Studying jellyfish fisheries: toward accurate national catch reports and appropriate methods for stock assessments

Abstract

People have been catching and eating jellyfish for centuries, predominantly in Asia. More recently, jellyfish fisheries have expanded around the globe, primarily for export to China and Japan. However, catch data for jellyfish remain scant. Many countries fishing for jellyfish do not explicitly report their catches to the Food and Agriculture Organization of the United Nations (FAO), and reported data are often problematic. Here, we estimate historic and current catches of jellyfish from 1950 to the present. We review past trends in global catch data for jellyfish and speculate on future catch potential. Research and management of jellyfish fisheries is inadequate, especially given the current rates of expansion and the unique challenges presented by jellyfish populations. Historically, jellyfish have been understudied, resulting in the current dearth of knowledge on jellyfish population dynamics and jellyfish fishery management. We discuss how jellyfish can be studied using straightforward adaptations of standard methods for size-based analysis of fish populations, and encourage researchers to rapidly scale up the study of these increasingly important animals.

Full text

In: Jellyfish ISBN: 978-1-63485-688-1
Editor: Gian Luigi Mariottini © 2017 Nova Science Publishers, Inc.
Chapter 15
STUDYING JELLYFISH FISHERIES:
TOWARD ACCURATE NATIONAL CATCH
REPORTS AND APPROPRIATE METHODS
FOR STOCK ASSESSMENTS
Lucas Brotz* and Daniel Pauly
Sea Around Us, Institute for the Oceans and Fisheries,
University of British Columbia, Vancouver, Canada
ABSTRACT
People have been catching and eating jellyfish for centuries, predominantly in Asia.
More recently, jellyfish fisheries have expanded around the globe, primarily for export to
China and Japan. However, catch data for jellyfish remain scant. Many countries fishing
for jellyfish do not explicitly report their catches to the Food and Agriculture
Organization of the United Nations (FAO), and reported data are often problematic. Here,
we estimate historic and current catches of jellyfish from 1950 to the present. We review
past trends in global catch data for jellyfish and speculate on future catch potential.
Research and management of jellyfish fisheries is inadequate, especially given the current
rates of expansion and the unique challenges presented by jellyfish populations.
Historically, jellyfish have been understudied, resulting in the current dearth of
knowledge on jellyfish population dynamics and jellyfish fishery management. We
discuss how jellyfish can be studied using straightforward adaptations of standard
methods for size-based analysis of fish populations, and encourage researchers to rapidly
scale up the study of these increasingly important animals.
Keywords: jellies, jellyfish fishery, scyphomedusae, Scyphozoa, zooplankton fisheries,
growth, mortality, length-frequency data
* Sea Around Us, Institute for the Oceans and Fisheries, 2202 Main Mall, University of British Columbia,
Vancouver, BC, V6T 1Z4, Canada. Email: lucasbrotz@gmail.com.
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Lucas Brotz and Daniel Pauly
314
INTRODUCTION
Jellyfish populations are increasing in numerous coastal regions of the globe [1], often
causing severe economic losses for industries such as tourism, fisheries, power generation,
and aquaculture [2]. As these problems are likely to increase [3], it has been suggested that
finding new uses for jellyfish as food and medicine could help to control jellyfish
populations, especially with regard to nuisance species [2, 4, 5]. While such a strategy as a
means to an end is flawed [6], the proliferation of jellyfish in some areas is already leading to
new fisheries and increased investigations of the use of jellyfish for nutrition, medicines, and
other applications (Table 1). Jellyfish have also been central to a number of breakthroughs in
fields such as biotechnology and biomedicine [e.g., 7, 8], as well as design engineering [9-
11]; however, these applications do not require large amounts of jellyfish, unlike some of the
possible uses listed in Table 1.
The idea of consuming jellyfish as food is relatively new in the Western Hemisphere, and
is often met with reactions ranging from surprise to disgust. However, eating jellyfish is
anything but novel, as people have been doing it for centuries. Consumption of jellyfish dates
back to at least 1,700 years ago [12]. Today, China continues to be the dominant producer and
consumer of jellyfish, representing approximately 60% of contemporary global capture
production [13], as well as the importation of jellyfish from many other countries. Indeed,
jellyfish are such a popular food item in China that many imitation products are now being
sold there that contain no actual jellyfish, but are artificially made using brown algae [14].
China is also the only country in the world to produce jellyfish through aquaculture using
large saltwater ponds, as well as employing hatchery programs whereby hundreds of millions
of juvenile jellyfish are cultured and released annually with the hopes of supplementing wild
stocks [14], a strategy that has only had partial success (see below).
Table 1. Examples of uses for jellyfish other than as food for humans
Category
Uses
Sample reference(s)
Agriculture
Livestock feeds
[15, 16]
Fertilizers
[17-24]
Insecticides
[25, 26]
Aquaculture
Finfish and shellfish feeds
[27-31]
Cosmetics
Gelatin/emulsifier
[32, 33]
Environmental monitoring
Pollution detection
[34, 35]
Fishing
Bait
[36-40]
Materials science
Absorbent polymers
[41, 42]
Cement additive
[16]
Nanoparticle filters
[43]
Pharmaceuticals
Antihypertensive peptides
[44, 45]
Anticoagulants
[46]
Antimicrobiotics
[47, 48]
Antioxidants
[49-53]
Bioactive compounds
[53-62]
Collagen
[63-70]
Mucins
[71, 72]

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315
Jellyfish populations are often subject to large interannual fluctuations in abundance. In
fact, changes in biomass of edible jellyfish are probably larger than for any other fishery [73].
Jellyfish are most often caught from small boats using dip-nets. However, a wide variety of
active and passive fishing gears are used in different areas of the world. At least 30 different
species of jellyfish have been identified as “edible” [13]; however, species that are primarily
targeted for food belong to the paraphyletic Order Rhizostomeae. Jellyfish belonging to this
group are typically less fragile than other jellies and will produce the desired crunchy texture
that is characteristic of edible jellyfish after processing. With the exception of Mexico, the
Food and Agriculture Organization of the United Nations (FAO) reports all jellyfish catches
as “Rhopilema spp,” which is incorrect in many cases. Combined with the fact that the
taxonomy of edible jellyfish is considerably confused [38], this makes it difficult to determine
exactly which jellyfish are being caught and eaten.
Although some may be eaten fresh in coastal areas [14], jellyfish are most often
processed in a stepwise salting procedure that takes weeks. The process may vary somewhat,
but typically involves soaking jellyfish in a variety of different mixtures of salt and alum
(usually potassium aluminum sulfate) in order to partially dehydrate the jellyfish,
decontaminate them, and produce the desired crunchy and crispy texture. Semi-dried (i.e.,
processed) jellyfish are then either sold at markets or packaged and shipped. Prior to
consumption, jellyfish are usually soaked in water to remove the salt and alum, and then
sliced, often blanched, and served as appetizer salads or as ingredients in other dishes. Ready-
to-eat jellyfish products are also available as snacks that can be consumed straight from the
package.
Desalted, processed edible jellyfish are typically 92-96% water and 3-7% protein,
primarily collagen. With only 36 kcal per 100 g serving [74], edible jellyfish have been
declared as a natural diet food. There is a long list of purported health benefits from eating
jellyfish according to Traditional Chinese Medicine [14, 75]; however, very few of these have
been tested using scientific criteria. In addition, there are health concerns about the
consumption of jellyfish related to the use of alum during processing, which contains
aluminum, detectable in the final product [76-78]. As such, whether the effects of eating
jellyfish are positive or negative for human health remains unresolved.
ESTIMATING THE GLOBAL CATCH OF JELLYFISH
At least 23 countries have been involved in jellyfish fisheries (Table 2). Some countries
(e.g., Turkey) appear to have abandoned their jellyfish fisheries, while others (e.g., Canada)
had test fisheries that were unsuccessful. There are 19 nations currently fishing for jellyfish,
with estimated current average landings of at least 900,000 tonnes annually [13]. Despite the
proliferation of jellyfish fisheries, catch data remain scant. Many countries do not explicitly
report their jellyfish catches to FAO, and even reported data are often problematic. As such, a
catch reconstruction of jellyfish landings from 1950 to the present was developed. Methods
were based on those that have been employed for a myriad of fisheries catch reconstructions
[79-81], whereby the following steps are followed:

Lucas Brotz and Daniel Pauly
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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;
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
reported catches (1) and interpolated missing data series (5).
Using this methodology, a global catch reconstruction for jellyfish was developed by
combining the catches of each country (Figure 1). Major findings are discussed below, and
details for all countries in the analysis can be found in [82].
As mentioned, China has the longest history of fishing for edible jellyfish, and is the
world’s largest producer. Estimating China’s catch is a challenge due to inconsistencies in
reporting. For example, FAO reports no catch prior to 1970; however, Dong et al. [83] report
annual landings dating back to 1957 from a variety of sources, including China Fishery
Statistical Yearbooks. The targeted species is Rhopilema esculentum, a conspicuous
rhizostome that fetches the highest price for edible jellyfish. Catch statistics for jellyfish in
China were also reported by Li et al. [84] from 1980 to 2012. For the period 1980-1990, it
appears that landings reported to FAO were for processed jellyfish, rather than wet weight.
As processed R. esculentum weighs only 15% of the original wet weight [84], we can assume
that landings from this period are only 15% of the true value, i.e., underreported more than
sixfold. When we examine Chinese landings from the 1950s, ‘60s, and ‘70s, they are of a
similar magnitude to those from the 1980s, and as such, it suggests that reported landings
prior to 1990 are similarly for processed jellyfish. While this is clearly a major assumption
with significant consequences for a catch reconstruction estimate, the current scale of
reported jellyfish landings in China is on the order of several hundred thousand tonnes. That
scale, combined with China’s long history of fishing jellyfish, would seem to justify the
assumption.
Catches of R. esculentum in China began declining in the 1970s, likely due to
overexploitation [83]. This led to extensive research on the life cycle and culturing of the
species [14], and in 1984, ephyrae were released into Chinese coastal waters with the hopes
of supplementing the wild stock and increasing the catch. For almost 2 decades, the hatchery
program continued to expand and was declared an economic success [14, 85]. However,
recent landings of this species have declined despite increased restocking programs, which
now release hundreds of millions of ephyrae annually. Around the turn of the century, another
rhizostome, the giant jellyfish Nemopilema nomurai, began increasing in abundance in East
Asian waters [86-88]. With declining catches of R. esculentum, jellyfish fishers in China
quickly turned their attention to N. nomurai, with landings on the order of hundreds of
thousands of tonnes in recent years [84]. Curiously, despite these massive catches, catches of
N. nomurai appear to be absent from FAO statistics.

Studying Jellyfish Fisheries
317
Table 2. Countries that are known to fish for jellyfish
Country
Dates
Australia
1995 - present
Bahrain
2004 - present
Canada
1984; 2002
China
<1950 - present
Ecuador
2013 - present
Honduras
2013 - present
India
1984 - present
Indonesia
<1950 - present
Iran
2010? - present
Japan
<1950 - present
Korea (South)
1980s? - present
Malaysia
<1950? - present
Mexico
2000 - present
Myanmar
1995? - present
Nicaragua
2008; 2013 - present
Pakistan
2007? - present
Philippines
1976 - present
Russia
2000 - present
Sri Lanka
1986 - present
Thailand
1970 - present
Turkey
1984 - 2006
U.S.A.
1993 - present
Vietnam
1990s - present
Figure 1. Estimated global jellyfish landings for two primary species in China and all species for other
countries.

Lucas Brotz and Daniel Pauly
318
Other countries such as Indonesia, Japan, and Malaysia also had fisheries for jellyfish by
the middle of the 20th century; however, the scale of these operations was small compared to
China’s. Thailand’s jellyfish fisheries began in the 1960s or 1970s, and have expanded to the
point where Thailand is now the world’s second largest producer. More recently, significant
fisheries for jellyfish have developed in India, Vietnam, and Mexico, resulting in a global
catch that has exceeded 500,000 tonnes since 1997 (Figure 1), ironically the same year when
world jellyfish catches “peaked” according to FAO statistics. FAO also reports (relatively
small) catches from several countries that are not known to have jellyfish fisheries, including
Namibia, the United Kingdom, and the Falkland Islands. We suspect that these reports are for
discarded jellyfish that are caught as bycatch in other fisheries. While such catches should
indeed be reported, they should also be differentiated from targeted landings. Clearly
reporting to and by FAO needs to improve, especially in the case of jellyfish.
STOCK ASSESSMENT FOR JELLYFISH FISHERIES
To understand and manage jellyfish fisheries, stock assessments are required which, if
only for expediency, ought to draw as much as possible from the existing toolkit of fishery
managers. These conceptual toolkits, and the mathematical models used to implement them,
were mostly derived, however, from the study of bony fishes. This has misled many jellyfish
specialists to assume, a priori, that standard models to describe the growth and mortality of
teleosts would not apply to jellyfish (as is also often, and equally falsely, assumed for squids
[89]). Instead, these specialists have studied jellyfish using a disparate array of concepts and
models, with the result that few generalities have emerged which could help in optimizing the
management of jellyfish fisheries or predicting growth and mortality patterns in unstudied
jellyfish taxa.
The von Bertalanffy growth function (VBGF) is one of the standard models of fishery
science; for length, the VBGF has the form
Lt = L∞·{1 – exp[-K(t – t0)]} (1)
where Lt is the length at age t, L∞ is the asymptotic length (roughly corresponding to the
maximum length in the population in question), K is a parameter of dimension time-1,
expressing how fast L∞ is approached, and t0 is the (usually negative) age at size = 0 (and not
discussed further here).
The seasonal oscillations in the growth of fish and invertebrates can be very marked
(Figure 2). A variant of the VBGF which accounts well for seasonal oscillation of growth in
length [90] has the form
Lt = L∞·{1 – exp-[K(t – t0) + S(t) – S(t0)]} (2)
where L∞, K and t0 are defined as in the standard VBGF (see above), and where S(t) =
(CK/2π)·sin π(t – ts) and S(t0) = (CK/2π)·sin π(t0 – ts). This equation involves two parameters
more than the standard VBGF: C and ts. Of these, the former is easier to visualize, as it
expresses the amplitude of the growth oscillations. When C = 0, the seasonally oscillating

Studying Jellyfish Fisheries
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VBGF reverts to the standard VBGF. When C = 0.5, the seasonal growth oscillations are such
that growth rate increases by 50% at the peak of the ‘growth season,’ i.e., in ‘summer,’ and,
briefly, declines by 50% in ‘winter.’ When C = 1, growth increases by 100%, i.e., doubles
during ‘summer,’ and becomes zero in the depth of ‘winter.’ In fishes, moreover, C = 1 when
the difference in mean monthly SST in the hottest summer month is about 10°C higher than
in the coldest month. For lower summer-winter differences, C is correspondingly lower.
The second new parameter, ts, expresses the time between t = 0 and the start of a sinusoid
growth oscillation. For visualization, it helps to define ts + 0.5 = WP (‘Winter Point’), which
expresses, as a fraction of the year, the period when growth is slowest. WP is often near 0.1
(i.e., early February) in the northern and 0.6 (early August) in the southern hemisphere, hence
the name. Note that it is not necessarily the alternation of high summer and low winter
temperatures which causes the seasonal oscillations of growth. Also note that the seasonally
oscillating VBGF cannot describe long periods of zero growth (and values of C > 1; but see
[91, 92]).
For weight, the VBGF takes the form
Wt = W∞ {1 – exp[-K(t – t0)]}b (3)
with W∞ being the weight corresponding to L∞ as obtained through a length-weight
relationship of the form W = aˑLb. (Seasonally oscillating forms of the VBGF for weight exist
[92], but are not discussed here).
Similarly, in fisheries research, the model most commonly used to represent the mortality
of fish is
Nt2 = Nt1ˑexp[Z(t2 – t1)] (4)
where Nt1 and Nt2 are numbers at time t1 and t2 and Z is the instantaneous rate of total
mortality, with Z = M+F, and M natural and F fishing mortality. The usefulness of this
representation of mortality lies not only in that Z can be readily separated into its components,
but also that the ratio M/K, which is an explicit parameter of various stock assessment
models, tends to be constant within taxa.
The parameters of the VBGF (asymptotic sizes and K) can be estimated by applying
length-frequency analysis (LFA; [96]) to jellyfish bell diameter (i.e., ‘length’) frequency data
[95]. To illustrate this, a selection of LFA methods, i.e., the ELEFAN method (Figure 2) and
Wetherall plots for growth estimation, and length-converted catch curves for mortality
estimation were applied to 34 sets of bell diameter frequency data of jellyfish. This led to the
estimates of parameters of the VBGF and estimates of mortality (notably natural mortality;
M) useful for modeling the life history of jellyfish.
Note that the crucial step in estimating the parameters of the VBGF does not consist of
the estimation of asymptotic bell diameter, for which the maximum size in a field sample
usually provides a good approximation, nor with the parameters describing the seasonality of
growth, which can be approximated from first principles. Rather, the crucial parameter of the
VBGF is K. How well this parameter is estimated can be assessed by plots such as those
shown in Figure 3, which are a standard feature of the ELEFAN procedure.

Lucas Brotz and Daniel Pauly
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Figure 2. Jellyfish growth curve fitting with ELEFAN. Panel A: Aurelia aurita from Tokyo Bay, Japan
in 1990-1992 (L/F data from [93]), with L∞ = 35.5 cm and K = 0.86 year -1 for fixed values of C = 0.5
and WP = 0.1. Panel B: First 3 of the 6-years’ L/F data of Catostylus mosaicus from Botany Bay,
Australia sampled between March 1990 and February 1998 [94] with L∞ = 37.0 cm and K = 0.60 year–1
for fixed values of C = 0.5 and WP = 0.7. These two growth curves were selected from thousands of
alternatives using a search algorithm in ELEFAN (see Figure 5 and text). (Modified from [95]).
The applications of Wetherall plots (Figure 4A) and catch curves (Figure 4B) yielded the
mean value of M/K for jellyfish that was estimated is about 3 year-1, about two times higher
than the values reported for fishes, which usually range between 1 and 2 year-1 [96]. This high
value of M/K may be due to, at least in some cases, shrinkages of the bells of jellyfish [97],
which could have biased the (fixed) interrelationships of number, size, and age, and which are
assumed in LFA. Note that when K is underestimated by ELEFAN or other LFA, M is also
underestimated (and conversely for overestimation), for which reason the above estimate of
M/K should be robust.
Figure 3. Examples of the goodness-of-fit estimator of ELEFAN in relation to K, as used to estimate
this parameter (and to assess the uncertainty associated with the point estimate) when the other
parameters of the seasonally oscillating VBGF (L∞, C, and WP) are known or assumed (see arrows).
Panel A: for Aurelia aurita (see Figure 2A); the best fitting K value is not very distinct from adjacent
values; hence, the best estimate of K (= 0.86 year−1) is uncertain. Panel B: Catostylus mosaicus (see
Figure 2B), for which K (= 0.60 year−1) is more reliably estimated. (Modified from [95]).

Studying Jellyfish Fisheries
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Figure 4. Methods to estimate M/K or Z from L/F data. Panel A: Wetherall Plot applied to a cumulative
bell diameters of Chrysaora melanaster from the Bering Sea, USA (inset; from [98]) sampled in
September 1996, 1997, and 1999. Only the lengths fully retained by the gear (straight section of graph;
bell diameters >25 cm) are used for the regression, which yielded L∞ = 56.3 cm and Z/K = 2.86. Panel
B: A catch curve applied to bell diameter data for Aurelia aurita in Tokyo Bay, Japan (inset; from
[93]), sampled from May 1990 to December 1992. Using the von Bertalanffy growth parameters in
Figure 2A, yields an estimate of Z = 2.95 year−1. (Modified from [95]).
Figure 5. Auximetric plot of (log10)K against re-scaled values of (log10)W∞ for the major groups of
jellyfishes, on a background of grey dots representing fishes (including two highlighted species,
Zoarces viviparous and Gadus morhua). As might be seen, the Aurelia aurita complex and Catostylus
mosaicus resemble small fishes in their growth pattern, but Chrysaora spp. and Phryllorhiza punctata
(and other species in [95]) may grow faster (higher K for a given W∞) than fishes. (Modified from
[95]).
Moreover, it was found that by scaling their asymptotic weight (W∞, a parameter of the
VBGF) to the weight they would have if they had the same water content as fish, most
jellyfish could be shown to grow at the same rate as small fishes. Thus, as in fish, the VBGF

Lucas Brotz and Daniel Pauly
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parameters K and W∞, when plotted in a double logarithmic (‘auximetric’) plot, tend to
cluster into ellipsoid shapes, which increase in area when shifting from species to genera,
families, etc. (Figure 5). This potentially provides a powerful tool for testing comparative
hypotheses on jellyfish life history. These results are compatible with the suggestion that the
VBGF is not only a convenient mathematical function for describing the growth of jellyfish,
but that it does so because their respiratory physiology makes this growth function, derived
from physiological considerations, the model of choice [89, 92].
The auximetric plot in Figure 5, finally, suggests that some jellyfish (Aurelia aurita
complex, Catostylus mosaicus), once their high water content is accounted for, have growth
patterns similar to small and very small fishes, such as guppies and anchovies. Others
(Phyllorhiza punctata, Chrysaora spp.) may grow faster than fishes (i.e., have higher values
of K for their value of W∞). However, the accuracy of the position of an organism on an
auximetric plot depends on the accuracy of the growth parameters, and in the case of jellyfish,
on a correct conversion to standard water content. Because of this, these results are still
preliminary. However, it is encouraging that, as in fish (here exemplified by Gadus morhua
and Zoarces viviparus, the different populations in a given species appear to form ellipsoid
clusters on an auximetric plot (see www.fishbase.org for more). Genera and higher taxa can
be expected, as well, to form such clusters, albeit larger ones. This suggests that the large
cluster for the Aurelia aurita complex would, indeed, include more than one species, as long
suggested by taxonomists [99].
CONCLUSION
An increasing number of countries have started fishing for jellyfish in recent years with
the hopes of profitably exporting catches to East Asia (see Table 2). Often, this follows
closely on the heels of collapses of more traditional fishery resources. While such ‘fishing
down’ [100] may seem like an obvious progression, new jellyfish fisheries face a number of
barriers to ‘success.’ With only a tiny fraction of the more than 1,200 species of jellyfish
preferred for consumption, new processing techniques will need to be developed if other
species are to meet some of the demand. Processing of jellyfish should also be improved to
eliminate the associated concerns with human and environmental health. In addition,
fluctuations due to changes in market demand should be considered for those hoping to
develop jellyfish fisheries to supply Asian markets (e.g., higher demand for Chinese New
Year celebrations).
Knowledge of the biology and ecology of most jellyfish species is limited, and as such,
implementing management strategies is a challenge, especially given the tendency for
jellyfish populations to fluctuate strongly [73]. Also, the reluctance of jellyfish researchers to
even attempt at using standard models of fish stock assessment derived from studies of teleost
fishes (as illustrated here) will have to be overcome, as there is no time to re-invent the wheel.
While fishing for jellyfish is likely to expand given the decline of fisheries around the
world [81] and the local increases in jellyfish blooms [1], it is unlikely to solve our jellyfish
problems [6] or feed the world.

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ACKNOWLEDGMENTS
Support is acknowledged from the Sea Around Us, funded by the Paul G. Allen Family
Foundation.
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