Notice for Google Chrome users:
If you are having trouble viewing or searching the PDF with Google Chrome, please download it here instead.

Page Metadata

Item Metadata


Media 24-ubc_2017_february_brotz_lucas.pdf [ 2.11MB ] Metadata JSON: 24-1.0340647.json JSON-LD: 24-1.0340647-ld.json RDF/XML (Pretty): 24-1.0340647-rdf.xml RDF/JSON: 24-1.0340647-rdf.json Turtle: 24-1.0340647-turtle.txt N-Triples: 24-1.0340647-rdf-ntriples.txt Original Record: 24-1.0340647-source.json Full Text 24-1.0340647-fulltext.txt Citation 24-1.0340647.ris

Full Text

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 051015202502004006008001,0001,2001950 1955 1960 1965 1970 1975 1980 1985 1990 1995 2000 2005 2010Number of Countries Fishing JellyfishJellyifsh Landings (000's tonnes)YearOthersVietnamIndonesiaIndiaThailandChina (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) 01002003004005006007008001950 1960 1970 1980 1990 2000 2010Jellyfish Landings (000's tonnes)YearFAO (Global)OthersIndonesiaVietnamIndiaThailandChina 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. 62 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 64 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, 68 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 71 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 72 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 77 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 79 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 80 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 82 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 83 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. 84 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). 85 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. 86 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 87 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 88 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 89 (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 90 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 91 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 92 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. 94 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 95 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). 96 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 97 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. 98 (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 99 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 100 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 101 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 102 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. 103 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. 104 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. 105 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 0501001502002503003504004505000 1 2 3 4 5 6 7 8 9 10 11Weight (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 References Abu Talib, A., G.H. Tan, & Y.A. Hamid 2003. Overview of the national fisheries situation with emphasis on the dermersal fisheries off the West Coast of Peninsular Malaysia, pp. 833-884 in G. Silvestre, L. Garces, I. Stobutzki, M. Ahmed, R. A. Valmonte-Santos, C. Luna, L. Lachica-Aliño, P. Munro, V. Christensen, & D. Pauly (eds.) Assessment, Management and Future Directions for Tropical Coastal Fisheries in Asian Countries. WorldFish Center Conference Proceedings 67. Addad, S., J.-Y. Exposito, C. Faye, S. Ricard-Blum, & C. Lethias 2011. Isolation, characterization and biological evaluation of jellyfish collagen for use in biomedical applications, Marine Drugs 9(6): 967-983. Álvarez-Tello, F.J. 2007. La pesquería de al medusa bola de cañón (Stomolophus meleagris) en la región de Kino-El Choyudo, Sonora, durante 2006, Master's thesis, Instituto Tecnológico de Guaymas, Sonora, Mexico (in Spanish). Álvarez-Tello, F.J., J. López-Martínez, & J. Rodríguez-Romero 2013. First record of the association between Stomolophus meleagris and Conchoderma cf virgatum in the Gulf of California, Hidrobiológica 23(1): 138-142 (in Spanish). Álvarez-Tello, F.J., J. López-Martínez, & D.B. Lluch-Cota 2016. Trophic spectrum and feeding pattern of cannonball jellyfish Stomolophus meleagris (Agassiz, 1862) from central Gulf of California, Journal of the Marine Biological Association of the United Kingdom 96(6): 1217-1227. Amaral, L., A. Raposo, Z. Morais, & A. Coimbra 2016. Is jellyfish ingestion safe in allergic patients? Preliminary results, Allergy 71(Special Issue, Supplement 102): 162. 128 Anonymous 1986. Sri Lanka opens markets for new species, Fishing News International, edition of December 1986. Anonymous 2005. Handbook on Fisheries Statistics 2004. Department of Animal Husbandry, Dairying and Fisheries, Ministry of Agriculture, Government of India, New Delhi, India, 166 pp. Anonymous 2006. Cookies made from giant jellyfish, Pink Tentacle. Accessed 3 October 2016. http://pinktentacle.com/2006/10/cookies-made-from-giant-jellyfish/ Anonymous 2007. Demand for jellyfish keeps olive ridley turtles starving in Orissa, India, Earthtimes.org, India, edition of December 5. Anonymous 2008a. Marine life hit by reclamation, Gulf Daily News, Bahrain, edition of November 25. Anonymous 2008b. Govt bid to save jellyfish, New Indian Express, Kendrapapra, India, edition of October 4. Anonymous 2013. Four firms found to be polluting Ha Long Bay, Viet Nam News, edition of April 9. Anonymous 2014. Jellyfish: beautiful, dangerous, delicious, CBS News, edition of July 13. Arai, M.N. 1988. Interactions of fish and pelagic coelenterates, Canadian Journal of Zoology 66(9): 1913-1927. Arai, M.N. 1997. A Functional Biology of Scyphozoa. Chapman and Hall, London, United Kingdom, 300 pp. Arai, M.N. 2005. Predation on pelagic coelenterates: a review, Journal of the Marine Biological Association of the United Kingdom 85(3): 523-536. Arai, M.N. 2009. The potential importance of podocysts to the formation of scyphozoan blooms: a review, Hydrobiologia 616: 241-246. 129 Armani, A., P. D'Amico, L. Castigliego, G. Sheng, D. Gianfaldoni, & A. Guidi 2012. Misalabeling of an "unlabelable" seafood sold on the European market: the jellyfish, Food Control 26: 247-251. Armani, A., L. Tinacci, A. Giusti, L. Castigliego, D. Gianfaldoni, & A. Guidi 2013. What is inside the jar? Forensically informative nucleotide sequencing (FINS) of a short mitochondrial COI gene fragment reveals a high percentage of mislabeling in jellyfish food products, Food Research International 54: 1383-1393. Asrial, E., A. Prajitno, E. Susilo, & G. Bintoro 2015a. RAPJELLYFISH method to evaluate the sustainability status of edible jellyfish resource management in the Saleh Bay, Indonesia, International Journal of Recent Scientific Research 6(7): 5190-5198. Asrial, E., A. Prajitno, E. Susilo, & G. Bintoro 2015b. Biology and blooms of the edible jellyfish (Crambione mastigophora) in the Saleh Bay, Indonesia, Journal of Biodiversity and Environmental Sciences 6(6): 356-369. Badre, S. 2014. Bioactive toxins from stinging jellyfish, Toxicon 91: 114-125. Balamurugan, E., B.V. Reddy, & V.P. Menon 2010. Antitumor and antioxidant role of Chrysaora quinquecirrha (sea nettle) nematocyst venom peptide against ehrlich ascites carcinoma in Swiss Albino mice, Molecular and Cellular Biochemistry 338(1-2): 69-76. Barzideh, Z., A. Abd Latiff, C.-Y. Gan, S. Benjakul, & A. Abd Karim 2014a. Isolation and characterisation of collagen from the ribbon jellyfish (Chrysaora sp.), International Journal of Food Science and Technology 49(6): 1490-1499. Barzideh, Z., A.A. Latiff, C.-Y. Gan, M.Z. Abedin, & A.K. Alias 2014b. ACE inhibitory and antioxidant activities of collagen hydrolysates from the ribbon jellyfish (Chrysaora sp.), Food Technology and Biotechnology 52(4): 495-504. 130 Belgorodsky, B., L. Fadeev, N. Hendler, E. Mentovich, M. Gozin, S. Richter, L. Reshef-Steinberger, R. Nudelman, & T. Gulakhmedova 2015. Jellyfish-derived polymer, United States Patent Application Pub. No. US 2015/0335014 A1, Provisional Application No. 61/749,396 filed on January 7, 2013, Ramot at Tel-Aviv University Ltd. November 26, 2015. Bland, A. 2014. Southern fishermen cash in on Asia's taste for jellyfish, The Salt, National Public Radio, edition of January 31. Box, G.E.P. 1976. Science and statistics, Journal of the American Statistical Association 71(356): 791-799. Brandon, M. & C.E. Cutress 1985. A new Dondice (Opisthobranchia: Favorinidae), predator of Cassiopea in southwest Puerto Rico, Bulletin of Marine Science 36(1): 139-144. Bridges, T. 2008. Demand for jellyfish aids state shrimpers, Sun-Sentinel, Florida, U.S.A., edition of January 6. Brodeur, R.D. 1998. In situ observations of the association between juvenile fishes and scyphomedusae in the Bering Sea, Marine Ecology Progress Series 163: 11-20. Brotz, L. 2011. Changing jellyfish populations: trends in Large Marine Ecosystems. Fisheries Centre Research Report 19(5), Fisheries Centre, University of British Columbia, Vancouver, Canada, 105 pp. Brotz, L., W.W.L. Cheung, K. Kleisner, E. Pakhomov, & D. Pauly 2012. Increasing jellyfish populations: trends in Large Marine Ecosystems, Hydrobiologia 690(1): 3-20. Brotz, L. & D. Pauly 2012. Jellyfish populations in the Mediterranean Sea, Acta Adriatica 53(2): 211-230. 131 Browne, J.G. & M.J. Kingsford 2005. A commensal relationship between the scyphozoan medusae Catostylus mosaicus and the copepod Paramacrochiron maximum, Marine Biology 146(6): 1157-1168. Bynum, R. 2003. Jellyfish help shrimpers to survive tough market, Rome News-Tribune, Rome, Georgia, U.S.A., edition of March 9. Calder, D.R. 1982. Life history of the cannonball jellyfish, Stomolophus meleagris, Biological Bulletin 162(2): 149-162. Cao, L., R. Naylor, P. Henriksson, D. Leadbitter, M. Metian, M. Troell, & W. Zhang 2015. China's aquaculture and the world's wild fisheries, Science 347(6218): 133-135. Carvalho-Saucedo, L., F. García-Domínguez, C. Rodríguez-Jaramillo, & J. López-Martinez 2010. Lipid variation in oocytes of the jellyfish Stomolophus meleagris from Las Guasimas Lagoon, Mexico, during gonadal development, Revista De Biología Tropical 58(1): 119-130 (in Spanish with English abstract). Carvalho-Saucedo, L., J. López-Martinez, F. García-Domínguez, C. Rodríguez-Jaramillo, & J. Padilla-Serrato 2011. Reproductive biology of the cannonball jellyfish Stomolophus meleagris in Las Guasimas Lagoon, Sonora, Mexico, Hidrobiológica 21(1): 77-88 (in Spanish with English abstract). Ceh, J., J. Gonzalez, A.S. Pacheco, & J.M. Riascos 2015. The elusive life cycle of scyphozoan jellyfish - metagenesis revisited, Scientific Reports 5(12037): 13 pp. Chancharern, P., N. Laohakunjit, O. Kerdchoechuen, & B. Thumthanaruk 2016. Extraction of type A and tybe B gelatin from jellyfish (Lobonema smithii), International Food Research Journal 23(1): 419-424. Chelsky, A., K.A. Pitt, A.J.P. Ferguson, W.W. Bennett, P.R. Teasdale, & D.T. Welsh 2016. Decomposition of jellyfish carrion in situ: Short-term impacts on 132 infauna, benthic nutrient fluxes and sediment redox conditions, Science of the Total Environment 566: 929-937. Chen, J.-W., X.-Y. Li, P. Qin, H.-P. Wang, X. Ma, F. Yin, & Q.-M. Wang 2016. Removal technology of aluminum in production process of jellyfish, Journal of Food Safety and Quality 7(4): 1681-1685 (in Chinese with English abstract). Chidambaram, L. 1984. Export oriented processing of Indian jelly fish (Muttai Chori, Tamil) by Indonesian Method at Pondicherry Region. Technical and Extension Series. Marine Fisheries Information Service, Central Marine Fisheries Research Institute, Indian Council of Agricultural Research, Cochin, India, 11-13 pp. Cho, S., J.-R. Ahn, J.-S. Koo, & S.-B. Kim 2014. Physicochemical properties of gelatin from jellyfish Rhopilema hispidum, Fisheries and Aquatic Sciences 17(3): 299-304. Chun, K.W., E. Damdinsuren, K. Yeom, & T. Ezaki 2011. Effect of jellyfish fertilizer application on seedling growth, Journal of the Japanese Society of Revegetation Technology 37(1): 155-158. CIESM 2010. New partnerships for blue biotechnology development - innovative solutions from the sea. Report on CIESM International Workshop, Monaco, 11-12 November 2010, The Mediterranean Science Commission, 44 pp. CMFRI 2007. Landings of jelly fish at Ongole, Central Marine Fisheries Research Institute Newsletter 114: 5. CMFRI 2009. Unusual heavy landings of jellyfish, Crambionella stuhlmanni at Pulicat landing centre, Chennai, Central Marine Fisheries Research Institute Newsletter 122: 14. CMFRI 2010. Seasonal jellyfish fishery in Jakhau, Gujarat, Cadalmin: Central Marine Fisheries Research Institute Newsletter 127: 10. 133 Coleman, A., S. Micin, P. Mulvay, & R. Rippingale 1990. The brown jellyfish (Phyllorhiza punctata) in the Swan-Canning estuary. Waterways Information No. 2. Swan River Trust, Perth, Australia, 4 pp. Coleman, N. 2004. Jellyfish fishery development and assessment 1999/138. Primary Industries Research Victoria, Marine and Freshwater Systems, Department of Primary Industries, Victoria, Australia, 120 pp. Coleman, R. 2010. Jellyfish, fluorescent proteins, Nobel Prizes and pioneers in histochemistry, Acta Histochemica 112(2): 113-117. Costanza, R. & B.C. Patten 1995. Defining and predicting sustainability, Ecological Economics 15(3): 193-196. Costello, J.H., S.P. Colin, B.J. Gemmell, J.O. Dabiri, & K.R. Sutherland 2015. Multi-jet propulsion organized by clonal development in a colonial siphonophore, Nature Communications 6: 8158. Cox, J. 2014. In the kitchen: moon jellyfish - How I became a medusivore, Edible Monterey Bay 12: 26-30. D'Amico, P., A. Armani, D. Gianfaldoni, & A. Guidi 2016. New provisions for the labelling of fishery and aquaculture products: difficulties in the implementation of Regulation (EU) n. 1379/2013, Marine Policy 71: 147-156. Dabiri, J.O. 2011. Jellyfish-inspired propulsion, Integrative and Comparative Biology 51: E29. Daly, M., M.R. Brugler, P. Cartwright, A.G. Collins, M.N. Dawson, D.G. Fautin, S.C. France, C.S. McFadden, D.M. Opresko, E. Rodriguez, S.L. Romano, & J.L. Stake 2007. The phylum Cnidaria: A review of phylogenetic patterns and diversity 300 years after Linnaeus, Zootaxa 1668: 127-182. Davis, P. 1982. Acceptable salted jellyfish produced, Australian Fisheries 41(3): 34. 134 DEH 2006. Asessment of the Queensland developmental jellyfish fishery. Department of the Envrironment and Heritage, Australian Government, 25 pp. Deng, C., L.-H. Tang, W. Chen, Q. Wang, & D. Lin 2009. Effects of collagen peptide from Cyanea nozakii on mouse immune function, Agricultural Science & Technology 10(2): 118-121. Derkus, B., Y.E. Arslan, A.T. Bayrac, I. Kantarcioglu, K.C. Emregul, & E. Emregul 2016. Development of a novel aptasensor using jellyfish collagen as matrix and thrombin detection in blood samples obtained from patients with various neurodisease, Sensors and Actuators B: Chemical 228: 725-736. DFA 2002a. Jellyfish exploratory survey, Trinity Bay. Project summary: FDP 424-5, Emerging Fisheries Development, Fisheries Diversification Program. Department of Fisheries and Aquaculture, Government of Newfoundland and Labrador, 4 pp. DFA 2002b. Production and marketing of jellyfish. Project summary: FDP 421-3, Emerging Fisheries Development, Fisheries Diversification Program. Department of Fisheries and Aquaculture, Government of Newfoundland and Labrador, 2 pp. Ding, J.-F., Y.-Y. Li, J.-J. Xu, X.-R. Su, X. Gao, & F.-P. Yue 2011. Study on effect of jellyfish collagen hydrolysate on anti-fatigue and anti-oxidation, Food Hydrocolloids 25(5): 1350-1353. Dong, J., L.X. Jiang, K.F. Tan, H.Y. Liu, J.E. Purcell, P.J. Li, & C.C. Ye 2009. Stock enhancement of the edible jellyfish (Rhopilema esculentum) in Liaodong Bay, China: a review, Hydrobiologia 616: 113-118. 135 Dong, Z., D. Liu, & J.K. Keesing 2014. Contrasting trends in populations of Rhopilema esculentum and Aurelia aurita in Chinese waters, pp. 207-218 in K. A. Pitt & C. H. Lucas (eds.) Jellyfish Blooms. Springer, Dordrecht, Netherlands. Dong, Z., Z. Liu, D. Liu, Q. Liu, & T. Sun 2016. Low genetic diversity and lack of genetic structure in the giant jellyfish Nemopilema nomurai in Chinese coastal waters, Marine Biology Research 12(7): 769-775. Dong, Z.J., D.Y. Liu, & J.K. Keesing 2010. Jellyfish blooms in China: Dominant species, causes and consequences, Marine Pollution Bulletin 60(7): 954-963. Doyle, T.K., G.C. Hays, C. Harrod, & J.D.R. Houghton 2014. Ecological and societal benefits of jellyfish, pp. 105-127 in C. H. Lucas & K. A. Pitt (eds.) Jellyfish Blooms. Springer, Dordrecht, Netherlands. Duarte, C.M., K.A. Pitt, C.H. Lucas, J.E. Purcell, S. Uye, K. Robinson, L. Brotz, M.B. Decker, K.R. Sutherland, A. Malej, L. Madin, H. Mianzan, J.M. Gili, V. Fuentes, D. Atienza, F. Pagès, D. Breitburg, J. Malek, W.M. Graham, & R.H. Condon 2013. Is global ocean sprawl a cause of jellyfish blooms?, Frontiers in Ecology and the Environment 11(2): 91-97. Duron-Dufrenne, M. 1987. Premier suivi par satellite en Atlantique d'une torue luth Dermochelys coriacea, Comptes Rendus Academie des Sciences Paris 304: 339-402 (in French). Elliott, A., V. Hobson, & K.W. Tang 2016. Balancing fishery and conservation: a case study of the barrel jellyfish Rhizostoma octopus in South Wales, ICES Journal of Marine Science doi:10.1093/icesjms/fsw157. Epstein, H.E., M.A. Templeman, & M.J. Kingsford 2016. Fine-scale detection of pollutants by a benthic marine jellyfish, Marine Pollution Bulletin 107(1): 340-346. 136 FAO 2015a. Global Capture Production, Fishery Statistical Collections, Fisheries and Aquaculture Department, Food and Agriculture Organization of the United Nations, Rome, Italy. Accessed 28 July 2015. http://www.fao.org/fishery/statistics/global-capture-production/en FAO 2015b. Fishery Commodities Global Production and Trade, Fishery Statistical Collections, Fisheries and Aquaculture Department, Food and Agriculture Organization of the United Nations, Rome, Italy. Accessed 24 September 2015. http://www.fao.org/fishery/statistics/global-commodities-production/en FAO 2016. Global Aquaculture Production, Fishery Statistical Collections, Fisheries and Aquaculture Department, Food and Agriculture Organization of the United Nations, Rome, Italy. Accessed 5 October 2016. http://www.fao.org/fishery/statistics/global-aquaculture-production/en Fisheries Victoria & MAFRI 2002. Developmental Fisheries Management Plan, Jellyfish (Catostylus mosaicus), 2003-2005. Fisheries Victoria and the Marine and Freshwater Resources Institute, Fisheries Division, Department of Natural Resources and Environment, Victoria, Australia, 15 pp. Fisheries Victoria 2006. Statement of management arrangements for the Victorian developmental jellyfish fishery (Catostylus mosaicus). Statement prepared for the Australian Department of Hertiage and Environment. Fisheries Victoria Division of the Department of Primary Industries, Victoria, Australia, 15 pp. Fleming, N.E.C., C. Harrod, D.C. Griffin, J. Newton, & J.D.R. Houghton 2014. Scyphozoan jellyfish provide short-term reproductive habitat for hyperiid amphipods in a temperate near-shore environment, Marine Ecology Progress Series 510: 229-240. Fowler, S.W., J.L. Teyssie, O. Cotret, B. Danis, C. Rouleau, & A. Warnau 2004. Applied radiotracer techniques for studying pollutant bioaccumulation in 137 selected marine organisms (jellyfish, crabs and sea stars), Nukleonika 49(3): 97-100. Frandsen, K.T. & H.U. Riisgard 1997. Size dependent respiration and growth of jellyfish, Aurella aurita, Sarsia 82(4): 307-312. Fukushi, K., N. Ishio, J. Tsujimoto, K. Yokota, T. Hamatake, H. Sogabe, K. Toriya, & T. Ninomiya 2004. Preliminary study on the potential usefulness of jellyfish as fertilizer, Bulletin of the Society of Sea Water Science, Japan 58(2): 209-217. Fukushi, K., J. Tsujimoto, & K. Yokota 2005. Determination of inorganic constituents in jellyfish, Bunseki Kagaku 54(2): 175-178 (in Japanese with English abstract). Gemmell, B.J., J.H. Costello, S.P. Colin, C.J. Stewart, J.O. Dabiri, D. Tafti, & S. Priya 2013. Passive energy recapture in jellyfish contributes to propulsive advantage over other metazoans, Proceedings of the National Academy of Sciences of the United States of America 110(44): 17904-17909. Gemmell, B.J., S.P. Colin, J.H. Costello, & J.O. Dabiri 2015. Suction-based propulsion as a basis for efficient animal swimming, Nature Communications 6: 8790. Georges, A. 1991. A management program for the sustainable harvest and export of the edible jellyfish Catostylus mosaicus. Prepared for Taiga Food Pty Ltd for submission to the Australian National Parks and Wildlife Service. Applied Ecology Research Group, University of Canberra. 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. Gopakumar, G., K. Madhu, R. Jayashankar, R. Madhu, J.K. Kizhakudan, J. Jose, B. Ignatius, P. Vijayagopal, S. Joseph, G. Mohamed, & M. Varghese 2008. Live feed research for larviculture of marine finfish and shellfish. Marine Fisheries Information Service (197), Technical and Extension Series. Central Marine 138 Fisheries Research Institute, Indian Council of Agricultural Research, Cochin, India, 1-6 pp. Gordon, M., C. Hatcher, & J. Seymour 2004. Growth and age determination of the tropical Australian cubozoan Chiropsalmus sp., Hydrobiologia 530: 339-345. Gordon, M. & J. Seymour 2012. Growth, development and temporal variation in the onset of six Chironex fleckeri medusae seasons: a contribution to understanding jellyfish ecology, PLoS ONE 7(2): e31277. Govindan, T.K. 1984. A novel marine animal with much export potential, Seafood Export Journal 16(7): 9-11. Graham, W.M., S. Gelcich, K.L. Robinson, C.M. Duarte, L. Brotz, J.E. Purcell, L.P. Madin, H. Mianzan, K.R. Sutherland, S. Uye, K.A. Pitt, C.H. Lucas, M. Bøgeberg, R.D. Brodeur, & R.H. Condon 2014. Linking human well-being and jellyfish: ecosystem services, impacts, and societal responses, Frontiers in Ecology and the Environment 12(9): 515-523. Graitcer, P. 2012. US Jellyfish land on Asian dinner tables, Voice of America, edition of March 20. Grocock, C. & S. Grainger 2006. Apicius - a critical edition with an introduction and an English translation of the Latin recipe text Apicius. Prospect Books, Devon, United Kingdom, 414 pp. Gul, S. & A.C. Morandini 2013. New records of scyphomedusae from Pakistan coast: Catostylus perezi and Pelagia cf. noctiluca, Marine Biodiversity Records 6(e86): 1-6. Gul, S., S. Jahangir, & A. Schiariti 2015. Jellyfish fishery in Pakistan, Plankton & Benthos Research 10(4): 220-224. Gul, S. & A.C. Morandini 2015. First record of the jellyfish Rhopilema hispidum from the coast of Pakistan, Marine Biodiversity Records 8(e30): 1-4. 139 Gusmani, L., M. Avian, B. Galil, P. Patriarca, & G. Rottini 1997. Biologically active polypeptides in the venom of the jellyfish Rhopilema nomadica, Toxicon 35(5): 637-648. Hamner, W.M. & R.M. Jenssen 1974. Growth, degrowth, and irreversible cell differentiation in Aurelia aurita, American Zoologist 14(2): 833-849. Hay, S. 2006. Marine ecology: gelatinous bells may ring change in marine ecosystems, Current Biology 16(17): R679-R682. He, J.R., L.M. Zheng, W.J. Zhang, & Y.S. Lin 2015. Life Cycle Reversal in Aurelia sp.1 (Cnidaria, Scyphozoa), PLoS ONE 10(12): e0145314. Heaslip, S.G., S.J. Iverson, W.D. Bowen, & M.C. James 2012. Jellyfish support high energy intake of leatherback sea turtles (Dermochelys coriacea): video evidence from animal-borne cameras, PLoS ONE 7(3): e33259. Heeger, T. 1998. Quallen. Gefährliche Schönheiten. Stuttgart, Wissenschaftliche Verlagsgesellschaft mbH, Stuttgart, Germany. Henderson, C.J. & R.S. Panush 1999. Diets, dietary supplements, and nutritional therapies in rheumatic diseases, Rheumatic Disease Clinics of North America 25(4): 937-968. Henschke, N., D.A. Bowden, J.D. Everett, S.P. Holmes, R.J. Kloser, R.W. Lee, & I.M. Suthers 2013. Salp-falls in the Tasman Sea: a major food input to deep-sea benthos, Marine Ecology Progress Series 491: 165-175. Henschke, N., J.D. Everett, A.J. Richardson, & I.M. Suthers 2016. Rethinking the role of salps in the ocean, Trends in Ecology & Evolution 31(9): 720-733. Hofmann, D.K. & M.G. Hadfield 2002. Hermaphroditism, gonochorism, and asexual reproduction in Cassiopea sp. - an immigrant in the islands of Hawai'i, Invertebrate Reproduction & Development 41(1-3): 215-221. 140 Holland, B.S., M.N. Dawson, G.L. Crow, & D.K. Hofmann 2004. Global phylogeography of Cassiopea: molecular evidence for cryptic species and multiple invasions of the Hawaiian Islands, Marine Biology 145(6): 1119-1128. Hong, H. 2002. The jellyfish and Rhopilema esculentum, The Biological Bulletin 37(2): 13-16 (in Chinese). Hossain, S.T., H. Sugimoto, N. Asagi, T. Araki, H. Ueno, M. Morokuma, & H. Kato 2013. The use of desalinated-dried jellyfish and rice bran for controlling weeds and rice yield, Journal of Organic Systems 8(1): 28-37. Hsieh, Y.H.P. & J. Rudloe 1994. Potential of utilizing jellyfish as food in Western countries, Trends in Food Science & Technology 5(7): 225-229. Hsieh, Y.H.P., F. Leong, & K.W. Barnes 1996. Inorganic constituents in fresh and processed cannonball jellyfish (Stomolophus meleagris), Journal of Agricultural and Food Chemistry 44(10): 3117-3119. Hsieh, Y.H.P., F.M. Leong, & J. Rudloe 2001. Jellyfish as food, Hydrobiologia 451(1-3): 11-17. Hsieh, Y.H.P. 2005. Use of jellyfish collagen (type II) in the treatment of rheumatoid arthritis, United States Patent No. 6,894,029, Auburn University. May 17, 2005. Huang, H.Y., T.K. Tso, C.H. Li, J.W. Cheng, & W.H. Shen 2005. Isolation and partial characterization of collagen from edible Rhopilema esculenta jellyfish, Faseb Journal 19(5): A1032-A1033. Huang, M., J. Hu, Y. Wang, & Z. Chen 1985. Preliminary study on the breeding habits of edible jellyfish in Hangzhou Wan Bay, Journal of Fisheries of China 9(3): 239-246 (in Chinese). Huang, Y. 1986. The processing of cannonball jellyfish (Stomolophus meleagris) and its utilization. Proceedings of the 11th Annual Tropical and Subtropical Fisheries 141 Technological Conference of the Americas, January 13-16, 1986. Tampa, Florida, U.S.A., 141-146 pp. Huang, Y., P. Christian, & D. Colson 1987. Harvest and preservation of a shrimp by-catch: cannonball jellyfish (Stomolophus meleagris). Proceedings of the 12th Annual Tropical and Subtropical Fisheries Technological Conference of the Americas, November 9-11, 1987. Orlando, Florida, U.S.A., 466-474 pp. Huang, Y. 1988. Cannonball jellyfish (Stomolophus meleagris) as a food resource, Journal of Food Science 53(2): 341-343. Hudson, R.J., N.F. Bridge, & T.I. Walker 1997. Feasibility study for establishment of a Victorian commercial jellyfish fishery. Marine and Freshwater Resources Institute, Fisheries Research Development Corporation, Queenscliff, Australia, 44 pp. Hussein, O.S. & O.I. Saleh 2014. Effect of soaking in jellyfish on some parameters of wheat and lentil seedlings, Journal of Environmental Science, Toxicology and Food Technology 8(4): 32-39. Hussein, O.S., R.M. Sayed, & O.I. Saleh 2015. Uses of jellyfish in pre sowing seeds treatment and pest control, American Journal of Experimental Agriculture 5(1): 60-69. I-chia, L. 2016. FDA to begin enforcing aluminum guidelines, Taipei Times, Taipei, Taiwan, edition of January 20. Imamura, K., D. Tsuruta, A. Tsuchisaka, T. Mori, C. Ohata, M. Furumura, & T. Hashimoto 2013. Anaphylaxis caused by ingestion of jellyfish, European Journal of Dermatology 23(3): 392-395. Inomata, N., K. Chin, & M. Aihara 2014. Anaphylaxis caused by ingesting jellyfish in a subject with fermented soybean allergy: possibility of epicutaneous 142 sensitization to poly-gamma-glutamic acid by jellyfish stings, Journal of Dermatology 41(8): 752-753. Jacquet, J.L. & D. Pauly 2008. Trade secrets: Renaming and mislabeling of seafood, Marine Policy 32(3): 309-318. James, D.B., E. Vivekanandan, & S. Srinivasarengan 1985. Menace from medusae off Madras with notes on their utility and toxicity, Journal of the Marine Biological Association of India 27(1 & 2): 170-174. Jarms, G. 2010. The early life of Scyphozoa with emphasis on Coronatae: a review with a list of described life cycles, Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg 45: 17-31. Jenkins, L. 2012. Reducing sea turtle bycatch in trawl nets: a history of NMFS turtle excluder device (TED) research, Marine Fisheries Review 74(2): 26-44. Jennings, S., M.J. Kaiser, & J.D. Reynolds 2001. Marine Fisheries Ecology. Blackwell Publishing, Malden, U.S.A., 417 pp. Jones, R.P. & J. Rudloe 1995. Harvesting & processing of Florida cannonball jellyfish. Report to the Florida International Affairs Commission, Florida Department of Commerce. Southeastern Fisheries Association, 28 + viii pp. Jones, S. 1960. Notes on animal associations. 2. The scyphomedusa, Acromitus ftagellatus and young Steroides leptolepis with the latter forming a vanguard, Journal of the Marine Biological Association of India 2(1): 51-52. Kailola, P.J., M.J. Williams, P.C. Stewart, R.E. Reichelt, A. McNee, & C. Grieve 1993. Australian Fisheries Resources. Bureau of Resource Sciences and the Fisheries Research and Development Corporation, Canberra, Australia, 422 pp. Kawabata, T., D.J. Lindsay, M. Kitamura, S. Konishi, J. Nishikawa, S. Nishida, M. Kamio, & H. Nagai 2013. Evaluation of the bioactivities of water-soluble extracts from twelve deep-sea jellyfish species, Fisheries Science 79(3): 487-494. 143 Kawahara, M., S. Uye, J. Burnett, & H. Mianzan 2006a. Stings of edible jellyfish (Rhopilema hispidum, Rhopilema esculentum and Nemopilema nomurai) in Japanese waters, Toxicon 48(6): 713-716. Kawahara, M., S. Uye, K. Ohtsu, & H. Izumi 2006b. Unusual population explosion of the giant jellyfish Nemopilemia nomurai in East Asian waters, Marine Ecology Progress Series 307: 161-173. Kelsey, M. 2009. Stinging intruders - invasive jellyfish removed from Kaunakakai wharf, The Molokai Dispatch, Kaunakakai, Hawaii, U.S.A., edition of June 23. Khong, N.M.H., F.M. Yusoff, B. Jamilah, M. Basri, I. Maznah, K.W. Chan, & J. Nishikawa 2016. Nutritional composition and total collagen content of three commercially important edible jellyfish, Food Chemistry 196: 953-960. Kikinger, R. 1992. Cotylorhiza tuberculata - life history of a stationary population, Marine Ecology 13(4): 333-362. Kim, D.H., J.N. Seo, W.D. Yoon, & Y.S. Suh 2012a. Estimating the economic damage caused by jellyfish to fisheries in Korea, Fisheries Science 78(5): 1147-1152. Kim, Y.-R., S.-W. Kim, E. Damdinsuren, T. Ezaki, & K.W. Chun 2012b. The effect of jellyfish (Nemopilema nomurai) fertilizer on tree growth in hillside erosion control works, Journal of Forest and Environmental Science 28(4): 227-231. Kimura, S., S. Miura, & Y.H. Park 1983. Collagen as the major edible component of jellyfish (Stomolophus nomurai), Journal of Food Science 48(6): 1758-1760. Kingsford, M.J. 1993. Biotic and abiotic structure in the pelagic environment - importance to small fishes, Bulletin of Marine Science 53(2): 393-415. Kingsford, M.J., K.A. Pitt, & B.M. Gillanders 2000. Management of jellyfish fisheries, with special reference to the Order Rhizostomeae, Oceanography and Marine Biology 38: 85-156. 144 Kitamura, M. & M. Omori 2010. Synopis of edible jellyfishes collected from Southeast Asia, with notes on jellyfish fisheries, Plankton & Benthos Research 5(3): 106-118. Klaiwong, T., P. Hutangura, S. Rutatip, P. Wongsa-Ngasri, & B. Thumthanaruk 2014. Comparative properties of pepsin hydrolyzed jellyfish protein from salted jellyfish, Journal of Agricultural Science and Technology B 4(7): 555-564. Kondo, Y., S. Ohtsuka, J. Nishikawa, E. Metillo, H. Pagliawan, S. Sawamoto, M. Moriya, S. Nishida, & M. Urata 2014. Associations of fish juveniles with rhizostome jellyfishes in the Philippines, with taxonomic remarks on a commercially harvested species in Carigara Bay, Leyte Island, Plankton & Benthos Research 9(1): 51-56. Kraeuter, J.N. & E.M. Setzler 1975. Seasonal cycle of Scyphozoa and Cubozoa in Georgia estuaries, Bulletin of Marine Science 25(1): 66-74. Krishnan, S. & P. Perumal 2013. Preparation and biomedical characterization of jellyfish (Chrysaora quinquecirrha) collagen from southeast coast of India, International Journal of Pharmacy and Pharmaceutical Sciences 5(3): 698-701. Kuthalingam, M.D.K., D.B. James, R. Sarvesan, P. Devadoss, S. Manivasagam, & P. Thirumilu 1989. A note on the processing of the jelly fish at Alambaraikuppam near Mahabalipuram. Marine Fisheries Information Service, Technical and Extension Series (98). Central Marine Fisheries Research Institute, Indian Council of Agricultural Research. Cochin, India, 8-10 pp. Landers, M. 2011. Coastal Georgia shrimpers turn to jellyfish to make money, Savannah Morning News, Savannah, Georgia, U.S.A., edition of June 7. Lebrato, M., K.A. Pitt, A.K. Sweetman, D.O.B. Jones, J.E. Cartes, A. Oschlies, R.H. Condon, J.C. Molinero, L. Adler, C. Gaillard, D. Lloris, & D.S.M. Billett 2012. 145 Jelly-falls historic and recent observations: a review to drive future research directions, Hydrobiologia 690(1): 227-245. Lebrato, M., P.d.J. Mendes, D.K. Steinberg, J.E. Cartes, B.M. Jones, L.M. Birsa, R. Benavides, & A. Oschlies 2013. Jelly biomass sinking speed reveals a fast carbon export mechanism, Limnology and Oceanography 58(3): 1113-1122. Leone, A., R.M. Lecci, M. Durante, & S. Piraino 2013. Extract from the zooxanthellate jellyfish Cotylorhiza tuberculata modulates gap junction intercellular communication in human cell cultures, Marine Drugs 11(5): 1728-1762. Leone, A., R.M. Lecci, M. Durante, F. Meli, & S. Piraino 2015. The bright side of gelatinous blooms: nutraceutical value and antioxidant properties of three Mediterranean jellyfish (Scyphozoa), Marine Drugs 13(8): 4654-4681. Li, C.P., H.H. Yu, S. Liu, R.E. Xing, Z.Y. Guo, & P.C. Li 2005. Factors affecting the protease activity of venom from jellyfish Rhopilema esculentum, Bioorganic & Medicinal Chemistry Letters 15(24): 5370-5374. Li, J., J. Ling, & J. Cheng 2014. On utilization of two edible macro-jellyfish and evaluation of the biomass of Nemopilema nomurai in China Sea, Marine Fisheries 36(3): 202-207 (in Chinese with English abstract). Li, J.R. & Y.H.P. Hsieh 2004. Traditional Chinese food technology and cuisine, Asia Pacific Journal of Clinical Nutrition 13(2): 147-155. Lilley, M.K.S., D. Thibault-Botha, & F. Lombard 2014. Respiration demands increase significantly with both temperature and mass in the invasive ctenophore Mnemiopsis leidyi, Journal of Plankton Research 36(3): 831-837. Liu, C.-s., S.-q. Chen, Z.-m. Zhuang, J.-p. Yan, C.-l. Liu, & H.-t. Cui 2015. Potential of utilizing jellyfish as food in culturing Pampus argenteus juveniles, Hydrobiologia 754(1): 189-200. 146 Liu, J.X., H.W. Qiu, & G.D. Zhuang 2009. Study on preparation technology of angiotensin-converting enzyme inhibitory peptides (ACEIP) from jellyfish and their antihypertensive effect in vivo, Food Science 30(11): 227-231 (in Chinese with English abstract). Liu, X., M.S. Zhang, A.R. Jia, Y.G. Zhang, H.M. Zhu, C. Zhang, Z.L. Sun, & C.H. Liu 2013. Purification and characterization of angiotensin I converting enzyme inhibitory peptides from jellyfish Rhopilema esculentum, Food Research International 50(1): 339-343. Liu, Y. 1973. Study on processing technology of jellyfish, Fisheries Science & Technology Information 1(7): 16-17 (in Chinese). Lluch-Cota, S.E., E.A. Aragón-Noriega, F. Arreguín-Sánchez, D. Aurioles-Gamboa, J.J. Bautista-Romero, R.C. Brusca, R. Cervantes-Duarte, R. Cortés-Altamirano, P. Del-Monte-Luna, A. Esquivel-Herrera, G. Fernández, M.E. Hendrickx, S. Hernández-Vázquez, H. Herrera-Cervantes, M. Kahru, M. Lavín, D. Lluch-Belda, D.B. Lluch-Cota, J. López-Martínez, S.G. Marinone, M.O. Nevárez-Martínez, S. Ortega-García, E. Palacios-Castro, A. Parés-Sierra, G. Ponce-Díaz, M. Ramírez-Rodríguez, C.A. Salinas-Zavala, R.A. Schwartzlose, & A.P. Sierra-Beltrán 2007. The Gulf of California: Review of ecosystem status and sustainability challenges, Progress in Oceanography 73(1): 1-26. López-Martinez, J. & J. Álvarez-Tello 2013. The jellyfish fishery in Mexico, Agricultural Sciences 4(6A): 57-61. López-Martínez, J. & J. Rodríguez-Romero 2008. First record of association of the blackfin jack Hemicaranx zelotes with the cannonball jellyfish Stomolophus meleagris in Kino Bay, Gulf of California, Hidrobiológica 18(2): 173-176 (in Spanish). 147 Lu, Z., Q. Dai, & Y. Yan 2003. Fishery biology of Cyanea nozakii resources in the waters of Dongshan Island, Chinese Journal of Applied Ecology 14(6): 973-976 (in Chinese with English abstract). Lucas, C.H., W.M. Graham, & C. Widmer 2012. Jellyfish life histories: role of polyps in forming and maintaining scyphomedusa populations, Advances in Marine Biology 63: 133-196. Lucas, C.H., S. Gelcich, & S.-I. Uye 2014. Living with jellyfish: management and adaptation strategies, pp. 129-150 in C. H. Lucas & K. A. Pitt (eds.) Jellyfish Blooms. Springer, Dordrecht, Netherlands. Magesh, S.J. & S. Coulthard 2004. Bloom or bust? Samudra Report No. 39, November, 15-20 pp. Mariottini, G.L. & L. Pane 2014. Cytotoxic and cytolytic cnidarian venoms. A review on health implications and possible therapeutic applications, Toxins 6(1): 108-151. Mariottini, G.L. & L. Brotz in press. Cnidarian venoms and alternative research methods: from cell damage to possible applications, in G. L. Mariottini (ed.) Jellyfish: Ecology, Distribution Patterns and Human Interactions. Nova Publishers, Hauppauge, New York, U.S.A. Masuda, A., T. Baba, N. Dohmae, M. Yamamura, H. Wada, & K. Ushida 2007. Mucin (Qniumucin), a glycoprotein from jellyfish, and determination of its main chain structure, Journal of Natural Products 70(7): 1089-1092. Matsumura, K., K. Kamiya, K. Yamashita, F. Hayashi, I. Watanabe, Y. Murao, H. Miyasaka, N. Kamimura, & M. Nogami 2005. Genetic polymorphism of the adult medusae invading an electric power station and wild polyps of Aurelia aurita in Wakasa Bay, Japan, Journal of the Marine Biological Association of the United Kingdom 85(3): 563-568. 148 MFD 2012. Handbook of fisheries statistics of Pakistan. Vol. 20. Marine Fisheries Department (MFD), Government of Pakistan, Fish Harbour, West Wharf, Karachi, Pakistan. Mianzan, H., J. Quiñones, S. Palma, A. Schiariti, E. Marcelo Acha, K.L. Robinson, & W.M. Graham 2014. Chrysaora plocamia: a poorly understood jellyfish from South America, pp. 219-236 in K. A. Pitt & C. H. Lucas (eds.) Jellyfish Blooms. Springer, Dordrecht, Netherlands. Mills, C.E. 1995. Medusae, siphonophores, and ctenophores as planktivorous predators in changing global ecosystems, ICES Journal of Marine Science 52(3-4): 575-581. Mills, C.E. 2001. Jellyfish blooms: are populations increasing globally in response to changing ocean conditions?, Hydrobiologia 451: 55-68. Miyajima, Y., R. Masuda, A. Kurihara, R. Kamata, Y. Yamashita, & T. Takeuchi 2011a. Juveniles of threadsail filefish, Stephanolepis cirrhifer, can survive and grow by feeding on moon jellyfish Aurelia aurita, Fisheries Science 77(1): 41-48. Miyajima, Y., R. Masuda, & Y. Yamashita 2011b. Feeding preference of threadsail filefish Stephanolepis cirrhifer on moon jellyfish and lobworm in the laboratory, Plankton & Benthos Research 6(1): 12-17. Mohammed, B. 2008. Taking the sting out of jellyfish, Gulf Daily News, Manama, Bahrain, edition of July 28. Mohan, S., S. Rajapackiam, & S. Rajan 2011. Unusual heavy landings of jellyfish Crambionella stuhlmanni and processing methods at Pulicat landing centre, Chennai. Marine Fisheries Information Service Technical and Extension Series. Marine Fisheries Information Service (208), Technical and Extension Series. Central Marine Fisheries Research Institute, Indian Council of Agricultural Research, Cochin, India, 27-29 pp. 149 Möller, H. 1980. Scyphomedusae as predators and food competitors of larval fish, Meeresforschung 28(2-3): 90-100. Moody, E. 2014. Jellyfish operation brings in catch, catches DHEC's attention, The Beaufort Gazette, Bluffton, South Carolina, U.S.A., edition of April 12. Mooney, C.J. & M.J. Kingsford 2016a. Statolith morphometrics as a tool to distinguish among populations of three cubozoan species, Hydrobiologia doi:10.1007/s10750-016-2949-6. Mooney, C.J. & M.J. Kingsford 2016b. Discriminating populations of medusae (Chironex fleckeri, Cubozoa) using statolith microchemistry, Marine and Freshwater Research doi:10.1071/MF16104. Morabito, R., S. Dossena, G. La Spada, & A. Marino 2014. Heavy metals affect nematocysts discharge response and biological activity of crude venom in the jellyfish Pelagia noctiluca, Cellular Physiology and Biochemistry 34(2): 244-254. Morandini, A.C., A. Schiariti, S.N. Stampar, M.M. Maronna, I. Straehler-Pohl, & A.C. Marques 2016. Succession of generations is still the general paradigm for scyphozoan life cycles, Bulletin of Marine Science 92(3). Morikawa, T. 1984. Jellyfish, FAO INFOFISH Marketing Digest 1(84): 37-39. Morishige, H., T. Sugahara, S. Nishimoto, A. Muranaka, F. Ohno, R. Shiraishi, & M. Doi 2011. Immunostimulatory effects of collagen from jellyfish in vivo, Cytotechnology 63(5): 481-492. Muhammed, F. & R. Sultana 2008. New record of edible jellyfish, Rhizostoma pulmo from Pakistani waters, Marine Biodiversity Records 1(e67): 1-3. Mujiono, N. 2010. Jellyfish (Crambionella sp.) fisheries around Cilacap waters, central Java (Cnidaria: Scyphozoa), Oseanologi dan Limnologi di Indonesia 36(1): 37-48. Muñoz-Vera, A., J.M.P. Castejón, & G. García 2016. Patterns of trace element bioaccumulation in jellyfish Rhizostoma pulmo (Cnidaria, Scyphozoa) in a 150 Mediterranean coastal lagoon from SE Spain, Marine Pollution Bulletin 110(1): 143-154. Murdock, Z. 2014. Jellyfish operations suspended pending state reviews, The Beaufort Gazette, Bluffton, South Carolina, U.S.A., edition of April 22. Murugan, A. & R. Durgekar 2008. Beyond the Tsunami: Status of Fisheries in Tamil Nadu, India: A Snapshot of Present and Long-term Trends. United Nations Development Programme and Ashoka Trust for Research in Ecology and the Environment, Bangalore, India, 75 pp. Naalir, M. 2008. Jelly fish exports becoming lucrative foreign exchange earner -Fisheries Minister, Daily News, Sri Lanka, edition of September 18, 2008. Nagai, T., W. Worawattanamateekul, N. Suzuki, T. Nakamura, T. Ito, K. Fujiki, M. Nakao, & T. Yano 2000. Isolation and characterization of collagen from rhizostomous jellyfish (Rhopilema asamushi), Food Chemistry 70(2): 205-208. Najem, J., S.A. Sarles, B. Akle, & D.J. Leo 2012. Biomimetic jellyfish-inspired underwater vehicle actuated by ionic polymer metal composite actuators, Smart Materials and Structures 21(9): 094026. NARA 2010. Press statement on recent occurrence of jellyfish on south west coast. National Aquatic Resources Research & Devlopment Agency, Sri Lanka. Nayak, P. 2002. Aluminum: impacts and disease, Environmental Research 89(2): 101-115. Nilsson, D.E., L. Gislen, M.M. Coates, C. Skogh, & A. Garm 2005. Advanced optics in a jellyfish eye, Nature 435(7039): 201-205. Nishikawa, J., N.T. Thu, T.M. Ha, & P.T. Thu 2008. Jellyfish fisheries in northern Vietnam, Plankton & Benthos Research 3(4): 227-234. Nishikawa, J., N.T. Thu, F.M. Yusoff, D.J. Lindsay, Mulyadi, N. Mujiono, S. Ohtsuka, & S. Nishida 2009. Jellyfish fisheries in Southeast Asia, with special reference 151 to those in Vietnam, Indonesia, and Malaysia, Kaiyo Monthly 41(7): 401-411 (in Japanese). Nishikawa, J., S. Ohtsuka, Mulyadi, N. Mujiono, D.J. Lindsay, H. Miyamoto, & S. Nishida 2015. A new species of the commercially harvested jellyfish Crambionella from central Java, Indonesia with remarks on the fisheries, Journal of the Marine Biological Association of the United Kingdom 95(3): 471-481. Nishimoto, S., Y. Goto, H. Morishige, R. Shiraishi, M. Doi, K. Akiyama, S. Yamauchi, & T. Sugahara 2008. Mode of action of the immunostimulatory effect of collagen from jellyfish, Bioscience Biotechnology and Biochemistry 72(11): 2806-2814. Nobel, J. 2014. Jellyfish: it's what's for dinner, Modern Farmer, Hudson, New York, U.S.A., edition of September 15. Ogimoto, M., K. Suzuki, J. Kabashima, M. Nakazato, & Y. Uematsu 2012. Aluminium content in foods with aluminium-containing food additives, Food Hygiene and Safety Science 53(1): 57-62 (in Japanese with English abstract). Ohta, N., M. Sato, K. Ushida, M. Kokubo, T. Baba, K. Taniguchi, M. Urai, K. Kihira, & J. Mochida 2009. Jellyfish mucin may have potential disease-modifying effects on osteoarthritis, BMC Biotechnology 9: 98. Okubo, Y., K. Yoshida, M. Furukawa, M. Sasaki, H. Sakakibara, T. Terakawa, & A. Akasawa 2015. Anaphylactic shock after the ingestion of jellyfish without a history of jellyfish contact or sting, European Journal of Dermatology 25(5): 491-492. Omori, M. 1978. Zooplankton fisheries of the world: a review, Marine Biology 48: 199-205. 152 Omori, M. 1981. Edible jellyfish in the far east waters: a brief review of the biology and fishery, Bulletin of the Plankton Society of Japan 28(1): 1-111 (in Japanese with English abstract). Omori, M. & E. Nakano 2001. Jellyfish fisheries in southeast Asia, Hydrobiologia 451(1-3): 19-26. Omori, M. & M. Kitamura 2004. Taxonomic review of three Japanese species of edible jellyfish, Plankton Biology and Ecology 51(1): 36-51. Omori, M. & M. Kitamura 2011. Seven species of edible jellyfish from Southeast Asia, Midoriishi 22: 1-7 (in Japanese). Ovchinnikova, T.V., S.V. Balandin, G.M. Aleshina, A.A. Tagaev, Y.F. Leonova, E.D. Krasnodembsky, A.V. Men'shenin, & V.N. Kokryakov 2006. Aurelin, a novel antimicrobial peptide from jellyfish Aurelia aurita with structural features of defensins and channel-blocking toxins, Biochemical and Biophysical Research Communications 348(2): 514-523. Ozer, N.P. & M.S. Celikkale 2001. Utilization possibilities of jellyfish Rhizostoma pulmo, as a food in the Black Sea, Journal of Food Science and Technology 38(2): 175-178. Page, J.W. 2015. Characterization of bycatch in the cannonball jellyfish fishery in the coastal waters off Georgia, Marine and Coastal Fisheries 7(1): 190-199. Palomares, M.L.D. & D. Pauly 2009. The growth of jellyfishes, Hydrobiologia 616: 11-21. Panda, S.K. & V.R. Madhu 2009. Studies on the preponderance of jellyfish in coastal waters of Veraval, Fishery Technology 46(2): 99-106. Patwa, A., A. Thiery, F. Lombard, M.K.S. Lilley, C. Boisset, J.F. Bramard, J.Y. Bottero, & P. Barthelemy 2015. Accumulation of nanoparticles in "jellyfish" mucus: a 153 bio-inspired route to decontamination of nano-waste, Scientific Reports 5: 11387. Pauly, D. 1995. Anecdotes and the shifting base-line syndrome of fisheries, Trends in Ecology & Evolution 10(10): 430-430. Pauly, D. 1998. Rationale for reconstructing catch time series, EC Fisheries Cooperation Bulletin 11(2): 4-7. Pauly, D., V. Christensen, J. Dalsgaard, R. Froese, & F. Torres 1998. Fishing down marine food webs, Science 279(5352): 860-863. Pauly, D., V. Christensen, S. Guénette, T.J. Pitcher, U.R. Sumaila, C.J. Walters, R. Watson, & D. Zeller 2002. Towards sustainability in world fisheries, Nature 418(6898): 689-695. Pauly, D., W. Graham, S. Libralato, L. Morissette, & M.L.D. Palomares 2009. Jellyfish in ecosystems, online databases, and ecosystem models, Hydrobiologia 616: 67-85. Pauly, D. & A. Greenberg (eds.) 2013. ELEFAN in R: A new tool for length-frequency analysis. Fisheries Centre Research Report 21(3), Fisheries Centre, University of British Columbia, Vancouver, Canada, iv + 46 pp. Pauly, D. 2016. On the importance of fisheries catches, with a rationale for their reconstruction, pp. 1-11 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. Pauly, D. & D. Zeller (eds.) 2016a. Global Atlas of Marine Fisheries: A Critical Appraisal of Catches and Ecosystem Impacts. Island Press, Washington, D.C., U.S.A., 497 pp. 154 Pauly, D. & D. Zeller 2016b. Catch reconstructions reveal that global marine fisheries catches are higher than reported and declining, Nature Communications 7: 10244. PCAMRD 2008. Small fishermen in the Malampaya Sound of Palawan benefit from jellyfish, The PCAMRD Waves, Philippine Council for Aquatic and Marine Research and Development, Department of Science and Technology, edition of January-March. Perera, L. 2008. Jellyfish exports need checking, The Sunday Times, Sri Lanka, edition of September 21. Perl, D.P. & A.R. Brody 1980. Alzheimer's disease - X-ray spectrometric evidence of aluminum accumulation in nerurofibrillary tangle-bearing neurons, Science 208(4441): 297-299. Pitcher, T.J. & D. Preikshot 2001. RAPFISH: a rapid appraisal technique to evaluate the sustainability status of fisheries, Fisheries Research 49(3): 255-270. Pitt, K. & M.J. Kingsford 2000. Reproductive biology of the edible jellyfish Catostylus mosaicus, Marine Biology 137: 791-799. Prabhu, M.S. 1954. The perch fishery by special traps in the area around Mandapam in the Gulf of Mannar and Palk Bay, Indian Journal of Fisheries 1: 94-129. Prieto, L., D. Astorga, G. Navarro, & J. Ruiz 2010. Environmental control of phase transition and polyp survival of a massive-outbreaker jellyfish, PLoS ONE 5(11): e13793. Pugh, P.R. 1989. Gelatinous zooplankton - the forgotten fauna, Progress in Underwater Science 14: 67-78. Purcell, J. 2012. Jellyfish and ctenophore blooms coincide with human proliferations and environmental perturbations, Annual Review of Marine Science 4: 209-235. 155 Purcell, J.E. 1985. Predation on fish eggs and larvae by pelagic cnidarians and ctenophores, Bulletin of Marine Science 37(2): 739-755. Purcell, J.E. 1997. Pelagic cnidarians and ctenophores as predators: selective predation, feeding rates, and effects on prey populations, Annales de l'Institut Océanographique 73(2): 125-137. Purcell, J.E. & M.N. Arai 2001. Interactions of pelagic cnidarians and ctenophores with fish: a review, Hydrobiologia 451(1-3): 27-44. Purcell, J.E., S. Uye, & W.T. Lo 2007. Anthropogenic causes of jellyfish blooms and their direct consequences for humans: a review, Marine Ecology Progress Series 350: 153-174. Rastogi, A., S. Biswas, A. Sarkar, & D. Chakrabarty 2012. Anticoagulant activity of moon jellyfish (Aurelia aurita) tentacle extract, Toxicon 60(5): 719-723. Richardson, A.J., A. Bakun, G.C. Hays, & M.J. Gibbons 2009. The jellyfish joyride: causes, consequences and management responses to a more gelatinous future, Trends in Ecology & Evolution 24(6): 312-322. Rigby, R.J. & M. Hafey 1972. Thermal properties of the collagen of jellyfish (Aurella coerulea) and their relation to its thermal behaviour, Australian journal of biological sciences 25(6): 1361-3. Ristroph, L. & S. Childress 2014. Stable hovering of a jellyfish-like flying machine, Journal of the Royal Society Interface 11(92): 20130992. Rizman-Idid, M., A.B. Farrah-Azwa, & V.C. Chong 2016. Preliminary taxonomic survey and molecular documentation of jellyfish species (Cnidaria: Scyphozoa and Cubozoa) in Malaysia, Zoological Studies 55(35): 19 pp. Robinson, K.L., J.J. Ruzicka, M.B. Decker, R.D. Brodeur, F.J. Hernandez, J. Quiñones, E.M. Acha, S. Uye, H. Mianzan, & W.M. Graham 2014. Jellyfish, forage fish, and the world's major fisheries, Oceanography 27(4): 104-115. 156 Roda, A. 2010. Discovery and development of the green fluorescent protein, GFP: the 2008 Nobel Prize, Analytical and Bioanalytical Chemistry 396(5): 1619-1622. Rottini, G., L. Gusmani, E. Parovel, M. Avian, & P. Patriarca 1995. Purification and properties of a cytolytic toxin in venom of the jellyifsh Carybdea marsupialis, Toxicon 33(3): 315-326. Rudloe, J. 1992. Jellyfish: a new fishery for the Florida Panhandle. Report prepared for The Apalachee Regional Planning Council, Blounstown, Florida. U.S. Department of Commerce, Economic Development Administration, EDA Project No. 04-06-03801, 35 pp. Ruiz, J., L. Prieto, & D. Astorga 2012. A model for temperature control of jellyfish (Cotylorhiza tuberculata) outbreaks: a causal analysis in a Mediterranean coastal lagoon, Ecological Modelling 233: 59-69. Rumpet, R. 1991. Some aspects of the biology and fishery of jellyfish found along the coast of Sarawak, Malaysia. Department of Fisheries, Ministry of Agriculture, Malaysia, 53 pp. Sal Moyano, M.P., A. Schiariti, D.A. Giberto, L.D. Briz, M.A. Gavio, & H.W. Mianzan 2012. The symbiotic relationship between Lychnorhiza lucerna and Libinia spinosa in the Río de la Plata (Argentina-Uruguay), Marine Biology 159(9): 1933-1941. Salinas-Zavala, C.A., D. Lluch-Belda, S. Hernández-Vazquez, & D.B. Lluch-Cota 1993. La aridez en el noroeste de México: un análisis de su variabilidad espacial y temporal, Atmósfera 11: 29-44 (in Spanish with English abstract). Santhana-Krishnan, G. 1984. Salted jelly fish, Seafood Export Journal 16(7): 23-26. Schiariti, A. 2008. Life history and population dynamics of Lychnorhiza lucerna - an alternative fishery resource?, PhD thesis, Universidad de Buenos Aires, Buenos Aires, Argentina (in Spanish). 157 Schiariti, A., M.P.S. Moyano, D.A. Giberto, & H.W. Mianzan 2012a. First record of the association between Lychnorhiza lucerna and Cyrtograpsus affinis, Latin American Journal of Aquatic Research 40(4): 1090-1093. Schiariti, A., E. Christiansen, A.C. Morandini, F.L. Silveira, D.A. Giberto, & H.W. Mianzan 2012b. Reproductive biology of Lychnorhiza lucerna: Individual traits related to sexual reproduction, Marine Biology Research 8(3): 255-264. Schiariti, A. & H. Mianzan 2013. La medusa Lychnorhiza lucerna como recurso pesquero alternativo en Argentina. Informe de Investigación Nº 36 INIDEP. Área: Pesquerías Pelágicas y Ambiente Marino. Proyecto: Ecología Pesquera, Argentina, 9 pp. (in Spanish). Schiariti, A., V. Fuentes, & H. Mianzan 2015. Pesquerías de medusas en Iberoamérica - ¿Alternativa en tiempos de crisis?, Investigación y Ciencia 465. SEAFDEC 2015. Southeast Asian Fisheries Development Center, Fishery Statistical Bulletins, Bangkok, Thailand. Accessed 5 June 2015. http://fishstat.seafdec.org/ Seo, J.-I., Y.-R. Kim, K. Otsuki, S.-W. Kim, K.-J. Yeom, J.-S. Yi, Y.-S. Ahn, T. Ezaki, & K.-W. Chun 2014. Contributions of jellyfish fertilizer to survival and growth of seedlings planted in a recently burned forest, Republic of Korea, Journal of the Faculty of Agriculture, Kyushu University 59(1): 181-189. Shamah, D. 2014. Israeli tech turns jellyfish into paper towels, Times of Israel, Jerusalem, Israel, edition of April 8. Shelton, P.A. & A.F. Sinclair 2008. It's time to sharpen our definition of sustainable fisheries management, Canadian Journal of Fisheries and Aquatic Sciences 65(10): 2305-2314. 158 Shih, C.T. 1977. A guide to the jellyfish of Canadian Atlantic waters. National Museum of Natural Sciences, National Museums of Canada, Ottawa, Canada, 90 pp. Shimomura, O. 1995. A short story of aequorin, Biological Bulletin 189(1): 1-5. Shimomura, O. 2005. The discovery of aequorin and green fluorescent protein, Journal of Microscopy 217(Pt 1): 3-15. Simpson, A. 2009. Japan hit by invasion of giant Nomura's jellyfish, The Telegraph. Accessed 3 October 2016. http://www.telegraph.co.uk/news/newstopics/howaboutthat/5876499/Japan-hit-by-invasion-of-giant-Nomuras-jellyfish.html Sloan, N.A. & C.R. Gunn 1985. Fishing, processing, and marketing of the jellyfish, Aurelia aurita, from southern British Columbia, Canadian Industry Report of Fisheries and Aquatic Sciences 157: 29 + v pp. Sloan, N.A. 1986. World jellyfish and tunicate fisheries, and the Northeast Pacific echinoderm fishery, Canadian Special Publication of Fisheries and Aquatic Sciences 92: 23-33. Sommer, U., H. Stibor, A. Katechakis, F. Sommer, & T. Hansen 2002. Pelagic food web configurations at different levels of nutrient richness and their implications for the ratio fish production: primary production, Hydrobiologia 484(1-3): 11-20. Song, E., S.Y. Kim, T. Chun, H.J. Byun, & Y.M. Lee 2006. Collagen scaffolds derived from a marine source and their biocompatibility, Biomaterials 27(15): 2951-2961. Soonthonvipat, U. 1976. Dried jelly fish, pp. 149-151 in K. Tiews (ed.) Fisheries Resources and their management in Southeast Asia. Westkreuz-Druckerei, West Berlin, Germany. 159 Su, X.R., C. Yang, C. Chen, & F. Chao 2008. Studies on hypotensive effect of cnidophorin of Stomolophus meleagris, Journal of Chinese Institute of Food Science and Technology 8(3): 42-45 (in Chinese with English abstract). Subasinghe, S. 1992. Jelly fish processing, INFOFISH International 4: 63-65. Suelo, L.G. 1988. Utilisation of the Australian jellyfish Catostylus sp. as a food product, Dissertation Abstracts International, B 48(11): 3167-3168. Swathilekshmi, P.S. 2011. Livelihood and level of aspiration of coastal fisher folk of Tamilnadu, Indian Journal of Social Research 52(1): 31-54. Sweetman, A.K., C.R. Smith, T. Dale, & D.O.B. Jones 2014. Rapid scavenging of jellyfish carcasses reveals the importance of gelatinous material to deep-sea food webs, Proceedings of the Royal Society B 281(1796): 20142210. Tahera, Q. & Q.B. Kazmi 2006. New records of two jellyfish medusae from Pakistani waters, Marine Biodiversity Records 1(e30): 1-4. Teh, L.C.L., D. Zeller, & D. Pauly 2015. Preliminary reconstruction of Thailand's fisheries catches: 1950-2010. Fisheries Centre Working Paper (2015-01), University of British Columbia, Vancouver, Canada, 14 pp. Templeman, M.A. & M.J. Kingsford 2010. Trace element accumulation in Cassiopea sp. from urban marine environments in Australia, Marine Environmental Research 69(2): 63-72. Thanh, C. 2011. Families ride crest of the wave in seafood, Viet Nam News, edition of October 17. Thomas, P.A. 1969. Goat fishes (Family Mullidae) of the Indian Seas, Marine Biological Association of India Memoirs III. Thumthanaruk, B., U. Rodsuwan, P. Chancharern, O. Kerdchoechuen, N. Laohakunjit, & G.W. Chism 2016. Physio-chemical properties of extruded 160 copolymer film, Journal of Food Processing and Preservation doi: 10.1111/jfpp.12808. Tian, T., Z. Chen, M. Wang, Y. Hu, & W. Wang 2016. Inbreeding and genetic diversity analysis in a hatchery release population and clones of Rhopilema esculentum based on microsatellite markers, Chinese Journal of Oceanology and Limnology. Tilves, U., J.E. Purcell, V.L. Fuentes, A. Torrents, M. Pascual, V. Raya, J.-M. Gili, & A. Sabatés 2016. Natural diet and predation impacts of Pelagia noctiluca on fish eggs and larvae in the NW Mediterranean, Journal of Plankton Research 38(5): 1243-1254. Towanda, T. & E.V. Thuesen 2006. Ectosymbiotic behavior of Cancer gracilis and its trophic relationships with its host Phacellophora camtschatica and the parasitoid Hyperia medusarum, Marine Ecology Progress Series 315: 221-236. Ueno, S., C. Imai, & A. Mitsutani 1995. Fine growth rings found in statolith of a cubomedusa Carybdea rastoni, Journal of Plankton Research 17(6): 1381-1384. Ullah, M.S., G.-S. Min, J. Dong, W.D. Yoon, & J.K. Choi 2015. First record of Rhopilema esculentum (Scyphozoa, Rhizostomae), edible jellyfish in Korea, Ocean and Polar Research 37(4): 287-293. USDA 2015. National Nutrient Database for Standard Reference, Release 28, United States Department of Agriculture, Agricultural Research Service, Nutrient Data Laboratory, Washington, D.C., U.S.A. Accessed 8 March 2016. Uye, S. 2008. Blooms of the giant jellyfish Nemopilema nomurai: a threat to the fisheries sustainability of the East Asian Marginal Seas, Plankton & Benthos Research 3(Suppl. 3): 125-131. Valiela, I., J.L. Bowen, & J.K. York 2001. Mangrove forests: One of the world's threatened major tropical environments, Bioscience 51(10): 807-815. 161 van Walraven, L., F. Driessen, J. van Bleijswijk, A. Bol, P.C. Luttikhuizen, J.W.P. Coolen, O.G. Bos, A. Gittenberger, N. Schrieken, V.T. Langenberg, & H.W. van der Veer 2016. Where are the polyps? Molecular identification, distribution and population differentiation of Aurelia aurita jellyfish polyps in the southern North Sea area, Marine Biology 163(8). Varghese, M., C. Kasinathan, & A. Gandhi 2008. Trap fishing in the Gulf of Mannar and Palk Bay. Marine Fisheries Information Service (197), Technical and Extension Series. Central Marine Fisheries Research Institute, Indian Council of Agricultural Research, Cochin, India, 7-8 pp. Vehling, J.D. 1977. Apicius - Cookery and Dining in Imperial Rome. Dover Publications, Toronto, Canada, 301 pp. Verity, P.G. & V. Smetacek 1996. Organism life cycles, predation, and the structure of marine pelagic ecosystems, Marine Ecology Progress Series 130(1-3): 277-293. von Bertalanffy, L. 1938. A quantitative theory of organic growth (Inquiries on growth laws. II), Human Biology 10(2): 181-213. Wakabayashi, K., R. Sato, H. Ishii, T. Akiba, Y. Nogata, & Y. Tanaka 2012. Culture of phyllosomas of Ibacus novemdentatus in a closed recirculating system using jellyfish as food, Aquaculture 330: 162-166. Wakabayashi, K., H. Sato, Y. Yoshie-Stark, M. Ogushi, & Y. Tanaka 2016. Differences in the biochemical compositions of two dietary jellyfish species and their effects on the growth and survival of Ibacus novemdentatus phyllosomas, Aquaculture Nutrition 22(1): 25-33. Wang, K. 2007. The use of untraditional sea food: the commercialization of Norwegian jellyfish, red sea cucumber and whelk. SINTEF Report, Trondheim, Norway, 108 pp. 162 Ward, J.M., J.E. Kirkley, R. Metzner, & S. Pascoe 2004. Measuring and assessing capacity in fisheries. 1. Basic concepts and management options. FAO Fisheries Technical Paper No. 433/1. Food and Agriculture Organization, Rome, Italy, 40 pp. Watson, R. & D. Pauly 2001. Systematic distortions in world fisheries catch trends, Nature 414(6863): 534-536. Wells, R.W. & R. Wellington 1992. The potential exploitation of the Brown Jelly Blubber, Catostylus mosaicus, in eastern Australian waters, Sydney Basin Naturalist 1: 57-61. Wetherall, J.A. 1986. A new method for estimating growth and mortality parameters from length-frequency data, Fishbyte, Newsletter of the Network of Tropical Fisheries Scientists, International Center for Living Aquatic Resources Management (ICLARM), 4(1): 12-14. Wong, W.W.K., S.W.C. Chung, K.P. Kwong, Y.Y. Ho, & Y. Xiao 2010. Dietary exposure to aluminium of the Hong Kong population, Food Additives and Contaminants Part A - Chemistry Analysis Control Exposure & Risk Assessment 27(4): 457-463. Wootton, M., K.A. Buckle, & D. Martin 1982. Studies on the preservation of Australian jellyfish (Catostylus spp.), Food Technology in Australia 34(9): 398-400. Wu, B.L. 1955. Rhopilema esculentum, The Biological Bulletin 4(4): 35-40 (in Chinese). Yakovlev, Y.M., P.A. Borodin, & E.V. Osipov 2005. The fishery of jellyfish Rhopilema in Peter the Great Bay, Rybnoe Khozyaistvo 5: 72-75 (in Russian). Yang, H. & F. Shuang 2015. Popular Chinese soups for clearing summer heat, Cooking Knowledge 7: 36-47 (in Chinese). 163 Ye, C.C. 2006. The resource and management strategy of jellyfish (Rhopilaema esculent) population in the Liaodong Bay, Fisheries Science 25(11): 591-592 (in Chinese). Yin, M.M., D. Liu, F. Xu, L. Xiao, Q.Q. Wang, B.L. Wang, Y.L. Chang, J.M. Zheng, X. Tao, G.Y. Liu, & L.M. Zhang 2016. A specific antimicrobial protein CAP-1 from Pseudomonas sp. isolated from the jellyfish Cyanea capillata, International Journal of Biological Macromolecules 82: 488-496. Yin, Q., D. Zhang, Y. Liu, & N. Jiang 2000. Study on processing technology of jellyfish, China Fisheries 27(11): 46 (in Chinese). Yoon, W.D., J.Y. Yang, M.B. Shim, & H.K. Kang 2008. Physical processes influencing the occurrence of the giant jellyfish Nemopilema nomurai around Jeju Island, Korea, Journal of Plankton Research 30(3): 251-260. You, K., C.H. Ma, H.W. Gao, F.Q. Li, M.Z. Zhang, Y.T. Qiu, & B. Wang 2007. Research on the jellyfish (Rhopilema esculentum) and associated aquaculture techniques in China: current status, Aquaculture International 15(6): 479-488. You, K., Y. Bian, C. Ma, X. Chi, Z. Liu, & Y. Zhang 2016. Study on the carry capacity of edible jellyfish fishery in Liaodong Bay, Journal of Ocean University of China 15: 471-479. Yu, H.H., X.G. Liu, X.L. Dong, C.P. Li, R.E. Xing, S. Liu, & P.C. Li 2005. Insecticidal activity of proteinous venom from tentacle of jellyfish Rhopilema esculentum, Bioorganic & Medicinal Chemistry Letters 15(22): 4949-4952. Yu, H.H., X.G. Liu, R.E. Xing, S. Liu, Z.Y. Guo, P.B. Wang, C.P. Li, & P.C. Li 2006. In vitro determination of antioxidant activity of proteins from jellyfish Rhopilema esculentum, Food Chemistry 95(1): 123-130. 164 Yu, H.H., R.F. Li, X.L. Dong, R.G. Xing, S. Liu, & P.C. Li 2014. Efficacy of venom from tentacle of jellyfish Nemopilema nomurai against the cotton bollworm Helicoverpa armigera, Biomed Research International. Yuferova, A.A. in press. The impact of different drying modes of scyphozoan jellyfish Rhopilema esculentum and Aurelia aurita on the protein and carbohydrate components in their composition and the possibility of their use as dried prepared food, Journal of Food Process Engineering. Yusoff, F.M., J. Nishikawa, & P. Kuppan 2010. Commercial jellyfish, a little known fishery industry in Malaysia, Fishmail 18: 8-12. Zeller, D., S. Booth, G. Davis, & D. Pauly 2007. Re-estimation of small-scale fishery catches for US flag-associated island areas in the western Pacific: the last 50 years, Fishery Bulletin 105(2): 266-277. Zeller, D., S. Harper, K. Zylich, & D. Pauly 2015. Synthesis of underreported small-scale fisheries catch in Pacific island waters, Coral Reefs 34(1): 25-39. Zhang, H., J.Y. Zhang, H.L. Wang, P.J. Luo, & J.B. Zhang 2016. The revision of aluminum-containing food addidive provisions in China, Biomedical and Environmental Sciences 29(6): 461-466. Zhang, J., R. Duan, L. Huang, Y. Song, & J.M. Regenstein 2014. Characterisation of acid-soluble and pepsin-solubilised collagen from jellyfish (Cyanea nozakii Kishinouye), Food Chemistry 150: 22-26. Zhang, W.-T., L.-H. Tang, W. Chen, D.-Q. Chen, & C. Deng 2008. Therapeutic effect of collagen from Cyanea nozakii on adjuvant arthritis in rats, Chinese Journal of Marine Drugs 27(4): 35-38 (in Chinese with English abstract). Zhong, X.M., J.H. Tang, & P.T. Liu 2004. A study on the relationship between Cyanea nozakii breaking out and ocean ecosystem, Modern Fisheries Information 19(3): 15-17 (in Chinese). 165 Zhuang, Y., L. Sun, & B. Li 2012a. Production of the angiotensin-I-converting enzyme (ACE)-inhibitory peptide from hydrolysates of jellyfish (Rhopilema esculentum) collagen, Food and Bioprocess Technology 5(5): 1622-1629. Zhuang, Y., L. Sun, Y. Zhang, & G. Liu 2012b. Antihypertensive effect of long-term oral administration of jellyfish (Rhopilema esculentum) collagen peptides on renovascular hypertension, Marine Drugs 10(2): 417-426. Zhuang, Y.L., L.P. Sun, X. Zhao, J.F. Wang, H. Hou, & B.F. Li 2009. Antioxidant and melanogenesis-inhibitory activities of collagen peptide from jellyfish (Rhopilema esculentum), Journal of the Science of Food and Agriculture 89(10): 1722-1727. Zhuang, Y.L., L.P. Sun, X. Zhao, H. Hou, & B.F. Li 2010. Investigation of gelatin polypeptides of jellyfish (Rhopilema esculentum) for their antioxidant activity in vitro, Food Technology and Biotechnology 48(2): 222-228. Zimmer, M. 2005. Glowing Genes: A Revolution in Biotechnology. Prometheus Books, Amherst, New York, U.S.A., 221 pp. Zlotnick, B.A., S. Hintz, D.L. Park, & P.S. Auerbach 1995. Ciguatera poisoning after ingestion of imported jellyfish: diagnostic application of serum immunoassay, Wilderness & Environmental Medicine 6(3): 288-294. 166 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


Citation Scheme: APAAPA (5th Edition)APA No DOI, No IssueBibTeXChicago, Author, DateIEEELexisNexis (Guide Lluelles, 7th Edition)Modern Language Association, With UrlTurabian, Full Note Bibliography

Citations by CSL (citeproc-js)

Usage Statistics

Share link (DOI):



Related Items

Continue lendo em https://open.library.ubc.ca/cIRcle/collections/ubctheses/24/items/1.0340647

Previous post Chat Commerce: WeChat E-Commerce Guide – Marketing China
Next post Live Commerce: Live Commerce: a tendência do varejo online