min1 in the swimming
and attaching groups, respectively, under the 0.20 m s1 water current.
For 30 DPB seahorses, the ventilation frequency decreased under the
high water current and was approximately 85 beats min 1, which
was not signicantly different from those of the seahorses at low or
mid water currents (Fig. 3). For all of the experimental seahorses, the
ventilation frequency of the swimming seahorses was higher than that
of the attaching seahorses. In addition, it was found that the younger
the seahorses, the higher their ventilation frequency (F(3, 118) = 19.1,
P b 0.05).
3.3. Feeding performance
Total feeding frequency (TFF) (one-way ANOVA, F(3, 92) = 8.36,
P b 0.05) and effective feeding frequency (EFF) (F(3, 92) = 18.80,
P b 0.05) were signicantly affected by the water current in each
group. The seahorses kept high feeding rates because of the high ratios
between the EFF and TFF (Fig. 4). Seahorses in the control group had the
highest EFF, which was 4.6 1.2 ind min 1 in the 3 DPB seahorses,
6.1 0.6 ind min1 in the 10 DPB seahorses, and 7.6 1.4 ind min1
in the 30 DPB seahorses, respectively. Three DPB seahorses could not
feed at the high water velocity, and a negative correlation was found between feeding frequency and water current. TFF and EFF of the swimming seahorses were signicantly higher than those of the attaching

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G. Qin et al. / Journal of Experimental Marine Biology and Ecology 461 (2014) 337343

100
AT

Percentage %

80

DR

60
DS

40

CS

20

RS

0
Control Low Mid

High

Control Low Mid

3 DPB

High

Control Low Mid

10 DPB

30 DPB

High

(seahorses)

Fig. 2. Swimming performances (AT, DR, DS, CS, and RS) in different ages of seahorses Hippocampus erectus (3, 10, and 30 DPB) under different water currents (control: 0 m s1; low:
0.02 0.01 m s1; mid: 0.10 0.02 m s1; high: 0.20 0.02 m s1). AT, attachment type; DR, driftage type; DS, downstream swimming type; CS, contranatant swimming type; RS,
ruleless swimming type; and DPB, day post birth.

seahorses, and the 30 DPB seahorses had the highest TFF and EFF among
the different seahorse groups (Fig. 4).
For different age groups of seahorses, the 30 DPB seahorses had the
highest daily food intake, which was 213.3 32.3 ind under the low
water current, and the 3 DPB seahorses had the lowest daily food intake.
For all of the seahorses, the daily food intake was the lowest under the
high water velocity (Fig. 5).
3.4. Growth and survival rates
The growth rates of the seahorses under different water currents
were varied. However, the seahorses had the highest nal wet weight
(one-way ANOVA, F(3, 76) = 24.22, P b 0.05), longest body length
(F(3, 76) = 15.02, P b 0.05), and highest survival rate (F(3, 76) = 31.11,
P b 0.05) under the low water current. All of the seahorses under the
high water current expired, and the seahorses in the low water current
had the highest survival rate, which was 23.7 1.2% after the 30 days of
culture (Table 1).
4. Discussion
The present study investigated the swimming and feeding behavior
of young seahorses under different owing water rates. Most previous

works afrmed that seahorses were likely to attach themselves to substrates (xed or oating) (Caldwell and Vincent, 2013; Luzzatto et al.,
2013). Moreover, it was not proper to utilize any classical model, such
as body and/or caudal n movements, or employ median and/or paired
n propulsion directly, to describe the swimming performance of
seahorses (Sfakiotakis et al., 1999). Seahorses never continue to swim
against water current and they always attempt to grasp surrounding
substrates. As a consequence, the cruising swimming speed and critical
swimming speed that were commonly used to describe the swimming
ability of sh were not suitable for seahorses (Fisher and Bellwood,
2002; Fisher et al., 2005). In the present study, the swimming ability
of seahorses became increasingly strong as they grew older, and the
30 DPB seahorses (36.1 4.4 mm) could swim against the water current or swim freely under the high water velocity. In contrast, the
3 DPB seahorses would attach themselves to the substrates, or they
would drift under the high water current. The swimming speed of the
3 DPB seahorses was approximately 46 cm s 1, which was much
slower than the average sustaining swimming speeds of the late-stage
pelagic larvae of coral reef sh (20.6 cm s1) (Stobutzki and
Bellwood, 1997). Therefore, the young seahorses were relatively poor
swimmers (Blake, 1976).
Water current is one of the most important factors affecting the
distribution of the pelagic larvae of sh (Norcross and Shaw, 1984).
10 DPB

3 DPB

150
120
90

120
90

30

30
Mid

Swimming
Attaching

30 DPB

150
120
90
60
30
Control Low

Mid

Control Low

High

High

VF in attaching seahorses min -1

Control Low

VF beats min-1

150

60

60

180

Attaching
Swimming

180

Attaching
Swimming

VF beats min-1

VF beats min-1

180

Mid

High

3 DPB
10 DPB
30 DPB

180
150
120
90
60
30
30

60

90

120

150

180

VF in swimming seahorses min-1

Fig. 3. Relationship of ventilation frequency (beats min1) between the swimming and attaching seahorses Hippocampus erectus (3, 10, and 30 DPB) under different water currents
(control: 0 m s1; low: 0.02 0.01 m s1; mid: 0.10 0.02 m s1; high: 0.20 0.02 m s1). VF, Ventilation frequency.

Ind min-1

G. Qin et al. / Journal of Experimental Marine Biology and Ecology 461 (2014) 337343

TFF
EFF

3 DPB, swimming

10 DPB, swimming

10 DPB, attaching

30 DPB, swimming

30 DPB, attaching

3 DPB, attaching

6
3

Ind min-1

0
9
6
3

Ind min-1

0
9
6
3
0
ControlLow Mid High

ControlLow Mid High

Fig. 4. Total feeding frequency (TFF) and effective feeding frequency (EFF) of the swimming and attaching seahorses Hippocampus erectus (3, 10, and 30 DPB) under different
water currents (control: 0 m s1; low: 0.02 0.01 m s1; mid: 0.10 0.02 m s1;
high: 0.20 0.02 m s1).

Daily food intake ind d -1

Newborn seahorses were recognized to take pelagic lives based on the

investigation of plankton samples (Foster and Vincent, 2004). Our observation showed that newborns could grasp substrates immediately
without the planktonic life, and this result is consistent with the report
of the settling phenomenon of Hippocampus fuscus (Golani and Fine,
2002). More than 40% of the 30 DPB seahorses were likely to grasp substrates, and the stronger and longer prehensile tails of larger seahorses
might partially contribute to this attachment. In addition, water velocity
could signicantly impact attachment for young seahorses. Of the
30 DPB seahorses in the high water current, 50.2% would grasp the substrates. In the ocean, swimming ability has been commonly recognized
to be important for dispersal (Fisher, 2005), and seahorses would be less
likely to demonstrate dispersal patterns similar to those of reef sh families with good swimming abilities (e.g., Holocentridae, Siganidae,
Acanthuridae, & Lutjanidae) (Fisher et al., 2005). Rafting of juveniles
and/or adults on drifting vegetation or other holdfasts was commonly
recognized to be an important dispersal mechanism for seahorses
(Boehm et al., 2013; Teske et al., 2007). The further investigation of
seahorse swimming and grasping performance in wild conditions
would contribute to understanding dispersal, as well as population
distribution of seahorses.
Adams et al. (2001) noted that ventilation frequency was signicantly elevated in seahorses either acutely or chronically exposed to
ammonia at a low concentration, and a positive relationship between
ventilation frequency and oxygen consumption was found. Moreover,

400
300

3 DPB
10 DPB
30 DPB

200
100
0

Control Low

Mid

High

Fig. 5. Daily food intake of seahorses Hippocampus erectus (3, 10, and 30 DPB) under
different water currents (control: 0 m s1; low: 0.02 0.01 m s1; mid: 0.10
0.02 m s1; high: 0.20 0.02 m s1).

341

an increase in the ventilation rates was accompanied by an increase in

the metabolic rates of adult seahorses (Aurlio et al., 2013). In the
present study, under the different water currents, the ventilation frequencies of different ages of seahorses were dramatically different,
with the younger seahorses having a higher frequency under the same
water current. Moreover, the seahorses that were swimming had a
higher ventilation frequency than those that were grasping. Ventilation
has been commonly used as an indicator for evaluating the physical
condition of organisms, and well-balanced ventilatory activity is fundamental to maintaining homoeostasis in terms of oxygen status, blood,
and tissue acidbase balance at precise set points (Martins et al., 2012).
The ventilation frequency of juvenile seahorses Hippocampus
abdominalis is 1037 beats min 1 (Adams et al., 2001), and that
of adult seahorses Hippocampus guttulatus is 1830 beats min 1
(Aurlio et al., 2013), which were all lower than that of young seahorses
H. erectus in the present study. Different experimental conditions, especially temperature, body conditions, and water ow velocity (Johnston
and Dunn, 1987) might directly contribute to the variances among
these seahorses.
Feeding can be signicantly affected by water current, and starvation
was the main reason for the death of sh larvae in high current conditions (Kato et al., 2008; Oshima et al., 2009). Feeding behavior has
been widely studied in seahorses under different environmental conditions (Gemmell et al., 2013; Lin et al., 2009c; Olivotto et al., 2008). Diet
composition and feeding periodicity of Hippocampus barbouri were
compared in illuminated and non-illuminated cages for juvenile and
adult seahorses (Garcia et al., 2012), and studies on prey selectivity
and nutrition under laboratory conditions were performed to identify
appropriate diets in aquaculture (Storero and Gonzalez, 2008). In the
present study, a negative correlation between EFF and water ow
velocity was found in both swimming and holding seahorses. Young
seahorses (3 DPB) could not feed, whereas, the 30 DPB seahorses had
high EFF, under high water ow velocity. Moreover, this study also
found that the swimming seahorses had higher feeding frequency
than that of holding seahorses, which seemed to be less affected by
water current during the feeding. This might be due to the unique
feeding behavior in seahorses, e.g., they usually feed on live diets and
utilize an ambushing strategy, which can greatly improve their feeding
success (James and Heck, 1994; Qin et al., 2012; Sheng et al., 2007; Van
Wassenbergh et al., 2009).
Water current would increase the encounter rate with prey and
decrease the prey escape response; however, the higher current could
also increase the predators' time and energy cost of swimming and
feeding on sh larvae (Kloppmann et al., 2002; MacKenzie and
Leggett, 1991). Moreover, grasping activity might be an effective strategy for seahorses to reduce the disturbances by the water current, so
increasingly more seahorses performed their feeding from the substrates as they grew older. In the present study, daily food intake efciency of the 30 DPB seahorses was 165213 ind per day, and the
young seahorses (3 DPB) had the lowest level of daily food intake.
This result is mainly due to the difference of digestive system, i.e., the
intestine of the 3 DPB seahorses is a short pipe which becomes more
complex in the 30 DPB seahorses (Lin et al., 2007; Storero and
Gonzalez, 2008).
High water currents can increase energy consumption, and reduce
the growth and survival rates of seahorses. Seahorses that cultured at
the mid water current rate had relatively higher feeding frequency
and daily food intake efciency, but they had lower growth and survival
rates during the 30 days of culture. Moreover, all seahorses expired
before the end of the experiment if they were cultured at the high
water current. This result is consistent with relatively low feeding and
daily food intake efciency, indicating that sh larva suffered from starvation and high energy consumption due to water current disturbance
(MacKenzie et al., 1994). Larva of Japanese ounder (Oshima et al.,
2009) and bluen tuna (Kato et al., 2008) also showed a similar phenomenon. In this study, seahorses placed at low water current had the

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G. Qin et al. / Journal of Experimental Marine Biology and Ecology 461 (2014) 337343

Table 1
Growth and nal survival rates of the 3 DPB seahorses Hippocampus erectus cultured under four water velocities (control, low, mid and high) after 30 days of culture.
Control
Initial body height (mm)
Initial wet weight (mg)
Final body height (mm)
Final wet weight (mg)
Condition factor (g cm3)
SGR (% d1)
Survival rate (%)

11.3
6.1
34.5
190.1
0.46
11.46
77.1

Low
1.2 a
1.3 a
3.6 a
60.3 b
0.03 a
1.33 a
5.5 b

11.5
6.5
36.3
210.3
0.44
11.59
84.1

Mid

1.7 a
1.8 a
4.6 a
40.6 a
0.05 b
1.30 a
3.4 a

11.3
6.4
28.9
150.3
0.62
10.52
23.7

High

1.5 a
.0.9 a
7.6 b
90.3 c
0.13 c
1.41 a
1.2 c

11.6 2.2 a
6.5 1.3 a

0d

Mean S.D. with different superscripts within a row are signicantly different (one-way ANOVA, P b 0.05).

highest growth and survival rates, and this might be due to the enhanced encounter rate between seahorses and prey at the appropriate
water current (Barlow, 1994; Dower et al., 1997). This result might
provide a new strategy for juvenile seahorse culture.
In conclusion, less than one third of the young seahorses H. erectus
attached to the substrate, and the rest swam if there was no water current, and their swimming performance was signicantly different under
the different water velocities. The ventilation rates decreased with the
increase of seahorse age under the high water current. A negative relationship between the high water current and the total feeding frequency (TFF) and effective feeding frequency (EFF) of seahorses was found,
and the young seahorses in the high water current suffered total mortality. This study provides a simulated scene for the habitation and survivorship of young seahorses in the wild.

Acknowledgments
Special thanks to the staff in the Seahorse Center of South China
Sea Institute of Oceanology, Chinese Academy of Sciences (SCSIO-CAS)
for their help at various stages of this study. This study was funded
by the Outstanding Youth Foundation in Guangdong Province
(S2013050014802), the National Science Fund for Excellent Young
Scholars (41322038), the Guangdong Oceanic and Fisheries Science and
Technology Foundation (A201201I03, A201208G02), and the Development of Special Biology Industry in Shenzhen City (HY20130205001). [SS]

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