Associations of fish juveniles with rhizostome jellyfishes in the

Plankton Benthos Res 9(1): 51–56, 2014
Plankton & Benthos
Research
© The Plankton Society of Japan
Associations of fish juveniles with rhizostome jellyfishes
in the Philippines, with taxonomic remarks on a
commercially harvested species in Carigara Bay,
Leyte Island
YUSUKE KONDO1, SUSUMU OHTSUKA1,*, JUN NISHIKAWA2, EPHRIME METILLO3,
HONORIO PAGLIAWAN4, SHOZO SAWAMOTO5, MITSUYASU MORIYA2,
SHUHEI NISHIDA2 & MAKOTO URATA1
1
Takehara Marine Science Station, Setouchi Field Science Center, Graduate School of Biosphere Science, Hiroshima
University, 5–8–1 Minato-machi, Takehara, Hiroshima 725–0024, Japan
2
Atmosphere and Ocean Research Institute, University of Tokyo, 5–1–5 Kashiwanoha, Kashiwa, Chiba 277–8564, Japan
3
Department of Biological Sciences, Mindanao State University-Iligan Institute of Technology, Andres Bonifacio Ave.,
Tibanga, Iligan City, The Philippines
4
College of Fisheries and Aquatic Sciences, Western Philippines University, Puerto Princesa City, Palawan, Philippines
5
Department of Marine Biology, School of Marine Science and Technology, Tokai University, 3–20–1 Orido, Shimizu,
Shizuoka 424–8610, Japan
Received 25 October 2013; Accepted 5 December 2013
Abstract: A wide variety of fish and invertebrates are associated with marine jellyfishes, and their interactions are
diverse. We preliminarily investigated symbionts on two species of rhizostome jellyfishes collected from Leyte and
Palawan Islands, the Philippines, in August 2013. The collected jellyfishes were Lobonemoides robustus Stiasny on
both islands and Acromitus maculosus Light on Palawan Island. Lobonemoides robustus is commercially harvested
on both islands, and seems to have been previously misidentified as Lobonema smithi Mayer. The associated fish juveniles on these hosts were identified as Alepes djedaba (Forsskål) and Carangoides equula (Temminck & Schlegel).
Alepes djedaba is closely associated with many species of scyphozoan jellyfish broadly in the southeastern Asian waters. The associations of carangids with jellyfish are also reviewed.
Key words: Acromitus maculosus, carangid, Lobonemoides robustus, rhizostome, symbiosis, the Philippines
Introduction
Symbiotic interactions are essential to understand the
marine ecosystem in general as well as the grazing and microbial food webs (Ohtsuka 2006). Recently much attention has been paid to symbioses in marine pelagic realms,
partly because many kinds of parasitoids and parasites
have great negative impacts on the population dynamics of
their host zooplankters (e.g. Ianora et al. 1987, Kimmerer
& McKinnon 1990, Gómez-Gutiérrez et al. 2003, Ohtsuka
et al. 2004, 2007, 2010, Coats & Bachvaroff 2012). Some
symbionts heavily depend on their host animals for sur* Corresponding author: Susumu Ohtsuka; E-mail, ohtsuka@hiroshimau.ac.jp
vival. For example, the population dynamics of gadids are
strongly influenced by the abundance of host jellyfishes,
because their juveniles are protected from visual predators
by the hosts (Lynam & Brierley 2007).
Cnidarians are regarded to be somewhat special as
hosts, due to the presence of venomous nematocysts, making symbioses tend to be such as in the mutualistic interactions between sea anemones and anemonefish (Mebs
1994). Large jellyfish can harbor a wide variety of symbionts such as dinoflagellates, cnidarians, helminths, crustaceans, ophiuroids and fishes (Mansueti 1963, Thiel 1970,
1978, Arai 1997, Purcell & Arai 2001, Ohtsuka et al. 2009,
2010, 2011). However, their symbiotic interactions are
highly diversified, ranging from phoresy through parasitoidism to mutualism (Ohtsuka et al. 2009). We have been
52
Y. Kondo et al.
Fig. 1. Location of the two sampling sites in the Philippines in
August 2013.
intensively investigating symbioses between jellyfish and
other organisms in Asian waters since 2008, and found that
juveniles of the shrimp scad Alepes djedaba (Forsskål) is
widely associated with many species of large jellyfish in
tropical Asian waters (Ohtsuka et al. 2010). In August
2013, we surveyed symbiotic relationships between two
species of rhizostomes and their symbionts at two sites in
the Philippines. The present paper deals with the symbiotic
communities on these hosts, with a taxonomic comment on
the Malampaya jellyfish. Two species of carangids, one of
which was identified as A. djedaba, were found on the jellyfishes. Mansueti (1963), Thiel (1970, 1978) and Purcell &
Arai (2001) comprehensively reviewed symbioses between
carangids and host jellyfishes in the world oceans. In the
Philippines, Light (1914) reported the association of juveniles of Caranx with Lobonema mayeri Light (=Lobonemoides robustus?) and Rhopilema hispidum (Vanhöffen)
(as R. visayana Light). Information on such associations in
Asian waters, however, is limited. Therefore, data published since the review of Purcell & Arai (2001) and our
original results on Asian Carangidae-jellyfish associations
are compiled here.
Materials and Methods
Sampling sites were located at two stations in the Philippines: Stn 1, Carigara Bay, Leyte Island (11°19′29.08″N,
124°42′49.54″E) on 23 August 2013; Stn 2, Malampaya
Fig. 2. A. Stained canal system of Lobonemoides robustus
(No. 3), abbreviations: iac, intra-circular anastomosing canal; irc,
interrhopalar canal; rc, rhopalar canal; B. Acromitus maculosus
(No. 2), exumbrellar view, part of umbrella cut off (No. 2); C.
Carangoides equula associated with A. maculosus (No. 5 in
Table 2); D. Alepes djedaba associated with A. maculosus (No. 1
in Table 2). Scale bars=1 cm (A); 2 cm (B); 0.5 cm (C, D).
Sound, Palawan Island (10°52′09.20″N, 119°24′51.78″E) on
24 August 2013 (Fig. 1). We rented a fishing boat at each
station. Each individual jellyfish was gently scooped with a
scoop-net (diameter 50 cm; mesh size 2 mm), and put in a
plastic bag together with their symbionts. Parts of the umbrella tissues of the jellyfish and the symbionts were preserved in 99.5% ethanol for stable isotopic analyses in the
future. At Stn 2, fishermen kindly kept us an individual of
the commercially harvested jellyfish (Lobonemoides robustus Stiasny) collected from a fixed net on 23 August
2013. Water temperature and salinity were measured near
the surface with a salinometer (Model 556 MPS, YSI).
Results and Discussion
At Stn 1 (Leyte Is.) the surface water temperature and
salinity were 30.3°C and 33.6, respectively. Only L. robustus was observed as a host. Omori & Nakano (2001) suggested that the fisheries target species in Carigara Bay
might be Lobonema smithi Mayer. However, we examined
the canal system, stained by methylene blue, and came to
the conclusion that it is Lobonemoides robustus (Fig. 2A).
The genus Lobonemoides is distinguished from the genus
Lobonema in that the intra-circular anastomosing canal
system does not communicate with the inter-rhopalar canals (Fig. 2A) (e.g. Kramp 1961, Kitamura & Omori 2010).
The large-size of the umbrella, numerous exumbrellar papillae and long marginal lappets of the collected specimens
are characteristic to L. robustus (Kitamura & Omori 2010).
53
Fish-jellyfish associations in Philippines
Table 1. Associations between Lobonemoides robustus and symbionts in Carigara Bay, Leyte Island.
Host indiv.
1
2
3
4
5
Umbrella diameter (cm)
Wet weight (kg)
54
5.2
48
4.4
46
5.0
38
3.2
29
1.7
Symbionts [number of indiv.;
range of body length in mm (mean±s.d.)]
Alepes djedaba [n=112; 4.3–28.4 (10.1±4.9)]
Charybdis feriata (megalopa) (n=1; 2.2*)
Alepes djedaba [n=35; 4.3–36.2 (10.9±6.3)]
Charybdis feriata (juvenile) (n=2; 3.9 ,4.7*)
Alepes djedaba [n=41; 5.3–32.8 (14.2±7.0)]
Charybdis feriata (megalopa) (n=1; 2.5*)
Alepes djedaba [n=49; 5.0–29.7 (13.0±6.5)]
Charybdis feriata (megalopa) (n=3; 2.0, 2.0, 2.2*)
Alepes djedaba [n=10; 9.6–40.5 (19.8±8.3)]
Charybdis feriata (juvenile) (n=3; 3.9, 4.6, 4.6*)
* carapace width
Table 2. Associations between Acromitus maculosus and symbionts in Malampaya Sound, Palawan Island.
Host
indiv.
1
2
3
4
5
Fig. 3. Body length of Alepes djedaba associated with Lobonemoides robustus collected from Leyte Island (Stn 1).
Measurements from five host individuals and the prevalence and intensity of symbiont occurrence are shown in
Table 1. Five individuals of L. robustus were captured,
ranging from 29–54 cm in umbrella diameter and 1.7–
5.2 kg in wet weight. All hosts harbored juveniles of A.
djedaba, with an average of 49.4 symbionts per medusa
(prevalence=100%, intensity=49.4). The juveniles ranged
from 4.3 to 40.5 mm in body length (n=247, mean±s.d.=
11.9±6.4 mm); the most abundant class was 5–10 mm
(51.8%) (Fig. 3).
At Stn 2 (Palawan Is.) the surface water temperature and
salinity were 29.2°C and 27.5, respectively. Two species of
rhizostomes, L. robustus and Acromitus maculosus Light
(Fig. 2B) were recorded. Symbionts were found only on A.
maculosus. An individual of L. robustus was collected on
the previous day by local fishermen and kept in a meshed
plastic bag at sea but any symbionts may have escaped.
The measurements of five individuals of A. maculosus and
prevalence and intensity of symbionts are shown in Table
2. Seven individuals of A. maculosus were collected, ranging from 4.5–16 cm in umbrella diameter and 132–532 g in
6
7
Umbrella
diameter (cm)
Wet weight (g)
16
532
16
527
13
332
13
327
11
317
11
317
4.5
132
Symbionts (number of indiv.;
body length in mm)
Alepes djedaba (n=2; 12.6, 16.5)
Carangoides equula (n=2; 18.3, 20.5)
None
None
Carangoides equula (n=1; 19.0)
None
None
wet weight. Among them only three (Nos 1,2,5) were hosts
to juveniles of A. djedaba and Carangoides equula (Temminck & Schlegel) (prevalence=42.9%). The mean intensity was much lower in A. maculosus (1.7) than in L. robustus at Stn 1 (49.4). The range of the body length of A.
djedaba juveniles fell within the values reported at Stn 1.
Juveniles of three taxonomic groups of fish (Carangidae,
Gadidae, Centrolophidae) are known to associate with
large jellyfish for school formation, predation avoidance
and food collection (Dahl 1961, Mansueti 1963, Thiel 1970,
1978, Brodeur 1998, Arai 1997, Purcell & Arai 2001, Masuda et al. 2008, Masuda 2009). Mansueti (1963) listed 23
species of carangids associated with medusae, belonging to
the Hydrozoa, Scyphozoa and Cubozoa. Purcell & Arai
(2001) added 3 species of carangids (Pseudocaranx dentex
(Bloch & Schneider), Trachurus novaezelandiae Richardson, Alepes djedaba) associated with jellyfish to the list of
Mansueti (1963). The host jellyfish belong to a variety of
taxonomic groups as follows: Hydrozoa (Physalia, Velella),
54
Y. Kondo et al.
Table 3. Symbioses between jellyfish and carangids. Data mainly published since Purcell & Arai (2001). Taxonomic group: R, Rhizostomeae; S, Semaeostomeae; C, Cubozoa; H, Hydrozoa.
Carangid species
Alepes djedaba (Forsskål)
Host jellyfish (taxonomic group)
Acromitus flagellatus (Haeckel) (R)
Acromitus maculosus Light (R)
Catostylus townsendi (Mayer) (R)
Chrysaoridae sp. (S)
Lobonemoides robustus Stiasny (R)
Phyllorhiza punctata von lendenfeld (R)
Alepes djedaba (as Caranx kalla)
Carangoides equula
(Temminck & Schlegel)
Carangoides malabaricus
(Bloch & Schneider) (as
Caranx malabaricus)
Caranx bartholomaei (Cuvier)
Caranx hippos (Linnaeus)
Decapterus maruadsi
(Temminck & Schlegel)
Rhopilema hispidum (Vanhöffen) (R)
Versuriga anadyomene (Maas) (R)
Cyanea nozakii Kishinouye (S)
Acromitus maculosus Light (R)
Gnathanodon speciosus (Forsskål)
Hemicaranx zelotes Gilbert
Porpita porpita (Linnaeus)
(as Porpita pacifica) (H)
Cyanea nozakii Kishinouye (S)
Stomolophus meleagris Agassiz (R)
Stomolophus sp. (R)
Aurelia aurita s. l. (S)
Nemopilema nomurai (Kishinouye)
(as Stomolophus nomurai) (R)
Chironex yamaguchii Lewis & Bentlage (C)
Stomolophus meleagris Agassiz (R)
Trachurus japonicus
(Temminck & Schlegel)
Aequorea coerulescens (Brandt) (H)
Aurelia aurita s. l. (S)
Trachurus lathami Nichols
Trachurus novaezelandiae
Richardson
Trachurus trachurus (Linnaeus)
Chrysaora pacifica (Goette)
(as Dactylometra pacifica) (S)
Pelagia noctiluca (Forsskål)
(as Pelagia panopyra) (S)
Nemopilema nomurai (Kishinouye) (R)
Morbakka virulenta (Kishinouye) (C)
Chrysaora lactea Eschscholtz (S)
Desmonema gaudichaudi (Lesson) (S)
Aequorea forskalea Péron & Lesueur (H)
Cubozoa (Chiropsalmus, Tamoya), Semaeostomeae (Aurelia, Chrysaora, Cyanea, Desmonema, Drymonema, Pelagia), and Rhizostomeae (Acromitus, Catostylus, Cephea,
Cotylorhiza, Mastigias, Mastigietta, Netrostoma, Phyllorhiza, Pseudorhiza, Rhizostoma, Rhopilema, Stomolophus,
Thysanostoma) (Mansueti 1963, Thiel 1970, 1978, Purcell
& Arai 2001, López & Rodriguez 2008, Cevik et al. 2011)
(see Table 3 also). Thiel (1970) pointed out that fish juveniles prefer rhizostomes to semaeostomes as hosts. Mansueti (1963) reported that these associations are only temporary, rarely extending for more than a few months, and
suggested that these may be classified as commensalism or
parasitism.
Data mainly published after these reviews and some of
our original observations on such associations are compiled in Table 3. In Asian waters, little data exists other
Locality
Source
Gulf of Thailand
Palawan Island
Gulf of Thailand
Gulf of Thailand
Gulf of Thailand,
Leyte Island
Mediterranean,
Turkey
Gulf of Thailand
Java Sea
Hong Kong
Palawan Island
Kondo et al. (unpublished)
Present study
Kondo et al. (unpublished)
Kondo et al. (unpublished)
Present study, Kondo et al.
(unpublished)
Cevik et al. (2011)
Ohtsuka et al. (2010)
Ohtsuka et al. (2010)
Morton (1972)
Present study
Karwar, India
Noble (1963)
Hong Kong
North Carolina
Gulf of Mexico
Genkai-nada, Japan
Genkai-nada, Japan
Morton (1972)
Rountree (1983)
Gunter (1938)
Shojima (1962)
Shojima (1962)
Okinawa, Japan
Gulf of California
Seto Inland Sea
Seto Inland Sea,
Japan Sea
Seto Inland Sea
Okada et al. (unpublished)
López-Martínez &
Rodríguez-Romero (2008)
Shojima (1962)
Shojima (1962),
Masuda et al. (2009)
Shojima (1962)
Seto Inland Sea
Shojima (1962)
Japan Sea
Seto Inland Sea
southern Brazil
northeastern New
Zealand
north-west Spain
Masuda et al. (2009)
Okada et al. (unpublished)
Bonaldo et al. (2004)
Kingsford et al. (2000)
Villegas-Ríos (2009)
than some studies on Alepes djedaba (Ohtsuka et al. 2010)
and Trachurus japonicus (Temminck & Schlegel) (Masuda
2009, Masuda et al. 2008). Alepes djedaba and T. japonicas have been observed on a range of tropical and subtropical Asian jellyfishes, respectively. Masuda (2009) clearly
revealed ontogenetic changes in the interactions between
host jellyfish and the development of T. japonicus: the host
jellyfish is suggested to play roles as a site for school formation for the early stages (mean standard length 11 mm),
as a hiding place for the next stages (19 mm), and finally as
a food source for the 38–50 mm stages. Alepes djedaba exhibited habitat segregation on the coasts of the Gulf of
Thailand where two hosts, L. robustus and Rhopilema hispidum, sympatrically occurred: small juveniles (mean
body length ca. 13 mm) preferred L. robustus, while larger
juveniles (20 mm) favoured R. hispidum (Kondo et al., per-
Fish-jellyfish associations in Philippines
sonal observation). Differences in swimming speed and
toxicity between these two hosts might be the cause of this
phenomenon, implying ontogenetic changes in the jellyfish-fish interaction as seen between T. japonicus and jellyfish (Masuda et al. 2008, Masuda 2009). Adaptive strategies of these associated carangids to jellyfish toxins should
be clarified in the future, as seen in sea anemone-anemonefish interactions (e.g. Dahl 1961, Mebs 1994). It is
worth noting that the association of the family Carangidae
with jellyfish seems to be confined mainly to the tribe Carangini (see Reed et al. 2002).
Megalopa larvae and juveniles of the Christ crab Charybdis feriata (Linnaeus), the former of which was identified based on Fielder et al. (1984), were found on L. robustus collected from Stn 1. The prevalence and intensity were
100% and 2.0, respectively (Table 1). This association was
also observed in L. robustus and R. hispidum found in the
Gulf of Thailand from July to October (Kondo et al., personal observation), suggesting that it directly settles on jellyfishes and stays on them until the death of the host.
Symbiosis of the ophiuroid Ophiocnemis marmorata
(Lamarck) on the rhizostome Rhopilema esculentum
Kishinouye has been reported from Palawan Island, the
Philippines, in March 2004 (Fujita & Namikawa 2006).
However, this association was not observed in the present
study. Since this ophiuroid is reported to be abundant on L.
robustus in the Gulf of Thailand in summer and fall and on
the western coast of the Malayan Peninsula in March (Ohtsuka et al. 2010), on L. robustus–O. marmorata association
may be found also in the Philippines in spring but not in
summer.
Acknowledgments
We express our sincere thanks to two anonymous reviewers for their constructive comments on the manuscript. The present study was partially supported by
grants-in-aid from the Japan Society of Promotion of Science (JSPS), awarded to SO (No. 25304031), to SN (No.
22380108) and to JN (No. 23405031), and by a grant of the
Asian Core Program of the JSPS.
References
Arai MN (1997) A Functional Biology of Scyphozoa. Chapman
& Hall, London, 316 pp.
Bonaldo RM, Krajewski JP, Sazima I (2004) Does the association of young fishes with jellyfishes protect from predation? A
report on a failure case due to damage to the jellyfish. Neotrop
Ichthyol 2: 103–105.
Brodeur RD (1998) In situ observations of the association between juvenile fishes and scyphomedusae in the Bering Sea.
Mar Ecol Prog Ser 163: 11–20.
Cevik C, Derici OB, Cevik F, Cavas L (2011) First record of
Phyllorhiza punctata von Lendenfeld, 1884 (Scyphozoa: Rhizostomeae: Mastigiidae) from Turkey. Aquat Invasions 6, Sup-
55
plement 1: S27–S28.
Coats DW, Bachvaroff TR (2012) Parasites of tintinnids. In: The
Biology and Ecology of Tintinnid Ciliates (eds Dolan JR,
Montagnes DJS, Agatha S, Coats DW, Stoecker DK). A John
Wiley & Sons, Ltd., Oxford, pp. 145–170.
Dahl E (1961) The association between young whiting, Gadus
merlangus, and the jelly-fish Cyanea capillata. Sarsia 3: 47–55.
Fielder DR, Greenwood JG & Campbell G (1984) The megalopa
of Charybdis feriata (Linnaeus) with additions to the zoeal
larvae descriptions (Decapoda, Portunidae). Crustaceana 46:
160–165.
Fujita T, Namikawa H (2006) New observations of Ophiocnemis
marmorata (Echinodermata: Ophiuroidea) associated with
Rhopilema esculentum (Cnidaria: Scyphozoa: Rhizostomeae)
in the Philippines and Japan. Mem Natn Sci Mus Tokyo 44:
31–37.
Gómez-Gutiérrez J, Peterson WT, De Robertis A, Brodeur RD
(2003) Mass mortality of krill caused by parasitoid ciliates.
Science 301: 339.
Gunter G (1938) Seasonal variations in abundance of certain estuarine and marine fishes in Louisiana, with particular reference to life histories. Ecol Monogr 8: 313–346.
Ianora A, Mazzocchi MG, Scotto di Carlo B (1987) Impact of
parasitism and intersexuality on Mediterranean populations of
Paracalanus parvus (Copepoda: Calanoida). Dis Aquat Org 3:
29–36.
Kimmerer WJ, McKinnon AD (1990) High mortality in a copepod population caused by a parasitic dinoflagellate. Mar Biol
107: 449–452.
Kingsford MJ, Pitt KA, Gillanders BM (2000) Management of
jellyfish fisheries, with special reference to the order Rhizostomeae. Oceanogr Mar Biol Annu Rev 38: 85–156.
Kitamura M, Omori M (2010) Synopsis of edible jellyfishes collected from Southeast Asia, with notes on jellyfish fisheries.
Plankton Benthos Res 5: 106–118.
Kramp PL (1961) Synopsis of the medusae of the world. J Mar
Biol Ass UK 40: 1–469.
Light SF (1914) Some Philippine Scyphomedusae, including two
new genera, five new species, and one new variety. Philipp J
Sci 9: 195–231.
Lynam CP, Brierley AS (2007) Enhanced survival of 0-group gadoid fish under jellyfish umbrellas. Mar Biol 150: 1397–1401.
López-Martínez J, Rodríguez-Romero J (2008) Primer registro
de la asociación del jurelillo negro Hemicaranx zelotes Gilbert
(Pisces: Carangidae) con la medusa bala de cañón Stomolophus meleagris Agassiz (Scyphozoa: Rhizostomatidae) en
Bahía de Kino, Golfo de California. Hidrobiológica 18: 161–
164.
Mansueti R (1963) Symbiotic behavior between small fishes and
jellyfishes, with new data on that between the stromateid, Peprilus alepidotus, and the Scyphomedusa, Chrysaora quinquecirrha. Copeia 1963: 40–80.
Masuda R (2009) Ontogenetic changes in the ecological function
of the association behavior between jack mackerel Trachurus
japonicus and jellyfish. Hydrobiologia 616: 269–277.
Masuda R, Yamashita Y, Matsuyama M (2008) Jack mackerel
Trachurus japonicus juveniles use jellyfish for predator avoidance and as a prey collector. Fish Sci 74: 276–284.
56
Y. Kondo et al.
Mebs D (1994) Anemonefish symbiosis: vulnerability and resistance of fish to the toxin of the sea anemone. Toxicon 32:
1059–1068.
Morton B (1972) The occurrence of Caranx kalla and C. malabaricus in association with the jellyfish, Cyanea nozakii. Copeia 1972: 873–875.
Noble A (1963) On the association between the fish, Caranx malabaricus Cuv. & Val. and the siphonophore, Porpita pacifica
Lesson. J Mar Biol Ass India 5: 142–143.
Ohtsuka S (2006) Importance of parasito-planktology for precise
understanding the marine ecosystem. Bull Plankton Soc Japan
53: 7–13. (in Japanese)
Ohtsuka S, Harada S, Shimomura M, Boxshall GA, Yoshizaki R,
Ueno D, Nitta Y, Iwasaki S, Okawachi H, Sakakihara T (2007)
Temporal partitioning: dynamics of alternating occupancy of a
host microhabitat by two different crustacean parasites. Mar
Ecol Prog Ser 348: 261–272.
Ohtsuka S, Hora M, Suzaki T, Arikawa M, Omura G, Yamada K
(2004) Morphology and host-specificity of the apostome ciliate
Vampyrophrya pelagica infecting pelagic copepods in the
Seto Inland Sea, Japan. Mar Ecol Prog Ser 282: 129–142.
Ohtsuka S, Koike K, Lindsay D, Nishikawa J, Miyake H, Kawahara M, Mulyadi, Mujiono N, Hiromi J, Komatsu H (2009)
Symbionts of marine medusae and ctenophores. Plankton Bentos Res 4: 1–13.
Ohtsuka S, Kondo Y, Sakai Y, Shimazu T, Shimomura M, Komai
T, Yanagi K, Fujita T, Nishikawa J, Miyake H et al. (2010) Insitu observations of symbionts on medusae occurring in Japan,
Thailand, Indonesia and Malaysia. Bull Hiroshima Univ Mus
2: 9–18.
Ohtsuka S, Kondo Y, Iwasaki S, Hayashi K (2011) Caridean
shrimps associated with the rhizostome jellyfish Netrostoma
setouchiana in the central part of the Seto Inland Sea, Japan.
Bull Hiroshima Univ Mus 3: 1–6. (in Japanese with English
abstract)
Omori M, Nakano E (2001) Jellyfish fisheries in southeast Asia.
Hydrobiologia 451: 19–26.
Purcell JE, Arai MN (2001) Interactions of pelagic cnidarians
and ctenophores with fish: a review. Hydrobiologia 451: 27–44.
Reed DL, Carpenter KE, deGravelle MJ (2002) Molecular systematics of the Jacks (Perciformes: Carangidae) based on mitochondrial cytochrome b sequences using parsimony, likelihood, and Bayesian approaches. Mol Phyl Evol 23: 513–524.
Rountree RA (1983) The ecology of Stomolophus meleagris, the
cannon ball jellyfish, and its symbionts, with special emphasis
on behavior. PhD thesis, University of North Carolina.
Shojima Y (1962) On the postlarvae and juveniles of carangid
fishes collected together with the jellyfishes. Contrib Seikai
Regional Fish Res 147: 48–58. (in Japanese with English abstract)
Thiel ME (1970) Das Zusammenleben von Jung- und Kleinfischen
mit Rhizostomeen (Scyphomedusae). Ber Deutsch Wiss Komm
Meeresforsch 21: 444–473.
Thiel ME (1978) Das Zusammenleben von Jung- und Kleinfischen
mit Semaeostomeen (Scyphomedusae). Mitt Hamb Zool Mus
Inst 75: 19–47.
Villegas-Ríos D (2009) Associative behaviour between Trachurus trachurus (Teleostei: Carangidae) and the hydromedusae Aequorea forskalea: a new relationship reported from
Galician waters. Mar Biodiv Rec 2: 1–3.