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JELLYFISH FISHERIES OF THE WORLD
by
Lucas Brotz
B.Sc., The University of British Columbia, 2000
M.Sc., The University of British Columbia, 2011
A DISSERTATION SUBMITTED IN PARTIAL FULFILLMENT OF
THE REQUIREMENTS FOR THE DEGREE OF
DOCTOR OF PHILOSOPHY
in
The Faculty of Graduate and Postdoctoral Studies
(Zoology)
THE UNIVERSITY OF BRITISH COLUMBIA
(Vancouver)
December 2016
© Lucas Brotz, 2016
ii
Abstract
Fisheries for jellyfish (primarily scyphomedusae) have a long history in Asia, where
people have been catching and processing jellyfish as food for centuries. More
recently, jellyfish fisheries have expanded to the Western Hemisphere, often driven
by demand from buyers in Asia as well as collapses of more traditional local finfish
and shellfish stocks. Despite this history and continued expansion, jellyfish fisheries
are understudied, and relevant information is sparse and disaggregated. Catches of
jellyfish are often not reported explicitly, with countries including them in fisheries
statistics as “miscellaneous invertebrates” or not at all. Research and management of
jellyfish fisheries is scant to nonexistent. Processing technologies for edible jellyfish
have not advanced, and present major concerns for environmental and human
health. Presented here is the first global assessment of jellyfish fisheries, including
identification of countries that catch jellyfish, as well as which species are targeted.
A global catch reconstruction is performed for jellyfish landings from 1950 to 2013,
as well as an estimate of mean contemporary catches. Results reveal that all
investigated aspects of jellyfish fisheries have been underestimated, including the
number of fishing countries, the number of targeted species, and the magnitudes of
catches. Contemporary global landings of jellyfish are at least 750,000 tonnes
annually, more than double previous estimates. Jellyfish have historically been
understudied, resulting in the current dearth of knowledge on population dynamics
and jellyfish fishery management. However, many of the tools used in traditional
fisheries science, such as length-frequency analysis, can be applied to jellyfish, as
demonstrated herein. Research priorities are identified, along with a prospective
outlook on the future of jellyfish fisheries.
iii
Preface
This dissertation represents a synthesis of existing information and original work
that was led by the author, and includes information contributed from numerous
collaborators. The work is presented here in its entirety in order to provide the entire
global overview. However, selected material has been extracted as contributions to 6
publications, including 2 peer-reviewed journal articles, 2 book chapters, and 2
report sections.
Specifically, selected information formed the basis of a peer-reviewed journal article
[Brotz, L., A. Schiariti, J. López-Martínez, J. Álvarez-Tello, Y.-H.P. Hsieh, R.P. Jones,
J. Quiñones, Z. Dong, A.C. Morandini, M. Preciado, E. Laaz, & H. Mianzan in press.
Jellyfish fisheries in the Americas: origin, state of the art, and perspectives on new
fishing grounds. Reviews in Fish Biology and Fisheries doi:10.1007/s11160-016-9445-y].
Information from Chapters 1-3 was used as the basis for a book chapter [Brotz, L.
2016. Jellyfish fisheries – a global assessment, pp. 110-124 in D. Pauly & D. Zeller
(eds.) Global Atlas of Marine Fisheries: A Critical Appraisal of Catches and
Ecosystem Impacts. Island Press, Washington, D.C., U.S.A.]; which also had an
abbreviated version that was published in a report [Brotz, L. 2014. Jellyfish fisheries
– a global assessment, pp. 77-81 in D. Pauly & D. Zeller (eds.) So long, and thanks for
all the fish: The Sea Around Us, 1999-2014 – A Fifteen-Year Retrospective, a report to
the Pew Charitable Trusts, University of British Columbia].
Material gleaned from this dissertation was also combined with additional
contributions and was published as a peer-reviewed book chapter [Brotz, L. &
D. Pauly in press. Studying jellyfish fisheries: toward accurate national catch reports
and appropriate methods for stock assessments in G.L. Mariottini (ed.) Jellyfish:
Ecology, Distribution Patterns and Human Interactions. Nova Publishers,
Hauppauge, New York, U.S.A.].
A summary of relevant information, especially using selections from Chapter 3, will
also be published as a report section [Brotz, L. in press. Jellyfish Fisheries in
R. Brodeur & S. Uye (eds.) Jellyfish blooms around the North Pacific Rim: Causes
and consequences, PICES Scientific Report No. 51].
iv
Finally, some conclusions and opinions related to this dissertation, especially from
Chapters 2 and 5, were also expressed in another peer-reviewed journal article
[Gibbons, M.J., F. Boero, & L. Brotz 2016. We should not assume that fishing jellyfish
will solve our jellyfish problem. ICES Journal of Marine Science 73(4): 1012-1018].
v
Table of contents
Abstract ................................................................................................................................... ii
Preface ..................................................................................................................................... iii
Table of contents .................................................................................................................... v
List of tables .......................................................................................................................... vii
List of figures ....................................................................................................................... viii
Acknowledgements .............................................................................................................. ix
1. Introduction .................................................................................................................... 1
2. From ocean to plate – aspects of jellyfish fisheries ................................................... 5
2.1. Ecology .................................................................................................................... 5
2.1.1. Target species ................................................................................................. 5
2.1.2. Life cycle ........................................................................................................ 10
2.2. Fisheries ................................................................................................................. 14
2.2.1. Fishing for jellyfish ...................................................................................... 14
2.2.2. Ecological implications................................................................................ 15
2.3. Use .......................................................................................................................... 21
2.3.1. Processing of edible jellyfish ...................................................................... 23
2.3.2. The edible product ....................................................................................... 28
2.3.3. Health effects of jellyfish consumption .................................................... 32
2.4. Management ......................................................................................................... 35
3. Reconstructing the global catch ................................................................................. 40
3.1. Methods ................................................................................................................. 42
3.1.1. Catch reconstruction .................................................................................... 42
3.1.2. Scaling factor ................................................................................................. 43
3.2. Results .................................................................................................................... 45
3.2.1. Estimating the contemporary global catch ............................................... 47
3.3. Countries fishing jellyfish ................................................................................... 53
3.3.1. Australia ........................................................................................................ 53
3.3.2. Bahrain ........................................................................................................... 57
3.3.3. Canada ........................................................................................................... 60
3.3.4. China .............................................................................................................. 62
3.3.5. Ecuador .......................................................................................................... 67
3.3.6. Honduras....................................................................................................... 68
3.3.7. India ............................................................................................................... 68
3.3.8. Indonesia ....................................................................................................... 73
3.3.9. Iran ................................................................................................................. 76
vi
3.3.10. Japan .............................................................................................................. 76
3.3.11. Korea (Republic of) ...................................................................................... 78
3.3.12. Malaysia ........................................................................................................ 79
3.3.13. Mexico ............................................................................................................ 82
3.3.14. Myanmar ....................................................................................................... 83
3.3.15. Nicaragua ...................................................................................................... 84
3.3.16. Pakistan ......................................................................................................... 85
3.3.17. Philippines .................................................................................................... 86
3.3.18. Russian Federation ...................................................................................... 87
3.3.19. Sri Lanka ........................................................................................................ 88
3.3.20. Thailand ......................................................................................................... 89
3.3.21. Turkey ............................................................................................................ 91
3.3.22. U.S.A. ............................................................................................................. 92
3.3.23. Vietnam ......................................................................................................... 97
3.3.24. Other countries ............................................................................................. 99
4. Growth of jellyfish in Mexico’s Gulf of California ................................................ 103
4.1. Length-frequency analysis and the ELEFAN software ................................ 104
4.2. Sampling of Stomolophus meleagris ................................................................... 105
4.3. Analysis and results ........................................................................................... 109
5. Conclusions ................................................................................................................. 117
5.1. Research priorities ............................................................................................. 119
5.2. Sustainability and the future of jellyfish fisheries ......................................... 120
References ........................................................................................................................... 127
Appendix A – Reconstructed jellyfish landings by country, 1950-2013 .................... 166
Appendix B – Supporting publications .......................................................................... 170
vii
List of tables
Table 1. Edible species of jellyfish in the Order Rhizostomeae ....................................... 8
Table 2. Non-rhizostome edible species of jellyfish (none at commercial scale) .......... 9
Table 3. Uses for jellyfish other than as food for humans .............................................. 23
Table 4. Countries known to fish jellyfish for human consumption ............................ 46
Table 5. Estimated contemporary annual jellyfish landings (2004-2013 mean) .......... 50
viii
List of figures
Figure 1. Life cycle of the cannonball jellyfish Stomolophus meleagris; based on
Calder (1982) ................................................................................................................. 11
Figure 2. Estimated global landings for two primary species in China and all
species for other countries ........................................................................................... 48
Figure 3. Locations of jellyfish fisheries in Southeast Asia; dark blue indicates
shelf (< 200 m); note that there are likely many additional locations that have
yet to be documented ................................................................................................... 49
Figure 4. Estimated global contemporary annual jellyfish landings (2004-2013)
and reported FAO catches (1950-2013) ...................................................................... 51
Figure 5. Jellyfish fisheries around the world; red circles indicate magnitude
category of catch in tonnes (see legend); circles are only approximately
representative of catch locations ................................................................................ 52
Figure 6. Major FAO areas .................................................................................................. 58
Figure 7. Map of the Gulf of California in Mexico; dark blue indicates shelf
(< 200 m); black circle indicates approximate location of Guaymas and Las
Guásimas coastal lagoon ........................................................................................... 108
Figure 8. Weatherall Plot using ELEFAN with L∞ = 18.4 cm ....................................... 110
Figure 9. L/F data plot for S. meleagris sampled Dec. 2015 - April 2016 appearing
to show two distinct cohorts of medusae ............................................................... 111
Figure 10. Response surface for the goodness-of-fit estimator using ELEFAN,
suggesting an estimate of K = 3.7 year-1 .................................................................. 112
Figure 11. Growth curve fitting using ELEFAN for S. meleagris sampled Dec. 2015
– April 2016; L∞ = 18.4 cm; K = 3.7 year-1.................................................................. 113
Figure 12. Length-weight relationship for S. meleagris ................................................. 114
Figure 13. Auximetric plot of various fish (small circles) and jellyfish species
including Stomolophus meleagris; based on Palomares & Pauly (2009) ............... 116
ix
Acknowledgements
I owe a debt of gratitude to my supervisor, Dr. Daniel Pauly, who has been a
continuous source of inspiration and always provided me with sage advice. I would
also like to thank my committee members, Drs. Evgeny Pakhomov, William W.L.
Cheung, and Ussif Rashid Sumaila, who were always there to give me guidance and
support. Special thanks are owed to current and former members of the Sea Around
Us, especially Christopher Hoornaert and Sarah Popov, who assisted with the
creation of maps, as well as Kyrstn Zylich and Eric Sy, who always dropped
everything (that was undoubtedly more important) to answer my questions. Thanks
also to Dr. Wilf Swartz for help with translation, Roberto Licandeo for helping to
enlighten me on the intricacies of R, and Dr. Maria (Deng) Palomares for assistance
with ELEFAN. Tanvi Vaidyanathan kindly provided information on jellyfish
fisheries in India, and Dr. Gabriel Reygondeau assisted with data queries. Thanks to
Dr. Antonella Leone, Dr. Stefano Piraino, and chef Gennaro Esposito for introducing
me to alternative preparations of edible jellyfish. I am indebted to Dr. Shin-ichi Uye
for hosting me in Japan and introducing me to the jellyfish fishery in the Ariake Sea.
Although insufficient, muchas gracias go out to Drs. Miguel Cisneros Mata, Gabriela
Montemayor, and Andrés Cisneros Montemayor for their help, data, and for hosting
me in Mexico. Their hospitality and kindness is unlimited. I would also like to give a
special thank you to all of the colleagues that contributed information on jellyfish
fisheries, especially Dr. Agustín Schiariti for being a valuable resource as well as
being a friend. And of course, many thanks to my family and friends for their
unrelenting encouragement and support.
1
1. Introduction
Jellyfish (herein referring to members of the Phylum Cnidaria, Subphylum
Medusozoa with a pelagic phase, primarily in the Class Scyphozoa) are notorious
for interfering with human activities and industries including fisheries, aquaculture,
and tourism, as well as power generation and desalination (Purcell et al. 2007; Lucas
et al. 2014). However, jellyfish (or ‘medusae’) are also considered traditional cuisine
in China, where they have been eaten for more than 1,700 years (Omori & Nakano
2001; Li & Hsieh 2004). Eating jellyfish continues to be very popular in China,
evidenced by the wide availability of not only edible jellyfish, but also the many
available imitation or artificial jellyfish products that are primarily made from
brown seaweeds (You et al. 2007). Consumption of jellyfish is also popular in Asian
countries other than China, such as Japan, Malaysia, Korea, Taiwan, and Singapore
(Kingsford et al. 2000; Hsieh et al. 2001; Omori & Nakano 2001). Interestingly,
cnidarians were also consumed in ancient Rome, as indicated by the Latin cookbook
Apicius (Vehling 1977), but whether the “sea nettles” referred to in the text are
indeed jellyfish or rather sessile anemones remains unresolved. Regardless, it is
amusing to note that the recipe suggests that when the cnidarians are served atop of
eggs in a type of omelette, “no one at the table will know what they are eating”
(Grocock & Grainger 2006).
2
Jellyfish fisheries are typically characterized by large interannual fluctuations in
abundance and biomass, as well as short fishing seasons of usually less than a few
months (Omori 1978; Omori & Nakano 2001). It has been suggested that rapid
changes in exploitable biomass of jellyfish are more of a concern than for any other
fishery (Kingsford et al. 2000), not least because the ecology is poorly understood.
These circumstances can cause instability of jellyfish fisheries and may prevent
fishers, stakeholders, and policy-makers from supporting development. A
contributing factor is that the species being targeted have complex life cycles and are
historically understudied organisms, making it difficult to model and predict
population dynamics and responses to fishing pressure. As a result, information and
research is lagging far behind the expansion of jellyfish fisheries, with potentially
negative consequences for both stakeholders and ecosystems. In the Western
Hemisphere, jellyfish have undergone a dramatic transition in some locations,
shifting from being a nuisance to a valuable fishery resource. In most cases, this
transition appears to have been preceded by declines of more traditional fisheries
resources such as finfish and shrimp. It remains uncertain if this transition should be
celebrated as an example of adaptability, or if it is yet another warning sign that we
are rapidly fishing down the food web (Pauly et al. 1998). The examination of
3
jellyfish fisheries around the globe presented herein will help to elucidate some of
the questions raised by the rapid development of jellyfish fisheries.
Despite the long history of jellyfish consumption, information on jellyfish fisheries is
sparse and disaggregated. In the scant available literature on the subject, there is
often conflicting information about the number of targeted species, as well as which
countries fish for jellyfish. Even basic data, such as the magnitudes of catches, differs
between sources. It is widely recognized that catch statistics are crucial for fisheries
management (Pauly 1998; Jennings et al. 2001; Pauly 2016). Clearly then, it follows
that such statistics should be as accurate as possible. The primary organization
compiling national and global fishery catch statistics is the Food and Agriculture
Organization of the United Nations (FAO). However, FAO is entirely dependent on
what individual countries report, with relatively little in the way of incentives or
enforcement to ensure accurate reporting. As such, catch statistics are often
underreported (Pauly & Zeller 2016a), or may even be over-reported in rare cases
(e.g., Watson & Pauly 2001). For jellyfish, the situation is even more confounded, as
many countries do not report their catches of jellyfish explicitly, including them
either as “miscellaneous marine invertebrates” or not at all. Fisheries for jellyfish
have heretofore not been reviewed on a global scale. It may be argued that the
review by Kingsford et al. (2000) is an exception; however, that review, while
4
excellent, focused primarily on the management of jellyfish fisheries and the
emergent fishery in Australia. While catch values from other countries were
included, they were simply quoted from FAO statistics. The present study is the first
global examination of jellyfish fisheries and their associated catches.
As jellyfish populations tend to exhibit dramatic interannual variation in abundance
and biomass even when they are not subject to exploitation (Brotz 2011), developing
management programs with a goal of sustainable jellyfish catches is sure to remain a
challenge. Nonetheless, many of the tools available to traditional fisheries science,
such as length-frequency analysis, can be adapted and applied to jellyfish. As
jellyfish fisheries continue to expand around the globe, the development of such
techniques will be important for informing management decisions.
5
2. From ocean to plate – aspects of jellyfish fisheries
Jellyfish may be targeted for a number of reasons, including for use in agriculture,
materials science, and pharmaceuticals; but by far the largest use of jellyfish is as
food for humans. A number of species are consumed, mostly from the scyphozoan
Order Rhizostomeae. Jellyfish have unique life cycles, adding to the complexity of
managing jellyfish fisheries. Consumption is primarily in Asia, whereas discussion
of eating jellyfish in the Western Hemisphere is typically met with reactions ranging
from surprise to disgust. While jellyfish fisheries have expanded throughout the
world in recent decades, the nascent fisheries are primarily serving growing Asian
markets. Consistent with Traditional Chinese Medicine, some research has
demonstrated the positive health effects associated with consuming edible jellyfish.
However, the chemicals used in jellyfish processing have also been shown to be
detrimental to human health.
2.1. Ecology
2.1.1. Target species
With the exception of Mexico, currently all catches of jellyfish reported by FAO are
classified as “Rhopilema spp.”, which is incorrect in many cases. The number of
identified species of edible jellyfish worldwide is unclear, and is typically
6
underestimated (e.g., Omori 1981; Sloan 1986; Hsieh & Rudloe 1994; Omori &
Nakano 2001; Armani et al. 2013), due in part to the taxonomy of edible jellyfish
being confused (Omori & Kitamura 2004; Kitamura & Omori 2010). A synthesis of
available information reveals approximately 35 species of jellyfish that have been
documented as being consumed by humans. The majority of these, including all
jellyfish fisheries operating at commercial scales, are from the scyphozoan Order
Rhizostomeae (Table 1). These jellyfish are typically large, with relatively tough and
rigid tissues. All members lack marginal tentacles, and instead have prominent oral
arms (sometimes called ‘legs’ or incorrectly referred to as ‘tentacles’). Rhopilema
esculentum is the most valuable species and is currently the choice for hatchery and
aquaculture operations in China (see 3.3.4 China). The giant jellyfish, Nemopilema
nomurai, is also widely exploited in East Asia, in much larger quantities than have
been reported until recently. Other rhizostomes may also be edible, as the diverse
order contains 92 extant species (Daly et al. 2007).
There are reports that humans also consume other types of jellyfish (Table 2). This
includes scyphozoans from the Order Semaeostomeae, such as Aurelia, Chrysaora,
and Cyanea; however, in most cases it appears that these species are less desirable
and are not currently targeted at commercial scales. There is also limited
information to suggest that cubozoans are consumed in some regions. Shih (1977)
7
reported that the people of the “Tawara” consume freshly caught or sun-dried
Tamoya sp. after boiling them. As the author refers to “Natives of Tawara in the
Pacific Ocean”, it is assumed the author was referring to Pacific atoll of Tarawa,
Kiribati. Purcell et al. (2007) noted that aboriginal peoples in Taitung, Taiwan also
eat cubomedusae.
Rhizostome jellyfish appear to be preferred for consumption as they produce the
desired crunchy and crispy texture that is characteristic of edible jellyfish products.
However, the documentation of other species, such as semaeostomes and
cubomedusae are testament to the fact that other types of jellyfish are indeed
“edible,” if not preferred. As such, the development of fisheries for the dozens of
non-rhizostome scyphozoans may be possible in the future, albeit with economic
challenges. Of course, jellyfish may also be targeted for a number of reasons other
than as for food for humans, thus increasing the total number of fished jellyfish
species (see 2.3 Use).
8
Table 1. Edible species of jellyfish in the Order Rhizostomeae
Family
Species
Country
Reference
Cassiopeidae
Cassiopea ndrosia
Philippines
Omori & Nakano (2001)
Catostylidae
Acromitus hardenbergi
Malaysia; Indonesia; Thailand
Nishikawa et al. (2009); Kitamura & Omori (2010)
Catostylus mosaicus
Australia
Fisheries Victoria (2006)
Catostylus perezi
Pakistan
Muhammed & Sultana (2008); Gul & Morandini (2013)
Catostylus tagi
Portugal
Amaral et al. (2016)
Crambione mastigophora
Indonesia
Omori & Nakano (2001); Kitamura & Omori (2010)
Crambionella annandalei
Myanmar
Kitamura & Omori (2010)
Crambionella orsini1
India; Sri Lanka
Kuthalingam et al. (1989); NARA (2010)
Crambionella helmbiru
Indonesia
Nishikawa et al. (2015)
Crambionella stuhlmanni
India
Kuthalingam et al. (1989); Mohan et al. (2011)
Cepheidae
Cephea cephea
Thailand
Omori & Nakano (2001)
Cotylorhiza tuberculata
Italy
pers. obs.
Lobonematidae
Lobonema smithi
China; India; Malaysia;
Philippines
Kingsford et al. (2000); Hong (2002); Murugan & Durgekar (2008);
Nishikawa et al. (2009)
Lobonemoides gracilis2
China; Philippines
Omori (1981); Hong (2002); Kitamura & Omori (2010)
Lobonemoides robustus
Indonesia; Myanmar; Vietnam;
Thailand; Philippines
Kitamura & Omori (2010)
Lychnorhizidae
Lychnorhiza lucerna
Argentina
Schiariti (2008)
Mastigiidae
Mastigias sp.
Thailand
Sloan & Gunn (1985)
Phyllorhiza punctata
Australia
Coleman et al. (1990); Kailola et al. (1993)
Rhizostomatidae
Rhizostoma octopus3
United Kingdom
Elliott et al. (2016)
Rhizostoma pulmo4
Turkey
Ozer & Celikkale (2001)
Rhizostoma sp.
India
Chidambaram (1984)
Rhopilema esculentum
China; India; Indonesia; Japan;
Korea; Malaysia; Thailand;
Russia; Vietnam
Omori (1978); Morikawa (1984); Sloan (1986); Kingsford et al.
(2000); Omori & Kitamura (2004); Yakovlev et al. (2005); Nishikawa
et al. (2008); Panda & Madhu (2009); Ullah et al. (2015)
Rhopilema hispidum
China; Indo.; Japan; Malaysia;
Pakistan; Thailand; Vietnam
Kingsford et al. (2000); Omori & Kitamura (2004); Muhammed &
Sultana (2008); Kitamura & Omori (2010); Gul & Morandini (2015)
Rhopilema nomadica
Turkey
Kingsford et al. (2000)
Rhopilema verrilli
U.S.A.
Rudloe (1992); Kingsford et al. (2000)
Rhizostomatidae?
(suspected unique sp.)
Indonesia; Malaysia
Omori & Nakano (2001); Kitamura & Omori (2010)
Stomolophidae
Nemopilema nomurai
China; Japan; Korea
Omori (1978); Morikawa (1984); Li et al. (2014)
Stomolophus meleagris
U.S.A.; Mexico; Nicaragua;
Ecuador; Honduras
Hsieh et al. (2001); López-Martinez & Álvarez-Tello (2013); this
study
1 May be a synonym of C. annandali (see Kitamura & Omori 2010); 2 May be a synonym of L. robustus (see Kitamura & Omori 2010); 3 edibility unconfirmed,
currently targeted for collagen; 4 also targeted for collagen in France
9
Table 2. Non-rhizostome edible species of jellyfish (none at commercial scale)
Class
Order
Family
Species
Country
Reference
Cubozoa
Carybdeida
Carybdeidae
Carybdea rastoni
Taiwan
Purcell et al. (2007)
Chiropsalmus sp.
Philippines
Heeger (1998)
Tamoya sp.
Tarawa, Kiribati
Shih (1977)
Scyphozoa
Coronatae
Periphyllidae
Periphylla periphylla
Norway
Wang (2007)
Semaeostomeae
Cyaneidae
Cyanea nozakii
China
Lu et al. (2003); Zhong et al. (2004); Dong et al. (2010)
Pelagiidae
Chrysaora pacifica
Japan
Morikawa (1984); Huang et al. (1987)
Chrysaora plocamia
Peru; Chile
this study
Pelagia noctiluca
?
Armani et al. (2013)
Ulmaridae
Aurelia aurita
Canada
DFA (2002a; b)
Aurelia labiata
Canada
Sloan & Gunn (1985)
Aurelia sp.
India; U.S.A.
Govindan (1984); Cox (2014)
10
2.1.2. Life cycle
Rhizostomes, which as mentioned, constitute the bulk of the edible species, have
several life history characteristics that may help to mitigate overfishing. The
canonical life cycle of these jellyfish is bipartite and metagenic, consisting of a sexual
pelagic medusoid phase and an asexual sessile polypoid phase (Figure 1). Medusae
are dioecious (i.e., gonochoristic) and females are typically highly fecund, producing
millions of eggs (e.g., Huang et al. 1985; Kikinger 1992). Fertilized eggs grow into
planulae, which are free-swimming and attach to hard substrates in a matter of
hours to days, subsequently transforming into polyps, or scyphistomae. Hard
substrate required for planulae settlement is essential habitat, of which natural
sources may be decreasing, as is the case with mangroves (Valiela et al. 2001), while
artificial habitat is increasing through anthropogenic substrates (Duarte et al. 2013).
Polyps of many species may asexually bud additional polyps (Lucas et al. 2012), or
may also produce or transform into cysts capable of resisting harsh environmental
conditions (Arai 2009). When conditions become favourable, polyps begin to
segment and asexually release ephyrae through a transverse fission process known
as strobilation. Each polyp may release numerous ephyrae and will often strobilate
more than once within the same season. Ephyrae join the plankton and grow rapidly
into medusae (Palomares & Pauly 2009), at which point they may be targeted for
11
fisheries. Sexually mature medusae are typically assumed to die off following
spawning due to senescence, disease, colder temperatures, or food limitation.
Figure 1. Life cycle of the cannonball jellyfish Stomolophus meleagris; based on Calder (1982)
12
As mentioned, this life cycle was historically deemed to be metagenic and bipartite,
indicating an alteration between two forms, namely, medusae and polyps. This was
thought to be heavily influenced by seasonality in temperate environments.
Recently, there has been considerable discussion of the scyphozoan life cycle in the
literature, often questioning the paradigm and terminology. Ceh et al. (2015) discuss
how medusae of Chrysaora plocamia may overwinter, potentially spending time in
deeper waters near the benthos, thereby going undetected in surface waters during
winter months. This has also been observed for other scyphomedusae, such as
Rhopilema verilli and Stomolophus meleagris in Georgia, U.S.A. (Kraeuter & Setzler
1975) as well as Aurelia labiata in British Columbia, Canada (pers. obs.). In addition,
limited evidence suggests that certain scyphozoans may sometimes “skip” either the
sexual or asexual phase of their life cycle. Aurelia in the laboratory have recently
been documented developing polyps directly from juvenile medusae and from
fragments of more mature medusae (He et al. 2015). Aurelia spp. have also been
shown to sometimes develop ephyrae directly from planulae, thereby occasionally
skipping the polypoid phase (Arai 1997; He et al. 2015). Some scyphozoan species,
such as Pelagia noctiluca and most members of the Order Coronatae have evolved to
the point where the sessile phase is entirely absent and are thus holoplanktonic. Of
13
course, the converse is also true, with many species having suppressed medusoid
stages.
Despite the extensive variety of scyphozoan life cycles and the myriad of
“exceptions” (Jarms 2010), i.e., less frequent features, Morandini et al. (2016) argue
that a “succession of generations” should still be considered the general paradigm
for scyphozoans. However, it seems important to underline the fact that for many
species, the polyps often survive after strobilation, and may strobilate more than
once within a season. Therefore, calling this a “succession of generations” seems
misguided, and perhaps it may be best to refer to the scyphozoan life cycle as
“polymorphic” (S. Piraino, Università del Salento, pers. comm., June 2016). In many
cases, it is also likely that asexual and sexual reproductive modes often overlap both
temporally and spatially. As such, management and research on jellyfish fisheries
may need to consider more complex models than the traditional one of a singular,
springtime cohort of medusae. Indeed, the peculiar life cycles of jellyfish may
provide buffers against overfishing, such as subsequent strobilation (and hence
recruitment) from surviving polyps, even without spawning adult medusae.
Nonetheless, overfishing of jellyfish stocks by catching medusae is plausible, and
appears to have occurred in some locations, such as China (Dong et al. 2014; also see
3.3.4 China) and the Salish Sea (Mills 2001; also see 3.3.22 U.S.A.).
14
2.2. Fisheries
2.2.1. Fishing for jellyfish
Fisheries for jellyfish are usually characterized by short fishing seasons of a few
months as well as dramatic interannual variations in catches (Omori 1978; Omori &
Nakano 2001). As such, jellyfish fisheries are often subject to instability; a concern
for fishers, managers, and other stakeholders alike. A wide variety of vessels may be
used for fishing jellyfish. While diesel-powered trawlers are used in select locations
(e.g., U.S.A.), jellyfish are most often fished from small (5-10 m) boats powered by
outboard engines operating relatively close to shore. These boats often have crews of
1 to 5 fishers and typically carry somewhere between 1 to 5 tonnes of catch when
fully loaded. Medusae are visually located in surface waters and caught using dip-
nets, also known as scoop nets. Large catches of jellyfish on the decks or in holds of
boats can result in concerns regarding vessel stability, and therefore it may be
important for vessels to have baffles in the hold to prevent the catch from shifting,
especially in rough conditions. Another concern for fishers is the possibility of stings
(Kawahara et al. 2006a), as jellyfish are often transferred out of nets by hand. Stings
from rhizostomes are generally irritating and painful, but most are non-lethal. A
possible exception is the giant jellyfish Nemopilema nomurai; however, more research
is needed to assess the threat of this species to humans (Kawahara et al. 2006a).
15
Although fishers can wear gloves to prevent stings, gloves are often not used as they
may be cumbersome, too hot, or unavailable.
A variety of other gears are also used to catch jellyfish including hooks, set-nets, gill
nets, drift nets, purse seines, beach seines, weirs, and trawl nets. In some cases,
combinations of gears are used to increase the quality and size of the catch. For
example, in the Ariake Sea (Kyushu, Japan) and Sarawak (Malaysia, Borneo),
jellyfish are concentrated using set-nets and then collected using dip-nets (Rumpet
1991).
2.2.2. Ecological implications
The ecological implications of fishing for jellyfish are largely unknown. Jellyfish are
historically understudied animals (Pugh 1989; Hay 2006), and published research on
jellyfish fisheries is scant. While jellyfish are often perceived as nuisance species,
they play a variety of important roles in ecosystems, often with very significant
impacts. Dip-nets, as well as nets with larger mesh sizes, may facilitate the
avoidance of bycatch and smaller medusae, and several countries have implemented
minimum size limits (MSLs). However, the effectiveness of MSLs in jellyfish
fisheries has not been evaluated scientifically (also see 2.4 Management). Larger mesh
sizes can also damage bigger and more valuable medusae, depending on the
methods and gear being used. Numerous species of fishes are known to associate
16
with jellyfish, presumably using the medusae as food and/or refugia from predators
(e.g., Jones 1960; Arai 1988; Kingsford 1993; Brodeur 1998; Purcell & Arai 2001;
López-Martínez & Rodríguez-Romero 2008; Mianzan et al. 2014). In addition, many
invertebrates associate with jellyfish, potentially benefitting from habitat, food,
refugia, and transportation (e.g., Brandon & Cutress 1985; Arai 2005; Browne &
Kingsford 2005; Towanda & Thuesen 2006; Sal Moyano et al. 2012; Schiariti et al.
2012a; Álvarez-Tello et al. 2013; Fleming et al. 2014). As such, the many close
associations between jellyfish and other species assure that bycatch concerns cannot
be eliminated entirely.
The use of other gears, such as set-nets and trawl nets may increase bycatch further,
and commercially important species may be unintentionally caught when
associating with jellyfish (Rumpet 1991; Panda & Madhu 2009). Bycatch in the trawl
fishery for cannonball jellyfish Stomolophus meleagris was examined in detail in
Georgia, U.S.A. In total, 133 tows were examined between 2005 and 2012. The
results, presented by Page (2015), show that 38 species of fish, as well as 3 species of
invertebrates (not including spider crabs Libinia spp., which are symbiotic with
S. meleagris) were recorded as bycatch. The most commonly observed bycatch were
harvestfish Peprilus paru (41%), cownose ray Rhinoptera bonasus (11%), Atlantic
bumper Chloroscombrus chrysurus (11%), butterfish Peprilus triacanthus (11%), and
17
blue crab Callinectes sapidus (7%). The 3 finfish species (harvestfish, Atlantic bumper,
and butterfish) are all known to associate with jellyfish, presumably using them as
refugia from predators, and potentially becoming ectoparasites that feed directly on
the medusae (Purcell & Arai 2001). As such, it is not surprising that these species
form a major component of the bycatch (Page 2015). A similar associative
relationship also explains the vast quantities of spider crabs that were caught as
bycatch. Other species that are known to associate with S. meleagris medusae but
were absent as bycatch may be due to the seasonality of the fishery and/or the ability
of species to escape the nets (e.g., carangids). Given that the top 5 bycatch species
(excluding spider crabs) comprised approximately 80% of all individuals caught, it
can be said that “the commercial cannonball jellyfish trawl fishery in Georgia is
dominated by a few recurring species and is minimal relative to the bycatch
associated with another important trawl fishery in the state – namely the
commercial food shrimp trawl fishery” (Page 2015). Indeed, 24% of the tows
analyzed contained zero bycatch (excluding spider crabs). Nonetheless, those
species comprising the majority of the bycatch can be caught in significant quantities
at times, and may be of commercial and/or ecological concern. As such,
management plans for the jellyfish fishery in the U.S.A. should take these species
18
into consideration if the fishery is to be scaled up in the future beyond the current
small fleet.
Other species caught as bycatch may also be of concern, even if they are less
abundant, such as sea turtles, many of which consume jellyfish. In the past, jellyfish
were so bothersome to shrimp fishers that modifications were made to trawl gear
that facilitated the exclusion of jellyfish while still permitting shrimp to travel into
the codend (Jones & Rudloe 1995). Essentially, a series of metal bars is used to divert
anything larger than the space between the bars to an escape hatch, whereas
anything smaller passes through to the codend. These device modifications proved
to also exclude sea turtles, and ultimately became known as turtle excluder devices
or ‘TEDs’ (Jenkins 2012). TEDs have saved countless numbers of sea turtles and have
been the basis for numerous awards (Landers 2011). As one would suspect based on
their original purpose, the use of TEDs dramatically reduces the catch of jellyfish,
often by more than 80% (Huang et al. 1987). As such, most jellyfish fishers in Georgia
opt to trawl in federal waters immediately adjacent to state waters, where TEDs are
not required (Page 2015). During the aforementioned bycatch study, a total of 13
protected species (11 sea turtles and 2 common bottlenose dolphins) were caught
during the 133 observed tows (which represented < 5% of all tows during the
period). While some animals caught as bycatch are released alive, tows routinely
19
exceed 1 hour in duration (average of 0.55 h), suggesting that mortality of air-
breathing species could be significant. There are ongoing efforts to modify TEDs
with spacing between the bars that is sufficient for jellyfish to pass through, but not
turtles (Page 2015).
Although jellyfish are sometimes perceived to be “trophic dead-ends,” whereby
their energy is not transferred to higher levels of the food web (e.g., Verity &
Smetacek 1996; Sommer et al. 2002), this perception is changing. Many sea turtles
will prey on jellyfish during some stage of their lives, and the leatherback sea turtle
Dermochelys coriacea is an obligate jellyfish predator, with individuals potentially
eating hundreds of kilograms of jellyfish in a single day (Duron-Dufrenne 1987;
Heaslip et al. 2012). As some populations of leatherbacks are critically endangered,
fishing for jellyfish in waters deemed critical habitat could be subject to restrictions
in some jurisdictions, and there are concerns that fisheries for jellyfish could deplete
important food resources for leatherbacks at local scales (Elliott et al. 2016). Recent
investigations are also revealing the importance of jellyfish as prey for more than
100 species of fish (Pauly et al. 2009). In addition, large blooms of jellyfish that die
and sink to the ocean floor (known as ‘jelly-falls’) have mainly been investigated for
their role in the biological pump, i.e., sequestering carbon to the benthos (Lebrato et
al. 2012; 2013). However, it is becoming apparent that gelatinous zooplankton may
20
instead/also be an important nutritional input for benthic animals (e.g., Henschke et
al. 2013; Sweetman et al. 2014; Chelsky et al. 2016; Henschke et al. 2016).
Jellyfish can also be voracious predators and often have very significant impacts on
the abundance, biomass, and size composition of zooplankton at lower trophic
levels (Möller 1980; Mills 1995; Purcell & Arai 2001). In addition, jellyfish are often
predators of eggs and larvae of fish, as well as in direct competition for food with
many fishes (Purcell 1985; 1997; Robinson et al. 2014; Tilves et al. 2016). Thus, the
consequences of removing jellyfish from the environment could have dramatic
effects on ecosystems, impacting organisms from phytoplankton through to top
predators.
Beyond their extensive roles in food webs, jellyfish also provide a number of
ecosystem services such as carbon transport, nutrient liberation, and ocean mixing
(Doyle et al. 2014). Given all of their influential roles in ecosystems, removing
jellyfish in large quantities is likely to have significant consequences. Unfortunately,
jellyfish are typically ignored or simplified in ecosystem models (Pauly et al. 2009),
and as such, the impacts of fishing them are not well understood.
21
2.3. Use
While the overwhelming majority of jellyfish caught are used as food for humans,
there are numerous other reasons why jellyfish may be targeted. In some cases,
jellyfish have been fished simply to remove them from locations where they are a
nuisance to tourism or other industries. Such efforts have proven effective in Hawaii
(Hofmann & Hadfield 2002; Kelsey 2009); however, these cases involved
Cassiopea spp., which are relatively sedentary (Holland et al. 2004). Cannonball
jellyfish (S. meleagris) have also been removed in the past from canals in Florida,
where they clogged the intake pipes of a nuclear power plant (Jones & Rudloe 1995).
Fishers have also been paid to remove Cotylorhiza tuberculata in Mar Menor in the
Mediterranean Sea, a species which appears to have increased largely due to
anthropogenic impacts (Brotz & Pauly 2012). While it appears that fishing of
medusae may have helped to reduce the jellyfish population there, it was an
extremely expensive program, and environmental conditions are likely more
influential of the population dynamics in question (Prieto et al. 2010; Ruiz et al. 2012).
Research on jellyfish has led to a better understanding of ocular vision (Nilsson et al.
2005), as well as two Nobel Prizes: one in 1913 for the discovery of anaphylaxis, and
another in 2008 for the discovery and development of green fluorescent protein (see
3.3.22 U.S.A.). Jellyfish have also informed the field of design engineering (Dabiri
22
2011; Najem et al. 2012; Ristroph & Childress 2014; Costello et al. 2015), where their
biomechanics are often mimicked due to their simple and efficient design (Gemmell
et al. 2013; 2015). Jellyfish are also being investigated for a wide variety of
applications including agriculture, materials science, and pharmaceuticals (Table 3).
Most of these technologies are in their infancy, and thus it will likely be sometime
before there is significant demand for jellyfish other than for food. The only current
instances of companies targeting wild jellyfish at commercial scales for purposes
other than for food are for the extraction of collagen and are located in France and
the United Kingdom (see 3.3.24 Other countries).
23
Table 3. Uses for jellyfish other than as food for humans
Category
Uses
Sample reference(s)
Agriculture
Livestock feeds
Hsieh & Rudloe (1994); CIESM (2010)
Fertilizers
Fukushi et al. (2004); Fukushi et al. (2005); Chun et al.
(2011); Kim et al. (2012b); Hossain et al. (2013); Hussein
& Saleh (2014); Seo et al. (2014); Hussein et al. (2015)
Insecticides
Yu et al. (2005); Yu et al. (2014)
Aquaculture
Finfish & shellfish
feeds
Gopakumar et al. (2008); Miyajima et al. (2011a); Miyajima
et al. (2011b); Wakabayashi et al. (2012); Liu et al. (2015);
Wakabayashi et al. (2016)
Cosmetics
Gelatin/emulsifier
Cho et al. (2014); Chancharern et al. (2016)
Environmental
monitoring
Pollution detection
Fowler et al. (2004); Templeman & Kingsford (2010);
Morabito et al. (2014); Epstein et al. (2016); Muñoz-Vera
et al. (2016)
Fishing
Bait
Prabhu (1954); Thomas (1969); Omori & Kitamura (2004);
Varghese et al. (2008); Mianzan et al. (2014)
Materials
science
Absorbent polymers
Shamah (2014); Belgorodsky et al. (2015)
Cement additive
CIESM (2010)
Copolymer films
Thumthanaruk et al. (2016)
Nanoparticle filtering
Patwa et al. (2015)
Pharmaceuticals
Antihypertensive
peptides
Zhuang et al. (2012a; b)
Anticoagulants
Rastogi et al. (2012)
Antimicrobiotics
Ovchinnikova et al. (2006); Yin et al. (2016)
Antioxidants
Yu et al. (2006); Zhuang et al. (2009); Balamurugan et al.
(2010); Zhuang et al. (2010); Leone et al. (2015)
Bioactive compounds
Rottini et al. (1995); Gusmani et al. (1997); Li et al. (2005);
Nishimoto et al. (2008); Morishige et al. (2011); Kawabata
et al. (2013); Leone et al. (2013); Badre (2014); Mariottini
& Pane (2014); Leone et al. (2015); Mariottini & Brotz (in
press)
Collagen
Nagai et al. (2000); Huang et al. (2005); Song et al.
(2006); Addad et al. (2011); Krishnan & Perumal (2013);
Barzideh et al. (2014a; b); Klaiwong et al. (2014); Zhang et
al. (2014); Derkus et al. (2016)
Mucins
Masuda et al. (2007); Ohta et al. (2009)
2.3.1. Processing of edible jellyfish
In some locations, jellyfish may be consumed fresh, as is occasionally the case with
Rhopilema esculentum and Nemopilema nomurai in coastal China (You et al. 2007; Yang
& Shuang 2015), or processed using oak leaves (Morikawa 1984); however, these are
rarities in comparison with the vast quantities that are processed for mass
consumption. Freshly caught jellyfish can spoil quickly, and therefore the catch is
24
usually brought to a local processing facility within a few hours. Initial signs of
spoilage include sliminess, colour change, and unpleasant smell. Sometimes the
catch is stored in containers on board the fishing vessels while in transit in order to
delay spoilage, with either seawater or a slurry of ice and alum – usually potassium
aluminum sulfate KAl(SO4)2. Most edible jellyfish species have prominent oral arms
rather than conspicuous tentacles. Depending on the species and the target market,
the oral arms may be disposed of or processed separately. The bell is usually more
expensive, and was historically valued at more than twice as much as the oral arms
(Hsieh & Rudloe 1994; Omori & Nakano 2001). However, demand for oral arms is
increasing in China (Kitamura & Omori 2010), and there are recent reported cases
where only the oral arms are sought, with the bell being disposed of at sea (e.g.,
Mohan et al. 2011).
Processing facilities range from seaside tents and shacks to large, industrialized
seafood processing factories. Facilities may employ dozens of labourers, including
many women and children who are sometimes the family members of the fishers.
Jellyfish are often scraped, sometimes with tools made of bamboo, to remove mucus
or the surface ‘skin’ if there are denticulations. The gastrovascular cavity and any
developed gonads are typically removed (Chidambaram 1984), which may also be
done near the end of processing (Rudloe 1992). The edges of the bell are sometimes
25
trimmed and removed (Govindan 1984; Santhana-Krishnan 1984; Nishikawa et al.
2008). Jellyfish are usually rinsed with seawater, which appears to be especially
important for species that produce a lot of mucus, such as cannonball jellyfish
Stomolophus meleagris (Jones & Rudloe 1995).
Processing typically involves soaking the jellyfish in large vats or tanks containing
different mixtures of salt and alum (which is granular, white, and odourless). These
mixtures may be dry or in brine solution, and jellyfish are soaked in different
mixtures for specified time periods. The salting process is intended to dehydrate and
preserve the jellyfish without denaturing the collagen, resulting in the desirable
firmness and crunchiness of the final product (Sloan & Gunn 1985). The alum will
also reduce the pH and act as a disinfectant (Hsieh et al. 2001), and processing
typically eliminates any remaining sting from nematocysts (Hsieh & Rudloe 1994).
Using only salt or alum alone does not result in a satisfactory product (Wootton et al.
1982). Heat is avoided during processing, as it will quickly denature the collagen
(Rigby & Hafey 1972). In some areas of Southeast Asia, such as Malaysia, Thailand,
and the Philippines, a small amount of soda (potentially NaHCO3 or NaOH) may be
added to facilitate additional dehydration, thereby increasing the crispiness of the
final product (Rumpet 1991; Rudloe 1992; Hsieh et al. 2001; PCAMRD 2008). In
certain locations, especially those exporting product to China, jellyfish may be
26
treated with mixtures containing hydrogen peroxide in order to bleach the product
white.
Jellyfish processing is stepwise and usually takes weeks before it is complete,
although acceptable products have been produced in as little as 8 days using
cannonball jellyfish (S. meleagris) which has a smaller maximum size than most
edible species (Huang 1988). Mechanical drying has also been investigated to reduce
processing time, but the resulting products were not satisfactory due to uneven
dehydration (Wootton et al. 1982; Huang 1988; Hudson et al. 1997). Research is also
expanding into drying techniques that result in a product that can be either partially
rehydrated or used as an additive. Yuferova (in press) demonstrated results that
were more promising for Rhopilema esculentum than Aurelia aurita, due to the
increased destruction of proteins and sugars in the latter.
As jellyfish processing is usually time-consuming and labour-intensive, it is a
potential limiting economic factor in countries where labour costs are high.
Processing techniques and formulas vary by region and species, and potentially
even by batch, so many Asian processors will employ “Jellyfish Masters” who make
adjustments to obtain an acceptable product, often keeping their formulas as
guarded secrets (Rudloe 1992; Jones & Rudloe 1995). Several different processing
protocols based on Japanese and Thai preferences are outlined in Sloan & Gunn
27
(1985), and numerous overviews of jellyfish processing are available in English (e.g.,
Soonthonvipat 1976; Wootton et al. 1982; Chidambaram 1984; Govindan 1984;
Santhana-Krishnan 1984; Huang 1988; Suelo 1988; Rumpet 1991; Rudloe 1992; Jones
& Rudloe 1995; Ozer & Celikkale 2001; Nishikawa et al. 2008), Spanish (e.g., Álvarez-
Tello 2007; Schiariti 2008; Schiariti & Mianzan 2013; Schiariti et al. 2015), and Chinese
(e.g., Wu 1955; Liu 1973; Yin et al. 2000). After stepwise processing in salt-alum
solutions, bells are often stacked in a pile and allowed to drain for several days
(Subasinghe 1992). Salt may be sprinkled on the bells before stacking, and the bells
are often rotated during dehydration to ensure evenness and proper drainage.
There are a number of concerns regarding pollution from effluent created by
processing facilities. Primarily, these concerns surround the disposal of huge
amounts of slime-salt wastewater created during the initial processing stages. This
issue has been the subject of recent debate in South Carolina, U.S.A., as companies
are looking to expand production but are being met with resistance and regulation
(see 3.3.22 U.S.A.). In most countries, the effluent from processing facilities is not
regulated. Research into the development of improved processing techniques that
minimize the harmfulness and toxicity of effluent should be a priority, and potential
solutions may have the added benefit of reducing costs as well as minimizing the
28
negative health effects associated with processing chemicals, such as aluminum (see
below).
2.3.2. The edible product
As stated by Hsieh et al. (2001), in China “jellyfish is more than a gourmet delicacy:
it is a tradition.” In many Asian countries, jellyfish is consumed in the home as well
as at restaurants, ceremonies, and banquets. While jellyfish are targeted and caught
in numerous countries around the world, these countries export nearly all of their
catch to Asia, usually after processing. Most consumption occurs in China, Japan,
Malaysia, South Korea, Taiwan, Singapore, Thailand, and Hong Kong, with demand
in Japan having increased especially in the 1970s (Omori & Nakano 2001). Smaller
markets for jellyfish exist in many cities with inhabitants of Asian descent, where
consumption typically involves jellyfish imported from China. Depending on the
original source, such jellyfish may have been re-exported, but no supply chain
tracing mechanisms are in place for jellyfish. While jellyfish as food continues to
remain a novelty for most people in the Western Hemisphere, increasingly
diversified urban populations have resulted in higher imports of jellyfish in many
major cities. Combined with a growing and wealthier population in China, global
demand for jellyfish is thus likely to continue to increase. Prices for jellyfish vary
29
widely depending on the product, but processed jellyfish can typically be found at
market fetching 2-10 USD/kg.
Both classically semi-dried and ready-to-eat jellyfish products are sold in a variety of
different presentations and packaging. These may include bulk bins, plastic jars, soft
plastic sealed bowls, and sealed plastic envelopes. Mislabelling of jellyfish products
appears rampant (as it does for fish products - Jacquet & Pauly 2008), and may
include a lack of adherence to local labelling regulations, a high rate of
misidentifying species, and even labelling of the product as a vegetable, such as
bamboo or tuber mustard (Armani et al. 2012; 2013). Unfortunately for consumers,
new seafood labelling regulations being implemented by the European Union (EU)
exclude invertebrates (D'Amico et al. 2016).
Processed jellyfish can weigh anywhere from 7 to 30% of the original wet weight,
depending on the species in question and the specific processing formula used
(Wootton et al. 1982; Huang 1986; Hsieh et al. 2001). Variation in the final moisture
content is typically dictated by different market preferences. For example, product
bound for Japanese markets tends to be much crunchier (i.e., lower water content)
than for Chinese markets. Products are often graded, and quality is determined
based on size, texture, and colour; with larger products that are whiter in colour and
of the preferred texture fetching the highest prices (Rudloe 1992). Colour tends to
30
turn from white to yellow to brown as the product ages, with the latter being
unacceptable. Jellyfish may be packed into crates or buckets for export, sent to local
auctions, or sold in small packages at retail markets. The shelf life of processed
jellyfish varies depending on the product and storage temperature. Huang (1988)
noted that product from S. meleagris can be stored for at least 6 months at 10°C.
Hsieh et al. (2001) stated that edible jellyfish products last up to a year at room
temperature, which can be extended to more than 2 years if kept cool. It is generally
reported that the product will spoil if frozen (Huang 1986; Rudloe 1992; Subasinghe
1992; Hsieh et al. 2001); however, freezing of processed jellyfish for storage is
possible for short periods (Govindan 1984; Santhana-Krishnan 1984; Kingsford et al.
2000; Ozer & Celikkale 2001). Eventually, frozen processed jellyfish will begin to dry
out and form wrinkles, negatively affecting the appearance and texture of the
product, and is therefore not recommended for prolonged periods (Huang 1986;
Rudloe 1992; Subasinghe 1992; Hsieh et al. 2001). A sample of Cotylorhiza tuberculata
that was frozen fresh near -80°C and then thawed before preparation by a chef is the
tastiest edible jellyfish the author has ever tried, demonstrating that traditional salt-
alum processing of jellyfish is not the only method available for future consumption
of jellyfish (also see 5.1 Research priorities). Fresh moon jellyfish (Aurelia sp.) has also
been declared as “not only edible, but actually quite delicious” and “best described
31
as a slightly salty oyster, with a clean flavour and more enjoyable texture” (Cox
2014).
Processed jellyfish has almost no intrinsic flavour, and is therefore usually served
with sauces that may include sesame oil, soy sauce, vinegar, and/or sugar. Edible
jellyfish has a surprising crunch, and the sensation of biting into the crisp ‘meat’ is
sometimes referred to as the product’s ‘sound.’ In preparation for consumption,
processed jellyfish is usually soaked for several hours or overnight, sometimes with
numerous water changes, in order to desalt and partially rehydrate the product.
Traditional recipes may even call for jellyfish to be soaked for several days (Wootton
et al. 1982). After soaking, the jellyfish may be scalded or blanched briefly in boiling
water, which forms ‘curls.’ Jellyfish is most often served at room temperature and
sliced into thin strips. It may be served with sliced vegetables and/or sliced meat as
an appetizer, salad, or soup. There are also many other Asian dishes that include
jellyfish as an ingredient. Ready-to-use or ready-to-eat jellyfish products have been
developed more recently, which, as their name suggests, do not require soaking or
scalding prior to eating. These products are sometimes packaged with spices and/or
sauces as a ready-to-eat snack.
Rehydrated edible jellyfish are primarily composed of water, accounting for 92-97%
of the weight, depending on the species, the type of product, and the processing
32
methods used. The main organic component is collagen (Kimura et al. 1983; Khong
et al. 2016), a connective tissue protein making up about 3-7% of the rehydrated
product weight (again, a value that varies according to the species in question and
the processing method used). Levels of fat, cholesterol, and carbohydrates are
extremely low or undetectable. Lipid content may increase in specimens with well-
developed gonads, but these are typically removed during processing. Tryptophan
has been identified as the limiting amino acid in some edible species (Kimura et al.
1983; Leone et al. 2015) but not others (Khong et al. 2016). With approximately 36
food calories per 100 g serving (USDA 2015), these characteristics have led to edible
jellyfish being declared as a natural diet food, comparable to many vegetables.
2.3.3. Health effects of jellyfish consumption
A number of inorganic constituents are detectable in processed jellyfish, the most
concerning of which is aluminum from the alum used in processing. Many salts and
minerals may be removed through soaking and scalding; however, aluminum has
been detected in the final edible product in significant quantities (Ogimoto et al.
2012; Zhang et al. 2016). Ready-to-eat jellyfish products are not rinsed by the
consumer prior to consumption, and have also been shown to have high aluminum
content (Wong et al. 2010; Armani et al. 2013). Consumption of aluminum has been
linked to a number of negative health effects, including neurobehavioural toxicity
33
and Alzheimer’s disease (Perl & Brody 1980; Nayak 2002). The links between
aluminum consumption and the associated negative health effects are still not well
understood, and further research is needed. Nonetheless, the development of
processing methods that avoid the use of aluminum for edible jellyfish is desirable
(Hsieh & Rudloe 1994), and should be a priority for the industry (see 5.1 Research
priorities). The United Nations and the European Union have both recently reduced
their limit for tolerable weekly intake of aluminum, and the Food and Drug
Administration of Taiwan has indicated that it will be ramping up monitoring and
enforcement of aluminum additives in food products such as jellyfish (I-chia 2016).
Additional additives may also be included in processed jellyfish, including
monosodium glutamate or potassium sorbate (Armani et al. 2012).
In contrast to the negative effects associated with aluminum found in edible jellyfish
products, there is also a long list of purported health benefits from consuming
jellyfish. Traditional Chinese Medicine (TCM) claims that eating jellyfish is
beneficial for curing arthritis and gout, decreasing hypertension, treating bronchitis,
alleviating back pain, curing ulcers and goiter, easing swelling, simulating blood
flow (especially during menstruation), remedying fatigue and exhaustion, softening
skin, aiding weight loss, improving digestion, and for treating cancer (Rudloe 1992;
Hsieh & Rudloe 1994; Jones & Rudloe 1995; Hsieh et al. 2001; You et al. 2016).
34
Australian Aborigines have used dried jellyfish powder to treat burns (Hsieh &
Rudloe 1994). Fishers in the Philippines believe that consumption of jellyfish will
increase resistance to hypertension, back pain, arthritis, and malaria (PCAMRD
2008). Very few clinical trials have been conducted to test these claims, and while
most remain neither proven nor disproven, many seem unlikely while some are
plausible. Hsieh et al. (2001) report the findings of a small study whereby several
rats were fed jellyfish collagen after being injected with an arthritis-inducing
reagent. Those rats that were fed jellyfish collagen reportedly showed significantly
reduced incidence, onset, and severity of arthritis in comparison to the control
group. More details are presented in Hsieh (2005); however, no human clinical data
are available. Zhang et al. (2008) also found that jellyfish collagen had positive effects
on rats with arthritis. In another study presented by Ding et al. (2011), mice were
administered various doses of jellyfish collagen hydrolysate. All groups that were
administered collagen showed a statistically significant increase in exercise tolerance
and glycogen levels, along with reductions in lactic acid and blood urea nitrogen. In
addition, mice that had been induced to represent aging showed antioxidative
effects when administered jellyfish collagen hydrolysate. Collagen peptides derived
from jellyfish may also have beneficial effects on immune functions in mice (Deng et
al. 2009). There are several studies reporting positive results for the treatment of
35
hypertension in rats using extracts from jellyfish (Su et al. 2008; Liu et al. 2009; 2013).
Despite these experiments, no clinical trials have been performed using humans and
edible jellyfish, and the effects of treating arthritis with collagen in humans have
generally been mixed (Henderson & Panush 1999). Therefore, it remains plausible
but unproven that consumption of jellyfish by humans has at least some of the
health benefits purported by TCM.
Some individuals may experience negative reactions soon after consuming
processed jellyfish, such as anaphylaxis (Imamura et al. 2013; Inomata et al. 2014;
Okubo et al. 2015); however, such cases appear to be rare. Mild allergic reactions to
the consumption of jellyfish may also occur, such as swelling of the mouth, but also
appear to be rare. In fact, preliminary results suggest that consumption of jellyfish
may be safe for some individuals with allergies to other seafoods (Amaral et al.
2016). There is a solitary case of ciguatera poisoning suspected to be caused by
consumption of jellyfish from American Samoa, although the details are vague
(Zlotnick et al. 1995).
2.4. Management
As mentioned, jellyfish populations typically exhibit dramatic interannual variation
in abundance and biomass (Brotz 2011). In fact, changes in biomass of edible jellyfish
are probably larger than for any other fishery (Kingsford et al. 2000). This presents
36
extremely large uncertainties for fisheries managers, makes predictions of future
catches difficult, and may prevent investment in infrastructure. There is also
evidence to suggest that discrete populations of medusae may exist at relatively
small spatial scales (Kingsford et al. 2000; Matsumura et al. 2005; Mooney &
Kingsford 2016a; b; van Walraven et al. 2016). This could make some stocks
vulnerable to overfishing, especially as fishers are likely to concentrate their effort in
areas that are closer to ports or processing facilities (Kingsford et al. 2000). However,
other species of edible jellyfish may have low genetic diversity across broad
geographic ranges (e.g., Dong et al. 2016). As such, management of jellyfish fisheries
will need to be informed by both general prescriptions addressing the unique
aspects of jellyfish, as well as the local conditions and species in question.
Nonetheless, many of the options for traditional fisheries management are available
to jellyfish fisheries, only a few of which have been employed.
In Australia, precautionary total allowable catches (TACs) have been implemented
(Fisheries Victoria & MAFRI 2002; Fisheries Victoria 2006), but only a small fraction
of the TACs have been utilized, presumably due to a lack of economic viability and
potentially onerous regulations. TACs for jellyfish fisheries appear to be rare in most
other countries; however, total catch may be limited by processing capacity where it
is regulated or industrialized, as is the case in the U.S.A. TACs can also be artificially
37
increased if portions of the jellyfish, such as the oral arms, are discarded at sea. You
et al. (2016) proposed a system of fishing quotas and a deficit quota buyback system
for the jellyfish fishery in Liaodong Bay, China. The fishery for Rhopilema esculentum
in the region has been subject to a dramatic increase in fleet size from hundreds of
boats to approximately 10 thousand, along with a concomitant reduction in the
length of the fishing season from months to hours over the course of only a few
decades (see 3.3.4 China). Combined with collapses of other fisheries in the region,
the situation has led to a consistent increase in illegal fishing of jellyfish, with many
fishers starting to target jellyfish weeks or even months ahead of the seasonal
openings. Despite increased monitoring in recent years, including GPS tracking and
video monitoring, illegal fishing of jellyfish in the area continues to be rampant, and
often includes far more violators than can be prosecuted. There are even examples of
collusion amongst groups of fishermen, whereby a second wave of fishers will agree
to pay the fines incurred by the first (You et al. 2016). Indeed, it seems that
management plans need to include input and support from fishers and other
stakeholders if they are to be effective, as monitoring and enforcement is expensive
and may even unrealistic in some cases, as illustrated by the situation in Liaodong
Bay.
38
Some countries have also implemented minimum size limits (MSLs) on medusae,
such as Australia, Mexico, and the U.S.A. The intent of MSLs is to prevent the
capture of medusae before they reach sexual maturity, as well as encouraging higher
fecundity, which typically increases with size (e.g., Coleman 2004; Schiariti et al.
2012b). However, there is no guarantee that medusae will spawn successfully at a
certain size, or that they will be in a location where planulae can find suitable
substrate for settlement. Conversely, medusae may reach sexual maturity over a
wide range of sizes, and maturation may be more related to environmental
conditions than size (Carvalho-Saucedo et al. 2010; 2011). Of course, a medusa’s size
is also related to environmental conditions, so the interplay amongst the
environment, a medusa’s size, and its state of sexual maturity are not well
understood. As such, MSLs are likely not enough to guarantee a sustainable jellyfish
fishery (admittedly, nor are they sufficient for finfish fisheries). In addition, larger
mesh sizes have the potential to damage medusae, depending on the species in
question and the gear used. Nevertheless, implementation of MSLs may be a useful
precautionary management technique, especially when knowledge of the target
organism’s life history is poor. MSLs can also have the added benefit of allowing
jellyfish to grow before being caught, which may result in more profit as larger
medusae typically fetch higher prices, unless of course natural mortality increases or
39
the jellyfish exhibit degrowth due to poor food availability or other environmental
conditions (e.g., Hamner & Jenssen 1974; Frandsen & Riisgard 1997; You et al. 2007;
Lilley et al. 2014). Additional research on such topics is essential, especially the
exploration of which management techniques are most appropriate for jellyfish
fisheries.
40
3. Reconstructing the global catch
As it is widely recognized that catch statistics are crucial for fisheries management
(Pauly 1998; Jennings et al. 2001), it follows that such statistics should be accurate in
order to properly inform the decisions dependent upon them. Indeed, “maintaining
catches is the raison d’être of fisheries science” (Pauly 2016). The primary
organization compiling national and global fishery catch statistics is the Food and
Agriculture Organization of the United Nations (FAO). This institution began
compiling fishery statistics in an annual yearbooks in 1950, soon after its founding,
part of the United Nations’ attempt to “quantify the world” (Ward et al. 2004).
However, FAO is entirely dependent on what individual countries report, and such
data are often problematic in a number of ways.
A common response to deficient catch datasets is to develop working groups and
intensive projects with the purpose of improving reporting at the regional and
national scale. Unfortunately, such projects are usually limited in their scope,
especially in time, negating the ability to identify longer-term changes in the data
(Pauly 1998). One exception is the Sea Around Us (www.seaaroundus.org), a research
program supported by The Pew Charitable Trusts and the The Paul G. Allen Family
Foundation. The Sea Around Us reconstructs, maintains, and updates a global
41
database of fisheries catches from every maritime country of the world back to 1950
(Pauly & Zeller 2016a). The reconstruction of this dataset has enabled many
significant conclusions at the global scale, including the facts that fisheries catches
are both of a larger magnitude and declining more rapidly than previously thought
(Pauly & Zeller 2016b). Important conclusions can also be drawn at the regional and
national scales by evaluation of long-term trends and comparison of neighbouring
jurisdictions. Such analyses can inform management decisions about the status of
fishery resources and how they might respond to changes in effort.
Catch reconstructions performed by members of the Sea Around Us and its
collaborators include catches disaggregated into different taxa. However,
information on jellyfish catches is often not uncovered during the catch
reconstruction process, as many countries do not record jellyfish explicitly. In
addition, the taxonomy of jellyfish is still confused (Omori & Kitmura 2004;
Kitamura & Omori 2010), confounding the situation further. Nonetheless, just as we
must overcome the psychological assumption that there is ‘no data’ available on
fisheries catches (Pauly 1998; 2016), as evidenced by the catch reconstructions
performed by the Sea Around Us, this is also true for jellyfish catches. As such, a
global catch reconstruction focused specifically on jellyfish will be a valuable input
for the Sea Around Us database, as well as helping to answer important questions
42
specific to jellyfish fisheries catches, including ‘how much?, ‘where?’, ‘what
species?’, and ‘what are the trends?’
3.1. Methods
3.1.1. Catch reconstruction
Jellyfish catches were estimated using an approach known as ‘catch reconstruction,’
which has been applied to all maritime countries of the world (Pauly & Zeller 2016a;
b). Regarding the jellyfish catches herein, the results are in fact landings, a subset of
catches, as no estimates of discarded bycatch of jellyfish from other fisheries were
made in this analysis. This was primarily due to the lack of availability of detailed
bycatch data for fisheries in most developing countries, as well as the fact that the
goal of this project was to estimate the quantity of jellyfish that are caught for
human consumption. As such, the true global ‘catch’ (including discards) of jellyfish
is likely much higher than what is estimated here.
The catch reconstruction methodology used was based on established procedures
(Zeller et al. 2007; 2015; Pauly & Zeller 2016a; b), and employed the following steps:
1. Identification and validation of existing reported catch time series (e.g., FAO
statistics);
2. Identification of countries and time periods not covered by (1), i.e., missing
catch data, via literature searches and consultations;
43
3. Search for available alternative information sources to supply the missing
catch data in (2), through extensive literature searches and consultations with
local experts;
4. Development of data anchor points in time for missing data items;
5. Interpolation for time periods between data anchor points for total catch;
6. Estimation of final total catch time series estimates for total catch, combining
verified reported catches (1) and interpolated missing data series (5).
Regarding the foundational information that was used as the primary basis for the
reconstruction indicated above in point (3), all sources are identified in the
individual country sections that follow (see 3.3 Countries fishing jellyfish). A wide
variety of sources were utilized, and most often include regional and national
fisheries catch datasets, import/export statistics, peer-reviewed publications, grey
literature, popular media articles, as well as direct communication with industry
stakeholders such as processors, brokers, and fisheries scientists.
3.1.2. Scaling factor
Given the stepwise methodology involving salt and alum that is typically employed
to process jellyfish, the result is a semi-dried product that is lighter in weight that
the original wet weight of the jellyfish catch. In some cases, the only information
available for a country’s jellyfish fishery is for processed product weight. In such
instances, estimates of the original wet weight of jellyfish caught can be calculated
using a scaling factor. Scaling factors can vary depending on the species in question
44
and the processing method used, which is often dictated by varying market
preferences (Mohammed 2008). The fraction of semi-dried product to wet weight
may range from 7% (Omori 1981; Morikawa 1984; Georges 1991) to more than 25%
(Wootton et al. 1982; M. Valle, Pesquera Mexico, pers. comm., Feb. 2014).
Intermediate values have also been reported, usually ranging from 14-20% (Huang
1986; Jones & Rudloe 1995; Fisheries Victoria & MAFRI 2002; DEH 2006). Therefore,
scaling factors may range anywhere from 4 to 15. In addition, jellyfish begin to lose
weight after being caught, as water continuously drains from their tissues. If jellyfish
are compressed, such as being piled on top of each other in a ship’s hold, this
process will be accelerated (Jones & Rudloe 1995). As such, simply defining the
‘true’ weight of jellyfish is difficult, thereby complicating the use of the traditional
fisheries catch metric of “wet weight.”
If parts of the jellyfish have been discarded, such as the oral arms, the wet weight of
the catch will be underestimated in these cases, with perhaps 35-40% of the weight
of the whole jellyfish ‘missing’ (Subasinghe 1992). While this is a common practice in
some regions (e.g., Pakistan, Malaysia), other regions process the entire jellyfish (e.g.,
Mexico). Without knowledge of specific instances of partial discarding, this was not
taken into account and therefore the estimated wet weights remain conservative.
45
In instances where specific scaling factors were reported (e.g., Bahrain), the stated
values were used to calculate the wet weight. However, if no specific value was
reported, the most conservative scaling factor of 4 was used (e.g., India).
3.2. Results
At least 23 countries have been involved in fishing jellyfish for human consumption
(Table 4). Some countries, (e.g., Turkey) appear to have abandoned their jellyfish
fisheries, while others (e.g., Canada) had test fisheries that were unsuccessful. Many
countries do not explicitly report their catches to FAO, including them as either
“miscellaneous marine invertebrates” or not at all. As such, the global catch of
jellyfish reported by FAO is a dramatic underestimate of the true landings. In
contrast, FAO also reports catches for several countries and territories where there
do not appear to be jellyfish fisheries present. These include the Falkland Islands
(Malvinas), Namibia, and the United Kingdom (also see 3.3.24 Other countries). It is
likely that the reported catches from these countries are in fact jellyfish caught and
discarded as bycatch from other fisheries. As such, they were not included in this
catch reconstruction, although the total for all such reports was only 3,205 tonnes
combined, and so would not have dramatically affected the results. While it is
positive that FAO reports these catches – as discards should still be considered part
46
of the total catch – it would be beneficial if they were identified as bycatch and/or
discards to differentiate them from targeted landings.
Table 4. Countries known to fish jellyfish for human consumption
Country
Dates
China
<1950 - present
Japan
<1950 - present
Indonesia
<1950 - present
Malaysia
<1950? - present
Thailand
1970 - present
Philippines
1976 - present
Korea (South)
1980s? - present
Canada
1984; 2002
Turkey
1984 - 2006
India
1984 - present
Sri Lanka
1986 - present
Vietnam
1990s - present
U.S.A.
1993 - present
Australia
1995 - present
Myanmar
1995? - present
Mexico
2000 - present
Russian Federation
2000 - present
Bahrain
2004 - present
Pakistan
2007? - present
Nicaragua
2008; 2013 - present
Iran
2010? - present
Ecuador
2013 - present
Honduras
2013 - present
Reconstructed landings for the world are presented in Figure 2 (see Appendix A for
all catches). Landings are dominated by China, with high variability, typically
fluctuating between 100 and 400 thousand tonnes through the second half of the 20th
century. Catches from Thailand began to make a significant contribution to the total
catch starting in the 1970s, and other countries such as Indonesia, India, and
Vietnam increased their production through the 1990s. Many of the fishing locations
are in Southeast Asia (Figure 3). Landings of China’s primary target species,
47
Rhopilema esculentum, began to taper off in the late 1990s, but soon after, fishing for
the giant jellyfish Nemopilema nomurai quickly escalated, offsetting the declining
catches. Indeed, increasing landings of N. nomurai have resulted in the estimated
global catch exceeding 1 million tonnes for the first time in 2013. Such values
demonstrate that jellyfish fisheries are clearly significant, exceeding the catches of
jellyfish of other fisheries at a global scale, such as lobsters (Pauly & Zeller 2016a;
miscellaneous marine crustaceans excluded). For additional details and discussion
on the catch reconstructions in each country, refer to 3.3 Countries fishing jellyfish.
3.2.1. Estimating the contemporary global catch
There are at least 21 countries currently fishing for jellyfish (Table 4); however, the
fisheries in Ecuador, Honduras, and Nicaragua are new and still developing, and
the magnitudes of catches in those countries in the near future remains unclear. For
other countries, contemporary catches were calculated by averaging the
reconstructed catches over the last 10 years of data (2004-2013). Combining these
estimates yields a mean contemporary global catch of more than 750,000 tonnes
annually (Table 5), which is more than double the amount based on FAO statistics
for the same period. This figure is likely an underestimate of the global jellyfish
catch, as it does not include jellyfish that is landed as part of the vast quantities of
trash fish from nontargeted fisheries in China that are delivered to factories to be
48
Figure 2. Estimated global landings for two primary species in China and all species for other countries
0
5
10
15
20
25
0
200
400
600
800
1,000
1,200
1950 1955 1960 1965 1970 1975 1980 1985 1990 1995 2000 2005 2010
Number of Countries Fishing Jellyfish
Jellyifsh Landings (000's tonnes)
Year
Others
Vietnam
Indonesia
India
Thailand
China (N. nomurai)
China (R. esculentum)
FAO (Global)
Number of countries
49
Figure 3. Locations of jellyfish fisheries in Southeast Asia; dark blue indicates shelf (< 200 m); note
that there are likely many additional locations that have yet to be documented
turned into ‘fishmeal’ (Cao et al. 2015). In addition, the global catch reconstruction
presented here does not include any bycatch or discards of jellyfish, which can be
huge, often resulting in losses to fishers of ten or hundreds of millions of dollars
annually (Purcell et al. 2007; Uye 2008; Kim et al. 2012a).
China continues to be the dominant player in the jellyfish market, responsible for
more than half of the global contemporary catch. Thailand and India follow, with
50
Vietnam and Indonesia rounding out the top 5 nations targeting jellyfish, which
collectively make up more than 90% of the global contemporary catch (Figure 4).
While fishing for jellyfish has clearly expanded around the globe (Figure 5), the
overwhelming majority of the catch continues to take place in the waters of only a
few countries (Figure 4).
Table 5. Estimated contemporary annual jellyfish landings (2004 -2013 mean)
Country
FAO landings
(tonnes)
Reconstructed
landings (tonnes)
China
212,343
428,973
Thailand
75,663
140,786
India
-
72,405
Vietnam
-
42,530
Indonesia
10,852
22,820
Mexico
18,725
13,124
Bahrain
2,278
10,022
Malaysia
6,017
6,087
U.S.A.
740
5,100
Japan
-
4,266
Myanmar
2,722
2,722
Iran
2,588
2,588
Sri Lanka
-
2,511
Pakistan
-
1,663
Russian Federation
293
293
Philippines
19
179
Korea (South)
-
114
Australia
4
5
Total
332,243
756,189
51
Figure 4. Estimated global contemporary annual jellyfish landings (2004-2013) and reported FAO
catches (1950-2013)
0
100
200
300
400
500
600
700
800
1950 1960 1970 1980 1990 2000 2010
Jellyfish Landings (000's tonnes)
Year
FAO (Global)
Others
Indonesia
Vietnam
India
Thailand
China
52
Figure 5. Jellyfish fisheries around the world; red circles indicate magnitude category of catch in tonnes (see legend); circles are only
approximately representative of catch locations
53
3.3. Countries fishing jellyfish
3.3.1. Australia
Several attempts to establish jellyfish fisheries in Australia have been for the most
part unsuccessful, and the current scale of the fishery continues to remain small. In
1980-1981, a product and market feasibility study was carried out by the Clarence
River Fishermen’s Co-operative at Iluka in New South Wales (Davis 1982). Around
the same time, processing methods were being investigated in the School of Food
Technology at the University of New South Wales. Wootton et al. (1982) reported
that a satisfactory edible product could be produced from Catostylus mosaicus caught
in Botany Bay. Nearly 10 years later, a report was prepared by Georges (1991) for the
targeting of C. mosaicus in New South Wales. It was proposed that medusae with a
minimum bell diameter of 20 cm would be caught using surface trawl gear. Fishing
would take place between December and June in a region limited by Broken Bay
and the border with Queensland. Ironically, it was suggested that the proposal
would contribute to the conservation of C. mosaicus by facilitating research and
management of the species. The author also stated that fishing for jellyfish was
unlikely to have a measurable effect on the ecosystem and may help to mitigate
nuisance to fishers and swimmers. Despite apparent interest, along with a solitary
report of C. mosaicus being targeted in Lake Illawarra in 1990 (Kailola et al. 1993),
54
immediate development of the Australian jellyfish fishery did not proceed in New
South Wales. This may have been due to criticism of the “management program”
developed by Georges (1991). Wells & Wellington (1992) published a scathing
commentary of the report, calling it “a pathetic example of how information can be
manipulated for a particular purpose without apparent concern for scientific
accuracy…” Indeed, the authors highlight numerous inconsistencies and
unsubstantiated claims contained in the report, and stated that they “seriously
caution allowing such an operation to proceed without a full and complete
understanding of both the ecological and economic implications of the industry on
the marine environment.”
Subsequent to these attempts, renewed interest in establishing a jellyfish fishery in
New South Wales prompted additional research and a new management plan,
culminating in a trial fishery of 10 tonnes which was processed and exported to Asia
in 1996 (Hudson et al. 1997). Catches ranged between 10 and 33 tonnes from 1995 to
1998, apparently limited by processing capacity (Coleman 2004). However, a fishery
was not established in New South Wales, and attention shifted to the neighbouring
state of Victoria.
Hudson et al. (1997) report on surveys made in 1997 that estimated the biomass of
C. mosaicus in several locations within Port Philip Bay. The authors suggested that
55
biomass and densities were sufficiently high to support a fishery. Subsequent
studies were made in the years following, and in 1998 one company was granted a
developmental permit to catch jellyfish for 3 years, which was extended for an
additional year (Coleman 2004). Later it was recommended that the permit be
renewed for an additional 3 years. The Total Allowable Catch (TAC) is set between
1,000 and 1,200 tonnes annually, a conservative limit representing approximately 10-
15% of the estimated biomass (Fisheries Victoria 2006). However, only a tiny fraction
of this TAC is caught in Australia, representing 0-1%. Small TACs of 100 tonnes
were also set for 3 other locations (Fisheries Victoria & MAFRI 2002); however, it
appears that fishing for jellyfish has been limited to Port Philip Bay (Fisheries
Victoria 2006).
Fishers targeting jellyfish in Australia are required to use dip-nets, along with
limited use of seine nets for corralling purposes only. Specific permits for catching
jellyfish are required, and fishers targeting other species are prohibited from
retaining jellyfish as bycatch, and are instead required to return them to the water
with the least possible damage (Fisheries Victoria 2006). A minimum size limit of
23 cm (bell diameter) is well above the 13 to 16 cm at which maturity is generally
achieved for the species (Pitt & Kingsford 2000). While a minimum catch of 150
56
tonnes was proposed for 2006 in order to encourage at least a minor level of activity
(Fisheries Victoria 2006), catches have continued to remain low.
With the exception of catches of 14 and 11 tonnes in 2008 and 2013 respectively,
catches have been 0, 1, or 2 tonnes since 1998 (FAO 2015a). Despite the potential for
a jellyfish fishery with annual catches of several thousands of tonnes (Fisheries
Victoria 2006), the jellyfish fishery in Australia does not appear to be expanding. As
mentioned, up to at least 2005, only a single development permit for fishing jellyfish
was being issued in the State of Victoria (Fisheries Victoria & MAFRI 2002). It is
unclear why the jellyfish fishery in Australia remains undeveloped despite
investments of approximately $500,000 (AUD) in equipment and personnel
(Fisheries Victoria & MAFRI 2002). Potential factors involved include poor
recruitment in some years (Fisheries Victoria & MAFRI 2002), economic viability,
and possibly onerous management regulations. The product resulting from
processed C. mosaicus is reportedly not as large or firm as jellyfish products from
other regions (Hudson et al. 1997), resulting in a ‘B-grade’ labelled product.
There have been attempts at exploiting jellyfish in several other locations in
Australia. Around 1989-1990, Phyllorhiza punctata was apparently being considered
for possible capture, processing, and export in the Swan River Estuary in Western
Australia near Perth; however, a fishery was not established (Coleman et al. 1990;
57
Kailola et al. 1993). In the Northern Territory, a test fishery began in 2000 to catch
C. mosaicus in the estuarine western section of the Gulf of Carpentaria, with a goal of
expanding processing capacity to 300 tonnes annually (Field 1999); however, it
remains unclear if this goal has been achieved. Fishing of C. mosaicus was also
proposed in Queensland, outlined in a 2006 report (DEH 2006). Developmental
permits were issued in 2006 for a total of 800 tonnes, divided between Moreton Bay
(200 tonnes), Tin Can Bay (200 tonnes), and the Gulf of Carpentaria (400 tonnes).
However, it remains unclear if any of this allowable catch was ever caught, and the
export permit for the fishery is currently expired.
Catches reported by FAO for Australia appear to be reasonably accurate, with a few
exceptions. As usual, the reported taxon is incorrect (Rhopilema spp. instead of
Catostylus mosaicus), and the reported FAO area where fishing occurs is strangely 58,
‘Indian Ocean, Antarctic’, rather than 57, ‘Indian Ocean, Eastern’ (Figure 6).
3.3.2. Bahrain
Bahrain apparently began fishing jellyfish around 2003 or 2004 (Mohammed 2008;
FAO 2015a). The target species is suspected to be Catostylus perezi, as this species is
abundant in the Persian Gulf and Arabian Sea (Gul & Morandini 2013; Gul et al.
2015), is the target of fisheries in nearby Pakistan (Gul et al. 2015), and resembles the
58
Figure 6. Major FAO areas
59
jellyfish being processed in photographs from Bahrain (see Mohammed 2008).
Apparently, part of the motivation to start a jellyfish fishery was to reduce stings to
fishers targeting finfish stocks, as well as to prevent them “scaring off fish”
(Mohammed 2008). The method used to catch jellyfish has not been reported, but
fishing appears to be doneby fishers who traditionally targeted finfishes (and still
do), with a jellyfish season from April to September (Mohammed 2008).
In 2007, a single processing company in Bahrain was reported to export 60
containers of processed jellyfish, each containing between 18 and 24 tonnes
(Mohammed 2008). If an average of 21 tonnes per container is assumed, this is
equivalent to 1,260 tonnes of processed jellyfish. As reported by Mohammed (2008),
processed jellyfish is 15% of its wet weight if the product is bound for China, or 8%
if bound for Japan. As the product is reportedly more popular in China, and since it
is cheaper to produce the product with higher water content, it was assumed that
two thirds of the product produced in 2007 was exported to China, with the
remaining third exported to Japan. This equates to 10,850 tonnes of jellyfish caught
by Bahrain in 2007, which is much larger than the 1,759 tonnes reported by FAO for
Bahrain in the same year. Interestingly, values quoted in the news article by
Mohammed (2008) for exports of processed jellyfish for 2005 and 2006 match almost
exactly those reported by FAO as landings. Therefore, it appears likely that the
60
values reported by FAO for jellyfish landings in Bahrain are actually for processed
jellyfish, and were similarly scaled up.
Catches apparently declined in Bahrain in 2008, and although it has been suggested
that this could be due to turbidity from land reclamation (Mohammed 2008), other
speculation points to the same processes having positive effects for jellyfish by
keeping predatory turtles away from the coast (Anonymous 2008a). According to
FAO (2015a), catches have continued to fluctuate since, with a relatively larger catch
in 2012. Bahrain’s jellyfish fishery appears to be persisting, with variable catches.
3.3.3. Canada
Canada has explored fisheries for jellyfish on the Atlantic and Pacific coasts.
However, both test fisheries did not continue, predominantly due to the fact that
they targeted Aurelia spp., a semaeostome for which there is limited demand.
Fisheries and Oceans Canada (also known as the Department of Fisheries and
Oceans, or DFO) explored the possibility of a fishery for Aurelia labiata in coastal
British Columbia in 1984. Sloan & Gunn (1985) present details for 11 dip-net and 2
seine fishing cruises conducted between August and November in the northern
Strait of Georgia. The total catch was 2.82 tonnes, which was then processed using 3
different protocols from potential Japanese buyers. Samples were provided to
61
Chinese fish wholesalers and to Japanese and Chinese restaurateurs in Vancouver.
The product was deemed unsuitable, based mainly on the poor texture that lacked
the preferred crunch. Ultimately, the test fishery for jellyfish in British Columbia did
not continue.
On Canada’s east coast, jellyfish frequently interfere with active and passive fishing
gears, making a targeted fishery for jellyfish desirable (DFA 2002a). As such, a test
fishery was implemented to understand the methods and costs involved in
producing jellyfish, and to evaluate the potential market (DFA 2002b). An estimated
49 tonnes of jellyfish were caught over a period of 2 weeks in September 2002 in
Newfoundland’s Trinity Bay; however, only about 1 tonne was retained, with the
rest being released at sea (DFA 2002a). A 50-foot shrimp beam trawl was used,
towed at approximately 1 knot. Catches consisted of approximately 90% Aurelia sp.
and 10% Cyanea capillata, with the latter reportedly being too delicate to handle. The
subsample of Aurelia retained for processing was stored onboard the ship in an
insulated container containing a slurry of slush ice and 1% alum. About 1.1 tonnes of
jellyfish were processed and samples were sent to China, Taiwan, and Florida,
U.S.A. for market testing (DFA 2002a). Due to a lack of demand for semaeostome
jellyfish, as well as unrefined handling and processing techniques, the test fishery
was discontinued.
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3.3.4. China
China has the longest history of fishing for jellyfish, dating back at least 1,700 years
(Omori & Nakano 2001), and is the world’s largest producer of jellyfish. China is
also the only country that farms jellyfish in aquaculture ponds (You et al. 2007), as
well as releasing juvenile medusae into the wild as part of a hatchery program
(Dong et al. 2009; Tian et al. 2016). Jellyfish grown in aquaculture operations were
not included as part of this analysis; however, production has averaged just over
50,000 tonnes annually for the period 2003-2014 (FAO 2016) and may increase in the
future if catches of the primary species from the wild continue to decline (see
below). There have been seasonal bans on trawling in Liaodong Bay in the Bohai Sea
since the 1980s with the intention of protecting polyp beds; however, some trawling
is allowed at certain times of year which may damage polyps and their habitat (Ye
2006). The primary species of interest in China is Rhopilema esculentum, as it is the
most in demand and the most valuable. However, declines in abundance,
presumably due to overfishing (Dong et al. 2014), along with increasing demand for
jellyfish, have resulted in the targeting of other species, including the giant jellyfish,
Nemopilema nomurai. China does report jellyfish catches to FAO; however, these
appear to be restricted to R. esculentum. In addition, catches appear to be
63
underreported in some years, resulting in dramatic underestimates for jellyfish
landings in China, and hence globally.
R. esculentum is targeted in nearly all of coastal China, including in the Bohai,
Yellow, East China, and South China Seas. Marine biologist Jack Rudloe travelled to
China in 1995 to learn about jellyfish fishing and processing in support of
establishing a fishery for cannonball jellyfish Stomolophus meleagris in the U.S.A. (J.
Rudloe, Gulf Specimen Marine Laboratory, pers. comm., April 2014). While visiting
cities including Qingdao, Weihi, Yantai, and Laizhou, he learned that many men
and women travelled from the south coast of China and followed the migrating
blooms of R. esculentum, catching and processing them at different locations along
the way. In addition, a fleet of 200 government-issued 50-foot wooden boats with
diesel engines fished the Gulf of Bohai everyday. Fishers left in the morning and
travelled 6 hours to reach the fishing grounds. Trawls or set-nets were fished
overnight for another 6 hours, and then catches were rushed back to the docks in
order to meet afternoon processing schedules. Omori (1981) also reports information
on fishing grounds and seasons in Chinese waters. You et al. (2016) describe a
dramatic increase in the number of fishing boats in Liaodong Bay in recent decades,
with a concomitant reduction in the length of the jellyfish fishing season. Prior to the
1980s, there were hundreds of boats and the season lasted approximately two
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months. This changed to several thousand boats in the 1980s, with a season of 1-2
weeks. By the 1990s, there were 8-9 thousand boats with a season lasting only 2-3
days. Today, there are more than 10 thousand boats fishing in Liaodong Bay,
resulting in a derby fishery that usually catches all of the available jellyfish in a
matter of hours. Consequently, this has led to regular instances of illegal fishing
(You et al. 2016; also see 2.4 Management).
Landings reported to FAO extend back to 1970, and averaged about 25,000 tonnes
through the 1970s and 1980s. Dong et al. (2014) present a useful summary of
R. esculentum in Chinese waters, along with catch statistics dating back to 1957,
gleaned from several sources including China Fishery Statistical Yearbooks. Up until
1997, the statistics presented are very consistent with those reported by FAO.
However, for the period from 1997-2006, it appears that statistics reported by FAO
were curiously underreported by approximately 15%. A more startling discrepancy
emerged with the publication by Li et al. (2014), which reports landings of
R. esculentum from 1980 to 2012. For the period 1980-1990, it appears that landings
reported by FAO were for processed jellyfish, rather than wet weight. As such,
landings reported for this period are only 15% of what they should be. Put another
way, these values are underestimated by more than 600%. Values reported by FAO
for the 1970s, and by Dong et al. (2014) for the 1960s and late 1950s are of the same
65
order of magnitude as those for the 1980s. Therefore, it is likely that all values
reported before 1991 are likely for processed jellyfish, and represent only 15% of the
true catch. Adopting this assumption, we find that catches of jellyfish in Chinese
waters have fluctuated between about 100,000 and 300,000 tonnes since 1950. While
this assumption is obviously very significant if it ultimately proves to be false, it
seems justified given that contemporary landings exceed these values, and there is a
long history of fishing for jellyfish in China. Indeed, there are even indications that
these scaled-up values may be underestimates, as Kingsford et al. (2000) notes that
one report claims a peak jellyfish catch in China of 700,000 tonnes in a single year.
As yet, that figure remains unconfirmed by other sources. There are also other major
discrepancies in Chinese catch statistics for jellyfish. For example, FAO (2015a),
Dong et al. (2014), and Li et al. (2014) all report landings of approximately 96,000
tonnes in 1991. In contrast, You et al. (2016) report a catch of 296,050 tonnes in 1991
for Liaodong Bay alone. As this value could not be verified, the more conservative
estimate was adopted; however, this is yet another example illustrating that the
reconstruction herein likely represents a minimum estimate, and landings of
jellyfish may be much higher in some years.
Catches of R. esculentum appear to have declined rapidly in the mid-1970s, which
has been attributed to overexploitation (Dong et al. 2014). This led to extensive
66
research on the culturing of R. esculentum, and in 1984, juvenile medusae were
released in the waters of Liaodong Bay with the hopes of enhancing the stock. This
hatchery program has continued each year, and has since expanded to include the
release of hundreds of millions of juvenile medusae into the coastal waters of
Liaoning and Shandong Provinces (Dong et al. 2014).
While the hatchery program has likely helped to enhance catches of R. esculentum in
Chinese waters (Dong et al. 2014), another rhizostome, the giant jellyfish Nemopilema
nomurai, has increasingly been a target for fishers in recent years. This species
appears to have increased in abundance in the Yellow and East China Seas in the
last decades (Yoon et al. 2008; Dong et al. 2010). Industrialized fishing for N. nomurai
reportedly began around 2000, with catches averaging about 125,000 tonnes until
2008 (J.-H. Cheng, East China Sea Fisheries Research Institute, pers. comm., Nov.
2013). Catches since 2009 have exceeded 200,000 and even 300,000 tonnes in some
years, with the most dramatic catch of more than half a million tonnes in 2013 (Li et
al. 2014). Combined with the catch of R. esculentum, this means that China landed
more than 750,000 tonnes of jellyfish in 2013. Curiously, catches of N. nomurai do not
appear to be reported to FAO at all.
There are also other species of jellyfish targeted for food in China, including
Lobonema smithi, Lobonemoides gracilis, Rhopilema hispidum, and Cyanea nozakii (see
67
Tables 1 and 2 for sources); however, the catches of these species are presumably
quite small in comparison to the two aforementioned species.
3.3.5. Ecuador
In 2013, Chinese dealers began promoting the possibility of catching, processing,
and exporting jellyfish (presumably Stomolophus meleagris) from Ecuadorian waters.
Shellfish fishers, who have been struggling to generate sufficient income, welcomed
the proposal. Approximately 100 small (~10 m) fiberglass and wooden boats began
fishing for jellyfish using modified gillnets and set-nets within and around the
Guayaquil Gulf Estuary. In 2014, an astounding 78,000 tonnes of jellyfish were
landed (most of which was caught in February and March), processed, and exported
to China, Japan, and Thailand (M. Perciado, Instituto Nacional de Pesca, pers.
comm., Jan. 2015). If such significant landings are maintained, it would catapult
Ecuador to being the 3rd largest producer of jellyfish worldwide (Table 5). While
studies are currently underway to evaluate the impacts of the fishery and to
establish management regulations, the fishery was completely closed from May to
September in 2014, as processing facilities were shuttered due to a lack of
environmental oversight. Many processing plants remained closed in 2015 as they
did not acquire the necessary permits, and the catch for 2015 was reduced to 9,135
tonnes as buyers turned their attention to importing jellyfish from Mexico (E. Laaz,
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Instituto Nacional de Pesca, pers. comm., Aug. 2016). It would seem that the future
scale of Ecuador’s jellyfish fishery remains to be determined.
3.3.6. Honduras
In 2007, samples of a species of Stomolophus (likely S. meleagris) were caught along
the Atlantic coast of southern Honduras (near Cauquira) using small boats with dip-
nets to test for exploitation potential. Ultimately, fishing grounds off Nicaragua
were selected for a test fishery in 2008 instead of Honduran waters (see 3.3.15
Nicaragua). This was primarily due to the fact that the region of Honduras is not
easily accessible by road, and any processing equipment and supplies have to be
brought into the port by boat. Such logistics proved to be inefficient for a product
that is relatively low-value, and therefore the future potential of a jellyfish fishery in
Honduras remains uncertain. However, FAO does report a catch of 50 tonnes for
Honduras in 2013, so perhaps some of the processing capacity concerns have been
addressed.
3.3.7. India
India is yet another country that has been catching jellyfish for decades, but is absent
from FAO jellyfish catch statistics. Fishers in numerous coastal states have been
actively involved in catching jellyfish including Tamil Nadu, Andhra Pradesh,
Odisha, and West Bengal. Fisheries for jellyfish have also been explored in the
69
Arabian Sea along India’s west coast, including the states of Karnataka and Gujarat,
and likely others.
Several species of jellyfish were initially reported to be targeted for processing and
export from India, including Rhizostoma sp. (Chidambaram 1984) and Aurelia sp.
(Govindan 1984). However, it is now believed that Crambionella stuhlmanni is the
primary species being exported (Kuthalingam et al. 1989; CMFRI 2009; Mohan et al.
2011), with Lobonema smithi also being targeted or caught and sold as bycatch
(Murugan & Durgekar 2008). In addition, Rhopilema esculentum is reported from
Indian waters (Panda & Madhu 2009), so it is likely that this valuable species is also
being exploited.
India’s jellyfish fishery presumably began in the 1980s. In 1984, at least 21 tonnes of
processed jellyfish were exported from India (Chidambaram 1984). This was
primarily in response to requests from the Japanese External Trade Organisation
(JETRO), which toured India in February 1984 and offered to import “any amount”
of jellyfish from India (Govindan 1984). This was a welcome proposition, as jellyfish
were a persistent nuisance to fishers in India due to clogged nets, spoiled catch,
fouled gear, and stings (Govindan 1984; James et al. 1985; Kuthalingam et al. 1989).
Several other countries were also reported to import jellyfish from India in the 1980s
70
including Thailand, Hong Kong, and Singapore (Chidambaram 1984; Kuthalingam
et al. 1989).
Exports from India to Japan continued sporadically through the 1980s, 1990s, and
2000s, with the annual tonnage of processed jellyfish product ranging from zero to
hundreds of tonnes (Omori & Nakano 2001; S.-I. Uye, Hiroshima University, pers.
comm., Oct. 2012). However, India’s total catch of jellyfish was more likely in the
hundreds of thousand of tonnes at its peak in the mid-2000s. The 2004 Handbook of
Fisheries Statistics from the Government of India (Anonymous 2005) indicates that
between 2000 and 2004, production of jellyfish commodities ranged between 7,723
and 30,866 tonnes, with an annual average of 22,515 tonnes. As the commodity is
reported as “Jellyfish, dried, salted or in brine” it was assumed that these values
represent semi-dried, processed jellyfish. As such, the values need to be scaled up to
obtain the original wet weight of the catch. Without knowing a specific scaling factor
for the species and processing method used, a conservative value of 4 was used (see
3.1.2 Scaling factor).
Even the limited statistics from the government Handbook are expected to be
underestimates of India’s total jellyfish catch due to non-reporting from many states.
For example, in 2000 and 2001, only the state of Gujarat reported landings of
jellyfish, whereas the totals for 2002 and 2003 include landings from Andhra
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Pradesh, Karnataka, and West Bengal. Jellyfish are also caught in other states, such
as Odisha (Anonymous 2007) and Tamil Nadu (Kuthalingam et al. 1989). Therefore,
the true magnitude of India’s jellyfish catch is likely much higher in some years.
Murugan & Durgekar (2008) report a catch of 7 million tonnes of jellyfish from the
Gulf of Mannar in 2007 alone; however, this figure is assumed to be erroneous as it
dramatically eclipses the entire global catch of jellyfish. Attempts to obtain
clarification of the estimate were unsuccessful, and therefore it was ignored in the
catch reconstruction.
The rapid expansion of jellyfish fisheries in India in the 2000s appears to have
resulted in some conflict in the region. Processing facilities are often makeshift tents
or huts that are temporary or abandoned between jellyfish fishing seasons
(Kuthalingam et al. 1989). Magesh & Coulthard (2004) chronicle how the number of
these huts quickly grew in the early 2000s, resulting in tension between
stakeholders. As middlemen from different regions and countries moved in to erect
the huts, local fishers felt left out of the industry and unable to benefit from the
export of jellyfish. Suspected pollution from the effluent of the processing huts
exacerbated the situation further, and in some cases, huts were destroyed. The
tension peaked after rapid expansion and very large landings in 2003. In a twist of
fate, the fishery in Tamil Nadu was completely wiped out by the devastating
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tsunami in 2004 (CMFRI 2009). Depending on the circumstances, processing of
jellyfish can be an important supplementary employment for fishers, and it has been
reported that more than one third of all fishers operating motorized boats in Tamil
Nadu were seasonally involved in the processing of jellyfish (Swathilekshmi 2011).
As with most jellyfish fisheries, landings in India are highly variable. As such, years
with high catches are often referred to as “unusual.” Examples include 2003 in Tamil
Nadu (CMFRI 2009), 2007 in Andhra Pradesh (CMFRI 2007), and 2009 in Tamil
Nadu (CMFRI 2009; Mohan et al. 2011). Interestingly, in 2009, fishers in northern
Tamil Nadu targeted only the oral arms, disposing of the bells at sea (Mohan et al.
2011).
Information on jellyfish fisheries along India’s west coast is even sparser. In Jakhau,
Gujarat, about 75 odis (fiberglass boats with inboard engines) were engaged in
gillnet fishing for jellyfish in 2010 (CMFRI 2010). Each boat has a capacity of
approximately 1 tonne per day, and there are two fishing seasons each year (April-
May and November-December). Landings from one season in 2010 in Jakhau were
approximately 800 tonnes, up from 250 tonnes in 2009 (CMFRI 2010).
There are also suggestions that exploitation of jellyfish in India is negatively
affecting olive ridley sea turtles (Lepidochelys olivacea) by reducing their prey
73
(Anonymous 2007; 2008b). This can also lead to conflicts of interest regarding the
mandates of different government departments (Murugan & Durgekar 2008).
More recently, it appears that many processing facilities for jellyfish in India have
closed due to a lack of buyers for the exported product (T. Vaidyanathan, University
of British Columbia, pers. comm., Aug. 2015). Fishers continue to catch large
amounts of jellyfish as bycatch, especially in shore seines. However, whereas this
bycatch was formerly processed and exported in the 2000s, it is now typically
discarded. It is unclear how much the country’s landings of jellyfish have declined,
as there were reportedly still processing facilities in some regions as recently as 2010
(e.g., CMFRI 2010); but it is assumed that recent catches in India are likely an order
of magnitude lower than the peak in the mid-2000s.
3.3.8. Indonesia
Indonesia is one of the world’s largest producers of jellyfish behind China and
Thailand, and targets jellyfish in many locations (see Figure 3). There is a long
history of jellyfish capture in Indonesia, with annual catches extending back to 1950,
when fisheries statistics were first tabulated (FAO 2015a). Despite this history,
catches of jellyfish in Indonesia are highly variable, with landings ranging in the
hundreds of tonnes in some years all the way up to a peak of 123,000 tonnes in 1995.
While this variability is likely a reflection of abundance, it may also have to do with
74
cultural misunderstandings and stakeholder relationships. Rudloe (1992) reported a
lack of trust between the Indonesian sellers and Chinese buyers, and noted that
“shipments of processed jellyfish sometimes have good product on top of the
batches and poor quality on the bottom.”
Indonesia reports jellyfish catches for both of its FAO areas (57, ‘Indian Ocean,
Eastern’ and 71, ‘Pacific, Western Central’; see Figure 6). However, there are other
sources of data that indicate the reported catch statistics are not accurate in some
years. For example, the Southeast Asian Fisheries Development Center (SEAFDEC)
has published Fishery Statistical Bulletins since 1978. While jellyfish catch statistics
are not available for every Southeast Asian country for every year, there are
numerous years where either catch statistics or export statistics (or both) are
available (see SEAFDEC 2015). Data on production of processed jellyfish are also
available from FAO for some years (see FAO 2015b). While export data was rarely
used in this analysis due to the possibility of importing and re-exporting, the export
statistics reported by SEAFDEC are identical to those reported as production of
processed jellyfish by FAO for the years where they are given. Therefore, it is
reasonable to assume that the export statistics reported by SEAFDEC for the period
1976-1990 represent jellyfish that were caught and processed in Indonesia. A recent
publication by Asrial et al. (2015a) also indicates that catches in Saleh Bay, Sumbawa
75
Island have exceeded 30,000 tonnes since 2010, an amount larger than the catches
reported by FAO in some years for the entire country, further suggesting
underreporting.
Numerous species of jellyfish are targeted in Indonesia. According to Kitamura &
Omori (2010), Lobonemoides robustus is caught in the Strait of Malacca along the east
coast of Sumatra from Medan to Lampung and Bangka Island. This species is also
targeted in the Makassar Strait off southeastern Kalimantan (Indonesian Borneo)
and along the north coast of the island of Java. The estuarine Acromitus hardenbergi is
caught in Sumatra in Tanjung Balai. Fishing for jellyfish also occurs along Java’s
southern coast (Nishikawa et al. 2009). A newly described species, Crambionella
helmbiru, is fished in Karangbolong and Cilacap from August to November (Mujiono
2010; Nishikawa et al. 2015). Further east, Crambione mastigophora is targeted near
Prigi and Muncar (Omori & Nakano 2001; Kitamura & Omori 2010), as well as in
Selah Bay, Sumbawa Island from October to December (Asrial et al. 2015a; b).
Rhopilema hispidum is also fished along the north (near Cirebon) and south (near
Kotabaru) coasts of Java, as well as southeastern Borneo. Fishing for jellyfish has
also been reported from Bacan Island for a yet to be described species known as the
‘Semi-China type,’ which is said to be similar to Rhopilema esculentum, but smaller
(Omori & Nakano 2001; Kitamura & Omori 2010). Several other locations in
76
Indonesia have reportedly caught jellyfish in the past, such as Nusa Tenggara, West
Sumatra, and Sulawesi (SEAFDEC 2015); however, these areas were not marked
explicitly on Figure 3, as specific locations where jellyfish are caught within these
regions could not be determined. Typical gears include dip-nets and drift nets
(Nishikawa et al. 2009) that are fished from outboard-powered boats with crews of 2
to 5 and capacities of 1 to 5 tonnes (Mujiono 2010).
3.3.9. Iran
Details of the jellyfish fishery in Iran are virtually nonexistent. Catches are reported
by FAO (2015a) starting in 2010, with landings averaging 3,451 tonnes annually
between 2010 and 2012. Target species are unknown; however, Catostylus perezi is
suspected to be the primary species targeted in neighbouring Pakistan (Gul et al.
2015), and possibly Bahrain, so it is likely that C. perezi is also caught in Iran. López-
Martinez & Álvarez-Tello (2013) report 300 tonnes of processed jellyfish products
exported to China from Iran in 2012, which likely represents the majority of Iran’s
catch.
3.3.10. Japan
Japan has a long history of targeting jellyfish, but has never explicitly reported
jellyfish catches to FAO. Omori (1981) notes that there are records of jellyfish fishing
in the Seto Inland Sea dating back to the late 1800s. Omori (1978) reported that
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Nemopilema nomurai is sometimes caught off the Hokuriku coast in the Sea of Japan,
and that Rhopilema esculentum is occasionally caught along the coasts of western
Japan. Omori (1981) also reports jellyfish fishing in Wakasa Bay, Mutsu Bay, as well
in the Sea of Japan off Tōhoku. Most of these reports are from the 1970s, and it
remains unclear if Japan continues to fish for jellyfish in these regions today. The
Japanese sea nettle, Chrysaora pacifica, has also reportedly been caught for food in
Japan (Morikawa 1984).
One region where fishing for jellyfish has persisted is the Ariake Sea, the largest bay
in the southern island of Kyūshū. Catches primarily consist of Rhopilema esculentum
and to a lesser extent, Rhopilema hispidum (S.-I. Uye, Hiroshima University, pers.
comm., Sept. 2012). Historically, catches were estimated to total about 1,000 tonnes
per year (Morikawa 1984), but processing in the region became more industrialized
in 1977 as jellyfish began to bloom in much higher abundances, and catches of 18,000
and 10,000 tonnes followed in 1978 and 1979 respectively (Omori 1981). However,
soon after 1979, jellyfish populations in the Ariake Sea returned to lower
abundances, and a few hundred tonnes were caught each year and sold in the local
fish market (S.-I. Uye, Hiroshima University, pers. comm., Sept. 2015). Similar
conditions persisted for several decades until 2011, when large jellyfish blooms
78
again led to increased capacity and landings exceeding 10,000 tonnes annually to at
least 2013.
Due to the recent increased proliferation of Nemopilema nomurai in the region
(Kawahara et al. 2006b; Uye 2008), one would expect that this species is also being
caught in Japan. While demand for N. nomurai appears to be relatively low in the
Japanese market, there have been recent attempts to develop them as an additive to
various foods such as candy, cookies, and ice cream (Anonymous 2006; Simpson
2009). Despite this, there are no available records for landings of N. nomurai in Japan.
Notwithstanding the fact that Japan produces and exports jellyfish, it also imports
jellyfish in even greater quantities. From the late 1980s through the 1990s, imports of
jellyfish to Japan averaged about 10,000 tonnes of processed product annually
(Omori & Nakano 2001). Through the 2000s, the average dropped slightly to around
8,000 tonnes annually (Information Office, Tokyo Customs House, data courtesy of
S.-I. Uye, Hiroshima University, Oct. 2012).
3.3.11. Korea (Republic of)
The Republic of Korea, also known as South Korea and hereafter referred to as
Korea, does not explicitly report catches of jellyfish to FAO. However, Omori (1981)
reports that fishing for jellyfish occurred historically in southwestern Korea, near
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Mokpo, insinuating that fishing for jellyfish in Korea may actually extend back for
centuries, rather than just decades. However, no additional information is readily
available on the timing or scale of this fishery. More recent fishing for jellyfish in
Korea also occurs along the southwestern coast near Muan and Gunsan (Omori
1981) as well as further north near Ganghwa and Ganghwado islands in Incheon
(C. Han, National Fisheries Research and Development Institute, pers. comm., Oct.
2015). Ullah et al. (2015) identified the species as Rhopilema esculentum. Fishing
typically occurs between September and November (Omori 1981). Catches appear to
be relatively small compared to other Asian countries, with an average of 31 tonnes
of processed jellyfish products produced annually between 1988 and 2011 according
to trade production statistics (FAO 2015b). However, López-Martinez & Álvarez-
Tello (2013) report 600 tonnes of processed jellyfish products exported to China from
Korea in 2012, suggesting that Korea’s jellyfish catches may be in the thousands of
tonnes, rather than hundreds.
3.3.12. Malaysia
Malaysia is one of the world’s largest producers of jellyfish behind China, Thailand,
and Indonesia. Interestingly, in some locations in Peninsular Malaysia, edible
jellyfish are available year-round (Yusoff et al. 2010; Khong et al. 2016), which is
relatively rare. FAO catch statistics for Malaysia start in 1981; yet Rudloe (1992) as
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well as Omori & Kitamura (2011) reported that commercial fishing for jellyfish
actually started in 1969. However, Nishikawa et al. (2009) and Yusoff et al. (2010)
indicate that fishing for jellyfish was introduced by the Japanese as early as the
1940s in Penang, and spread to Bagan Datoh by the 1950s. Khong et al. (2016) also
state that the “jellyfish fishery in Malaysia is a century-old industry that has been
carried on for at least three generations.” It is difficult to estimate catches through
the 1950s and 1960s, but operations were presumably small, with landings in the
hundreds of tonnes. Catches reported by the Southeast Asian Fisheries Development
Centre (SEAFDEC) extend back to 1976; however, there are indications that these
values may be underestimates. For example, SEAFDEC (2015) reports a catch of 441
tonnes of jellyfish for Malaysia in 1979, whereas Omori (1981) reports a catch of
3,400 tonnes for Malaysia in the same year. Ironically, SEAFDEC (2015) also reports
exports of 604 tonnes of processed jellyfish by Malaysia in the same year, which
suggests that the estimate by Omori (1981) is probably accurate. Regardless, capture
statistics reported for Malaysia by FAO (2015a) and SEAFDEC (2015) are consistent
after 1982, suggesting accuracy, with the exception of accounting for illegal,
unregulated, and unreported catches (IUU).
Fishers in Malaysia catch jellyfish in both the Indian and Pacific Oceans (see Figure
3). The major fishing grounds and target species are outlined by Kitamura & Omori
81
(2010). Lobonemoides robustus is targeted in the Straits of Malacca off the Malaysian
Peninsula, mainly between Bagan Datoh and Kuala Selangor, as well as near Klang.
The estuarine Acromitus hardenbergi is also caught near Bagan Datoh, Teluk Intan,
and the island of Pangkor, often using set-nets to take advantage of the tidal
currents (Nishikawa et al. 2009). At least one other unidentified species of jellyfish
known as the ‘Semi-China type’ is also caught in the Straits of Malacca, with fishing
grounds near Penang and Teluk Intan (Omori & Nakano 2001). Rhopilema hispidum is
targeted on the southern peninsula near Kukup, Johor (Nishikawa et al. 2009; Khong
et al. 2016) where the oral arms are often discarded at sea (Yusoff et al. 2010). Fishing
for jellyfish also occurs along the eastern peninsula near Kota Bharu (Omori 1981)
and Terengganu, although catches there are reportedly highly variable, with the
market being nearly saturated in 1986, followed by very low production in the years
following (Rudloe 1992).
Jellyfish are also caught in Malaysian Borneo, with several locations in Sarawak. In
fact, the relatively large catch reported by Malaysia in 1997 of more than 50,000
tonnes may have been due primarily to catches from Sarawak, which were
apparently 49,665 tonnes that year (Abu Talib et al. 2003). Detailed information on
jellyfish fisheries in Sarawak is presented by Rumpet (1991) based on visits to the
region in 1998. Interviews with jellyfish fishers suggested a possible decline in catch
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per unit effort (CPUE) through the 1980s, suspected to be due to a combination of
factors including increased demand incentivising more fishers, as well as lower
abundances of jellyfish. The primary target species, known as the ‘white type,’ is
generally caught in Southern Sarawak and is suspected to be L. robustus (Kitamura &
Omori 2010; Rizman-Idid et al. 2016). A secondary species, the ‘red type,’ is mostly
caught further north near Matu in the Division of Mukah, and is potentially
Rhopilema esculentum (Omori & Nakano 2001; Rizman-Idid et al. 2016). The primary
gears used to catch the white type are scoop nets, hooks, and trawl nets, while the
red type is often targeted using drift nets, bag nets, set-nets, and trawl nets (Rumpet
1991). Lobonema smithi is also targeted in Malaysia, although apparently not at
commercial scale (Khong et al. 2016).
3.3.13. Mexico
Development of Mexico’s jellyfish fishery began in 2000 in the Gulf of Mexico off the
state of Tabasco, targeting cannonball jellyfish Stomolophus meleagris. However, the
fishery shifted to the Gulf of California in 2001, principally to the shallow coastal
waters along the state of Sonora. A summary of the fishery is provided by López-
Martinez & Álvarez-Tello (2013). Average annual catches are 10,000 to 15,000 tonnes,
but may vary from 1,000 tonnes to a peak of more than 30,000 tonnes in 2015. The
fishery started relatively small, with about 70 small boats (‘pangas’), each with a
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crew of 2 or 3 fishers dip-netting for jellyfish. In 2010, management measures were
approved that would set a minimum size limit (MSL), restrict gears, and limit
fishing effort, among others. However, the scale of the fishery continued to escalate,
partially due to a lack of enforcement, and in 2013, over 1,000 pangas fished for
jellyfish, with the season lasting only 5 days. In response, additional management
measures and enforcement are currently being investigated and implemented, and
while catches continue to remain relatively large, the future of Mexico’s jellyfish
fishery remains uncertain.
3.3.14. Myanmar
Available information on Myanmar’s jellyfish fishery is scant. FAO (2015a) reports
catches commencing in 1995 and continuing to the present, with a mean annual
catch of just over 2,000 tonnes. However, in 2005, FAO reports the exact same value
for the wet weight of jellyfish caught – 1,976 tonnes – as it does for exports of semi-
dried processed jellyfish from Myanmar (FAO 2015a; b). FAO export and catch data
appear to be more reconcilable for 2008-2011, although export data from SEAFDEC
(2015) suggests exports were 30-80% higher than production for the period 2002-
2006. The possibility of importing and re-exporting is possible, and as such, only the
more conservative capture values reported by FAO (2015a) and SEAFDEC (2015)
were used in the catch reconstruction.
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At least several species of jellyfish are caught in Myanmar. Lobonemoides robustus is
targeted in the Bay of Bengal, both in the north in Rakhine State and in the south
near Myeik (Kitamura & Omori 2010). Crambionella annandalei is targeted near
Sittwe, where Rhopilema hispidum also occurs (Omori & Nakano 2001).
3.3.15. Nicaragua
In 2008, 205 tonnes of jellyfish (a species of Stomolophus, most likely S. meleagris)
were caught and processed in Tuapi, near the city of Puerto Cabezas on Nicaragua’s
Atlantic coast. There were approximately 34 small wooden and fiberglass boats
involved in the fishery, with a typical capacity of about 1.5 tonnes. Fishermen used
dip-nets with a 2-inch mesh size. Bells and oral arms were processed separately,
yielding 57 tonnes of processed jellyfish that was exported to Asia. The fishery did
not continue in subsequent years, potentially due to a combination of inferior
product quality and regulatory obstacles imposed by local authorities. However,
interest in catching jellyfish in Nicaraguan waters was recently renewed, and an
estimated 660 tonnes and 1,955 tonnes of jellyfish were caught in 2013 and 2014
respectively (J. Álvarez-Tello, Centro de Investigaciones Biológicas del Noroeste,
pers. comm., May 2015).
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3.3.16. Pakistan
Pakistan is yet another nation that does not explicitly report its jellyfish catches to
FAO. Data is scant; however, the government handbook of fisheries statistics (MFD
2012) does report catches for the period 2007-2009 with an average annual catch of
1,663 tonnes. While this is only for one of two targeted species, it is the primary
species. Although Rhizostoma pulmo and Catostylus mosaicus were previously
reported as the target jellyfish species in Pakistan (Tahera & Kazmi 2006;
Muhammed & Sultana 2008), it now appears that these were misidentifications, and
the target species are in fact Catostylus perezi and Rhopilema hispidum, with the former
being caught in much larger quantities (Gul & Morandini 2013; 2015).
Gul et al. (2015) discuss what little information is available regarding the jellyfish
fishery in Pakistan. C. perezi is fished in Baluchistan, while R. hispidum is fished in
Sindh province. Gill nets and set-nets are the typical gears used. It is unclear when
the jellyfish fishery in Pakistan began, but currently at least 8 to 10 companies
regularly process jellyfish there. In contrast with jellyfish fisheries in some other
countries, the fishery in Pakistan primarily only uses the subumbrellar portions of
the jellyfish, with the bell typically being discarded at sea. It is suspected that
Pakistan’s jellyfish fishery is growing, as López-Martinez & Álvarez-Tello (2013)
report 1,600 tonnes of jellyfish product exported to China from Pakistan in 2012.
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3.3.17. Philippines
Fishing for jellyfish in the Philippines presumably began in the mid-1970s, as both
FAO and SEAFDEC first report catches beginning in 1976. The catch statistics
reported by both organizations are virtually identical; however, SEAFDEC reports
exports from the Philippines that are an order of magnitude higher than the catches
(hundreds of tonnes versus tens of tonnes respectively). While the possibility of re-
exporting exists, FAO (2015b) reports nearly identical exports as well as the same
values for production of semi-dried jellyfish, indicating that the actual wet weight
landings are likely at least 4 times higher than indicated by the exports (see 3.1.2
Scaling factor), and much higher than the catch values reported. This suggests that
catches in some years are probably in the thousands of tonnes, a full two orders of
magnitude higher than reported. When production statistics of processed jellyfish
were available, they were scaled up to estimate the wet weight of the catch;
however, due to the high variability of the catches, in years with no export statistics,
only the reported catches were used in the catch reconstruction.
Important fishing grounds in the Philippines include San Miguel Bay in southern
Luzon Island as well as Carigara Bay off Samar Island. Jellyfish are also targeted
around Palawan Island, with fisheries along the east coast of the island as well as in
Malampaya Sound and near Port Barton. Fisheries in all of these locations are
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presumably for Lobonemoides robustus (Omori & Nakano 2001; Kitamura & Omori
2010). Lobonema smithi has also been reported as a target species (e.g., PCAMRD
2008); however, it is likely that these specimens are also Lobonemoides robustus
(Kondo et al. 2014). Over 1,500 people benefit from jellyfish fishing in Malampaya
Sound alone (more than 10% of the population in the region), including fishermen as
well as women and children involved in processing (PCAMRD 2008). However,
reports in the late 2000s suggested that overfishing had led to a decline of jellyfish in
the area, and fishing for jellyfish was suspended in October 2008 in Malampaya
Sound as Japanese importers focused more on jellyfish from Indonesia, where
catches are larger and shipping costs are lower (PCAMRD 2008; Kitamura & Omori
2010). Jellyfish are likely targeted in many more regions in the Philippines than are
indicated in Figure 3, such as Mindanao (SEAFDEC 2015); however, specific
locations of where additional jellyfish are caught could not be readily determined.
3.3.18. Russian Federation
Information on the recent development of a jellyfish fishery in Russian waters is
chronicled by Yakovlev et al. (2005). Prior to 1999, the jellyfish Rhopilema esculentum
was rare along the Primorsky Coast. However, in 1999, R. esculentum arrived in Peter
the Great Gulf in large abundances, presumably carried there by shifting currents, as
other rare marine species started showing up around the same time as well. As
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R. esculentum is the most valuable jellyfish in Asia, a few firms began catching and
processing R. esculentum in Peter the Great Gulf for export to China. According to
Yakovlev et al. (2005), 2,000 tonnes were landed in 2000, followed by 346 tonnes in
2001. These values are in contrast with FAO statistics, which report no catch in 2000,
and a catch of 142 tonnes in 2001. According to FAO, catches in Russia have
continued since 2003 at < 1,000 tonnes annually.
3.3.19. Sri Lanka
Sri Lanka is yet another country catching jellyfish that is not found in FAO’s jellyfish
catch statistics. Sri Lanka began catching and exporting jellyfish no later than 1986
(Anonymous 1986). By 2008, the fishery had grown to the point where nearly 20,000
fishers targeted jellyfish, with export values in the millions of U.S. dollars (Naalir
2008). There appears to be considerable controversy over the jellyfish fishery in Sri
Lanka, with environmentalists expressing concern over ecological impacts,
overfishing of jellyfish, and a lack of regulation; while the Minister of Fisheries
supports the economic benefits of exporting processed jellyfish to China (Perera
2008).
Fishing grounds are reportedly near Panama and Komariya in the Ampara District,
and in the Kirinda area in the Hambantota District (Perera 2008). In 2009, one of the
two primary target species was identified as belonging to the genus Crambionella
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(NARA 2010). Catch statistics are not readily available; however, an article in 2008
indicated that “more than 100 to 150 tons of jellyfish are processed daily” for “only a
few weeks of the year” (Perera 2008). If an average of 125 tonnes is assumed to be
processed daily for 21 days, it can be estimated that Sri Lanka catches approximately
2,625 tonnes of jellyfish annually. To estimate the catch, this amount was scaled
linearly from zero between 1985 and 2008, and carried forward from 2008. This
estimate for recent catches is also consistent with the figure reported by López-
Martinez & Álvarez-Tello (2013) of 800 tonnes of processed jellyfish product
exported to China from Sri Lanka in 2012.
3.3.20. Thailand
Thailand is the world’s second largest producer of jellyfish, behind China. Jellyfish
are caught both in the Pacific (Gulf of Thailand) and the Indian Ocean (Andaman
Sea). Catches in the Pacific were first reported by FAO in 1970, with catches from the
Indian Ocean following in 1978 (FAO 2015a). However, Soonthonvipat (1976) noted
in the mid-1970s that Thailand was targeting jellyfish both in the Gulf of Thailand
and the Andaman Sea, and wrote at the time that these activities began “only within
the past decade.” Initially, the product was exported solely to Japan; however,
production for both domestic markets and export to several countries increased
rapidly (Soonthonvipat 1976). Unlike many countries, total catches reported for
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Thailand appear to be consistent between FAO (2015a) and SEAFDEC (2015), with
the exception of 2010, where landings reported by the latter were three times that of
the former (110,000 and 37,133 tonnes respectively). However, in several years, the
production of processed jellyfish reported by FAO (2015b) is higher than what
would be expected from the reported catch. While the possibility of re-exports exists,
this is unlikely to account for the differences in the values, and the catches are
assumed to be underreported by FAO in those years.
In addition to the commercial jellyfish fisheries in Thailand that are primarily
exporting catches to other countries in Asia, it is likely that jellyfish are also targeted
in Thailand by artisanal and subsistence fisheries for local use. Estimates for these
sectors have been made by Teh et al. (2015), and were included in the reconstruction.
Estimates of foreign illegal fishing of jellyfish in Thai waters were also borrowed
from Teh et al. (2015); however, as jellyfish are likely not caught in direct proportion
to other fished groups, only 50% of the foreign fishing estimates for Myanmar and
Malaysia were used in order to remain conservative.
A large proportion of the jellyfish catch in Thailand is suspected to be Cephea cephea;
however, this has yet to be verified. Both Lobonemoides robustus and Rhopilema
hispidum are presumably caught in the Gulf of Thailand near Rayong and Samut
Sakhon (Omori & Nakano 2001; Kitamura & Omori 2010). L. robustus is also targeted
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in the Andaman Sea near Ranong and Phuket. The estuarine Acromitus hardenbergi is
caught at the mouth of the Trat River, in both Nong Khan Song and Tha Phrik
(Kitamura & Omori 2010). Soonthonvipat (1976) indicated that the targeted species
in Thailand is Rhopilema esculentum; however, this has not been confirmed by
updated studies of edible jellyfish species in Southeast Asia, where the taxonomy is
still considerably confused (Omori & Kitamura 2004; Kitamura & Omori 2010).
Fishing for jellyfish has also been reported near Songkhla, further south in the Gulf
of Thailand (Rudloe 1992).
Interestingly, Soonthonvipat (1976) suggested that there seemed to be a decline of
jellyfish in the Andaman Sea as early as the 1970s, but noted that “the question
regarding overfishing of jellyfish could not be answered.” Rudloe (1992) reported
that there is some government regulation of jellyfish fisheries in Thailand, including
closure of the fishery during the last several weeks of the season to allow for
spawning.
3.3.21. Turkey
Turkey began catching jellyfish in 1984 and continued until 2006, which is the last
year with reported landings. Catches over this time period ranged from zero to 4,000
tonnes, with an annual average of approximately 1,200 tonnes. Catches reported by
FAO appear to agree with those from Turkey’s State Institute of Statistics, with the
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latter providing a spatialized breakdown of the catches. For the years where spatial
data could readily be obtained, the majority of the landings came from the Eastern
Black Sea (46%) and the Marmara Sea (31%), with a smaller proportion of the catches
coming from the Mediterranean Sea (11%), the Western Black Sea (7%), and the
Aegean Sea (4%).
Information on this jellyfish fishery is scant, but it is suspected that the targeted
species was Rhizostoma pulmo (Ozer & Celikkale 1998; Omori & Nakano 2001; Ozer &
Celikkale 2001). The markets for Turkish jellyfish are unclear. Like jellyfish fisheries
in most countries, much of the product may have been exported to Asia; however, it
is also suspected that some of the product was used to feed farm animals such as
chickens (Hsieh & Rudloe 1994; CIESM 2010).
3.3.22. U.S.A.
In the southeastern United States of America, both along the Atlantic coast and in
the Gulf of Mexico, cannonball jellyfish or ‘jellyballs’ (Stomolophus meleagris) have
traditionally been a nuisance for shrimp fishers and power plants, but are now
commercially exploited. There have been numerous attempts to establish fisheries
for S. meleagris in the U.S.A., with varying degrees of success. The first attempt was
reportedly in Medart, Florida in the 1970s for export to Taiwan; however, the
venture failed, partially due to the reluctance of fishers to provide the product
93
(Rudloe 1992). Interest was renewed in the late 1980s, both in Florida and Georgia.
At the time, processing techniques were being investigated by Huang (1986; 1988) at
the University of Georgia. In 1991, development of a jellyfish fishery was officially
launched through a grant from the U.S. Department of Commerce (USDC). Under
the grant, marine scientist Jack Rudloe traveled to Malaysia and Thailand to
investigate jellyfish fishing and processing methods. Outlined in Rudloe (1992), a
jellyfish fishery was proposed for the Florida Panhandle in the northern Gulf of
Mexico, where commercial fisheries had suffered dramatic declines due to
overfishing and rapid coastal development. In addition, jellyfish were considered a
nuisance to swimmers, shrimp fishers, and power plants. The initial report
concluded that a jellyfish fishery could be developed in Florida; however, several
challenges would have to be overcome including economic viability, processing
knowledge, labour costs, and pollution from processing facilities. An additional
challenge proved to be the size of the product. Cannonball jellyfish rarely exceed
19 cm in bell diameter, but jellyfish products fetching the highest prices at the time
were 30 cm or more. Nevertheless, it was thought that a superior product could be
produced from cannonball jellyfish and the exploration of the fishery in Florida
continued.
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In 1993, a small amount (~1 tonne) of jellyfish were caught in the Atlantic near
Fernandina Beach, Florida, processed, and sent to potential buyers in Asia (Jones &
Rudloe 1995). While the overall quality of the jellyfish was rated high, the darker
colour of Atlantic cannonball jellyfish was not preferred. Several additional samples
were then produced by various parties in 1993 and 1994 using cannonball jellyfish
from the Gulf of Mexico, which are whiter in colour. After some success,
approximately 15-30 tonnes were caught, processed, and shipped to Korea.
Challenges for the fishery continued, including higher labour costs and an
unfamiliarity with the species in Asia. In 1995, catches exceeding 90 tonnes were
caught using seine gear. Through the 1990s, a jellyfish fishery in Florida did not
expand due economics (Bynum 2003); however, jellyfish have reportedly been
caught near the Florida panhandle, and then transported to Georgia for processing
(Bridges 2008). Nonetheless, fishing of jellyfish in Florida waters continues, with
boats operating out of both Apalachicola and Port Saint Joe, with a combined annual
catch of approximately 1,100 tonnes.
The state of Georgia has also established a small jellyfish fishery. It began in the
1990s, with a solitary processing plant located in Darien (Graitcer 2012). Since 1998,
licenses for catching jellyfish have been limited to 6-12 fishers (Page 2015), mainly
due to limited processing capacity. The fishers involved are shrimpers that
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occasionally convert their boats to fish for jellyfish, and are reportedly thankful for
their newfound opportunity (Bynum 2003; Landers 2011). The jellyfish fishery in
Georgia transitioned from an experimental to a recognized fishery in 2013, and
continues to operate at processing capacity with the possibility of future expansion.
FAO statistics for the U.S.A. show an average of 574 tonnes of jellyfish landed
annually for the period 1999-2013; however, the processing capacity in Georgia
alone is reported to be on the order of 450 tonnes per week (Bland 2014). Indications
are that exports exceed 1,000 tonnes of semi-dried product annually, suggesting
average annual landings of roughly 4,000 tonnes (Anonymous 2014; Nobel 2014).
This makes jellyfish the 3rd largest fishery in Georgia by weight, behind shrimp and
blue crab (Page 2015). Most fishing occurs between November and May in federal
waters adjacent to Doboy Sound (Page 2015).
Entrepreneurs in the state of South Carolina are eager to start catching, processing,
and exporting cannonball jellyfish. However, development plans have been
hampered due to concerns over pollution from processing facilities. Proposals have
been put forth in Beaufort and Colleton Counties, with capacities in excess of 2,000
tonnes per week (Bland 2014). While approximately 6 tonnes of jellyfish were
apparently landed and processed at a temporary facility in 2014, these operations
have ceased pending further review (Moody 2014; Murdock 2014).
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There was also a historical fishery for jellyfish in Washington State’s Puget Sound,
but instead of targeting jellyfish for food, this fishery sought the hydromedusan
Aequorea victoria for research on bioluminescence. The tale is chronicled by
Shimomura (1995), and includes the isolation of luminescent proteins ‘aequorin’ and
‘green fluorescent protein’ (GFP) in 1962 and 1979 respectively. GFP, which absorbs
ultraviolet light and emits a green glow without the addition of any chemical
additives, has proven to be an invaluable genetic marker, resulting in a veritable
revolution in biotechnology (Zimmer 2005). This immense contribution to science
was recognized in 2008, when the Nobel Prize in Chemistry was awarded for the
discovery and development of GFP (Coleman 2010; Roda 2010). A. victoria has not
been harvested from Puget Sound since the 1990s, as GFP is now synthesized in the
laboratory. However, during the course of researching the luminescent proteins of
A. victoria, it has been estimated that a total of 1 million medusae were collected over
the ~25 year period in Friday Harbor (Zimmer 2005). Taking the estimate of
Shimomura (1995) of 50 g for a typical specimen, this equates to a total catch of
approximately 50 tonnes, or 2 tonnes per year. This is miniscule in the context of
most jellyfish fisheries, which may catch tens of thousands or even hundreds of
thousands of tonnes of jellyfish in a single season. However, it is worth noting that
even these small annual catches may have affected the population of A. victoria
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around Friday Harbor. Upon Osamu Shimomura’s arrival on San Juan Island in
1961, the medusae were reportedly “abundant” and provided a “constant stream”
flowing past the docks. However, it was observed that the abundance of this species
began to decline in the 1990s (Mills 2001) and has since “almost completely
disappeared from the area” (Shimomura 2005). The effects of this apparent
overfishing do not appear to be widespread, as Aequorea populations in the nearby
waters of British Columbia, Canada can form extensive blooms with occasional
densities of 1-2 medusae/m3 (pers. obs.).
3.3.23. Vietnam
Vietnam is yet another country known to catch jellyfish but remains absent from
FAO’s jellyfish statistics. The jellyfish fishery began in the early 1990s, introduced by
Chinese importers (Nishikawa et al. 2009). Details on the jellyfish fisheries of
northern Vietnam, especially those in the Gulf of Tonkin near Thanh Hoa, were
investigated by Nishikawa et al. (2008) though interviews with processing plant
owners and fishers, field sampling, and examination of local statistics. As with most
jellyfish fisheries, large interannual variations were reported. Fishing for jellyfish
can be very profitable for fishers in Vietnam, as well as their families that assist with
pre-processing on beaches. Despite a short fishing season of less than 2 months,
income from jellyfish may support fishers for the rest of the year. Nishikawa et al.
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(2008) estimate that 800,000 to 1,200,000 individual medusae may be caught each
year in the Thanh Hoa region alone. In addition, the authors report statistics for 2
major government-run processing companies, one for 1995 to 2005 and the other for
2000 to 2005, with combined mean annual exports of approximately 4,500 tonnes of
semi-dried product. The values can be conservatively scaled up (see 3.1.2 Scaling
factor) to provide a minimum estimate of Vietnam’s jellyfish catches since 1995. Of
course, the estimated values are likely underestimates, as there are additional
private companies that do not report their exports (Nishikawa et al. 2008). For
example, four enterprises in Quang Ninh province were found to be illegally
discharging wastewater into the UNESCO World Heritage Site of Ha Long Bay in
2013 in connection with jellyfish processing (Anonymous 2013); however, the
relative scale of these operations remains unclear.
Thanh (2011) reports that the town of Diem Dien in Thai Binh province, which
accepts jellyfish caught from neighbouring provinces such as Quang Ninh, Hai
Phong, and Nam Dinh, processes approximately 15,000 tonnes of jellyfish annually.
It is unclear how much overlap there is between the figures reported by Nishikawa
et al. (2008) and Thanh (2011). As such, only the values reported by Nishikawa et al.
(2008) were used in the catch reconstruction. However, the reported values are
encouragingly of similar magnitude, and serve to highlight that Vietnam has
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emerged as a major player in the global jellyfish market. To extrapolate jellyfish
production beyond 2005, a linear regression model was used, which generally
reflects the pattern of apparent growth in Vietnam’s jellyfish landings between 1995
and 2005, based on the available minimum estimates.
Regarding the target species, Lobonemoides robustus is presumably caught near Nha
Trang and in Cam Ranh Bay in southern Vietnam, as well as near the island of Phu
Quoc in the Gulf of Thailand. Further north, Rhopilema hispidum is targeted in the
Gulf of Tonkin, near Thanh Hoa and Haiphong (Omori & Nakano 2001; Kitamura &
Omori 2010). Rhopliema esculentum is also caught in northern Vietnam, albeit in
much lower abundances than R. hispidum (Nishikawa et al. 2009). Gears may include
dip-nets and drift gill nets, and other species of jellyfish may also be caught as
bycatch, including Cyanea, Chrysaora, and Sanderia spp.; however, these non-target
species are not processed for export (Nishikawa et al. 2009).
3.3.24. Other countries
As mentioned, FAO reports jellyfish catches for the Falkland Islands (Malvinas),
Namibia, and the United Kingdom. However, these countries and territories are not
known to have jellyfish fisheries and it is likely that the reported catches are in fact
discarded bycatch from other fisheries. Indeed, the FAO fishing areas reported for
jellyfish catches by the United Kingdom are not part of the United Kingdom’s
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Exclusive Economic Zone (EEZ), namely 41, ‘Atlantic, Southwest’ and 48, ‘Atlantic,
Antarctic’ (see Figure 6). In addition to the countries discussed above, a number of
other nations have been reported to export jellyfish. For example, North Korea,
Hong Kong, Singapore, and Taiwan all exported jellyfish to Japan in the 1980s and
1990s (Omori & Nakano 2001). Cambodia can also be added to this list for a solitary
export to Japan in 2001 (S.-I. Uye, Hiroshima University, pers. comm., June 2014).
However, it is unclear if these countries caught the jellyfish in question in their own
waters, or if they are simply re-exports.
Given the expanding jellyfish fisheries in Middle-Eastern countries such as Iran,
Bahrain, and Pakistan, it seems likely that there may also be fisheries for jellyfish in
the United Arab Emirates (U.A.E.), Saudi Arabia, and Oman. Unfortunately no
information on jellyfish fisheries in these countries is readily available, other than
the fact that López-Martinez & Álvarez-Tello (2013) report that 1,000 tonnes of
processed jellyfish was exported to China from the U.A.E. in 2012. Due to the
possibility of re-exports, this account was not included in the reconstruction;
however, this may be evidence that yet another country is developing a jellyfish
fishery.
Lychnorhiza lucerna has also been investigated as a possible fishery in Argentina
(Schiariti 2008), and numerous specimens have been caught around the Río de la
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Plata along the northern coast of Buenos Aires province. These catches have been
processed by scientists and fishers in order to investigate the quality of the product.
As L. lucerna interferes with tourism as well as fisheries for finfish and shrimp in the
region (Schiariti 2008; Nagata et al. 2009), there is interest in targeting this species.
However, this is considerable uncertainty regarding how much jellyfish can be
produced from the area on a consistent basis. As buyers from Asia necessitate a
minimum to be involved, this uncertainty has heretofore prevented the fishery from
developing. Until significant investment is made to overcome the ecological and
economic knowledge gaps, a jellyfish fishery in Argentina will likely remain
undeveloped.
There have been recent attempts to exploit Chrysaora plocamia along the coasts of
Peru, particularly near Pisco, for export to China. While there have been stakeholder
meetings and commissioned reports, the fishery has not developed, mainly due to
the fact that the target species is a semaeostome, and is therefore less desirable.
However, there is potential for development of this fishery given the dramatic
abundances of this species, which can sometimes approach the biomass of small
pelagic fishes in the region (Mianzan et al. 2014; Quiñones et al. 2015). Large blooms
of C. plocamia are often a costly nuisance to fishers, aquaculture operations,
desalination plants, tourism, and other industries (Quiñones et al. 2013; Mianzan et
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al. 2014), signifying that many would welcome a targeted fishery in the area.
Similarly large abundances of C. plocamia also occur in northern Chile, suggesting
that if a fishery were to be established in Peru, expansion to Chile would also be a
possibility (Palma 2011).
Norway also explored the possibility of exploiting the mesopelagic Perhiphylla
periphylla (Wang 2007). Despite dramatic increases in abundance of this species in a
number of Norwegian fjords (Brotz 2011), a commercial jellyfish fishery has not
developed.
Javenech, a pharmaceutical company in France, lands approximately 3 to 4 tonnes of
Rhizostoma plumo annually from the Bay of Biscay for processing into collagen.
Rhizostoma octopus is also commercially exploited for high-grade medical collagen in
Wales, United Kingdom using gill nets with a mesh size of 5 × 5 cm. The
unregulated fishery began in 2014, with landings of 4.3 tonnes in 2015 (Elliott et al.
2016). Currently, these two fisheries are assumed to be the only commercial-scale
operations fishing jellyfish for a use other than human consumption.
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4. Growth of jellyfish in Mexico’s Gulf of California
Understanding the growth and mortality of fish populations is essential for
predicting how stocks will respond to fishing pressure, and thus for incorporation in
ecosystem models and the implementation of regulations to ensure sustainable
fisheries. Traditionally, estimates of growth were most often made using age-based
methods, whereby the age of captured fishes were estimated by examining their
otolith structures (or less often scales), which exhibit annual growth markings.
However, such markings are less evident in tropical fishes, as seasonality is less
pronounced in the tropics. These methods are also not applicable to most marine
invertebrates, such as jellyfish, which lack otoliths and often do not live beyond one
year. Some cubomeduse exhibit daily growth markings in their statoliths (e.g., Ueno
et al. 1995; Gordon et al. 2004; Gordon & Seymour 2012); however, such markings
have not been identified in other taxonomic classes of jellyfish. As a result, past
studies of jellyfish growth tend to lack standardized methods, instead reporting
instantaneous growth rates in terms of percent growth, making comparison within
and between species virtually impossible (Palomares & Pauly 2009). Thankfully,
size-based methods, which examine the lengths or weights of a sample of animals,
have also proven to be effective, and are often much easier to implement than age-
based methods.
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Medusae of Stomolophus meleagris were examined from Mexico’s Gulf of California,
where there is an expanding fishery for them. Data were analyzed in order to
identify growth patterns, which will help inform fisheries management decisions
and contribute to the understanding of how stocks will respond to fishing pressure.
4.1. Length-frequency analysis and the ELEFAN software
The ELEFAN software, short for Electronic Length-Frequency Analysis, was
developed by Dr. Daniel Pauly more than 30 years ago and was recently updated as
a stand-alone application in the open-source programming language R (see Pauly &
Greenberg 2013). The primary inputs of ELEFAN are length-frequency (L/F) data,
which are then used to estimate parameters of the von Bertalanffy Growth Function
(VBGF), namely the asymptotic length L∞ and the curvature of the growth curve K.
The standard, non-seasonal VBGF (see von Bertalanffy 1938) takes the form:
Lt = L∞·(1 – e-K(t – t0))
where Lt is the predicted length at age t, L∞ is the asymptotic length (roughly
corresponding to the maximum length in the population in question), K is the
curvature parameter of dimension time-1 that expresses how fast L∞ is approached,
and t0 is the (usually negative) age at which size = 0.
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There is a concern that jellyfish size is not always related to age. In fact, jellyfish may
exhibit degrowth under adverse environmental conditions, such as poor food
availability (e.g., Hamner & Jenssen 1974; Frandsen & Riisgard 1997; You et al. 2007;
Lilley et al. 2014). However, as the medusae included in this analysis are relatively
young, they are expected to exhibit continued growth over the study period. Of
course, all models are wrong (Box 1976), but length-frequency analysis has proven
to be an informative and useful tool for studying jellyfish growth patterns
(Palomares & Pauly 2009).
4.2. Sampling of Stomolophus meleagris
The Gulf of California, also known as the Sea of Cortés, is a productive and variable
body of water separating the Mexican mainland and the Baja California Peninsula
(22-32° N and 105-107° W). It is approximately 1,130 km long and 80-209 km wide
(Lluch-Cota et al. 2007). The peninsular shoreline is mostly rocky with sandy
stretches and virtually no riverine input, while in contrast the continental shore is
characterized by long sandy beaches, muddy bays, and large coastal lagoons with
considerable freshwater input. The shelf is wider along the continental shore (Figure
7), and the depth of the Gulf increases towards its mouth. The environment in the
Gulf of California is strongly seasonal, with most rainfall occurring during the
summer and fall (Salinas-Zavala et al. 1993). Fishing is the most important human
106
activity in the Gulf, with a variety of fisheries including both highly industrialized
pelagic and coastal artisanal fleets. The most important targeted taxa include
shrimp, small pelagic fishes such as sardines, herring, and anchovies, as well as
squids and tunas (Lluch-Cota et al. 2007). As discussed, there is also a rapidly
developing fishery for cannonball jellyfish Stomolophus meleagris since 2001 (see
3.3.13 Mexico).
S. meleagris has a polymorphic life cycle including fertilized egg and planula, as well
as various polypoid and medusoid phases (Calder 1982; Figure 1). Small medusae
typically appear in the coastal lagoons in the Gulf of California in December,
suggesting that polyps are likely located in the lagoons. The medusae grow rapidly
through December, January, and February, and are then targeted by the fishery in
March through to April or May after migrating out of the lagoons. Medusae appear
to be virtually nonexistent between June and Novemeber (Álvarez-Tello et al. 2016).
Individual S. meleagris medusae were collected by handheld trawl nets in 2014, 2015,
and 2016 by members of the Instituto Nacional de Pesca. Most samples were taken
near Guaymas, including in the Las Guásimas coastal lagoon (Figure 7) using a 2 m
long trawl net with 50 × 50 cm square steel frame opening and ¼-inch stretched
mesh. The date, time, latitude, longitude, and water temperature were also recorded
for each sample. In 2016, smaller medusae were also sampled from lagoons using
107
the same trawl gear with 1 mm mesh. Specimens were also measured each year from
local processing plants. Individual bell diameters were measured using a vernier
caliper. More than 20,000 individual medusae were sampled in total.
108
Figure 7. Map of the Gulf of California in Mexico; dark blue indicates shelf (< 200 m); black circle
indicates approximate location of Guaymas and Las Guásimas coastal lagoon
109
4.3. Analysis and results
All samples were pooled to create a Wetherall Plot (Wetherall 1986), which can
provide an estimate of L∞ by examining the differences between the mean and
maximum values of L within a sample. Using all of the sample data, the Wetherall
Plot yielded an estimate of L∞ = 18.4 cm (Figure 8). Samples from December 2015 –
March 2016 were binned into biweekly increments and used to generate a length-
frequency plot (Figure 9), which appears to represent two distinct cohorts. These
data, representing more than 5,000 individual specimens, were selected given that
the sampling method used targeted small medusae and would be useful for
estimating growth parameters. ELEFAN was then used to estimate the goodness-of-
fit based on various values of K (Figure 10), suggesting an estimate of K = 3.7 year-1.
Values of K can vary widely for jellyfish, from as low as 0.45 year-1 for some
specimens of Aurelia aurita to as high as 4.69 year-1 for selected Phyllorhiza punctata
(Palomares & Pauly 2009). Even within a genus, K may range from less than 1 year-1
to nearly 4 year-1, as is the case for Aurelia spp. (Palomares & Pauly 2009). Therefore,
the estimate of K = 3.7 year-1 for S. meleagris suggests rapid growth, and appears to
reasonably represent the observed growth patterns in the population (Figure 11).
110
Figure 8. Weatherall Plot using ELEFAN with L∞ = 18.4 cm
111
Figure 9. L/F data plot for S. meleagris sampled Dec. 2015 - April 2016 appearing to show two
distinct cohorts of medusae
112
Figure 10. Response surface for the goodness-of-fit estimator using ELEFAN, suggesting an
estimate of K = 3.7 year-1
113
Figure 11. Growth curve fitting using ELEFAN for S. meleagris sampled Dec. 2015 – April 2016;
L∞ = 18.4 cm; K = 3.7 year-1
In order to compare between different taxa, it is useful to compare values of K and
W∞, the asymptotic weight (rather than L∞). This was accomplished by examining
the length-weight relationship for S. meleagris (Figure 12). In total, 136 specimens
114
were measured for their length (L; bell diameter) and weight (W), which was then
used to estimate parameters (a and b) of the length-weight equation:
W = a·Lb
For S. meleagris, it was found that W = 1.19·L2.58 which explained 95% of the variance.
The estimated value of b = 2.58 is consistent, although not equivalent, with that
found by Álvarez-Tello et al. (2016) of b = 2.8106 for the same species. More samples
of larger medusae in both studies might help to narrow the discrepancy.
Figure 12. Length-weight relationship for S. meleagris
0
50
100
150
200
250
300
350
400
450
500
0 1 2 3 4 5 6 7 8 9 10 11
Weight (g)
Bell diameter (cm)
115
Using the derived parameters for the length-weight relationship, L∞ = 18.4 cm can be
converted to W∞ = 2.18 kg. In order to compare with other taxa, such as fish, jellyfish
can be normalized to the water content of fishes. S. meleagris is approximately 96%
water (Hsieh et al. 1996), whereas fish typically have a mean water content of 75%
(Palomares & Pauly 2009). As such, W∞ for S. meleagris can be re-scaled by the ratio
of the dry weights, 25/4 = 6.25, and thus W∞(norm) = 348 g. Taking log10(K) = 0.57 and
log10(W∞(norm)) = 2.54, these values can then be compared with other taxa on an
auximetric plot (Figure 13). As can be seen from Figure 13, S. meleagris grows
similarly to Phyllorhiza punctata, which is a more rapid growth pattern than for fish
(higher K for a given W∞). This is in contrast with other jellyfish, such as the Aurelia
aurita complex and Catostylus mosaicus, whose growth patterns tend to resemble
those of small fishes (Palomares & Pauly 2009).
It was found that S. meleagris from the Gulf of California grow according to a pattern
that can be described with VBGF parameters K = 3.7 year-1 and a maximum bell
diameter of 18.4 cm. This is comparable to other large jellyfish, and exceeds the rate
at which most fishes approach their maximum sizes, even when they are normalized
for their differing water content. Using L/F analysis to estimate the growth
parameters of S. meleagris demonstrates that a tool from fisheries science can be
adapted for application to jellyfish. Such analyses are useful for understanding the
116
growth of jellyfish, which as mentioned, is important for understanding responses
to fishing pressure though estimations of mortality. In addition, such studies may
also help contribute to taxonomy and understanding of jellyfish evolution by
comparing and contrasting the growth patterns observed across and between
species. Indeed, we must continue to examine these understudied animals using all
available tools, especially those that have already been evaluated and established in
traditional fisheries science.
Figure 13. Auximetric plot of various fish (small circles) and jellyfish species including
Stomolophus meleagris; based on Palomares & Pauly (2009)
117
5. Conclusions
All investigated aspects of jellyfish fisheries proved to be previously
underestimated. This includes the number of countries that have explored fishing
for jellyfish (at least 28) and the number of edible species (exceeding 35). The
magnitudes of catches have been dramatically underestimated, with contemporary
global landings exceeding 750,000 tonnes annually, and a global catch exceeding 1
million tonnes in 2013. These estimates are more than double the values published
by FAO (2015a). Research and management of jellyfish fisheries is lagging far
behind the expansion of the industry, despite the availability of traditional fisheries
science methods such as length-frequency analysis, which can be applied to jellyfish.
While jellyfish are often perceived as a nuisance, they can also be a valuable
commodity and are savoured as a delicacy in many places. Perhaps as awareness of
jellyfish as human food continues to spread, perceptions of jellyfish will also change.
From an Anglocentric point of view, since they are not truly “fish” (itself a
paraphyletic term), there have been suggestions to step away from the moniker of
“jellyfish” and instead refer to them as “jellies” or “gelatinous zooplankton.”
However, given that some of the established tools available from fisheries science
can be applied to jellyfish, perhaps it is time to put the “fish” back into “jellyfish,”
118
and include them in the realm of “fisheries.” Indeed, the global catches of jellyfish
already eclipse those of many groups of taxa targeted by established fisheries, such
as lobsters (Pauly & Zeller 2016a; miscellaneous marine crustaceans excluded).
Moreover, despite the considerable information and research on the fishery for
Antarctic krill (Euphausia superba) the catches presented herein are even greater,
making jellyfish the largest zooplankton fishery on the planet. Clearly much more
attention needs to be paid to research on jellyfish and their associated fisheries. After
examining some of the peculiarities that characterize jellyfish fisheries, a number of
management implications and research priorities emerge.
Due to the poor understanding of jellyfish population dynamics, management
decisions for jellyfish fisheries should be adaptive and will likely vary from year to
year, or even within a single season. Given the extreme variability of jellyfish
populations, along with additional factors that contribute to high uncertainty in
jellyfish fisheries (Kingsford et al. 2000), ensuring long-term sustainability of jellyfish
fisheries will likely be difficult. As such, managers should consider employing
conservative strategies that may include catch limits, size limits, adaptive
management, harvest control rules, the precautionary principle, ecosystem-based
management, and the protection of critical habitat, especially for polyps. Combined
with economic drivers, concerns related to processing technologies (see below), as
119
well as minimizing bycatch and competition with species of concern (e.g., Elliott et
al. 2016), management of jellyfish fisheries will surely continue to be a challenge.
5.1. Research priorities
Jellyfish fisheries are clearly growing and expanding faster than research and
regulations on the subject. As such, there are a variety of knowledge gaps that
should be a priority for researchers and managers that include, but are not limited
to:
Estimates of medusae abundance in regions where fishing occurs or is
proposed to occur;
Surveys to locate (and potentially protect) important polyp habitat;
Investigations on the linkages between polyp density and medusae
abundance;
Studies on local populations of jellyfish (every species is different, and there
are potential important differences even within species);
Investigations on the use of models for jellyfish fisheries (e.g., are traditional
models for finfish applicable to jellyfish fisheries?);
Monitoring and tracking of medusae to identify the factors that control
aggregations and mixing of stocks;
Genetic analyses to determine discrete stocks and mixing of populations;
Investigations of ephyrae growth and survival;
The development of processing technologies should also be a top priority, given
both the environmental and human health concerns. Large quantities of effluent are
120
generated as a byproduct of jellyfish processing, which are too often not dealt with
in an environmentally responsible way. As discussed, edible jellyfish may also
contain concerning amounts of aluminum (Wong et al. 2010; Ogimoto et al. 2012;
Armani et al. 2013; Zhang et al. 2016), the consumption of which is linked to a
number of negative health effects, including neurobehavioural toxicity and
Alzheimer’s disease (Perl & Brody 1980; Nayak 2002). The development of new
processing technologies that either reduce the aluminum content in the edible
products (e.g., Chen et al. 2016) or eliminate the use of alum altogether is desirable
(Hsieh & Rudloe 1994). Such research could easily be undertaken by the vast array
of food scientists in industry and academia. As an example, Cotylorhiza tuberculata
that was frozen fresh at ultra-low temperature (close to –80°C) and then
reconstituted by a professional chef with a small amount of sugar and vinegar was
delicious (pers. obs.). As the seafood industry already has significant infrastructure
for freezing, storage, and distribution of food, this may provide an alternative to
chemical processing of jellyfish in some places, subject to economic viability.
5.2. Sustainability and the future of jellyfish fisheries
While assessing the ‘sustainability’ of a system (or fishery) can be a complicated task
(Costanza & Patten 1995; Shelton & Sinclair 2008), it is generally accepted that
fisheries have rarely been sustainable (Pauly et al. 2002). If we consider a simplistic
121
definition of a sustainable fishery as one that produces consistent, sustained catches
(or yields), it is interesting to consider jellyfish fisheries. Jellyfish populations are
highly variable, even when they are not subject to exploitation (Brotz 2011).
Therefore, it cannot be expected that catches would be consistent on short timescales
of several years. However, the catch reconstruction herein makes it possible to
evaluate whether or not catches are being sustained by inspecting the landings from
countries that have been fishing for jellyfish for decades. Thailand has been catching
jellyfish since the 1970s and has contemporary annual catches that can exceed
100,000 or even 200,000 tonnes in some years, comparable with historic maximum
catches. Indonesia has also been targeting jellyfish for decades, and still has annual
catches that usually exceed 20,000 tonnes, which although is not approaching
maximal historic levels, is considerable nonetheless (for specific values, see Appendix
A). A more recent example is Vietnam, which began catching jellyfish in the mid-
1990s, and has been steadily increasing landings, which now exceed 50,000 tonnes
annually. Of course, the ‘sustainability’ of the jellyfish fisheries in these countries
could be masked by increasing geographic expansion, improved technology, or the
fact that stocks may not be fully exploited. While these examples suggest that
sustainable jellyfish fisheries might be possible, there are also examples that appear
to be anything but. For example, overfishing of medusae is suspected to be the
122
primary cause for the decline of Rhopilema esculentum in China (Dong et al. 2014),
suggesting that while the polymorphic life cycle of edible jellyfish (Figure 1) likely
provides a buffer against overfishing, it should not be viewed as a total safeguard.
Indeed, catches of R. esculentum continue to decline in China, despite bans on
trawling in Liaodong Bay to protect polyp habitat (Ye 2006), as well as hatchery
programs that rear and release hundreds of millions of juvenile medusae in coastal
waters each year (see 3.3.4 China).
One of the few studies to evaluate the sustainability of a jellyfish fishery is presented
by Asrial et al. (2015a) for Saleh Bay, Indonesia. A Gordon-Schaefer surplus
production model was generated using catches and effort (number of scoop nets)
since 2006. The authors determined that the maximum sustainable yield (MSY) for
Crambione mastigophora in the region is 33,261 tonnes annually, with a maximum
sustainable fishing effort (fMSY) of 3,917 scoop nets and a corresponding catch per
unit effort (CPUE) of 8.49 tonnes per net. Given that catches in the region have
exceeded 30,000 tonnes since 2010 (Asrial et al. 2015a), the authors conclude that the
jellyfish fishery in Saleh Bay is “fully exploited,” and effort is actually exceeding the
sustainable level by 63%. As such, the authors recommend reducing the effort and
catches in the region. Asrial et al. (2015a) also used a modified version of the
RAPFISH technique (see Pitcher & Preikshot 2001) to evaluate the fishery based on a
123
number of bio-ecological, economic, technological, and social indicators. Their
overall conclusion was that the jellyfish fishery can be classified as “quite
sustainable,” which is neither the best nor worst possible ranking, and
improvements could be made through management interventions.
While several jellyfish fisheries have a long history in Asia, those in the Western
Hemisphere have only developed recently, with varying degrees of success. For
example, jellyfish fisheries in the U.S.A. and Mexico have proven to be a boon for
local fishers, whereas no market has yet developed for jellyfish from Argentina,
Peru, or Canada. There appear to be a number of factors that are conducive to
success for new jellyfish fisheries in the short term, and several additional
recommendations that may help to ensure establishment of sustainable jellyfish
fisheries in the longer term. Firstly, not just any species of jellyfish will do. There are
more than 1,400 species of jellyfish worldwide (Purcell 2012), but fewer than 40 of
those have been documented as being consumed by humans (see 2.1.1 Target species).
In fact, the number of jellyfish species that are part of major jellyfish fisheries around
the world number fewer than 20, and are all rhizostomes. While it is conceivable
that consumption of semeaostomes and other types of jellyfish may increase in the
future, demand for non-rhizostome jellyfish currently remains very low and is likely
124
a major reason why experimental jellyfish fisheries in some countries – such as
Canada and Peru – were not successful.
Secondly, attention must be paid to the processing of jellyfish. In order ensure
economic success, specific details regarding the nuances of jellyfish processing
should come from potential buyers, likely in Asia. As discussed, the method and
materials used in the processing of jellyfish can vary greatly, so potential exporters
need to work closely with buyers to deliver a suitable product. Jellyfish processing is
typically labour intensive, so the time and effort required will have to be factored
into the economics of any operation, especially in regions where labour costs are
high. Jellyfish fisheries in the U.S.A. appear to have overcome this obstacle by a
combination of the development of shorter processing times through technical
advances and the use of smaller medusae, as well as partial industrialization of
processing. Moreover, the significant environmental and human health concerns
regarding the contemporary use of processing chemicals needs to be addressed (see
5.1 Research priorities). In addition, the development of alternative processing
technologies could provide multiple benefits for jellyfish fisheries, including
expansion beyond rhizostome species, reduction of costs, and the development of
new markets.
125
To ensure success of jellyfish fisheries in the longer-term, cooperation between
stakeholders appears to be key. In addition to the collaboration between processors
and buyers mentioned above, fishers, managers, and researchers all need to work
together to help ensure the sustainability of jellyfish fisheries. If it is hoped that
jellyfish can fill some of the void left by the collapses of more traditional fisheries,
much more research will be required if repeating history is to be avoided.
Understanding of jellyfish population dynamics remains extremely poor, and as
such, the development of management strategies for jellyfish fisheries continues to
be a challenge. Collection of even the most basic fishery data, such as catch amounts,
dates, and locations remains meager, greatly limiting the advancement of research
and development of management plans. A shift towards ecosystem-based
management would contribute to building knowledge of the interactions between
the resource and the environment, as well as helping to quantify the impacts of
developing jellyfish fisheries. Fluctuations in market demand also present additional
challenges, and it is clear that economic considerations will have to be added to the
list of relevant concerns for jellyfish fisheries.
As jellyfish populations are increasing in many areas of the world (Brotz et al. 2012),
it is likely that humans will look for new ways to exploit them. Indeed, the
development of jellyfish fisheries for food and medicines has been proposed as a
126
possible strategy to deal with increasing jellyfish blooms (e.g., Purcell et al. 2007;
Richardson et al. 2009). Although increasing abundances of jellyfish will bring some
benefits to humans including jellyfish fisheries (Doyle et al. 2014), it has been
submitted that the costs associated with the negative impacts of jellyfish blooms will
outpace any increased revenues (Graham et al. 2014). Such an attitude can also serve
to ignore the root causes of the problem. Increasing jellyfish blooms have been
linked to numerous anthropogenic factors, such as overfishing, coastal
development, shipping, global warming, and pollution (Purcell et al. 2007;
Richardson et al. 2009; Duarte et al. 2013). If we simply adapt to a new normal
instead of addressing and correcting the underlying causes, our baseline will shift
(Pauly 1995), and ultimately, jellyfish may be the only seafood left. Even if a fishery
for jellyfish can be developed, any stakeholders that end up profiting from such an
endeavour will desire a sustained income, rather than seeing the resource
eradicated, so we cannot simply fish away our jellyfish problems (Gibbons et al.
2016). Indeed, it is only by increasing our understanding of these understudied
creatures through collaboration between fishers, managers, researchers, processors,
brokers, and buyers that we will be able to minimize the impacts and maximize the
opportunities offered by future jellyfish blooms.
127
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Appendix A – Reconstructed jellyfish landings by country, 1950-2013
Year
Total Est.
FAO
Aus.
Bahrain
Canada
China
Ecuador
Honduras
India
Indo.
Iran
Japan
Korea (S)
1950
216,887
300
215,487
300
1,000
1951
218,287
1,700
215,487
1,700
1,000
1952
218,487
1,900
215,487
1,900
1,000
1953
218,487
1,900
215,487
1,900
1,000
1954
218,687
2,100
215,487
2,100
1,000
1955
218,687
2,100
215,487
2,100
1,000
1956
218,787
2,200
215,487
2,200
1,000
1957
218,687
2,100
215,487
2,100
1,000
1958
311,567
2,200
308,267
2,200
1,000
1959
248,620
2,100
245,420
2,100
1,000
1960
392,274
2,100
389,073
2,100
1,000
1961
102,667
2,800
98,767
2,800
1,000
1962
183,573
2,900
179,573
2,900
1,000
1963
99,873
3,000
95,773
3,000
1,000
1964
138,880
3,100
134,680
3,100
1,000
1965
178,187
3,500
173,587
3,500
1,000
1966
367,040
3,800
362,140
3,800
1,000
1967
145,367
3,600
140,667
3,600
1,000
1968
85,707
3,800
80,807
3,800
1,000
1969
267,003
4,200
260,380
4,200
1,000
1970
348,804
59,100
337,333
6,000
1,000
1971
118,137
24,500
106,667
6,000
1,000
1972
121,681
24,600
110,667
5,300
1,000
1973
476,771
129,900
388,000
4,800
1,000
1974
152,901
24,900
147,120
432
1,000
1975
127,316
26,985
114,060
5,507
1,000
1976
73,401
36,763
34,207
8,886
1,000
1977
203,187
95,619
77,687
8,886
9,500
1978
132,212
68,158
24,293
7,440
18,000
1979
174,244
70,314
86,447
5,744
10,000
1980
72,025
12,249
59,920
5,896
311
1981
173,321
52,561
119,773
12,352
311
167
Year
Total Est.
FAO
Aus.
Bahrain
Canada
China
Ecuador
Honduras
India
Indo.
Iran
Japan
Korea (S)
1982
269,336
132,737
117,213
8,952
311
1983
349,604
232,963
72,000
23,842
311
1984
289,607
67,771
3
243,073
1,817
10,224
311
1985
470,355
108,027
406,053
3,634
7,500
311
1986
273,889
126,587
129,200
5,452
19,048
311
1987
486,579
132,580
389,893
7,269
8,656
311
1988
306,790
92,479
213,987
9,086
28,880
311
140
1989
335,909
103,662
232,960
10,903
22,672
311
1990
338,358
75,219
261,160
12,720
7,668
311
4
1991
222,037
166,284
96,009
14,537
16,844
311
668
1992
424,700
353,838
228,459
16,355
10,440
311
204
1993
219,687
191,797
132,572
18,172
26,443
311
76
1994
292,694
213,221
113,354
19,989
16,152
311
128
1995
389,253
338,761
22
171,905
21,806
123,076
311
152
1996
398,827
326,850
22
265,325
23,623
17,080
311
244
1997
579,604
486,393
32
400,483
25,440
17,719
311
56
1998
604,281
464,423
23
430,784
27,258
8,176
311
60
1999
637,060
475,751
0
402,206
29,075
32,652
311
132
2000
794,493
467,512
0
459,869
30,892
29,516
311
100
2001
652,990
365,241
0
456,297
41,124
30,836
311
40
2002
720,782
344,815
5
1
413,113
164,756
60,096
311
32
2003
709,388
365,439
5
423,884
123,464
90,649
311
52
2004
546,825
190,933
5
1,300
330,716
123,464
18,396
311
16
2005
685,728
289,049
9
3,083
412,703
123,464
14,604
311
168
2006
778,676
345,424
1
362
356,692
109,281
8,172
311
24
2007
551,952
242,597
1
10,850
348,868
95,098
26,780
311
252
2008
782,385
396,402
14
1,354
361,626
80,915
6,800
311
68
2009
801,009
353,358
1
13,921
427,878
66,732
20,000
311
148
2010
688,383
271,474
1
329
326,322
52,549
30,519
7,918
311
172
2011
751,939
359,865
1
4,492
418,264
38,366
40,000
810
14,598
40
2012
868,181
332,950
2
58,767
543,126
24,183
32,115
1,625
13,958
2,400
2013
1,101,521
397,965
11
5,758
763,539
50
10,000
30,818
< 0.5
11,934
124
168
Year
Total Est.
FAO
Malay.
Mexico
Myan.
Nic.
Pakistan
Phil.
Russia
Sri
Lanka
Thailand
Turkey
USA
Vietnam
1950
216,887
300
100
1951
218,287
1,700
100
1952
218,487
1,900
100
1953
218,487
1,900
100
1954
218,687
2,100
100
1955
218,687
2,100
100
1956
218,787
2,200
100
1957
218,687
2,100
100
1958
311,567
2,200
100
1959
248,620
2,100
100
1960
392,274
2,100
100
1961
102,667
2,800
100
1962
183,573
2,900
100
1963
99,873
3,000
100
1964
138,880
3,100
100
1965
178,187
3,500
100
1966
367,040
3,800
100
1967
145,367
3,600
100
1968
85,707
3,800
100
1969
267,003
4,200
1,423
1970
348,804
59,100
1,423
3,047
1971
118,137
24,500
1,423
3,047
1972
121,681
24,600
1,423
3,291
1973
476,771
129,900
1,423
81,548
1974
152,901
24,900
1,423
2,926
1975
127,316
26,985
1,423
5,326
1976
73,401
36,763
1,423
12
27,874
1977
203,187
95,619
5,172
280
101,663
1978
132,212
68,158
3,564
1,228
77,687
1979
174,244
70,314
3,400
68,654
1980
72,025
12,249
2,576
600
2,723
1981
173,321
52,561
2,328
2,002
36,555
1982
269,336
132,737
10,754
1,572
130,534
1983
349,604
232,963
18,715
2,316
232,420
1984
289,607
67,771
18,000
91
14,303
1,785
1985
470,355
108,027
12,997
5
38,136
1,719
1986
273,889
126,587
15,853
34
114
101,076
2,802
169
Year
Total Est.
FAO
Malay.
Mexico
Myan.
Nic.
Pakistan
Phil.
Russia
Sri
Lanka
Thailand
Turkey
USA
Vietnam
1987
486,579
132,580
25,737
88
228
54,394
3
1988
306,790
92,479
27,195
55
342
24,974
1,821
1989
335,909
103,662
42,036
86
457
25,311
1,173
1990
338,358
75,219
20,530
64
571
34,222
1,109
1991
222,037
166,284
12,094
1,956
685
78,933
1992
424,700
353,838
15,623
2,404
799
149,542
564
1993
219,687
191,797
16,293
1,012
913
23,114
781
1
1994
292,694
213,221
10,138
600
1,027
130,159
814
23
1995
389,253
338,761
7,692
1,812
2,496
1,141
52,263
487
90
6,000
1996
398,827
326,850
19,902
3,426
604
1,255
56,131
904
10,000
1997
579,604
486,393
53,811
1,412
112
1,370
67,559
900
10,400
1998
604,281
464,423
11,802
432
139
1,484
106,564
1,750
5,100
10,400
1999
637,060
475,751
7,182
1,000
2,660
1,598
139,541
1,203
5,100
14,400
2000
794,493
467,512
9,036
1,000
1,736
2,000
1,712
233,414
900
5,100
18,908
2001
652,990
365,241
10,299
1,200
1,100
1,860
346
1,826
77,091
2,000
5,100
23,560
2002
720,782
344,815
6,648
8,000
1,300
450
1,940
46,330
500
5,100
12,200
2003
709,388
365,439
5,863
1,600
1,500
40
44
2,054
27,927
4,000
5,100
22,896
2004
546,825
190,933
4,041
8,000
1,700
23
53
2,168
26,952
1,000
5,100
23,580
2005
685,728
289,049
5,285
13,048
1,976
1,472
67
2,283
55,420
544
5,100
46,192
2006
778,676
345,424
4,909
15,172
2,000
92
79
2,397
238,348
1,017
5,100
34,719
2007
551,952
242,597
4,068
2,000
2,200
2,114
116
105
2,511
14,081
5,100
37,497
2008
782,385
396,402
5,846
6,752
2,410
205
1,756
17
81
2,625
266,232
5,100
40,274
2009
801,009
353,358
4,013
10,389
2,710
1,118
17
237
2,625
202,757
5,100
43,052
2010
688,383
271,474
5,249
16,887
3,180
1,663
19
312
2,625
189,398
5,100
45,829
2011
751,939
359,865
3,738
16,755
3,420
1,663
15
671
2,625
152,774
5,100
48,607
2012
868,181
332,950
11,980
20,988
3,690
6,400
13
698
2,625
89,126
5,100
51,385
2013
1,101,521
397,965
11,745
21,253
3,935
660
6,400
9
622
2,625
172,776
5,100
54,162
170
Appendix B – Supporting publications
Several of the publications using material extracted from this dissertation contain
supporting information that was not included herein. These publications are
available for download from the following links:
Brotz, L., A. Schiariti, J. López-Martínez, J. Álvarez-Tello, Y.-H.P. Hsieh, R.P. Jones,
J. Quiñones, Z. Dong, A.C. Morandini, M. Preciado, E. Laaz, & H. Mianzan in
press. Jellyfish fisheries in the Americas: origin, state of the art, and perspectives
on new fishing grounds. Reviews in Fish Biology and Fisheries.
http://link.springer.com/article/10.1007/s11160-016-9445-y
Brotz, L. 2016. Jellyfish fisheries – a global assessment, pp. 110-124 in D. Pauly & D.
Zeller (eds.) Global Atlas of Marine Fisheries: A Critical Appraisal of Catches and
Ecosystem Impacts. Island Press, Washington, D.C., U.S.A.
https://www.islandpress.org/book/global-atlas-of-marine-fisheries
Gibbons, M.J., F. Boero, & L. Brotz 2016. We should not assume that fishing jellyfish
will solve our jellyfish problem. ICES Journal of Marine Science 73(4): 1012-1018.
http://icesjms.oxfordjournals.org/content/73/4/1012.full
