Marine Pollution Bulletin 150 (2020) 110728

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Marine Pollution Bulletin

journal homepage: www.elsevier.com/locate/marpolbul

Risk screening of the potential invasiveness of non-native jellyfishes in the T

Mediterranean Sea
⁎
Nurçin Killia, Ali Serhan Tarkana,b, , Sebastian Kozicb, Gordon H. Coppb,c,d, Phil I. Davisonc,d,
Lorenzo Vilizzib
a
Muğla Sıtkı Koçman University, Faculty of Fisheries, Department of Basic Sciences, 48000, Muğla, Turkey
b
Department of Ecology and Vertebrate Zoology, Faculty of Biology and Environmental Protection, University of Łódź, Łódź, Poland
c
Centre for Environment Fisheries and Aquaculture Science, Lowestoft, UK
d
Centre for Environment and Sustainability, Bournemouth University, Poole, UK

A R T I C LE I N FO A B S T R A C T

Keywords: The aim of the present study was to risk screen 45 jellyfish species (30 hydromedusae, 14 scyphomedusae, one
Cubomedusae cubomedusa) for their potential invasiveness in the Mediterranean Sea to aid managers in making informed
Hydromedusae decisions on targeting appropriate species for management. Using the Aquatic Species Invasiveness Screening Kit
Scyphomedusae (AS-ISK), calibrated basic and climate-change threshold assessment scores of 6.5 and 12.5, respectively, were
Lessepsian
identified for distinguishing reliably between species that pose ‘low-to-medium’ and ‘high’ risk of becoming
Global warming
invasive in the risk assessment area. Using these thresholds, 16 species were classified as high risk, 23 as medium
risk and six as low risk under current climate conditions. Whereas, under future climate conditions, 13, 30 and
two species, respectively, were classified as high, medium and low risk, respectively. Upside-down jellyfish
Cassiopea andromeda, Australian spotted jellyfish Phyllorhiza punctata, sea nettle Chrysaora quinquecirrha and
Rhopilema nomadica were the highest-scoring species, with the maximum increase in risk score under predicted
climate change conditions being achieved by C. andromeda.

1. Introduction
Jellyfishes play a key role in marine food webs as predators of
zooplankton, thereby reducing an important resource for larval fin-
fishes (Arai, 2005), and as predators of fish eggs and juveniles (Purcell
and Arai, 2001). By way of example, the hydromedusan, Portuguese
man-of-war Physalia physalis, is a species native to the Atlantic Ocean
that has invaded the Mediterranean Sea and reached the coasts of
Sardinia, probably via ballast water transfers (Prieto et al., 2015). This
species can consume up to 120 fish eggs and larvae per day (Purcell and
Arai, 2001). Similarly, considerable numbers of fish eggs and larvae
have been found in the stomachs of the scyphomedusan, Atlantic sea
nettle Chrysaora quinquecirrha (Möller, 1984; Purcell et al., 1994). On
the other hand, jellyfishes are consumed by at least 124 fish species,
including those of the families Centrolophidae, Molidae, Scombridae,
Scorpaenidae and Stromateidae (Harbison, 1993), as well as by 34
other marine organisms (Pauly et al., 2009).
Jellyfish blooms can be facilitated by eutrophication, overfishing,
bottom trawling, aquaculture, increasing human encroachment on
coastal areas, and global warming (Arai, 2001; Mills, 2001; Purcell,
2005; Hay, 2006; Graham and Bayha, 2008). The high densities asso-
ciated with jellyfish blooms are known to cause economic losses to
fishers through net damage, to clog the intake pipes of power-plant
cooling systems, and to be responsible for detrimental effects on
tourism and human health (Shimomura, 1959; Matsueda, 1969;
Yasuda, 1988; Rajagopal et al., 1989; Burnett, 2001; Graham et al.,
2003). Furthermore, some jellyfishes native to the Mediterranean Sea
such as Pelagia noctiluca can be highly traumatogenic to humans (Stein
et al., 1989).
Several non-native (NN) jellyfishes have entered the Mediterranean
Sea via human-made migration routes, such as the opening of the Suez
Canal, which has resulted in the introduction of several Indo-Pacific
species (cf. Lessepsian migration). Of these, scyphozoan jellyfishes have
been especially successful and, favoured by global warming conditions,
have been able to increase their reproductive success (Boero et al.,
2008). More generally, thermophilic jellyfishes are positively affected
by increases in water temperature, which facilitates range expansion.
Whereas, cool-water jellyfishes are negatively affected by increases in
water temperature through decreased habitat suitability (Boero et al.,
2016). Currently, > 400 hydrozoan, 20 scyphozoan and one cubozoan

⁎
Corresponding author at: Muğla Sıtkı Koçman University, Faculty of Fisheries, Department of Basic Sciences, 48000 Muğla, Turkey.
E-mail address: serhan@mu.edu.tr (A.S. Tarkan).

https://doi.org/10.1016/j.marpolbul.2019.110728
Received 15 July 2019; Received in revised form 5 November 2019; Accepted 7 November 2019
Available online 25 November 2019
0025-326X/ © 2019 Elsevier Ltd. All rights reserved.
N. Killi, et al.
jellyfishes are known to be distributed across the Mediterranean Sea
(Bouillon et al., 2004), and amongst the Lessepsian scyphozoans, up-
side-down jellyfish Cassiopea andromeda, the erythraean jellyfish Coty-
lorhiza erythraea, Marivagia stellata, Australian spotted jellyfish Phyl-
lorhiza punctata and the nomad jellyfish Rhopilema nomadica are
especially abundant (Galil et al., 1990, 2017). Of these, R. nomadica is
known to undergo blooms in the Eastern Mediterranean coasts of
Turkey, where it may represent up to 60% of the total catch of trawls,
purse seines and gillnets (Turan et al., 2011). Notably, of the 423 cni-
darian jellyfish species (i.e. ‘true jellyfish’ sensu Falkenhaug, 2014) that
occur in the Mediterranean Sea (Bouillon et al., 2004; İşinibilir and
Yılmaz, 2016; Gülşahin, 2017; Yilmaz et al., 2017; Öztürk et al., 2018),
57 (13.5%) are non-native (Zenetos et al., 2010, 2012; Galil et al., 2016;
Yilmaz et al., 2017; Zenetos et al., 2017; Langeneck et al., 2019).
The aim of the present study was to determine the potential inva-
siveness of the medusa forms of non-native jellyfishes in the
Mediterranean Sea and to assess whether predicted global warming
conditions may contribute further to their dispersal. To this end, a risk-
screening procedure was undertaken to identify which of the various
NN jellyfishes present in the Mediterranean Sea are likely to be invasive
and cause ecological and economic impacts, thereby aiding environ-
mental managers and policy makers in the decision-making process to
implement appropriate management and conservation measures for
protection of the Mediterranean Sea and its native species.
2. Materials and methods
2.1. Risk screening
In total, 45 species of NN jellyfish from three taxonomic Classes (i.e.
30 hydromedusae, 14 scyphomedusae and one cubomedusa) were
screened for their potential invasiveness in the Mediterranean Sea,
henceforth the Risk Assessment (RA) area (Table 1; Fig. 1). For the
purposes of the present study, jellyfishes that have a medusa stage were
selected preferentially because of the greater likelihood of movement
and spread by currents. Whilst several literature sources were consulted
(i.e. Zenetos et al., 2010, 2012) and searches for Internet-based re-
sources implemented (i.e. Web of Science, supplemented by Goo-
gleScholar), the final selection of NN jellyfishes for screening was based
upon the following criteria (number and relative percentage of species
in brackets): (1) native and NN species already present in the Medi-
terranean Sea, hence ‘extant’ (37 species, 82.2%), and (2) ‘horizon’ NN
species likely to enter the Mediterranean Sea via the Suez Canal (Red
Sea) and the Strait of Gibraltar (Atlantic Ocean) or via ballast water (8
species, 17.8%) (Table 2).
The risk screening process was undertaken to identify potentially
invasive species with respect to the RA area using the Aquatic Species
Invasiveness Screening Kit (AS-ISK), which is available for free down-
load at www.cefas.co.uk/nns/tools/ (Copp et al., 2016a). The AS-ISK,
which is a second-generation adaptation of the Australian Weed Risk
Assessment decision-support tool (Pheloung et al., 1999), was created
by revising the questions of the generic screening module of the Eur-
opean Non-native Species in Aquaculture Risk Analysis Scheme (Copp
et al., 2016b) and by incorporating them into the architecture of the
Fish Invasiveness Screening Kit v2 (Lawson Jr et al., 2013). During this
process, the AS-ISK was made to comply with the ‘minimum standards’
(Roy et al., 2018) for the assessment of NN species with regard to the
2014 EC Regulation on the prevention and management of invasive
alien species (EU Regulation No. 1143/2014). The AS-ISK consists of 55
questions: the first 49 questions cover the biogeography/historical and
biology/ecology aspects of the taxon under assessment, and comprise
the Basic Risk Assessment (BRA); the other six questions require the
assessor to predict how future climatic conditions are likely to affect the
BRA with respect to risks of introduction, establishment, dispersal and
impact, and these comprise the Climate Change Assessment (CCA).
For the CCA, two studies on likely future climate scenarios for
2
Marine Pollution Bulletin 150 (2020) 110728
Turkey were used (Demir et al., 2008; www.mgm.gov.tr/FILES/iklim/
iklim-degisikligi-projeksiyon2015.pdf), with most scenarios predicting
an increase of 0.5–1 °C in air temperature over the next 50 years. From
these sources of information, the expected increase in water tempera-
ture (Tw) was calculated based on its relationship with air temperature
(Ta) as: Tw = 3.47 + 0.898 Ta (after Erickson and Stefan, 1996).
For each question in AS-ISK, the assessor must provide a response,
justification and level of confidence (see below), and the screened
species eventually receives both a BRA and a BRA+CCA (composite)
score (respectively ranging from −20.0 to 68.0 and from −32.0 to
80.0). As in its ancestral decision-support tool, the WRA, AS-ISK
scores < 1 suggest that the species is unlikely to become invasive in the
RA area, and is therefore classified as ‘low risk’ (Pheloung et al., 1999);
on the contrary, higher scores classify the species as posing either a
‘medium risk’ or a ‘high risk’ of becoming invasive, the threshold value
to distinguish between medium and high risk levels is typically ob-
tained through RA area-specific ‘calibration’, which is subject to the
availability of a representative sample size (i.e. number of screened
taxa) – this was recently estimated to be n = 15–20 (Vilizzi et al.,
2019). The ranked levels of confidence (1 = low; 2 = medium;
3 = high; 4 = very high) associated with each question-related re-
sponse in AS-ISK mirror the confidence rankings recommended by the
Intergovernmental Panel on Climate Change (IPCC, 2005; Copp et al.,
2016a), and from these levels of confidence, a confidence factor (CF) is
computed (as detailed in the following section).
Risk screening assessments were undertaken by the first author,
who is knowledgeable in the biology and ecology of the jellyfishes of
the Mediterranean Sea. Whilst it is preferable for each species to be
assessed by two or more assessors, this was not possible in the present
study and risk screening studies based on a single assessor are not un-
common (Vilizzi et al., 2019).
2.2. Statistical analysis
Following computation of the BRA and BRA+CCA scores, Receiver
Operating Characteristic (ROC) curve analysis (Bewick et al., 2004) was
used to assess the predictive ability of AS-ISK to discriminate between
jellyfishes posing a high risk and those posing a medium or low risk of
being invasive for the RA area. For ROC curve analysis to be im-
plemented, the species has to be categorised a priori in terms of their
documented invasiveness (i.e. non-invasive or invasive). However,
unlike fishes, for which ‘global’ a priori categorisations for invasiveness
have been possible thanks to the availability of online databases (e.g.
Glamuzina et al., 2017; Li et al., 2017; Tarkan et al., 2017a, 2017b;
Dodd et al., 2019), in the present study an ‘integrated approach’ was
adopted due to the limited literature on, and thus overall more limited
knowledge of, jellyfish biology, ecology and invasiveness-related in-
formation. The present approach relied on a preliminary consultation of
SeaBaseLife (www.sealifebase.org) and the Global Invasive Species
Database (GISD: www.iucngisd.org/gisd/) for any reference to the in-
vasiveness status of each species. If no reference was available, then a
GoogleScholar literature search was performed using the keywords
‘invasive’, ‘invasiveness’ and ‘impact’ (together with that of the species)
and, if at least one peer-reviewed literature reference in support was
found, then this was taken as ‘sufficient evidence’ for categorising the
species as a priori invasive.
Statistically, a ROC curve is a graph of sensitivity vs 1 – specificity
(or alternatively, sensitivity vs specificity), where in the present context
sensitivity and specificity will be the proportion of invasive and non-
invasive jellyfishes, respectively, that are in agreement with the experts'
assessments. A measure of the accuracy of the calibration analysis is the
Area Under the Curve (AUC), which typically ranges between 0.5 and
1.0, and the closer to 1.0 the better the ability (of the AS-ISK tool) to
differentiate between invasive and non-invasive species (for more de-
tails see e.g. Tarkan et al., 2017a, 2017b). Following ROC analysis, the
best AS-ISK threshold value that maximises the true positive rate (i.e.
N. Killi, et al. Marine Pollution Bulletin 150 (2020) 110728

Table 1
Non-native jellyfish species screened for their risk of invasiveness in the Mediterranean Sea with the Aquatic Species Invasiveness Screening Kit (AS-ISK). To
determine invasiveness status (N = non-invasive; Y = invasive) for a priori categorisation (see Table 2), SeaBaseLife (www.sealifebase.org) and the Global Invasive
Species Database (GISD: http://www.iucngisd.org/gisd/) were consulted, and if no information was provided an additional GoogleScholar search of the peer-
reviewed literature was made and the (generally) most recent reference provided if an impact has been documented. Also provided are the literature sources
reporting alien jellyfish species distribution in the Mediterranean Sea (tick marks and dashes denoting presence and absence, respectively, in the corresponding lists).
See also Fig. 1.
Class/Family/Species SeaLifeBase GISD GoogleScholar Invasive Zenetos et al., 2010 Zenetos et al., 2012

Cubozoa
Carukiidae
Carukia barnesi N – – N – –
Hydrozoa
Aequoridae
Aequorea conica – – No evidence N √ –
Aequorea globosa – – Mizrahi et al. (2015) Y – √
Aequorea vitrina – – Yilmaz et al. (2017) Y – –
Bougainvilliidae
Nubiella mitra – – No evidence N √ –
Campanulariidae
Campanularia morgansi – – No evidence N – √
Clytia hummelincki N – N √ –
Clytia linearis – – Corriero et al. (2016) Y √ √
Clytia mccradyi – – Zenetos et al. (2012) Y √ –
Cirrholoveniidae
Cirrholovenia tetranema – – Y √ –
Corymorphidae
Corymorpha annulata – – No evidence N √ –
Corymorpha bigelowi – – No evidence N √ –
Corynidae
Coryne eximia – – İşinibilir et al. (2010) N √ –
Cytaeididae
Paracytaeis octona – – No evidence N √ –
Eirenidae
Eirene viridula – – No evidence N √ √
Halicreatidae
Haliscera bigelowi – – No evidence N – √
Laodiceidae
Laodicea fijiana – – No evidence N √ –
Lovenellidae
Eucheilota paradoxica – – No evidence N √ –
Eucheilota ventricularis – – No evidence N – √
Magapiidae
Kantiella enigmatica N – – N √ –
Moerisiidae
Moerisia carine – – No evidence N √ –
Moerisia inkermanica – – Rodriguez et al. (2014) Y √ –
Olindiidae
Gonionemus vertens Y – – Y √ –
Olindias singularis – – No evidence N √ –
Scolionema suvaense – – No evidence N √ –
Pandeidae
Halitiara inflexa – – No evidence N – √
Physaliidae
Physalia physalis – – Prieto et al. (2015) N – –
Rhopalonematidae
Amphogona pusilla – – Vidjak et al. (2018) Y √ –
Arctapodema australis – – Gravili (2016) Y √ –
Tetrorchis erythrogaster – – No evidence N √ –
Trichydridae
Trichydra pudica N – – N √ –
Scyphozoa
Cassiopeidae
Cassiopea andromeda Y – – Y √ –
Cassiopea xamachana – – No evidence N – –
Cepheidae
Cephea cephea – – No evidence N – –
Cotylorhiza erythraea – – Boero et al. (2016) Y – –
Marivagia stellata – – Galil and Goren (2014) Y √ –
Netrostoma setouchianum – – No evidence N – –
Mastigiidae
Phyllorhiza punctata – Y – Y √ –
Pelagiidae
Chrysaora fulgida N – – N – –
Chrysaora quinquecirrha Y – – Y – –
Mawia benovici – – Avian et al. (2016) N – –
Rhizostomatidae
Rhopilema esculentum – – No evidence N – –
(continued on next page)

3
N. Killi, et al.
Table 1 (continued)
Class/Family/Species SeaLifeBase GISD GoogleScholar
Rhopilema nomadica – – Yahia et al. (2013)
Rhopilema verrilli N – – N
Stomolophus meleagris – – No evidence
jellyfishes categorised a priori as invasive and classified as such) and
minimises the false positive rate (i.e. jellyfishes categorised a priori as
non-invasive but classified as invasive) was determined using Youden's
J statistic; whereas, the ‘default’ threshold of 1 was set to distinguish
between low risk and medium risk species (see Risk screening). Notably,
jellyfishes categorised a priori as invasive but classified as non-invasive
will be ‘false negatives’, whereas those categorised a priori as non-in-
vasive and also classified as non-invasive will be ‘true negatives’. ROC
analysis was carried out with package pROC (Robin et al., 2011) for
R × 64 v3.0.3 (R Development Core Team, 2016) using 2000 bootstrap
replicates for the confidence intervals of specificities, which were
computed along the entire range of sensitivity points (i.e. 0 to 1, at 0.1
intervals), and with the addition of a smoothed mean ROC curve. The
threshold intervals are presented using the statistically-appropriate
brackets (“]” and “[“; www.mathwords.com/i/interval_notation.htm).
Based on the confidence level (CL) allocated to each response for a
given species (see Risk screening), an overall confidence factor (CF) was
computed as:
∑ (CLQi)/(4 × 55) (i = 1,…, 55)
where CLQi is the CL for Question i (Qi), 4 is the maximum achievable
value for certainty (i.e. ‘very certain’) and 55 is the total number of
questions comprising the AS-ISK tool. The CF ranges from a minimum
of 0.25 (i.e. all 55 questions with certainty score equal to 1) to a
maximum of 1 (i.e. all 55 questions with confidence level equal to 4).
Two additional confidence factors were computed, namely the CFBRA
and the CFCCA based on the 49 Qs of the BRA and the 6 Qs of the CCA,
respectively.
Differences between the mean CL for the BRA and CCA (CLBRA and
CLCCA, respectively) and between the mean CF for the BRA and CCA
(CFBRA and CFCCA, respectively) were tested by permutational (uni-
variate) analysis of variance (PERANOVA) based on a one-factor design
(i.e. Component, with two levels: BRA and CCA). Analysis was carried
out in PERMANOVA+ for PRIMER v6, with normalisation of the data
and using a Bray-Curtis dissimilarity measure, 9999 unrestricted per-
mutations of the raw data (Anderson et al., 2008), and with statistical
effects evaluated at α = 0.05.
3. Results
The ROC curves for the BRA resulted in an AUC of 0.6289
(0.4444–0.8134 95% CI) and for the BRA+CCA in an AUC of 0.6356
(0.4483–0.8228 95% CI) (Fig. 2a,b). These AUCs indicated that AS-ISK
was able to discriminate between non-invasive and invasive jellyfishes
for the RA area, though with a caveat for the relatively low mean values
(i.e. < 0.7). Youden's J provided a threshold of 6.5 for the BRA and 12.5
for the BRA+CCA, which were used for calibration of the AS-ISK risk
outcomes. Accordingly, the BRA threshold allowed to distinguish be-
tween medium risk species with scores within the interval [1.0, 6.5[,
and high risk species with scores within ]6.5, 68.0]; the BRA+CCA
threshold allowed to distinguish between medium risk species with
scores within [1.0, 12.5[, and high risk species with scores within ]12.5,
80.0]. Species classified as low risk were those with BRA scores within
[−20.0, 1.0[and BRA+CCA scores within [−32.0, 1.0[(note the re-
verse bracket notation indicating in all cases an open interval).
Of the 45 jellyfish species assessed, based on the BRA threshold, 16
(35.6%) were classified as high risk, 23 (51.1%) as medium risk, and six
(13.3%) as low risk. Based on the BRA+CAA threshold, 13 (28.9%)
4
Marine Pollution Bulletin 150 (2020) 110728
Invasive Zenetos et al., 2010 Zenetos et al., 2012
Y √ –
– –
N √ –
were classified as high risk, 30 (66.7%) as medium risk, and two (4.4%)
as low risk. For the BRA, amongst the species categorised a priori as
invasive, eight were correctly classified as high risk (true positives),
whereas Cotylorhiza erythraea was the only false negative. Amongst the
species categorised a priori as non-invasive, five were correctly classi-
fied as low risk (true negatives), and eight were false positives. For the
BRA+CAA, amongst the species categorised a priori as invasive, seven
were correctly classified as high risk (true positives), and there were no
false negatives; amongst the species categorised a priori as non-in-
vasive, two were correctly classified as low risk (true negatives), and six
were false positives. Finally, for the BRA, of the 23 medium risk species
17 were categorised a priori as non-invasive and six as invasive; for the
BRA+CAA, of the 30 medium risk species 22 were categorised a priori
as non-invasive and eight as invasive (Table 2; combined AS-ISK report
for the screenedspecies in Supplementary material I).
For the BRA, the highest-scoring species (score ≥ 19, chosen as a
relative very high risk ‘sub-threshold’) were Cassiopea andromeda,
Phyllorhiza punctata, Chrysaora quinquecirrha and Rhopilema nomadica.
And the same was true for the BRA+CCA, but with Cassiopea andro-
meda having the highest and Rhopilema nomadica the lowest score, and
with Chrysaora quinquecirrha and Phyllorhiza punctata attributed inter-
mediate scores (Table 2).
The CCA resulted in an increase in the BRA score for 22 (48.9%) of
the screened species, whereas for the remaining 23 species there was no
change from the BRA to the BRA+CCA score (delta = 0: Table 2), nor
were there any species for which the BRA+CCA score resulted in a
decrease relative to the BRA score. Notably, Cassiopea andromeda was
the only species achieving the highest possible (positive) change in
score of 12, whereas for the other species the delta value ranged from 2
to 6 (Table 2).
The CL (over all 55 Qs) was 2.370 ± 0.033 SE, the CLBRA
2.402 ± 0.035 SE, and the CLCCA 2.107 ± 0.044 SE (hence, in all
cases indicating medium to high confidence), and the CLBRA was sig-
nificantly higher than the CLCCA (F# 1,88 = 27.51, P < 0.001). Similarly,
mean values for CF = 0.593 ± 0.008 SE and CFBRA = 0.601 ± 0.009
SE were higher than the mean value for the CFCCA = 0.527 ± 0.011
SE, and the mean CFBRA was significantly higher than the mean CFCCA
(same significance values as for the CLBRA vs CLCCA comparison due to
the two indices being related). In all cases, the narrow standard errors
indicated overall similarity in CLs and CFs across the species assessed.
4. Discussion
In this first application of AS-ISK to the jellyfishes of the
Mediterranean Sea, approximately one third (16) of the 45 screened
species in total were classified as posing a high risk of being invasive
(Table 2). Overall, common traits can be identified in the highest
scoring species. Thus, the four species considered to pose the greatest
risk (BRA scores > 19) all possessed the following traits: (i) relatively
large bodies (> 15 cm diameter) in their medusa life stage, predators
on zooplankton, and reported to form prolific blooms under certain
environmental conditions; (ii) known NN occurrence in the Medi-
terranean Sea, with the exception of Chrysaora quinquecirrha; and (iii)
history of known invasiveness in other regions. Notably, all of these
other criteria are directly related to AS-ISK questions, namely questions
4.01 “Is it likely that the taxon will smother one or more native taxa
(that are not threatened or protected)?”, 2.03 “Is the taxon already
present outside of captivity in the RA area?” and 3.01 “Has the taxon
N. Killi, et al. Marine Pollution Bulletin 150 (2020) 110728

Fig. 1. Distribution map of the native and non-native (NN) jellyfish species screened with the Aquatic Species Invasiveness Screening Kit (AS-ISK) for the
Mediterranean Sea. 1: Indo-Pacific Ocean; 2: Mediterranean Sea; 3: North Atlantic Ocean; 4: South Atlantic Ocean; 5: Gulf of Mexico; 6: South Pacific Ocean; 7: North
Pacific Ocean. See also Table 1 (the base map was taken and modified from https://commons.wikimedia.org).

become naturalised (established viable populations) outside its native
range?”, respectively (Copp et al., 2016a). Finally, the BRA scores of the
majority of these species increased under the predicted climate change,
as indicated by their BRA+CCA scores – these species tended to ori-
ginate from warmer regions (primarily the Indo-Pacific).
The highest-scoring species was Cassiopea andromeda, which was
first reported in the Mediterranean Sea for Cyprus (Maas, 1903), and it
has since spread to the coasts of Israel, Lebanon, Turkey, Greece and
Malta (Galil et al., 1990; Bilecenoğlu, 2002; Schembri et al., 2010;
Zenetos et al., 2010). Currently, C. andromeda has established popula-
tions in the south-western Aegean Sea and along the Mediterranean
coasts of Turkey (Çevik et al., 2006; Özgür and Öztürk, 2008; Gülşahin

5
N. Killi, et al. Marine Pollution Bulletin 150 (2020) 110728

Table 2
Non-native (NN) jellyfish species assessed with AS-ISK for the Mediterranean Sea. For each species, the taxonomic class (C = Cubomedusae; H = Hydromedusae;
S = Scyphomedusae), selection criterion [Crit.: E (Extant) = native and NN species already present in the Mediterranean Sea; H (Horizon-scanning) = NN species
likely to enter the Mediterranean Sea via the Suez Canal (Red Sea) and the Strait of Gibraltar (Atlantic Ocean) or via ballast water], a priori categorisation (N = non-
invasive; Y = invasive), Basic Risk Assessment (BRA), and BRA plus Climate Change Assessment (BRA+CCA) scores and corresponding risk outcomes, difference
(Delta) between BRA+CCA and BRA scores, Confidence Level (CL) and Confidence Factor (CF) (see text for explanation) for all questions (Total) and separately for
the BRA and CCA components of the risk assessment. Risk outcomes are based on a threshold of 6.5 for the BRA (Low: score within interval [−20, 1[; Medium: [1,
6.5[; High:]6.5, 68]) and of 12.5 for the BRA+CCA (Low: [−32, 1[; Medium: [1, 12.5[; High:]12.5, 80]) (note the reverse bracket notation indicating in all cases an
open interval). See also Fig. 2. Combined AS-ISK report for the assessed species in Supplementary material I.
Species name Common name Class Crit. Inv. Assessment component Confidence

BRA BRA + CCA CL CF

Score Outcome Score Outcome Delta Total BRA CCA Total BRA CCA

Aequorea conica – H E N 3.0 Medium 3.0 Medium 0.0 2.24 2.27 2.00 0.56 0.57 0.50
Aequorea globosa – H E Y 7.0 High 9.0 Medium 2.0 2.36 2.41 2.00 0.59 0.60 0.50
Aequorea vitrina – H E Y 3.0 Medium 3.0 Medium 0.0 2.24 2.27 2.00 0.56 0.57 0.50
Amphogona pusilla – H E Y 4.0 Medium 4.0 Medium 0.0 2.24 2.27 2.00 0.56 0.57 0.50
Arctapodema australis – H E Y 4.0 Medium 4.0 Medium 0.0 2.24 2.27 2.00 0.56 0.57 0.50
Campanularia morgansi – H E N −1.0 Low −1.0 Low 0.0 2.29 2.33 2.00 0.57 0.58 0.50
Carukia barnesi box jelly C H N 6.0 Medium 12.0 Medium 6.0 2.36 2.41 2.00 0.59 0.60 0.50
Cassiopea andromeda upside-down jellyfish S E Y 26.0 High 38.0 High 12.0 3.00 3.06 2.50 0.75 0.77 0.63
Cassiopea xamachana upside-down jelly S H N 5.0 Medium 5.0 Medium 0.0 2.25 2.29 2.00 0.56 0.57 0.50
Cephea cephea Crowned jellyfish S H N 0.0 Low 4.0 Medium 4.0 2.31 2.35 2.00 0.58 0.59 0.50
Chrysaora fulgida – S H N 5.0 Medium 5.0 Medium 0.0 2.42 2.47 2.00 0.60 0.62 0.50
Chrysaora quinquecirrha Sea nettle S H Y 24.0 High 28.0 High 4.0 2.31 2.35 2.00 0.58 0.59 0.50
Cirrholovenia tetranema – H E Y 5.0 Medium 5.0 Medium 0.0 2.11 2.12 2.00 0.53 0.53 0.50
Clytia hummelincki – H E N 10.0 High 14.0 High 4.0 2.44 2.49 2.00 0.61 0.62 0.50
Clytia linearis – H E Y 10.0 High 14.0 High 4.0 2.44 2.49 2.00 0.61 0.62 0.50
Clytia mccradyi – H E Y 10.0 High 14.0 High 4.0 2.44 2.49 2.00 0.61 0.62 0.50
Corymorpha annulata – H E N 4.0 Medium 10.0 Medium 6.0 2.56 2.51 3.00 0.64 0.63 0.75
Corymorpha bigelowi – H E N 4.0 Medium 10.0 Medium 6.0 2.56 2.51 3.00 0.64 0.63 0.75
Coryne eximia – H E N 11.0 High 13.0 High 2.0 2.65 2.73 2.00 0.66 0.68 0.50
Cotylorhiza erythraea – S E Y 0.0 Low 4.0 Medium 4.0 2.25 2.27 2.17 0.56 0.57 0.54
Eirene viridula – H E N 4.0 Medium 10.0 Medium 6.0 2.56 2.51 3.00 0.64 0.63 0.75
Eucheilota paradoxica – H E N 5.0 Medium 5.0 Medium 0.0 2.11 2.12 2.00 0.53 0.53 0.50
Eucheilota ventricularis – H E N 5.0 Medium 5.0 Medium 0.0 2.11 2.12 2.00 0.53 0.53 0.50
Gonionemus vertens Clinging jellyfish H E Y 5.0 Medium 5.0 Medium 0.0 2.11 2.12 2.00 0.53 0.53 0.50
Haliscera bigelowi – H E N 8.0 High 14.0 High 6.0 2.36 2.41 2.00 0.59 0.60 0.50
Halitiara inflexa – H E N 5.0 Medium 5.0 Medium 0.0 2.11 2.12 2.00 0.53 0.53 0.50
Kantiella enigmatica – H E N 5.0 Medium 5.0 Medium 0.0 2.11 2.12 2.00 0.53 0.53 0.50
Laodicea fijiana – H E N −1.0 Low −1.0 Low 0.0 2.29 2.33 2.00 0.57 0.58 0.50
Marivagia stellata – S E Y 3.0 Medium 9.0 Medium 6.0 2.31 2.35 2.00 0.58 0.59 0.50
Mawia benovici – S E N −1.0 Low 1.0 Medium 2.0 2.29 2.33 2.00 0.57 0.58 0.50
Moerisia carine – H E N 11.0 High 13.0 High 2.0 2.65 2.73 2.00 0.66 0.68 0.50
Moerisia inkermanica – H E Y 11.0 High 13.0 High 2.0 2.65 2.73 2.00 0.66 0.68 0.50
Netrostoma setouchianum – S H N −2.0 Low 2.0 Medium 4.0 2.22 2.24 2.00 0.55 0.56 0.50
Nubiella mitra – H E N 14.0 High 14.0 High 0.0 2.40 2.45 2.00 0.60 0.61 0.50
Olindias singularis – H E N 4.0 Medium 10.0 Medium 6.0 2.56 2.51 3.00 0.64 0.63 0.75
Paracytaeis octona – H E N 5.0 Medium 5.0 Medium 0.0 2.11 2.12 2.00 0.53 0.53 0.50
Phyllorhiza punctata Australian spotted jellyfish S E Y 25.0 High 27.0 High 2.0 2.96 3.06 2.17 0.74 0.77 0.54
Physalia physalis Portuguese man-of-war H E N 12.0 High 12.0 Medium 0.0 2.49 2.55 2.00 0.62 0.64 0.50
Rhopilema esculentum Flame jellyfish S H N 14.0 High 14.0 High 0.0 2.45 2.51 2.00 0.61 0.63 0.50
Rhopilema nomadica – S E Y 19.0 High 23.0 High 4.0 2.67 2.76 2.00 0.67 0.69 0.50
Rhopilema verrilli mushroom jellyfish S H N 3.0 Medium 3.0 Medium 0.0 2.25 2.29 2.00 0.56 0.57 0.50
Scolionema suvaense – H E N 5.0 Medium 5.0 Medium 0.0 2.11 2.12 2.00 0.53 0.53 0.50
Stomolophus meleagris cannonball jellyfish S E N 11.0 High 11.0 Medium 0.0 2.69 2.78 2.00 0.67 0.69 0.50
Tetrorchis erythrogaster – H E N 4.0 Medium 4.0 Medium 0.0 2.24 2.27 2.00 0.56 0.57 0.50
Trichydra pudica – H E N 5.0 Medium 5.0 Medium 0.0 2.11 2.12 2.00 0.53 0.53 0.50

and Tarkan, 2012). This species has a symbiotic relationship with algae
(zooxanthellae) and a commensal relationship with the crustacean
mysid Idiomysis tsurnamali (Ludwig, 1969; Bacescu, 1973). The high AS-
ISK score reflects the species' ability to tolerate low-nutrient environ-
ments (Niggl and Wild, 2009), its documented invasiveness elsewhere
(e.g. Hawaii: Graham and Bayha, 2008) and adverse effects caused by
large swarms of zooplankton communities (Purcell, 2012), and its po-
tential adverse effects on human health caused by its painful sting,
ultimately resulting in negative socio-economic effects on tourism
(Graham and Bayha, 2008; Öztürk and İşinibilir, 2010). Owing to its
ability to tolerate low-nutrient environments (Niggl and Wild, 2009),
the warm-water C. andromeda is well suited to the Mediterranean Sea,
where further propagation and proliferation can be expected, and
especially so under predicted global warming conditions. This is re-
flected in its CCA score of 12, the maximum achievable in AS-ISK.
The second highest-scoring species, Chrysaora quinquecirrha, is dis-
tributed along the coasts of the Atlantic and Indian oceans as well as the
western Pacific. This species was also recorded along the eastern coasts
of the Americas from New England (USA) to Brazil (Calder, 1972).
Genetic and morphological differences indicate that Chrysaora spp. are
distributed in American estuaries and the Gulf of Mexico, whereas this
is the case of Chrysaora chesapeakei (Bayha et al., 2017). Chrysaora
quinquecirrha was included in the present study as a horizon-scanning
species that has not yet been recorded in the Mediterranean, and it
attained a high BRA score due to its propensity to form large blooms in
its native range of the western Atlantic, with documented negative

6
N. Killi, et al.
Fig. 2. (a) Receiver Operating Characteristic (ROC) curve (solid line) for the
Basic Risk Assessment (BRA) of the NN jellyfish species screened with AS-ISK
for the Mediterranean Sea. Smoothing line and confidence intervals of specifi-
cities (grey area) are also provided. (b) Same for the BRA+CCA (Climate
Change Assessment). See also Table 2.
impacts on tourism (beach closures: Burnett, 2001), industry (clogging
of pipes: Lynam et al., 2006) and fisheries (Duffy et al., 1997). Chry-
saora quinquecirrha can play an important role in structuring zoo-
plankton communities (Purcell, 1992; Purcell and Decker, 2005) and
was found to exert high predation pressure on copepods in Chesapeake
Bay, USA (Purcell, 1992). Although native to colder regions, C. quin-
quecirrha polyp and egg production increases in warmer waters (Purcell
and Decker, 2005), hence its risk of establishment in the Mediterranean
under predicted climate change is reflected in its elevated CCA score.
Finally, an overall tolerance for low oxygen levels has also been re-
ported for this species (Condon et al., 2001).
The third highest-scoring species, Phyllorhiza punctata, was first
observed in the Mediterranean Sea in 1965 and has since been reported
7
Marine Pollution Bulletin 150 (2020) 110728
from both its western and eastern parts (Galil et al., 1990, 2009; Abed-
Navandi and Kikinger, 2007). This species prefers coastal lagoons
(García, 1990), lives with symbiotic zooxanthellae (García and Durbin,
1993), and in the absence of the latter feeds on copepods and other
zooplankton (Graham et al., 2003). The high-risk score attributed to P.
punctata was in large part the result of its establishment, large blooms
and adverse impacts elsewhere as an invasive NN species – this includes
an estimated USD 10 million loss to shrimp fisheries in the Gulf of
Mexico, where fishery closure was forced by net clogging (Graham
et al., 2003), and a reduction in zooplankton levels with potential
knock-on effects on local fisheries in Puerto Rico (García and Durbin,
1993). Under future climate conditions, this species may or may not
benefit from warmer water temperatures, as reflected in the limited
increase in the BRA+CCA score relative to the BRA score (Table 2).
The fourth highest-scoring species, Rhopilema nomadica, has estab-
lished populations that are known to cause large swarms in the
Levantine Basin of the Mediterranean Sea (Galil et al., 1990; Kıdeys and
Gücü, 1995). It is believed that this species entered the RA area by
currents and ship ballast waters via the Suez Canal (Galil et al., 1990).
Blooms of R. nomadica have been recorded from Adana, Antalya, İs-
kenderun and Mersin in Turkey and often cause clogging of fishing nets
(Öztürk and İşinibilir, 2010; Turan et al., 2011). Also, several cases of
hospitalisation have been reported as a result of its traumatogenic ef-
fects (Gülşahin, 2017). As a warm-water species, global warming con-
ditions might be expected to benefit R. nomadica (CCA score equal to 4).
Clearly, global warming could exert a similar effect on all species, but a
rise in water temperature could increase the likelihood of the species
moving outside of its native range to more northern (or southern) lo-
cations, i.e. depending upon the hemisphere in question (Walther et al.,
2002).
Amongst the other high-risk species (BRA scores from 7.0 to 14.0),
there are several that have shown high rates of dispersal throughout the
Mediterranean Sea. Amongst these, Clytia linearis was the first suc-
cessful Lessepsian hydrozoan to enter the Mediterranean via the Suez
Canal, subsequently spreading into the Aegean Sea (Çınar et al., 2014)
and then throughout the Mediterranean and into the Atlantic Ocean
(Boero et al., 2005). Similarly, the hydrozoan Coryne eximia, which has
a global distribution (Schuchert, 2005), is believed to have elevated
dispersal potential due to its ability to deposit its eggs on mussels (Puce
et al., 2003), and it can also act as host to digenean parasites of fishes
(Briz et al., 2012). Four of the other high-risk species already estab-
lished in the Mediterranean include: Clytia mccradyi, which was re-
corded from several coastal areas in the western and eastern Medi-
terranean Sea (Segura-Puertas and Damas-Romero, 1997; Canchè-
Canchè and Castellanos, 2005; Segura-Puertas et al., 2010); Clytia
hummelincki, which was first recorded in 1997 and established in the
Adriatic Sea (Boero et al., 2005; Zenetos et al., 2010); Haliscera bigelowi,
which is native to the Indo-Pacific and southern oceans, and reported
occasionally in the Adriatic Sea at depths of 600–800 m (Thuesen and
Childress, 1994; Gili et al., 1998); and the cannonball jellyfish Stomo-
lophus meleagris, which is also present in the Mediterranean (Zenetos
et al., 2010) and creates large blooms that adversely affect local fish-
eries, e.g. in East China and the Yellow Sea (Ding and Cheng, 2007).
Of the species classified as medium risk (Table 2), the box jelly
Carukia barnesi could pose a major health risk if introduced into the
Mediterranean Sea. In extreme cases, this species' traumatogenic effects
can result in fatal cerebral haemorrhage and hypertension ≈30 min
following sting contact (Barnes, 1964; Fenner et al., 1986). Native to
coasts of Thailand and Australia (De Pender et al., 2006; Underwood
and Seymour, 2007), this species is difficult to detect due to its small
size and transparency (Underwood and Seymour, 2007).
Amongst the species that were found to pose a low risk of inva-
siveness (Table 2), the scyphozoan Mawia benovici is known for its
blooms in the Adriatic Sea (Piraino et al., 2014). However, the natural
distribution area for this species remains uncertain, although it is
thought to have spread as a result of shipping and aquaculture activities
N. Killi, et al.
(Galil, 2012). Whereas, the low-risk classification of the scyphozoan
Cotylorhiza erythraea is perplexing given its a priori categorisation as
invasive (hence, making it a false negative). This species has only re-
cently been recorded for the Mediterranean Sea (Galil et al., 2017), and
despite its preference for high temperatures and salinities, it can only
produce one larva per polyp per year – a reproductive strategy that
contributes to its low score and thus low-risk classification.
The overall medium-to-high confidence rankings for the assess-
ments in the present study contrast the higher (i.e. medium-high to
high) confidence levels found in other AS-ISK applications on fresh-
water and marine fishes (Glamuzina et al., 2017; Tarkan et al., 2017a,
2017b; Bilge et al., 2019; Dodd et al., 2019) and freshwater in-
vertebrates (Paganelli et al., 2018; Semenchenka et al., 2018). This
discrepancy can be explained by the limited information available on
the biology/ecology of jellyfishes relative to that available on fishes (cf.
FishBase: Froese and Pauly, 2017) and invertebrates (cf. RA area spe-
cific studies). This is the case for some of the jellyfishes assessed in the
present study, though some species are quite well studied, particularly
those that attracted the highest scoring. However, this was not always
necessarily the case, as some less studied jellyfishes, such as Cassiopea
andromeda, could yield high scores. Also, larger availability of literature
information generally results in higher confidence rankings, though not
necessarily higher risk scores. In this respect, a detailed study of the NN
freshwater fish species, pumpkinseed Lepomis gibbosus, in England
found limited evidence of impacts (Copp et al., 2010). For most species,
the potential (or likely) impacts of climate change are poorly known,
and as such the CCA questions are more likely to receive low confidence
rankings than the BRA ones. This discrepancy in confidence rankings
between BRA and BRA+CCA scores has been reported in the initial
applications of AS-ISK (Glamuzina et al., 2017; Paganelli et al., 2018;
Semenchenka et al., 2018). And it also emphasises the dynamic nature
of NN species risk analysis (Yemshanov et al., 2009), whereby risk
screenings and full assessments need to be regularly updated to in-
corporate new information (see Vilizzi et al., 2019), which will modify
the outcome risk score and consequently the level of confidence in the
assessment.
Computationally, the classification of some species (both BRA and
BRA+CCA scores) contrasted the expert-based, a priori categorisation
of the jellyfishes. This was the case of flame jellyfish Rhopilema escu-
lentum, which was categorised as non-invasive but attracted a BRA
score of 14.0 (Table 2). This species forms blooms and causes many
stinging cases in the south Asia, but blooms are controlled in China,
where it is harvested for human consumption (Dong et al., 2010).
However, blooms of this species in another region may cause serious
problems due to an absence of human harvest. However, the a priori
categorisation of species may be biased, being based purely on locality-
specific information or derived simply from the knowledge base of (i.e.
the extent of literature search by) the source from which the a priori
categorisation itself was derived (see Onderdonk et al., 2010). This is
particularly relevant in the present study, which adopted an expert-
based a priori categorisation of NN jellyfishes that was constrained by a
limited amount of information on the invasion histories (i.e. invasive-
ness status) of the species screened.
Clearly, if translated into management actions, then the con-
sequences of ‘rejecting’ a non-invasive species (false positive) vs those
of ‘accepting’ an invasive species will ultimately have to be weighed in
terms of economic benefits vs losses. Overall, the present assessments
resulted in a high level of accuracy for the AS-ISK tool (i.e. a larger
proportion of true vs false negatives and of true vs false positives).
Therefore, in future studies of NN jellyfishes limited to a small number
of species, which prevents local calibration to identify ad hoc thresh-
olds, the thresholds set in the present study for the Mediterranean Sea
could be employed in AS-ISK applications in other RA areas. Whereas,
at a broader scale, the current thresholds may contribute to the re-
finement of overall thresholds for marine invertebrates in general.
8
Marine Pollution Bulletin 150 (2020) 110728
5. Policy and management implications
Overall, the outcomes of the present study are expected to con-
tribute to the development and implementation of management mea-
sures to prevent further spread and/or introduction of high-risk NN
jellyfishes. This is especially important for the Mediterranean Sea,
which is subject to increasing levels of biological invasion (Galil et al.,
2018) and where the biological and societal impacts of some NN jel-
lyfishes could be underestimated in the absence of basic information.
There are numerous global examples of serious economic impacts of
jellyfish blooms; for example, some 50 t of Rhopilema nomadica had to
be removed from power plant intakes (Verner, 1984), whilst Japanese
fishers lost millions of dollars as a result of Nomura's jellyfish Nemopi-
lema nomurai blooms (Uye, 2008).
The management implications of the present study are therefore
especially relevant for: (i) prioritising controls on pathways (e.g. ballast
water) that could be used by new jellyfish invaders, or aid spread of
established species within the Mediterranean; (ii) informing citizens
and municipalities about the potential human-induced impacts, such as
coastal developments, on jellyfish blooms; (iii) implementation of early
warning systems; (iv) regulation of coastal construction activities; and
(v) conservation of amenity values for the tourism industry. For ex-
ample, salmon farms are not recommended for establishment in tidal
zones, which are prone to jellyfish blooms (Graham et al., 2001). In
Australia and other regions, which have suffered from detrimental jel-
lyfish blooms, local managers and various forms of the media (news,
social) are working to minimise jellyfish stings. To this end, tourists are
informed about dangerous jellyfishes (e.g. Physalia physalis and box
jellyfish Chironex fleckeri) and warned to swim behind stinger nets
(Harrison et al., 2004; Lucas et al., 2014). Also, early warning systems
and net barriers for Pelagia noctiluca blooms were applied in some
beaches of Spain and France. The Internet is also being used in the
Europe to share photos and information on jellyfish blooms (Lucas
et al., 2014). Collaboration with research institutes, municipalities, ci-
tizens and the tourism sector is therefore crucial to inform about jel-
lyfish outbreaks, which are known to occur irregularly both spatially
and temporally. For this reason, several citizen-oriented programmes
have been activated to share information about jellyfish blooms, such as
CIESM JellyWatch Programme, Monterey Bay Aquarium Research In-
stitute Jellywatch Programmes (JelliesZone) and UK Marine Con-
servation Society Jellyfish Survey.
Early warning systems should be implemented for controlling water
quality via long-term monitoring studies as well as aerial and satellite
surveys, all supported by modelling of sea current patterns (Elzeir et al.,
2005; Houghton et al., 2006). As coastal constructions may increase the
spread of polyps by favouring their settling on scaffoldings, marina
ropes and shipwrecks, regulatory measures need to be implemented,
including the use of environmentally friendly anti-fouling ship paints,
to avoid settlement and dispersion of polyps in marina areas (Guenther
et al., 2010). Also, since tourism activities can be disrupted by jellyfish
blooms (especially in the case of traumatogenic species), so early-
warning measures for bathers and the closure of beaches are re-
commended (Gershwin et al., 2010).
Declaration of competing interests
The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to influ-
ence the work reported in this paper.
Acknowledgements
This study was supported by Muğla Sıtkı Koçman University
Scientific Research Project Office under project number 17/114. The
participation of GHC and PID was supported by Cefas as part of their
contribution to the ICES WGITMO initiative for the global trial of AS-
N. Killi, et al.
ISK.
Appendix A. Supplementary data
Supplementary data to this article can be found online at https://
doi.org/10.1016/j.marpolbul.2019.110728.
References
Abed-Navandi, D., Kikinger, R., 2007. First record of the tropical scyphomedusa
Phyllorhiza punctata von Lendenfeld, 1884 (Cnidaria: Rhizostomeae) in the Central
Mediterranean Sea. Aquat. Invasions 2, 302–312.
Anderson, M.J., Gorley, R.N., Clarke, K.R., 2008. Permanova for Primer: Guide to
Software and Statistical Methods. PRIMER–E Ltd, Plymouth.
Arai, M.N., 2001. Pelagic coelenterates and eutrophication: a review. Hydrobiologia 451,
69–87.
Arai, M.N., 2005. Predation on pelagic coelenterates: a review. J. Mar. Biol. Assoc. UK 85,
523–536.
Avian, M., Ramšak, A., Tirelli, V., D'Ambra, I., Malej, A., 2016. Redescription of Pelagia
benovici into a new jellyfish genus, Mawia, gen. Nov., and its phylogenetic position
within Pelagiidae (Cnidaria: Scyphozoa: Semaeostomeae). Invert. System. 30,
523–546.
Bacescu, M., 1973. A new case of commensalism in the Red Sea: the mysid Idiomysis
tsurnamali n. sp. with the Coelenterata Megalactis and Cassiopea. Rom. J. Biol. – Zool.
18, 3–7.
Barnes, J.H., 1964. Cause and effect in Irukandji stingings. Med. J. Aust. 1, 897–904.
Bayha, K.M., Collins, A.G., Gaffney, P.M., 2017. Multigene phylogeny of the scyphozoan
jellyfish family Pelagiidae reveals that the common US Atlantic sea nettle comprises
two distinct species (Chrysaora quinquecirrha and C. chesapeakei). PeerJ 5, e3863.
Bewick, V., Cheek, L., Ball, J., 2004. Statistics review 13: receiver operating characteristic
curves. Crit. Care 8, 508.
Bilecenoğlu, M., 2002. Türkiye kıyılarında ilk kez rastlanan bir denizanası türü: Cassiopea
andromeda (Forsskal, 1775) (Cassiopeidae, Scyphozoa) [A jellyfish species found for
the first time in Turkey coast: Cassiopea andromeda (Forsskal, 1775) (Cassiopeidae,
Scyphozoa)]. Underwater World Magazine 72, 42–43 (in Turkish).
Bilge, G., Filiz, H., Yapıcı, S., Tarkan, A.S., Vilizzi, L., 2019. A risk screening study on the
potential invasiveness of Lessepsian fishes in the south-eastern coasts of Anatolia.
Acta Ichthyol. Piscat. 49, 23–31.
Boero, F., Di Camillo, C., Gravili, C., 2005. Phantom aliens in Mediterranean waters.
Aquat. Invasions 3, 21–22.
Boero, F., Bouillon, J., Gravili, C., Miglietta, M.P., Parsons, T., Piraino, S., 2008.
Gelatinous plankton: irregularities rule the world (sometimes). Mar. Ecol. Prog. Ser.
356, 299–310.
Boero, F., Brotz, L., Gibbons, M.J., Piraino, S., Zampardi, S., 2016. Impacts and effects of
ocean warming on jellyfish, pp. 213–237. In: Laffoley, D., Baxter, J.M. (Eds.),
Explaining Ocean Warming: Causes, Scale, Effects and Consequences. IUCN, Gland,
Switzerland.
Bouillon, J., Medel, M.D., Pagés, F., Gili, J.M., Boero, F., Gravili, C., 2004. Fauna of the
Mediterranean Hydrozoa. Sci. Mar. 68, 1–449.
Briz, L.M.D., Martorelli, S.R., Genzano, G.N., Mianzan, H.W., 2012. Parasitism
(Trematoda, Digenea) in medusae from the southwestern Atlantic Ocean: medusa
hosts, parasite prevalences, and ecological implications. Hydrobiologia 690,
215–226.
Burnett, J.W., 2001. Medical aspects of jellyfish envenomation: pathogenesis, case re-
porting and therapy. Hydrobiologia 451, 1–9.
Canché-Canché, V.E., Castellanos-Osorio, I., 2005. Medusas (Cnidaria) de la Bahía de la
Ascensión, Quintana Roo, México (1997). Hidrobiológica 15 (1), 65–72.
Calder, D.R., 1972. Development of the sea nettle Chrysaora quinquecirrha (Scyphozoa,
Semaeostomeae). Chesap. Sci. 13, 40–44.
Çevik, C., Erkol, I.L., Toklu, B., 2006. A new record of an alien jellyfish from the Levantine
coast of Turkey – Cassiopea andromeda (Forsskål, 1775) [Cnidaria: Scyphozoa:
Rhizostomea]. Aquat. Invasions 1, 196–197.
Çınar, M.E., Yokeş, M.B., Açık, Ş., Bakir, A.K., 2014. Checklist of Cnidaria and Ctenophora
from the coasts of Turkey. Turk. J. Zool. 38, 677–697.
Condon, R.H., Decker, M.B., Purcell, J.E., 2001. Effects of low dissolved oxygen on sur-
vival and asexual reproduction of scyphozoan polyps (Chrysaora quinquecirrha).
Hydrobiologia 451, 89–95.
Copp, G.H., Stakenas, S., Cucherousset, J., 2010. Aliens versus the natives: interactions
between introduced pumpkinseed and indigenous brown trout in small streams of
Southern England. Am. Fish. Soc. Symp. 73, 347–370.
Copp, G.H., Vilizzi, L., Tidbury, H., Stebbing, P.D., Tarkan, A.S., Miossec, L., Goulletquer,
P., 2016a. Development of a generic decision-support tool for identifying potentially
invasive aquatic taxa: AS-ISK. Manag. Biol. Invasion 7, 343–350.
Copp, G.H., Godard, M.J., Russell, I.C., Peeler, E.J., Gherardi, F., Tricarico, E., Macleod,
A., Cowx, I.G., Nunn, A.D., Occhipinti-Ambrogi, A., Savini, D., Mumford, J., Britton,
J.R., 2016b. European non-native species in aquaculture risk analysis scheme–a
summary of assessment protocols and decision support tools for use of alien species in
aquaculture. Fisheries Manag. Ecol. 23, 1–11.
Corriero, G., Pierri, C., Accoroni, S., Alabiso, G., Bavestrello, G., Barbonei, E., Bastianini,
M., Bazzoni, A.M., Aubry, F.B., Boero, F., Buia, B.C., Cabrini, M., Camatti, E.,
Cardone, F., Cataletto, B., Vietti, R.C., Cecere, E., Cibic, T., Colangelo, P., De
Olazabal, A., D'onghia, G., Finotto, S., Fiore, N., Fornasaro, D., Fraschetti, S., Gambi,
M.C., Giangrande, A., Gravili, C., Guglielmo, R., Longo, C., Lorenti, M., Lugliè, A.,
9
Marine Pollution Bulletin 150 (2020) 110728
Maiorano, P., Mazzocchi, M.G., Mercurio, M., Mastrototaro, F., Mistri, M., Monti, M.,
Munari, C., Musco, L., Nonnis-Marzano, C., Padedda, B.M., Patti, F.P., Petrocelli, A.,
Piraino, S., Portacci, G., Pugnetti, A., Pulina, S., Romagnoli, T., Rosati, I., Sarno, D.,
Satta, C.T., Sechi, N., Schiaparelli, S., Scipione, B., Sion, L., Terlizzi, A., Tirelli, V.,
Totti, C., Tursi, A., Ungaro, N., Zingone, A., Zupo, V., Basset, A., 2016. Ecosystem
vulnerability to alien and invasive species: a case study on marine habitats along the
Italian coast. Aquat. Conserv. 26, 392–409.
De Pender, A.M., Winkel, K.D., Ligthelm, R.J., 2006. A probable case of Irukandji syn-
drome in Thailand. J. Travel Med. 13, 240–243.
Demir, İ., Kılıç, G., Coşkun, M., 2008. PRECIS Bölgesel İklim Modeli ile Türkiye İçin İklim
Öngörüleri: HadAMP3 SRES A2 Senaryosu (PRECIS Regional climate model and
climate predictions for Turkey: HadAMP3 SRES A2 Scenario). In: IV. Atmosfer
Bilimleri Sempozyumu, Bildiriler Kitabı, 365–373. İTÜ Uçak ve Uzay Bilimleri
Fakültesi Meteoroloji Mühendisliği Bölümü, 25–28 Mart 2008, İstanbul, Turkey.
Ding, F.Y., Cheng, J.H., 2007. Dynamic distribution of Stomolophus meleagris in the East
China Sea region. Journal of Fishery Sciences of China 14, 83–89.
Dodd, J.A., Vilizzi, L., Bean, C.W., Davison, P.I., Copp, G.H., 2019. At what spatial scale
should risk screenings of translocated freshwater fishes be undertaken – river basin
district or climo-geographic designation? Biol. Conserv. 230, 122–130.
Dong, Z., Liu, D., Keesing, J.K., 2010. Jellyfish blooms in China: dominant species, causes
and consequences. Mar. Pol. Bul. 60, 954–963.
Duffy, J.T., Epifanio, C.E., Fuiman, L.A., 1997. Mortality rates imposed by three scy-
phozoans on red drum (Sciaenops ocellatus Linnaeus) larvae in field enclosures.
Journal of Experimental Marine Biology and Ecology 212 (1), 123–131.
Elzeir, E., Hansen, I.S., Hay, S., 2005. Predicting jellyfish outbreaks around Shetland
using MIKE 3. In: WIT Trans. Built Env. vol 78 WIT Press.
Erickson, T.R., Stefan, H.G., 1996. Correlations of Oklahoma Stream Temperatures with
Air Temperatures. University of Minnesota, St Anthony Falls Laboratory, Project
Report No. 398.
Falkenhaug, T., 2014. Review of jellyfish blooms in the Mediterranean and Black Sea.
Mar. Biol. Res. 10, 1038–1039.
Fenner, P.J., Williamson, J., Callanan, V.I., Audley, I., 1986. Further understanding of,
and a new treatment for, “Irukandji” (Carukia barnesi) stings. Med. J. Aust. 145,
569–574.
Froese, F., Pauly, D., 2017. Fish Base. www.fishbase.org, Accessed date: 18 August 2018.
Galil, B.S., 2012. Truth and consequences: the bioinvasion of the Mediterranean Sea.
Integr. Zool. 7, 299–311.
Galil, B.S., Spanier, E., Ferguson, W.W., 1990. The Scyphomedusae of the Mediterranean
coast of Israel, including two Lessepsian migrants new to the Mediterranean. Zool.
Med. Leiden. 64, 95–105.
Galil, B.S., Shoval, L., Goren, M., 2009. Phyllorhiza punctata von Lendenfeld, 1884
(Scyphozoa: Rhizostomeae: Mastigiidae) reappeared off the Mediterranean coast of
Israel. Aquat. Invasions 4, 381–389.
Galil, B.S., Douek, J., Mienis, H.K., Rinkevich, B., 2016. Comments on “the Mediterranean
Sea as a gateway for invasion of the Red Sea: the case of the indo-West Pacific head-
shield slug Chelidonura fulvipunctata Baba, 1938” by Manuel António E. Malaquias,
Andrea Zamora-Silva, Dyana Vitale, Andrea Spinelli, Sergio De Matteo, Salvatore
Giacobbe, Deneb Ortigosa and Juan L. Cervera. Aquat. Invasions 11 (4), 351–354
2016.
Galil, B.S., Gershwin, L.A., Zorea, M., Rahav, A., Rothman, S.B., Fine, M., Lubinevsky, H.,
Douek, J., Paz, G., Rinkevich, B., 2017. Cotylorhiza erythraea Stiasny, 1920
(Scyphozoa: Rhizostomeae: Cepheidae), yet another erythraean jellyfish from the
Mediterranean coast of Israel. Mar. Biodivers. 47, 229–235.
Galil, B.S., Marchini, A., Occhipinti-Ambrogi, A., 2018. East is east and west is west?
Management of marine bioinvasions in the Mediterranean Sea. Estuar. Coast. Shelf
Sci. 201, 7–16.
Galil, B.S., Goren, M., 2014. Metamorphoses: bioinvasions in the Mediterranean Sea. In
The Mediterranean Sea Springer, Dordrecht, pp. 463–478.
García, J.R., 1990. Population dynamics and production of Phyllorhiza punctata (Cnidaria:
Scyphozoa) in Laguna Joyuda, Puerto Rico. Mar. Ecol. Prog. Ser. 64, 243–251.
García, J.R., Durbin, E., 1993. Zooplanktivorous predation by large scyphomedusae
Phyllorhiza punctata (Cnidaria: Scyphozoa) in Laguna Joyuda. J. Exp. Mar. Biol. Ecol.
173, 71–93.
Gershwin, L.A., De Nardi, M., Winkel, K.D., Fenner, P.J., 2010. Marine stingers: review of
an underrecognised global coastal management issue. Coast. Manage. 38, 22–41.
Gili, J.M., Bouillon, J., Pagès, F., Palanques, A., Puig, P., Heussner, S., 1998. Origin and
biogeography of the deep-water Mediterranean hydromedusae including the de-
scription of two new species collected in submarine canyons of Northwestern
Mediterranean. Sci. Mar. 62, 113–134.
Glamuzina, B., Tutman, P., Nikolić, V., Vidović, Z., Pavličević, J., Vilizzi, L., Copp, G.H.,
Simonović, P., 2017. Comparison of taxon-specific and taxon-generic risk screening
tools to identify potentially invasive non-native fishes in the River Neretva catchment
(Bosnia and Herzegovina and Croatia). River Res. Appl. 33, 670–679.
Graham, W.M., Pagès, F., Hamner, W.H., 2001. A physical context for gelatinous zoo-
plankton aggregations: a review. Hydrobiologia 451, 199–212.
Graham, W.M., Bayha, K.M., 2008. Biological invasions by marine jellyfish. In: Nentwig,
W. (Ed.), Biological Invasions, Ecological Studies (Analysis and Synthesis). vol. 193.
Springer Berlin Heidelberg, pp. 239–255.
Graham, W.M., Martin, D.L., Felder, D.L., Asper, V.L., Perry, H.M., 2003. Ecological and
economic implications of a tropical jellyfish invader in the Gulf of Mexico. Biol.
Invasions 5, 53–69.
Gravili, C., 2016. Zoogeography of Hydrozoa: past, present and a look to the future. In:
The Cnidaria, Past, Present and Future. Springer, Cham, pp. 95–107.
Guenther, J., Misimi, E., Sunde, L.M., 2010. The development of biofouling, particularly
the hydroid Ectopleura larynx, on commercial salmon cage nets in mid-Norway.
Aquaculture 300, 120–127.
N. Killi, et al.
Gülşahin, N., 2017. Jellyfish blooms and injuries in Turkey. In: Mariottini, Gian Luigi
(Ed.), Jellyfish: Ecology, Distribution Patterns and Human Interactions. Nova Science
Publishers, USA, pp. 149–162.
Gülşahin, N., Tarkan, A.N., 2012. Occurrence of the alien jellyfish Cassiopea andromeda
(Scyphozoa: Rhizostomeae: Cassiopeidae) in Hisarönü Bay, Muğla, Turkey. Biharean
Biol. 6, 132–133.
Harbison, G.R., 1993. The Potential of Fishes for the Control of Gelatinous Zooplankton.
vol. L,74. ICES (International Council for the Exploration of the Sea) CMS, pp. 1–10.
Harrison, S.L., Leggat, P.A., Fenner, P.J., Durrheim, D.N., Swinbourne, A.L., 2004.
Reported knowledge, perceptions, and behavior of tourists and North Queensland
residents at risk of contact with jellyfish that cause “Irukandji Syndrome”. Wild.
Environ. Med. 15, 4–10.
Hay, S., 2006. Marine ecology: gelatinous bells may ring change in marine ecosystems.
Curr. Biol. 16, 679–682.
Houghton, J.D.R., Doyle, T.K., Davenport, J., Hays, G.C., 2006. Developing a simple,
rapid, method for identifying and monitoring jellyfish aggregations from the air. Mar.
Ecol. Prog. Ser. 314, 159–170.
IPCC, 2005. In: Metz, B., Davidson, O., de Coninck, H., Loos, M., Meyer, L. (Eds.),
Working Group III of the Intergovernmental Panel on Climate Change, 2005. IPCC,
2005: IPCC Special Report on Carbon Dioxide Capture and Storage.
İşinibilir, M., Yılmaz, İ.N., 2016. Jellyfish species of the Sea of Marmara (pp. 390-400).
2016 In: Özsoy, E., Çağatay, M.N., Balkıs, N., Balkıs, N., Öztürk, B. (Eds.), The Sea of
Marmara; Marine Biodiversity, Fisheries, Conservation and Governance. Turkish
Marine Research Foundation (TUDAV), Publication No: 42, Istanbul, TURKEY (957
pages).
Kıdeys, A.E., Gücü, A.C., 1995. Rhopilema nomadica: A Lessepsian scyphomedusan new to
the Mediterranean coast of Turkey. Isr. J. Zool. 41, 615–617.
Langeneck, J., Crocetta, F., Doumpas, N., Giovos, I., Piraino, S., Boero, F., 2019. First
record of the non-native jellyfish Chrysaora cf. achlyos (Cnidaria: Pelagiidae) in the
Mediterranean Sea. Bioinvasions Rec 8, 608–613.
Lawson Jr., L.L., Vilizzi, L., Hill, J.E., Hardin, S., Copp, G.H., 2013. Revisions of the Fish
Invasiveness Screening Kit (FISK) for its application in warmer climatic zones, with
particular reference to peninsular Florida. Risk Anal. 33, 1414–1431.
Li, S., Chen, J., Wang, X., Copp, G.H., 2017. Invasiveness screening of non-native fishes
for the middle reach of the Yarlung Zangbo River, Tibetan Plateau, China. River Res.
Appl. 33, 1439–1444.
Lucas, C.H., Gelcich, S., Uye, S.I., 2014. Living with jellyfish: Management and adaptation
strategies. In: Jellyfish Blooms. Springer, Dordrecht, pp. 129–150.
Ludwig, F.D., 1969. Die zooxanthellenbei Cassiopea Andromeda Eschscholtz 1829 (Polyp-
Stadium) und ihre Bedeutung für die strobilation. Zoologische Jahrbücher. Abteilung
für Anatomie und Ontogenie der Tiere Abteilung für Anatomie und Ontogenie der
Tiere 86, 238–277.
Lynam, C.P., Gibbons, M.J., Axelsen, B.E., Sparks, C.A., Coetzee, J., Heywood, B.G.,
Brierley, A.S., 2006. Jellyfish overtake fish in a heavily fished ecosystem. Curr. Biol.
16, 492–493.
Maas, O., 1903. Die Scyphomedusen der Siboga Expedition — Siboga Expedition 1901.
vol. 11. pp. 1–91.
Matsueda, N., 1969. Presentation of Aurelia aurita at thermal power station. Bulletin of
the Marine Biological Station Asamushi 13, 191–197.
Mills, C.E., 2001. Jellyfish blooms: are populations increasing globally in response to
changing ocean conditions? Hydrobiologia 451, 55–68.
Mizrahi, G., Shemesh, E., van Ofwegen, L., Tchernov, D., 2015. First record of Aequorea
macrodactyla (Cnidaria, Hydrozoa) from the Israeli coast of the eastern
Mediterranean Sea, an alien species indicating invasive pathways. Neobiota 26, 55.
Möller, H., 1984. Reduction of a larval herring population by jellyfish predator. Science
224, 621–622.
Niggl, W., Wild, C., 2009. Spatial distribution of the upside-down jellyfish Cassiopea sp.
within fringing coral reef environments of the Northern Red Sea: implications for its
life cycle. Helgoland Mar. Res. 64, 281–287.
Onderdonk, D.A., Gordon, D.R., Fox, A.M., Stocker, R.K., 2010. Lessons learned from
testing the Australian weed risk assessment system: the devil is in the details. Plant
Prot. Q. 25, 79–85.
Özgür, E., Öztürk, B., 2008. A population of the alien jellyfish, Cassiopea andromeda
(Forsskål, 1775) [Cnidaria: Scyphozoa: Rhizostomea] in the Ölüdeniz Lagoon,
Turkey. Aquat. Invasions 3, 423–428.
Öztürk, B., İşinibilir, M., 2010. An alien jellyfish Rhopilema nomadica and its impacts to
the Eastern Mediterranean part of Turkey. J. Black Sea/Mediterranean Environment
16, 149–156.
Öztürk, B., Topaloğlu, B., Sümen, S., Turan, C., İşinibilir, M., Aktaş, Ş., Özen, Ş., 2018.
Jellyfish of the Black Sea and Eastern Mediterranean Waters. Turkish Marine
Research Foundation (TUDAV) Publication No. 48, Istanbul, TURKEY (75 pages).
Paganelli, D., Pandolfi, A., Sconfietti, R., Marchini, A., Vilizzi, L., 2018. Potential inva-
siveness by non-indigenous macrozoobenthos in the secondary hydrographic system
of a temperate-climate river catchment. Ecol. Indic. 88, 274–281.
Pauly, D., Graham, W., Libralato, S., Morissette, L., Palomares, M.D., 2009. Jellyfish in
ecosystems, online databases, and ecosystem models. Hydrobiologia 616, 67–85.
Pheloung, P.C., Williams, P.A., Halloy, S.R., 1999. A weed risk assessment model for use
as a biosecurity tool evaluation plant introductions. J. Environ. Manag. 57, 239–251.
Piraino, S., Aglieri, G., Martell, L., Mazzoldi, C., Melli, V., Milisenda, G., Scorrano, S.,
Boero, F., 2014. Pelagia benovici sp. nov. (Cnidaria, Scyphozoa): a new jellyfish in the
Mediterranean Sea. Zootaxa 3794, 455–468.
Prieto, L., Macías, D., Peliz, A., Ruiz, J., 2015. Portuguese Man-of-War (Physalia physalis)
in the Mediterranean: a permanent invasion or a casual appearance? Sci. Rep.-UK 5,
11545.
Puce, S., Bavestrello, G., Azzini, F., Cerrano, C., 2003. On the occurrence of Coryne eximia
Allman (Cnidaria, Corynidae) in the Mediterranean Sea. Ital. J. Zool. 70, 249–252.
10
Marine Pollution Bulletin 150 (2020) 110728
Purcell, J.E., 1992. Effects of predation by the scyphomedusan Chrysaora quinquecirrha on
zooplankton populations in Chesapeake Bay, USA. Mar. Ecol. Prog. Ser. 87, 65–76.
Purcell, J.E., 2005. Climate effects on formation of jellyfish and ctenophore blooms: a
review. J. Mar. Biol. Assoc. UK 85, 461–476.
Purcell, J.E., 2012. Jellyfish and ctenophore blooms coincide with human proliferations
and environmental perturbations. Annu. Rev. Mar. Sci. 4, 209–235.
Purcell, J.E., Arai, M.N., 2001. Interactions of pelagic cnidarians and ctenophores with
fish: a review. Hydrobiologia 451, 27–44.
Purcell, J.E., Decker, M.B., 2005. Effects of climate on relative predation by scyphome-
dusae and ctenophores on copepods in Chesapeake Bay during 1987–2000. Limnol.
Oceanogr. 50, 376–387.
Purcell, J.E., Nemazie, D.A., Dorsey, S.E., Houde, E.D., Gamble, J.C., 1994. Predation
mortality of bay anchovy (Anchoa mitchilli) eggs and larvae due to scyphomedusae
and ctenophores in Chesapeake Bay. Mar. Ecol. Prog. Ser. 114, 47–58.
R Core Team, 2016. R: A Language and Environment for Statistical Computing. R
Foundation for Statistical Computing, Vienna, Austria.
Rajagopal, S., Nair, K.V.K., Azariah, J., 1989. Some observations on the problem of jelly
fish ingress in a power station cooling system at Kalpakkam, east coast of India.
Mahasagar 22, 151–158.
Robin, X., Turck, N., Hainard, A., Tiberti, N., Lisacek, F., Sanchez, J.C., Müller, M., 2011.
PROC: an open-source package for R and S+ to analyze and compare ROC curves.
BMC Bioinformatics 12, 77.
Rodriguez, C.S., Pujol, M.G., Mianzan, H.W., Genzano, G.N., 2014. First record of the
invasive stinging medusa Gonionemus vertens in the southern hemisphere (Mar del
Plata, Argentina). Lat. Am. J. Aquat. Res. 42, 653–657.
Roy, H.E., Rabitsch, W., Scalera, R., Stewart, A., Gallardo, B., Genovesi, P., Essl, F.,
Adriaens, T., Bacher, S., Booy, O., Branquart, E., Brunel, S., Copp, G.H., Dean, H.,
D'hondt, B., Josefsson, M., Kenis, M., Kettunen, M., Linnamagi, M., Lucy, F.,
Martinou, A., Moore, N., Nentwig, W., Nieto, A., Pergl, J., Peyton, J., Roques, A.,
Schindler, S., Schönrogge, K., Solarz, W., Stebbing, P.D., Trichkova, T.,
Vanderhoeven, S., van Valkenburg, J., Zenetos, A., 2018. Developing a framework of
minimum standards for the risk assessment of alien species. J. Appl. Ecol. 55,
526–538.
Schembri, P.J., Deidun, A., Vella, P.J., 2010. First record of Cassiopea andromeda
(Scyphozoa: Rhizostomeae: Cassiopeidae) from the central Mediterranean Sea. Mar.
Biodivers. Rec. 3, e6.
Schuchert, P., 2005. Species boundaries in the hydrozoan genus Coryne. Mol. Phylogenet.
Evol. 36, 194–199.
Segura-Puertas, L., Damas-Romero, M., 1997. Variación estacional de la comunidad de
medusas en la laguna Bojórquez, Cancún, México. Hidrobiologica 7, 59–64.
Segura-Puertas, L., Franco-Gordo, C., Suárez-Morales, E., Gasca, R., Godínez-Domínguez,
E., 2010. Summer composition and distribution of the jellyfish (Cnidaria: Medusozoa)
in the shelf area off the central Mexican Pacific. Rev. Mex. Biodivers. 81, 103–112.
Semenchenka, V., Lipinskaya, T., Vilizzi, L., 2018. Risk screening of non-native macro-
invertebrates in the major rivers and associated basins of Belarus using the aquatic
species invasiveness screening kit. Manag. Biol. Invasion 9, 127–136.
Shimomura, T., 1959. On the unprecedented flourishing of ‘Eshizen-Kurage’, Stomolophus
nomurai (Kishinouye), in the Tsushima Warm Current regions in autumn, 1958.
Bulletin of the Japan Sea Regional Fisheries Research Laboratory 7, 85–107.
Stein, M., Marraccini, J.V., Rothschild, N.E., Burnett, J.W., 1989. Fatal Portuguese man-
o'-war (Physalia physalis) envenomation. Ann. Emerg. Med. 18, 312–315.
Tarkan, A.S., Vilizzi, L., Top, N., Ekmekçi, F.G., Stebbing, P.D., Copp, G.H., 2017a.
Identification of potentially invasive freshwater fishes, including translocated spe-
cies, in Turkey using the aquatic species invasiveness screening kit (AS-ISK). Int. Rev.
Hydrobiol. 102, 47–56.
Tarkan, A.S., Sarı, H.M., İlhan, A., Kurtul, I., Vilizzi, L., 2017b. Risk screening of non-
native and translocated freshwater fish species in a Mediterranean-type shallow lake:
Lake Marmara (West Anatolia). Zool. Middle East 63, 48–57.
Thuesen, E.V., Childress, J.J., 1994. Oxygen consumption rates and metabolic enzyme
activities of oceanic California medusae in relation to body size and habitat depth.
Biol. Bull. 187, 84–98.
Turan, C., Gürlek, M., Özbalcılar, B., Yağlıoğlu, D., Ergüden, D., Öztürk, B., Güngör, M.,
2011. Jellyfish bycatch data by puse seine, trawl and net fisheries during
March–April 2011 in the Mediterranean coasts of Turkey, (pp. 1–7) In First National
Workshop on Jellyfish and Other Gelatinous Species in Turkish Marine Waters,
Bodrum, 20–21 May 2011. Turkish Marine Research Foundation, (In: Turan, C.,
Öztürk, B. eds.) (Istanbul, Turkey).
Underwood, A.H., Seymour, J.E., 2007. Venom ontogeny, diet and morphology in Carukia
barnesi, a species of Australian box jellyfish that causes Irukandji syndrome. Toxicon
49, 1073–1082.
Uye, S.I., 2008. Blooms of the giant jellyfish Nemopilema nomurai: a threat to the fisheries
sustainability of the East Asian Marginal Seas. Plankton Benthos Res 3 (Supplement),
125–131.
Verner, B., 1984. Jellyfish flotation by means of bubble barriers to prevent blockage of
cooling water supply and a proposal for a semi-mechanical barrier to protect bathing
beaches from jellyfish. In: Proceedings of the Workshop Jellyfish Blooms, pp. 1–10
Athens. Oct 31–Nov 4 1983.
Vilizzi, L., Copp, G.H., Adamovich, B., Almeida, D., Chan, J., Davison, P.I., Dembski, S.,
Ekmekçi, F.G., Ferincz, Ă., Forneck, S.C., Hill, J.E., Kim, J.E., Koutsikos, N., Leuven,
R.S.E.W., Luna, S.A., Magalhães, F., Marr, S.M., Mendoza, R., Mourão, C.F., Neal,
J.W., Onikura, N., Perdikaris, C., Piria, M., Poulet, N., Puntila, R., Range, I.L.,
Simonović, P., Ribeiro, F., Tarkan, A.S., Troca, D.F.A., Vardakas, L., Verreycken, H.,
Vintsek, L., Weyl, O.L.F., Yeo, D.C.J., Zeng, Y., 2019. A global review and meta-
analysis of applications of the fish invasiveness screening kit. Rev. Fish Biol. Fisher.
29, 529–568.
Walther, G.R., Post, E., Convey, P., Menzel, A., Parmesan, C., Beebee, T.J., Fromentin,
N. Killi, et al.
J.M., Hoegh-Guldberg, O., Bairlein, F., 2002. Ecological responses to recent climate
change. Nature 416 (6879), 389.
Yahia, M.N.D., Yahia, O.K.D., Gueroun, S.K.M., Aissi, M., Deidun, A., Fuentes, V., Piraino,
S., 2013. The invasive tropical scyphozoan Rhopilema nomadica Galil, 1990 reaches
the Tunisian coast of the Mediterranean Sea. Bioinvasions Rec. 2, 319–323.
Yasuda, T., 1988. Unusually gregarious occurrences of jellyfishes in Japanese waters.
Saishu to Shiiku 50, 338–346.
Yemshanov, D., Koch, F.H., McKenney, D.W., Downing, M.C., Sapio, F., 2009. Mapping
invasive species risks with stochastic models: a cross-border United States-Canada
application for Sirex noctilio Fabricius. Risk Anal. 29, 868–884.
Yilmaz, I.N., Isinibilir, M., Vardar, D., Dursun, F., 2017. First record of Aequorea vitrina
Gosse, 1853 (Hydrozoa) from the Sea of Marmara: a potential invader for the
Mediterranean Sea. Zool. Middle East 63, 178–180.
Zenetos, A., Gofas, S., Verlaque, M., Çinar, M.E., Garcia Raso, J.E., Bianchi, C.N., Morri,
C., Azzurro, E., Bilecenoğlu, M., Froglia, C., Siokou, I., Violanti, D., Sfriso, A., San
Martin, G., Giangrande, A., Katağan, T., Ballesteros, E., Ramos-Espla, A.A.,
11
Marine Pollution Bulletin 150 (2020) 110728
Mastrototaro, F., Ocana, O., Zingone, A., Gambi, M.C., Streftaris, N., 2010. Alien
species in the Mediterranean by 2010. A contribution of European Union's Marine
Strategy Framework Directive (MFSD). Part I. Spatial distribution. Mediterr. Mar. Sci.
11, 381–493.
Zenetos, A., Gofas, S., Morri, C., Rosso, D., Violanti, D., Raso, G., Çınar, M.E., Almogi-
Labin, A., Ateş, A.S., Azzurro, E., Ballesteros, E., Bianchi, C.N., Bilecenoğlu, M.,
Gambi, M.C., Giangrande, A., Gravili, C., Hyams-Kaphzan, O., Karacle, P.K.,
Katsanevakis, S., Lipej, L., Mastrototaro, F., Mineur, F., Pancucci-Papadopoulou,
M.A., Espla, A.R., Salas, C., San Martin, G., Sfriso, A., Streftaris, N., Verlaque, M.,
2012. Alien species in the Mediterranean Sea by 2012. A contribution to the appli-
cation of European Union's Marine Strategy Framework Directive (MSFD). Part 2.
Introduction trends and pathways. Mediterr. Mar. Sci. 13, 328–352.
Zenetos, A., Çinar, M.E., Crocetta, F., Golani, D., Rosso, A., Servello, G., Shenkar, N.,
Turon, X., Verlaque, M., 2017. Uncertainties and validation of alien species catalo-
gues: the Mediterranean as an example. Estuar. Coast. Shelf Sci. 191, 171–187.