Aquaculture Nutrition
2016 22; 25–33
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1 1
1
Graduate School of Marine Science and Technology, Tokyo University of Marine Science and Technology, Minato, Tokyo,
Japan; 2 Faculty of Life Sciences, Toyo University, Itakura, Oura, Gunma, Japan; 3 Faculty of Nutritional Science,
Sagami Women’s University, Sagamihara, Kanagawa, Japan
Phyllosomas (planktonic larvae) of slipper lobsters cling
onto and feed on jellyfish under both natural and labora-
tory conditions. Phyllosomas of Ibacus novemdentatus are
capable of feeding on various jellyfish species including
venomous stingers; however, the range of jellyfish species
capable of supporting the growth and survival of phylloso-
mas is unknown. Seventeen (12 for the first and five for the
second trials) and 18 (13 for the first and five for the sec-
ond trials) phyllosomas were fed exclusively on the jellyfish
Aurelia aurita and Chrysaora pacifica, respectively. Aurelia
aurita-fed phyllosomas metamorphosed into the nisto stage
(postbenthic larvae)  54 days after hatching, whereas
C. pacifica-fed phyllosomas did not. Major nutritional
compositions such as amino acids, fatty acids and minerals
were compared between the two jellyfish species. The pro-
portion of each major nutritional component was not sig-
nificantly different between the two jellyfish species,
suggesting that C. pacifica was not nutritionally inferior to
A. aurita. Therefore, the observation that the C. pacifica-
fed phyllosomas did not metamorphose into the nisto
stage was not because of major nutritional compositions
but due to other factors such as the lack or excess of other
minor nutrients, or the species-specific texture of the
jellyfish.
KEY WORDS: amino acids, fatty acids, growth, lobster, minerals,
phyllosoma
Received 7 January 2014; accepted 19 June 2014
Correspondence: K. Wakabayashi, Tokyo University of Marine Science
and Technology, 4–5–7 Konan, Minato, Tokyo 108–8477, Japan.
E-mail: kaoriw@kaiyodai.ac.jp
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ª 2015 John Wiley & Sons Ltd
doi: 10.1111/anu.12228
1,2 3 1
Mass production of juveniles in spiny (palinurid) and slip-
per (scyllarid) lobsters is a critical step for the commercial
production and aquaculture of these species. One of the
principal issues hampering successful mass production of
juvenile lobsters is the limited information available on
suitable dietary regimes for growth and survival during the
long phase of planktonic larvae (phyllosomas) (Cox &
Johnston 2003; Cox et al. 2011). To this end, many
researchers have focused on an understanding of phylloso-
ma development. The complete development of phylloso-
mas from hatching to settlement has been reported in 10
(Phillips & Matsuda 2011) of 32 (Groeneveld et al. 2013;
Jeffs et al. 2013; Phillips et al. 2013) and 11 (Vijayakuma-
ran & Radhakrishnan 2011) of 33 (Spanier & Lavalli 2013)
economically important species of spiny and slipper lob-
sters, respectively. However, the mass production of juve-
nile lobsters has only been achieved in a slipper lobster
Thenus orientalis (Mikami & Kuballa 2007).
There is abundant evidence that wild phyllosomas associ-
ate with and feed on jellyfish (reviewed by Booth et al.
2005; Sekiguchi et al. 2007; Wakabayashi & Tanaka 2012).
For example, phyllosomas of the genus Ibacus and Scylla-
rus are known to cling usually onto the cnidarian jellyfish
(e.g. Herrnkind et al. 1976; Ates et al. 2007), and those of
Parribacus antarctics collected from the north of Bermuda
possessed several types of cnidarian nematocysts in their
faecal matter (Sims & Brown 1968). In addition, those of
Panulirus japonicus and Scyllarides squammosus collected
from Atlantic and Pacific waters utilized cnidarians and
urochordates as food, according to the DNA analysis of
their gut contents (Suzuki et al. 2006). Laboratory experi-
ments also suggest that gelatinous zooplankton is a suitable
diet for phyllosomas (e.g. Goldstein & Nelson 2011). In a
series of prey feeding trials, Saunders et al. (2012) found
that phyllosomas of Panulirus cygnus preferred chaetogn-
aths and salps to krill, and Wakabayashi et al. (2012b)
found that those of Ibacus novemdentatus showed a higher
survival rate when fed on jellyfish rather than Manila clam,
Ruditapes philippinarum. Morphological analyses have also
suggested that mouthparts and internal organs of phylloso-
mas are adapted for masticating and digesting the soft tis-
sues of gelatinous zooplankton (Nishida et al. 1990; Cox
& Johnston 2003; Simon et al. 2012).
The next major step towards the mass production of lob-
ster juveniles will be the identification of essential dietary
requirements that influence the growth and survival of
phyllosomas (Cox & Johnston 2003). A key element is the
study of the biochemical compositions of potential feed-
stock (Jones et al. 1997; Wang & Jeffs 2013). All gelatinous
zooplankton including jellyfish may not always be suitable
for the growth and survival of phyllosomas because their
biochemical compositions vary depending on the species,
collection dates and locality (Lucas 1994; Nelson et al.
2000; Wang & Jeffs 2013). Kittaka (2005) suggested that
the growth and survival of phyllosomas were affected by
differences in the nutritional value of dietary jellyfish
although dietary regimes used in that study were altered
during the course of the feeding cycle. Here, we examined
the growth and survival of phyllosomas fed exclusively on
the jellyfish Aurelia aurita or Chrysaora pacifica (both are
scyphozoans) to clarify the effect of different dietary jelly-
fish species on phyllosoma development. The major nutri-
ents in both jellyfish species were determined, and the
impact of the nutritional values of these jellyfish on the
growth and survival of phyllosomas is discussed.
Aurelia aurita and C. pacifica were captured using scoop
nets from Tokyo Bay from April to August, 2012, on the
training ships Seiyo-maru and Hiyodori of the Tokyo Uni-
Table 1 Numbers of Ibacus novemdentatus phyllosomas used for each experiment
No. of phyllosomas used for experiments
Trial Date of hatching Aurelia aurita
1 15 April 2012 12
2 19 April 2012 5 5
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versity of Marine Science and Technology (TUMSAT). Jel-
lyfish bell diameters ranged 100–260 mm for A. aurita and
80–200 mm for C. pacifica. All the jellyfish were transported
to the laboratory immediately after collection. In the same
method described by Purcell et al. (2013), the jellyfish for
phyllosomal feeding trials were maintained in a 500-L tank
(density of one individual per 5 L) with recirculating sea
water without feeding. For biochemical analyses, the jelly-
fish were washed with freshwater immediately after trans-
port to the laboratory and freeze-dried under vacuum
pressure (10–20 °C and 5 9 100–5 9 10 1 Pa) using a freeze
dryer (VD-800R; Taitec Co., Tokyo, Japan).
Ibacus novemdentatus was chosen for this study because
phyllosomas of this species were known to feed on various
jellyfish species (Wakabayashi et al. 2012a). A total of four
egg-bearing females were caught off Kitaura, Miyazaki Pre-
fecture, on April 5, 2012, and transported to the laboratory
of TUMSAT. The females were maintained at 23 °C with
recirculating sea water until phyllosoma hatching. We
made duplicate feeding trials in this study (Table 1). The
phyllosomas originated from two females that released
thousands of phyllosomas on April 15 and 19, 2012,
respectively. A day after hatching, a total of 25 and 10
phyllosomas, which were swimming near the water surface
with intact appendages, were chosen as specimens for the
respective trials.
The newly hatched phyllosomas were kept individually in
PVC mesh cases (width, 70 mm; length, 70 mm; and
height, 140 mm) and placed on a perforated plate 50 mm
from the bottom of the tank (width, 600 mm; length,
300 mm; and height, 360 mm). The tank contained approx-
imately 30 L of sea water and was equipped with an exter-
nal filter (Eheim 2234 ecco comfort; Eheim GmbH & Co.
KG, Deizisau, Germany) to clean water and provide a
No. of phyllosomas
metamorphosed successfully
Chrysaora pacifica A. aurita C. pacifica
13 4 0
2 0
Aquaculture Nutrition, 22; 25–33 ª 2015 John Wiley & Sons Ltd
 gentle flow (50 mL s 1). Approximately 10 L of sea water
was exchanged weekly; water temperature and salinity were
kept at 23 °C and 34 gL 1, respectively. The phyllosomas
were reared under natural laboratory light conditions.
Phyllosomas were divided into two experimental groups:
12 for A. aurita and 13 for C. pacifica from the April 15
batch and 5 for A. aurita and 5 for C. pacifica from the
April 19 batch (Table 1). They were fed exclusively with
A. aurita or C. pacifica. Immediately before feeding, both
jellyfish species were cut into pieces approximately twice of
the carapace weight of the phyllosomas. Eighteen sets of
two mesh cases joined together, one in which the phylloso-
ma was fed exclusively on A. aurita or the other which the
phyllosoma was fed exclusively on C. pacifica, were ran-
domly arranged in the tank to minimize variations in
experimental conditions. Instars and number of surviving
phyllosomas were recorded daily.
The improved Dumas method (Anderson & Shubin 1957)
was used to determine total carbon and nitrogen in jellyfish
using gas chromatography (SUMIGRAPH NC-220; Sumi-
ka Chemical Analysis Service, Osaka, Japan). A calibration
curve was obtained with aspartic acid (Sumika Chemical
Analysis Service). Crude protein was calculated using the
nitrogen conversion factor of 6.25 (AOAC 1995), total lipid
was determined as total fatty acids, and ash was calculated
by the difference between initial and final weight of the
samples.
Total amino acid composition was determined according to
the method of Ogushi & Harada (1997) with minor modifi-
cations as follows. Samples were hydrolysed using HCl
(3 mg freeze-dried jellyfish sample in 1 mL of 6 M HCl
with 0.02% of 2-methylmercaptoethanol at 110 °C for
24 h) in vacuum hydrolysis tubes (Thermo Fisher Scientific
Inc., Waltham, MA, USA). After evaporation to remove
HCl, protein hydrolysates were dissolved in 1–2 mL of
0.02 N HCl and purified using DISMIC 0.45 lm filters
(DISMIC-13CP series; Advantec MFS, Inc., Dublin, CA,
USA). The hydrolysate solution was analysed using an
amino acid analyzer (L-8900; Hitachi High-Technologies
Co., Tokyo, Japan). Amino acids mixture standard solu-
tion, Type H (Wako Pure Chemical Industries, Ltd.,
Osaka, Japan), was used for making a calibration curve to
identify and quantify amino acids.
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Aquaculture Nutrition, 22; 25–33 ª 2015 John Wiley & Sons Ltd
Fatty acid composition was determined by gas chromatog-
raphy as described by the AOCS official method Ce 1b-89
(AOCS 1998) with minor modifications as follows. Capric
acid methyl ester was added as an internal standard to
accurately weigh total lipid in approximately 1 g of freeze-
dried jellyfish sample. The mixture was saponified with
1.25 M HCl in methanol and methylated at 100 °C for 3 h
to obtain fatty acid methyl esters (FAMEs). The FAMEs
obtained were dissolved in n-hexane (Wako Pure Chemical
Industries) and dehydrated with sodium sulfate (Wako
Pure Chemical Industries). The dehydrated FAMEs were
analysed using a gas chromatograph (TRACE GC Ultra;
Thermo Fisher Scientific Inc.) equipped with a mass spec-
trometer (DSQ II; Thermo Fisher Scientific Inc.), an auto-
sampler (AS-3000; Thermo Fisher Scientific Inc.) and a
Supelcowax 10 fused silica capillary column
(30 m 9 0.25 mm i.d. 9 0.25 lm film thickness; Sigma-
Aldrich Co., St. Louis, MO, USA). Helium gas was used
as a carrier at a constant flow rate of 20 mL min 1. The
injector temperature was 250 °C. The column oven temper-
ature programme was as follows: an initial temperature of
50 °C for 5 min, followed by an increase from 50 to
250 °C at a rate of 10 °C min 1. The final hold time was
30 min. Fatty acids were identified by comparing their
retention times with those of authentic standard com-
pounds (Fatty acids and their methyl esters Kit; GL Sci-
ence, Tokyo, Japan).
For mineral analysis, microwave digestion of the freeze-
dried samples was performed under nitrogen (Wako Pure
Chemical Industries) using the ETHOS-1 microwave diges-
tion system (Milestone Inc., Shelton, CT, USA). Levels of
Ca, Mg, Na and K were determined by atomic absorption
photometry (ANA-182; Tokyo Koden, Tokorozawa,
Japan), and those of B, Al, Mn, Fe, Cu, Zn, As, Mo, Cd,
Hg, Pb, V and Cr were determined by ICP-MS (X series 2;
Thermo Fisher Scientific Inc.).
Significant differences of larval duration between A. aurita-
fed and C. pacifica-fed phyllosomas were determined using
Student’s t-test. Survival rates were calculated using the
Kaplan-Meier method (Kaplan & Meier 1958). The log-
rank test was applied to analyse difference of survival rates
between A. aurita-fed phyllosomas and C. pacifica-fed
phyllosomas. Only the results of first trial were used for the
analysis because the sample number for second trial was
small.
For the analysis of biochemical compositions, all samples
were analysed in triplicate. Data are presented as
mean  standard deviation (n = 3). Significant differences
(P < 0.05) were determined using Student’s two-sided
t-test.
Four phyllosomas (33.3%) from the April 15 batch and
two phyllosomas (40.0%) from the April 19 batch fed with
A. aurita metamorphosed into nistos by the 54th day after
hatching; the shortest metamorphosis time was 41 days
(Fig. 1). On the other hand, no nistos were obtained from
the C. pacifica-fed phyllosomas (Table 2). The survival rate
of C. pacifica-fed phyllosomas was not significantly differ-
ent from that of A. aurita-fed phyllosomas until the 25th
day when most of the phyllosomas were in the fifth instar
(P = 0.91, Fig. 1). Nevertheless, after the 30th day, the sur-
vival rate in C. pacifica-fed phyllosoma dropped signifi-
cantly (P < 0.05, Fig. 1) because of the failure of moulting
from the fifth to the sixth (final) instar. Overall, growth of
the C. pacifica-fed phyllosomas gradually slowed after the
third instar. The duration of the moulting interval tended
to be longer in C. pacifica-fed phyllosomas than that in
A. aurita-fed phyllosomas although no significant differ-
ences were detected except the duration from the second
instar to the third instar (Table 2). Four C. pacifica-fed
phyllosomas reached the sixth instar, and one of them
attempted to metamorphose into nisto after 55 days of
hatching; however, the attempt failed.

The quantities of crude protein, total lipid, total carbon

Figure 1 Percentage survival of Ibacus novemdentatus phyllosomas
fed exclusively on Aurelia aurita (single line) or Chrysaora pacifica
(double line). Each percentage is the mean of two experiments.
Arrow and grey bar indicate the average and range of time when
metamorphosis took place, respectively. No C. pacifica-fed phyllos-
omas underwent metamorphosis, which is why there are no arrows
given here.
Table 2 Average duration (in days) from hatching to the achieve-
ment in each instar in Ibacus novemdentatus phyllosoma fed exclu-
sively on Aurelia aurita or Chrysaora pacifica
and total nitrogen were higher in C. pacifica than in A. au-
rita, whereas that of ash was significantly lower in C. pacif-
ica than in A. aurita (Table 3).
The quantity of total amino acids was significantly
higher in C. pacifica than in A. aurita. In addition,
C. pacifica showed significantly higher levels of individual
amino acids than A. aurita except for phenylalanine
(Appendix 1). Although the quantities of each amino acid
were higher in C. pacifica, the percentage composition of
them were similar between the two jellyfish species (Fig. 2,
Table 3 Proximate composition of the jellyfish species, Aurelia au-
rita and Chrysaora pacifica (mg gDW 1; mean  standard devia-
tion; n = 3)
Jellyfish

Instar A. aurita C. pacifica Chemical parameters A. aurita C. pacifica

2 5.9  1.2 6.6  1.6 Crude protein 34.87  b

2.83 75.31  19.96a
3 10.5  1.7a 12.4  2.4b Total lipids 4.26  0.88b 7.15  1.94a
4 15.8  2.0 16.9  2.4 Ash 761.9  9.96a 690.5  9.17b
5 21.4  3.2 24.7  4.3 Carbohydrates 199.0  11.86b 227.1  11.44a
6 30.7  5.4 36.3  6.7 Total carbon 20.47  1.83b 42.28  10.93a
Nisto 47.7  4.4 N.A. Total nitrogen 5.58  0.45b 12.05  3.19a
Amino nitrogen 5.19  0.25b 9.82  0.39a
No nistos were obtained from C. pacifica-fed phyllosomas.
Different superscript letters indicate a significant difference Different superscript letters on each line indicate a significant
(P < 0.05). The duration of the third instar was significantly dif- difference (P < 0.05). Carbohydrates correspond to the organic
fered between A. aurita and C. pacifica. matter minus crude protein, total lipids and ash.
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Aquaculture Nutrition, 22; 25–33 ª 2015 John Wiley & Sons Ltd
 Figure 2 Average percentage of total amino acid composition of the jellyfish species Aurelia aurita (grey bars) and Chrysaora pacifica
(dotted bars) (n = 3). Asterisks (*) indicate significant differences (P < 0.05). Essential amino acids (EAA) for crustaceans are underlined.
EAA shows the sum of the nine kinds of EAA.

Appendix 1). For both species, the major amino acid was
glycine, which accounted for approximately 15% of total
amino acids, followed by glutamine and proline. Essential
amino acids (EAA) seemed to be higher in A. aurita than
in C. pacifica although no significant differences were
detected (Fig. 2, Appendix 1). More than 90% of the total
nitrogen was amino nitrogen in A. aurita, whereas 81.5%
in C. pacifica (Table 3).
Among five essential fatty acids (EFA), the quantities of
C20:4 and C22:6 were significantly higher in C. pacifica
than in A. aurita, whereas that of C20:5 was higher in
A. aurita than in C. pacifica (Fig. 3, Appendix 2). On the
other hand, the quantities of C18:2, C18:3 and the total
EFA were not significantly different between the two spe-
cies. No significant differences were detected between the
quantities of polyunsaturated, monounsaturated and satu-
rated fatty acids. Moreover, the percentage composition of
each fatty acid was similar between the two species (Fig. 3,
Appendix 2).
Although the quantities of ash and major minerals were
significantly lower in C. pacifica than in A. aurita, the
percentage composition of each major mineral was similar
between the two species (Appendix 3). In addition, no sig-
nificant differences were observed in the quantities of trace
minerals.
The quantities and proportion of prey and its biochemical
composition are major factors in the selection of prey by
predators (Schoener 1971; Wang & Jeffs 2013). Lipids are
especially important nutritional components for phylloso-
mas since the accumulation and storage of such energy
constituents are critically linked to the non-feeding nisto or
puerulus stage (Jeffs et al. 2001). Although proteins and
lipids were more abundant in C. pacifica than in A. aurita,
I. novemdentatus phyllosomas were able to survive by
exclusive feeding on A. aurita as well as C. pacifica. Indeed,
we confirmed that the proportion of major nutritional com-
ponents was basically similar between the two species,
which means both A. aurita-fed and C. pacifica-fed phyllos-
omas could utilize the similar major nutritional compo-
nents for their growth and survival. We also confirmed
that both A. aurita and C. pacifica contained amino acids

Figure 3 Average percentage of total fatty acid composition of the jellyfish species Aurelia aurita (grey bars) and Chrysaora pacifica (dotted
bars) (n = 3) Asterisks (*) indicate significant differences (P < 0.05). Others include C11:0, C12:0, C17:1, C18:4, C19:0, C20:0, C20:1, C20:3
and C22:1. C18:2 and C18:3 are shown here although the levels of both were < 2% of the total. Essential fatty acids (EFA) for crustaceans
are underlined, and EFA shows the sum of the five kinds of EFA. Polyunsaturated fatty acids (PUFA), monounsaturated fatty acids
(MUFA) and saturated fatty acids (SFA) also show the sum of PUFA, MUFA and SFA, respectively.
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Aquaculture Nutrition, 22; 25–33 ª 2015 John Wiley & Sons Ltd
and fatty acids essential for crustacean nutrition, support-
ing previous studies of A. aurita (Fukuda & Naganuma
2001; Kittaka 2005). Moreover, six essential minerals (Ca,
Cu, K, Mg, Se and Zn), which must be supplied in the diet
for crustaceans (Davis & Gatlin 1996), were found in both
jellyfish species. Phosphate, also essential for crustaceans,
had previously been found in both A. aurita and C. pacif-
ica (Fukushi et al. 2004). All of the nutritional data are
presented as dry weight, and no moisture content data are
provided here; however, we here assume that moisture con-
tent of both jellyfish species was not different, approxi-
mately 96% as reported by Fukushi et al. (2004). These
results suggest that a diet containing larger quantities of
major nutritional components is not always suitable for
growth and survival of phyllosoma.
Phyllosomas fed exclusively on C. pacifica could not
complete metamorphosis into the nisto stage, whereas
those that fed on A. aurita could. The same result was
obtained from a duplicate experiment. Although the num-
ber of tested phyllosomas was small, it may not be inap-
propriate to conclude that A. aurita is better than
C. pacifica as food of the phyllosomas. Survival rate of
the A. aurita-fed phyllosomas was similar to that
observed in our previous experiments (Wakabayashi et al.
2012b) as well as the negative effects on the survival of
the C. pacifica-fed phyllosomas (data not shown). Consid-
ering the failure of moulting and the rapid decrease of
survival rate only in the late instars of C. pacifica-fed
phyllosomas, a minor difference between A. aurita and
C. pacifica may have critically influenced the growth and
survival of the late instar phyllosomas. We did not ana-
lyse minor nutrients such as phospholipids, sterol and
vitamins, which are all important for crustaceans (Kanaz-
awa 2000), nor the clusters of related nutritional ele-
ments. These minor but essential elements might affect
less for the survival of the late instar phyllosomas in
C. pacifica than in A. aurita. It is also possible that a
sublethal factor was accumulated in phyllosomas by feed-
ing jellyfish. Heavy metals (such as Cd, Pb, As and Hg)
included in food can have acute toxicity to lobsters
(Bryan 1971; Canli & Furness 1995), and crustacean lar-
vae are considered to be greatly sensitive to metal toxicity
(Ahsanullah & Arnott 1978). Although the composition
of heavy metals was similar between both jellyfish species,
accumulation rates of these metals into phyllosomas were
not examined in this study. Consumption rates between
A. aurita-fed and C. pacifica-fed phyllosomas should be
compared, and quantitative analyses of minor nutrients in
these two jellyfish species and accumulated sublethal fac-
tors in each instar of phyllosomas should be examined in
the future.
Similar results have been reported for two species of
mussel, Mytilus coruscus and M. galloprovincialis, both
considered suitable diets for P. japonicus phyllosomas
(Takeuchi 2010; Murakami et al. 2011). Mytilus gallopro-
vincialis-fed phyllosomas in the final instar could com-
pletely metamorphose into puerulus but not M. coruscus-
fed phyllosomas. There were few differences in biochemical
composition between the two species (Takeuchi 2010).
Takeuchi (2010) suggested that the most important charac-
teristic of the diet was the texture rather than the biochemi-
cal composition. Similarly, the differences in the texture
between the jellyfish species used in this study may have
affected phyllosoma feeding (e.g. mastication of diets and
absorption of nutrients). Further studies on the relationship
between food texture and phyllosoma development are
needed to clarify this issue.
Our results do not rule out C. pacifica as a dietary sup-
plement for phyllosoma. We have previously reported that
I. novemdentatus phyllosomas could metamorphose into
nisto if they fed on both C. pacifica and A. aurita (Waka-
bayashi et al. 2012b). Considering their ability to prey not
only on jellyfish but also on Manila clams (Takahashi &
Saisho 1978), I. novemdentatus phyllosoma appear to be an
opportunistic feeder as reported in phyllosomas of other
species (Jeffs et al. 2004). In the natural environment, phyl-
losomas are teleplanic, and this extended planktonic phase
may benefit from feeding on a wide variety of zooplankton
in the open sea where prey organisms are scarce (Jeffs et al.
2004; Sekiguchi et al. 2007; Chow et al. 2011). However,
this is not a suitable strategy for commercial production of
juvenile lobsters. These results suggested that the diets nec-
essary for phyllosomas to maintain high growth rates and
survival and to complete metamorphosis were specific and
that A. aurita was a suitable diet for I. novemdentatus phyl-
losomas.
The authors thank Dr. Hiroshi Nagai, Dr. Haruto Ishii,
and the captains and crew members of the Seiyo-maru and
Hiyodori, Tokyo University of Marine Science and Tech-
nology (TUMSAT), for collecting jellyfish. We also thank
Dr. Seiji Nagasaka, Toyo University, for his kind help with
analysing mineral composition. This study was supported
by Grants-in-Aid for JSPS Fellows to K.W. (No.
25•10983), the Programme for Promotion of Basic and
Applied Researches for Innovations in Bio-oriented
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Aquaculture Nutrition, 22; 25–33 ª 2015 John Wiley & Sons Ltd
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Appendix 1 Amino acid composition of the jellyfish species, Aurelia aurita and Chrysaora pacifica (mean  standard deviation; n = 3)
1
mg gDW % in total amino acids

Amino acid A. aurita C. pacifica A. aurita C. pacifica

Aspartic acid 3.54  0.16 b

6.10  0.39a
9.4  0.2a
8.6  0.2b
Threonine 1.90  0.08b 3.22  0.12a 5.0  0.2a 4.5  0.0b
Serine 1.76  0.11b 3.27  0.18a 4.6  0.1 4.6  0.1
Glutamic acid 5.21  0.21b 9.83  0.29a 13.8  0.3 13.9  0.1
Glycine 5.49  0.24b 11.76  0.48a 14.5  0.6b 16.6  0.3a
Alanine 2.54  0.16b 4.65  0.23a 6.7  0.3 6.6  0.1
Cystein 0.19  0.04b 0.31  0.02a 0.5  0.1 0.4  0.0
Valine 1.37  0.03b 2.57  0.03a 3.6  0.2 3.6  0.1
Methionine 0.57  0.07b 1.32  0.11a 1.5  0.1b 1.9  0.1a
Isoleucine 1.20  0.05b 2.31  0.04a 3.2  0.2 3.3  0.1
Leucine 1.69  0.80b 3.98  0.06a 4.4  2.0 5.6  0.1
Tyrosine 1.10  0.06b 2.15  0.05a 2.9  0.0b 3.0  0.1a
Phenylalanine 1.66  0.80 1.80  0.36 4.4  2.1 2.5  0.4
Lysine 2.58  0.13b 4.56  0.05a 6.8  0.1 6.4  0.3
Histidine 0.44  0.03b 0.97  0.03a 1.2  0.0b 1.4  0.1a
Arginine 2.62  0.09b 4.54  0.30a 6.9  0.2a 6.4  0.2a
Proline 3.93  0.15b 7.56  0.18a 10.4  0.3 10.7  0.2
Essential amino acids 14.02  1.20b 25.27  0.95a 36.6  1.6 34.8  0.3
Total 37.77  2.00b 70.90  2.72a 100 100

Different superscript letters on each line indicate a significant difference (P < 0.05).

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Aquaculture Nutrition, 22; 25–33 ª 2015 John Wiley & Sons Ltd
 Appendix 2 Fatty acid composition of the jellyfish species, Aurelia aurita and Chrysaora pacifica (mean  standard deviation; n = 3)
1
mg gDW % in total fatty acids

Fatty acid A. aurita C. pacifica A. aurita C. pacifica

C11:0 0.002  0.001 0.004  0.001 <0.1  0.0 <0.1  0.0

C12:0 0.019  0.009 0.031  0.015 0.4  0.1 0.4  0.1
C14:0 0.122  0.029b 0.262  0.077a 2.8  0.1b 3.7  0.1a
C15:0 0.089  0.023b 0.191  0.042a 2.1  0.3b 2.7  0.2a
C16:0 0.947  0.144 1.043  0.158 22.5  2.2a 14.9  1.8b
C16:1 0.190  0.026b 0.414  0.064a 4.5  0.7 5.9  1.0
C17:0 0.225  0.031 0.323  0.104 5.4  1.0 4.5  0.8
C17:1 0.033  0.015 0.040  0.014 0.8  0.2 0.6  0.1
C18:0 0.817  0.162b 1.247  0.161a 19.2  0.2 18.1  4.2
C18:1 0.262  0.056 0.381  0.253 6.2  0.3 5.1  3.2
C18:2 0.056  0.018 0.056  0.055 1.3  0.2 0.7  0.5
C18:3 0.038  0.019 0.059  0.028 0.9  0.3 0.8  0.4
C18:4 0.081  0.012 0.118  0.026 1.9  0.4 1.7  0.3
C19:0 0.029  0.004 0.074  0.031 0.7  0.2 1.0  0.2
C20:0 0.012  0.001 0.043  0.032 0.3  0.0 0.6  0.3
C20:1 0.026  0.002 0.080  0.078 0.6  0.2 1.0  0.8
C20:3 0.004  0.002 0.010  0.010 <0.1  0.0 0.1  0.1
C20:4 0.336  0.086 0.983  0.373a 7.8  0.4b 13.5  1.7a
C20:5 0.760  0.273 1.104  0.429 17.5  3.0 15.2  2.1
C22:1 0.004  0.001 0.028  0.009a 0.1  0.0b 0.4  0.0a
C22:6 0.055  0.015 0.232  0.080a 1.3  0.2b 3.2  0.3a
Unidentified 0.156  0.037 0.419  0.141a 3.6  0.4b 5.8  1.3a
Essential fatty acids 1.245  0.403 2.435  0.925 28.7  3.8 33.5  3.5
Polyunsaturated fatty acids 1.330  0.411 2.562  0.955 30.7  3.6 35.3  3.4
Monounsaturated fatty acids 0.516  0.094 0.944  0.363 12.2  1.0 13.0  3.0
Saturated fatty acids 2.261  0.371 3.220  0.574 53.4  3.3 45.9  5.9
Total 4.263  0.882 7.145  1.935 100 100

Different superscript letters on each line indicate a significant difference (P < 0.05).

Appendix 3 Mineral composition of the jellyfish species, Aurelia aurita and Chrysaora pacifica (mean  standard deviation; n = 3)

mg gDW 1
or lg gDW 1
% in total minerals

Mineral A. aurita C. pacifica A. aurita C. pacifica

Major minerals (mg gDW 1)

Ca 4.583  0.173a 4.164  0.061b 1.076  0.058 1.154  0.075
Na 373.109  18.987a 317.661  18.580b 87.452  0.241 87.828  0.714
Mg 32.459  1.323a 27.284  0.040b 7.612  0.177 7.559  0.409
K 16.457  0.709a 12.480  0.267b 3.861  0.162 3.459  0.231
Total 426.608  20.588a 361.589  18.325b 100 100
Trace minerals (lg gDW 1)
B 7.759  2.995 8.475  3.739
Al 0.312  0.161 0.279  0.142
V 0.016  0.004 0.024  0.009
Cr 0.186  0.171 0.357  0.322
Mn 0.159  0.037 0.116  0.049
Fe 0.599  0.313 1.427  0.605
Cu 0.136  0.049 0.210  0.035
Zn 4.004  0.947 3.644  4.062
As 1.371  0.133 1.413  0.076
Se 5.490  0.565 5.251  0.242
Mo 0.021  0.007 0.035  0.020
Cd 0.002  0.001 0.006  0.004
Hg 0.023  0.011 0.020  0.007
Pb 0.009  0.002 0.007  0.004

Different superscript letters on each line indicate a significant difference (P < 0.05).

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Aquaculture Nutrition, 22; 25–33 ª 2015 John Wiley & Sons Ltd
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