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Manuscript_60dc489875b2fdfeb350dad6467b8b2c

1 Refugia area for the ctenophore Mnemiopsis leidyi A.

2 Agassiz 1865 in the Berre Lagoon (southeast France): the
3 key to its persistence

4 Guillaume Marchessaux1*, Vincent Faure2, Cristèle Chevalier1, Delphine Thibault1

6 1. Aix Marseille Univ, Univ Toulon, CNRS, IRD, MIO, Marseille, France
7 2. GIPREB Syndicat Mixte, Berre-l’Etang, France
8 * corresponding author: guillaume.gmarchessaux@gmail.com

10 For submission to Progress in Oceanography as an article.

11 15 February 2020

12 Corrected manuscript: 2 June 2020

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© 2020 published by Elsevier. This manuscript is made available under the Elsevier user license
https://www.elsevier.com/open-access/userlicense/1.0/
26 Abstract
27 The invasive ctenophore, Mnemiopsis leidyi has been proliferating in lagoons and
28 coastal areas around Europe for almost 20 years but the role and the impact of its presence
29 in sink ecosystems is still not completely understood. In France, Mnemiopsis leidyi is present
30 in Mediterranean lagoons and estuaries along La Manche Sea. Mnemiopsis was first
31 recorded in the Berre Lagoon in early 2000. This lagoon has been highly perturbated for
32 years, with a large volume of freshwater inflow through natural rivers and a succession of
33 large hydroelectric power plants, inducing important eutrophication. Legislation has been
34 implemented to improve the health status of the lagoon since 1994. A long-term study was
35 undertaken in 2010 to measure Mnemiopsis population dynamics, and to identify the main
36 drivers of its persistence in this highly anthropogenic lagoon. In 2011 and 2012, during
37 extreme winter conditions, populations of this ctenophore were not observed for months. Its
38 re-appearance later in year could be linked to either a new introduction from the
39 Mediterranean Sea or the existence of retention areas where individuals sought refugia.
40 Following measurement of biochemical conditions (i.e. chlorophyll a), plankton biomass and
41 Mnemiopsis populations structure (eggs, cydippid larvae/transitional phase and adult) in
42 different areas of the lagoon, as well as the lagrangian modelling of “particles” distribution
43 (i.e. Ichthyop), we highlighted the seasonal patterns in the population structure, the level of
44 available carbon always above the minimal for the survival of Mnemiopsis (24 µgC L-1) and
45 the potential refugia area role the Vaine sub-basin could play. Populations from the Vaine
46 sub-basin probably serve as source populations for the rest of the lagoon by advective
47 transport in spring.

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49 Keywords: invasive species, Mediterranean lagoon, Lagrangian transport, long-term

50 monitoring.

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59 1. Introduction
60 Non-indigenous species (NIS) constitute a major source of biological pollution, as
61 some become invasive and have strong ecological and economic impacts on biodiversity,
62 ecosystem functioning (e.g. competition, predation) and human activities (e.g. fisheries,
63 industrial complex, and tourism) (Katsanevakis et al., 2014; Tsiamis et al., 2018).

64 Among NIS, the ctenophore Mnemiopsis leidyi is ranked within the top 100 marine bio
65 invaders by Lowe et al. (2007). Mnemiopsis leidyi originates from estuaries and coastal
66 waters along the Americas East Coast from the Chesapeake Bay down to Argentina, as well
67 as in the Gulf of Mexico (Groupe of Experts on the Scientific Aspects of Marine
68 environmental Protection: GESAMP, 1997). This species has spread using ballast waters to
69 Europe, following two main pathways (Reusch et al., 2010; Ghabooli et al., 2011; Bolte et al.,
70 2013; Ghabooli et al., 2013; Bayha et al., 2015; Shiganova et al., 2019). The first one to the
71 Black Sea, connected to the Gulf of Mexico seeding population, occurred in 1982 (Pereladov,
72 1988), the invader then rapidly spread through the whole Black Sea and the Caspian Sea
73 (Kideys and Niermann, 1993, 1994; Vinogradov et al., 1995; Shiganova, 1993, 1998; Ivanov
74 et al., 2000; Bilio et al., 2004; Shiganova et al., 2019) and was found at the turn of the
75 century in various Mediterranean basins (Boero et al., 2009; Galil et al., 2009; Shiganova
76 and Malej, 2009; Fuentes et al., 2009, 2010). The second invasion into the North and Baltic
77 seas was linked to the seeding population of the East Coast of the USA (Faasse and Bayha,
78 2006; Hansson, 2006; Javidpour et al., 2006; Boersma et al., 2007; Janas and Zgrundo,
79 2007; Oliveira, 2007; Tendal et al., 2007; Van Ginderdeuren et al., 2012; Antajan et al., 2014;
80 Shiganova et al., 2019).

81 The presence of this NIS has put pressure on the local environment both at ecological
82 and economic level (Vinogradov et al., 1995; GESAMP, 1997; Purcell et al., 2001a). In fact,
83 M. leidyi presents high reproductive rates (Reeve et al., 1989; Condon and Steinberg, 2008;
84 Jaspers et al., 2015; Marchessaux, 2019), high predation capacities on zooplankton
85 communities (Kremer, 1979; Feigenbaum and Kelly, 1984; Purcell and Decker, 2005;
86 Condon and Steinberg, 2008), and has proven to impact economically both fish and shellfish
87 communities (Purcell, 1991; Purcell et al., 1994).

88 Around 2000, M. leidyi was observed for the first time in several coastal lagoons
89 along the French coast of the Mediterranean Sea: from the Berre Lagoon in the East to the
90 Bages-Sigean Lagoon in the West (Delpy et al., 2016). The Berre Lagoon is the largest (155
91 km2, 0.98 x 109 m3, 7 m average depth) semi-enclosed brackish lagoon in France located in
92 the southeast, near Marseille. It became during the 20th Century a hot spot of petrochemical

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 93 industries, as well as an area of urban expansion and intensive agriculture, which strongly
94 modified the environmental conditions both terrestrial and aquatic. In 1966, EDF (French
95 electricity company) implemented a series of hydro-electric power plants along the Durance
96 canal with the last one located at Saint-Chamas in the North of the lagoon. Freshwater was
97 discharged as needed, and with no control at all into the lagoon (Capuzzo, 1980). The
98 biological conditions of the lagoon quickly deteriorated, high level of eutrophication (Minas,
99 1974, 1976; Caddy, 2000; Gouze et al., 2008; Warner, 2012), significant drops (down to 4) in
100 salinity, and a strong decrease in specific diversity (only 2 species of zooplankton, the rotifer
101 Brachionus plicatilis (Muller, 1786) and the invasive copepod Acartia tonsa (Dana, 1849))
102 were reported for decades (Gaudy and Viñas, 1985; Pagano and Gaudy, 1986; Arfi, 1991;
103 Gaudy, 1992; Gaudy et al., 1995; Cervetto et al., 1999). Actions were then taken to restore
104 the state of the lagoon, with the Barnier Plan in 1994. Then in 2004 a judgment of the EU
105 Court of Justice forced the French Government to put in place its obligations in regards with
106 the state of the lagoon, followed in 2005 by a formal notice including threat of financial
107 penalty. In 2006, the implementation of the new regulation on the release, i.e. freshwater
108 inputs lowered down to 1.2 x 106 m3 y-1 and salinity kept within allocated values and time
109 (being most of the time > 15) was finally put in place. These efforts occurred at the same
110 time as the introduction and development of M. leidyi in the lagoon.

111 Several surveys (i.e. the structure and the functioning of the benthic ecosystem
112 through measurements of the Zostera meadow, and benthic diversity, and abundance; for
113 the pelagic ecosystem nutrients, suspended matters, phytoplankton diversity, and
114 abundance) were then conducted by the GIPREB Syndicat Mixte (local public authority
115 whose goal is to restore the Berre lagoon) to follow the response of the ecosystems to these
116 restrictions (GIPREB pers. com.). Zooplankton community structure and functioning surveys
117 were also undertaken from 2008 onwards. Since 2010, M. leidyi dynamics were also studied
118 at the three stations (SA1, SA2 and SA3) located in the main section of the Berre Lagoon.
119 Seasonal patterns in terms of abundance and community structure indicated that
120 temperature influenced the life cycle of M. leidyi with cydippid larvae observed in winter,
121 transitional stages in winter/spring and adults from spring to autumn. About abundances, no
122 clear differences were reported for these 3 stations (Delpy et al., 2012), but strong temporal
123 variations with no specimen being sampled for several weeks and months after a strong cold
124 event (T < 6 °C) were reported in 2011 and 2012. Reappearance of Mnemiopsis within the
125 main sector of the lagoon could be either linked to a seeding population from outside the
126 lagoon or from specific areas within the lagoon. We decided then in October 2015 to add
127 new sampling locations giving a better coverage of the different environmental conditions
128 existing within the lagoon to try to clarify and understand Mnemiopsis population connectivity

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129 and potential locations of seeding population. To reach this goal we propose here an
130 approach combining a Lagrangian particle transport model (using Ichthyop) coupled with in
131 situ measurements.

132 2. Materials and methods

133 2.1. Study area
134 The study area is composed of two sectors: the Berre main lagoon (maximum depth
135 9m) and the Vaine sub-basin (maximum depth 5m) to the east separated by a sand shoal (~
136 1.5 m). Inflows of freshwaters are all located in the northern part of the main basin with two
137 small rivers, the Touloubre, and the Arc (Figure 1), and the large industrial channel of the
138 Durance supplying several hydroelectric power plants with the last one located on the shore
139 of the lagoon in the vicinity of the city of St-Chamas (4 km from the town center). The lagoon
140 is connected to the Mediterranean Sea in the southwestern part through the Caronte
141 channel. Low semi-diurnal tides (~ 30 cm) are observed, but the water level is mainly
142 affected by barometric conditions or large release of freshwater (Alekseenko et al., 2013a,b).

143 Bi-monthly samplings were conducted at SA2 (from January 2010 onwards), then
144 from 16 December 2016 to 27 September 2017 at an additional four coastal stations in the
145 Berre basin (from North to South Istres, EDF, Arc and 3 frères), one in Vaine sub-basin
146 (Rognac) and one over the sand shoal separating the two basins (Anse de Berre) (Table 1).

147

148 2.2. Environmental parameters

149 Meteorological data (wind direction and speed) were measured at the Météo France
150 station in Marignane every 3 hours between 2010 and 2017.

151 Inflows of freshwater (m3 s-1) coming through the Power plant were provided by the
152 EDF (French Electricity Company). For rivers (Arc and Touloubre), inflows (m3 s-1) were
153 extracted from www.hydrofrance.fr.

154 Environmental parameters (temperature and salinity) were measured from 2010 to
155 2016 by EDF company in continuous at SA2 from 5 moored Seabird® SBE37 CTD located at
156 1 m, 4 m, 5.3 m, 6.3 m, and 7.3 m respectively. In 2016-2017, environmental parameters
157 (temperature, salinity, fluorescence) were recorded with a CTD probe (Seabird® 63) handled
158 from bottom to the top for each station.

159 Water samples were collected at the surface with a 5 L sampling bottle, then
160 transferred into 2 L plastic containers and kept in the dark. Volumes of 25 mL or 50 mL of
161 water were filtered separately onto 10 µm and 3 µm polycarbonate filters and then GF/F.

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162 Filters were then frozen (- 20 °C until further analysis). Chlorophyll a was extracted using
163 acetone (90 %), in the dark and at – 20 °C overnight. Chlorophyll a was measured by
164 fluorimetry (Turner Trilogy®) following the acidification method according to Yentsh and
165 Menzel (1963), modified by Holm-Hansen et al. (1965). Phytoplankton biomass was
166 estimated in terms of carbon by applying a conversion factor of 50 to the chlorophyll a
167 concentration (Banse, 1977; Bamstedt et al., 2000).

168

169 2.3. Zooplankton

170 Zooplankton was collected using a modified conical plankton net (80 µm mesh size, 1
171 m long and 50 cm opening) mounted with a flowmeter. A vertical tow was performed over the
172 whole water column at a speed of ~ 1 m s-1. Samples for taxonomic identification were
173 preserved in buffered sea water-formalin solution (4 % final concentration). After
174 homogenization, a sub-sample (2 % to 10 %) was taken using a micropipette and placed in a
175 Dollfuss dish and counted under a stereomicroscope (Leica M165-C® stereomicroscope).
176 The different taxa are identified down to genus or species levels when possible using
177 reference books (Rose, 1933; Trégouboff and Rose, 1957; Razouls et al., 2010-2018) and
178 validated against World Register of Marine Species (WoRMS, 2019). The abundance of each
179 taxon (individuals m-3; ind m-3) was estimated from the counts, the fraction counted, and the
180 volume of water sampled by the net. Main taxa were also measured, and their carbon mass
181 determined according to allometric equations (Marchessaux, 2019) literature. Individual taxa
182 carbon content was multiplied by the number of individuals of each taxa collected per net.
183 Total biomass (µgC L-1) of zooplankton was calculated then by summing all the individual
184 taxa biomass.

185 The planktonic biomass was estimated at only SA2 from net tows. A modified conical
186 net (80 µm mesh size, 1 m long and 50 cm opening) was deployed vertically between the
187 bottom and surface at a speed of ~ 1 m s-1. The sample volume (m3) by the net was
188 calculated from the depth of the sampling and the opening area of the net. The content of the
189 net’s collector was collected in a bottle containing a large volume of filtered lagoon water and
190 kept cool and dark until return to the laboratory. The plankton was then fractionated into size
191 classes using a sieve column (80 µm, 200 µm, and 500 µm). After rinsing with fresh water to
192 remove as much salt as possible, the contents of each sieve were recovered on a pre-
193 weighed filter (Labover Precisa 40SM-200A® microbalance, accuracy ± 1 mg) and pre-
194 calcined (450 °C for 4 h). The filters were then placed in an oven (60 °C) for 72 h and
195 reweighed. The planktonic biomass (mgDW m-3; DY: Dry Mass) was calculated from the

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196 difference in mass between the filter+plankton mass and the initial mass of the filter, relative
197 to the volume of water sampled.
198

199 2.4. Mnemiopsis leidyi abundance

200 Individuals of Mnemiopsis leidyi were collected with a modified Regent net (700 µm
201 mesh size, 2 m long and 75 cm opening area). The net was towed horizontally at the surface
202 for 2 to 5 min at ~ 2 knots. Sampled volume was recorded using a flowmeter. The total
203 number of Mnemiopsis was counted and abundance (ind m-3) estimated from the volume
204 sampled by the net.

205 Eggs and larvae of Mnemiopsis were sampled using a conical net mounted with a
206 200 µm mesh size, and with a 30 cm opening diameter hauled through the water column.
207 Young stages of Mnemiopsis were difficult to count on site but as they were largely degraded
208 by formalin solution, individuals were then fixed with acid Lugol (3 %) (Gorokhova and
209 Lehtiniemi, 2009; Sullivan and Gifford, 2009). In the laboratory, sub-samples (2 % to 50 % of
210 the total sample) were analysed and the number of eggs and larvae (cydippid and
211 transitional stages) counted. Abundances (ind m-3) were then calculated using the water
212 volume sampled.

213

214 2.5. Hydrodynamic modelling

215 The Ichthyop tool developed by Lett et al. (2008) was used to understand the role of
216 hydrodynamic forcing (winds, currents, temperature, salinity) on the spatial and temporal
217 distribution of Mnemiopsis leidyi. Thus, particles (here M. leidyi) moved in the environment
218 according to a Lagrangian model based on hydrodynamic circulation scenario defined by
219 Noveltis (Hydrodynamic modelling of the Berre Lagoon and the associated environments;
220 reference: NOV-7272-NT-4156) integrated by GIPREB in the TELEMAC 3D model. This
221 scenario was mainly discriminated by winds direction and speed.

222 The GIPREB configuration of TELEMAC3D (version v6p2) was forced by atmospheric
223 conditions (winds, pressure, humidity, and temperature), astronomic tides, freshwater inflows
224 by rivers, Wastewater Treatment Plant (WWT) inflows, and the hydroelectric power plant
225 characteristics (both in terms of flows and temperature). The open boundary conditions
226 (temperature, salinity and water level) were located between the Mediterranean Sea and the
227 Gulf of Fos, as to force the water level with the available data from the in-situ tide gauge (in
228 the Caronte Canal). The water column was divided into 25 meshes based on hybrid sigma
229 coordinates (Figure 2): 20 planes positioned according to depth (z, between bottom and
230 surface) and 5 sigma planes which were distributed along the depth in function of the

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231 percentage of the water thickness between -2m and the surface. Hydrodynamics circulation
232 was calculated on an unstructured grid of TELEMAC3D (resolution from 4 m to 1500 m, 4128
233 nodes). However, the 3D simulation outputs were interpolated for each depth on a regular
234 grid of 100m resolution before integration in Ichthyop.

235 Ichthyop was forced by outputs originating from a non-hydrostatic simulation based
236 on the whole year 2007 (time step of the outputs is one hour). For these simulations, forcing
237 data were realistic (i.e. measured data) except for salinity and temperature at the open
238 boundary conditions (coming from the outputs of a simulation from the SYMPHONIE model).
239 Four weather scenarios were included in our analysis (Table 2) based on the most frequent
240 wind conditions.

241 The lagoon was then divided into five (Figure 3) according to their hydrological
242 conditions. Area 1, the northernmost area, presented a semi-permanent eddy (Alekseenko et
243 al., 2013a,b) and was influenced in its South-Eastern part by the freshwater plume of the
244 hydroelectric power plant. Area 2 was directly under the influence of the EDF plant and the
245 Arc inflows. Area 3 has an intermediate position influenced by both the freshwater plume
246 from area 2 and the seawater signature at depth from the Caronte Canal). Area 4 was
247 characterized by seawater input (South-West) and the maximum bottom depth (max 9 m).
248 Finally, Area 5, the Vaine sub-basin, was physically separated from the main lagoon by a
249 shoal and its water circulation follows a semi-permanent eddy (Alekseenko et al., 2013a,b).
250 In Ichthyop, 1,000 particles (considered to be enough to characterized transport
251 dynamics) were integrated into the model, distributed between the bottom and the surface at
252 each station. The duration of the simulation was set to three days with a time step of 10
253 minutes. The outputs were in NetCDF format and contain GPS coordinates and depth of
254 each point at each time step. NetCDF files were integrated into R Studio.

255

256 2.6. Data analysis

257 Tree-dimensional plots for temperature, salinity, chlorophyll a concentrations and
258 winds were made with OceanDataView using the Diva gridding method.

259 The trajectories of particles were extracted from NetCDF files with R studio and were
260 projected on a map of the lagoon using QGIS Lyon 12.5. From the model outputs, the
261 percentage contribution of the particles was defined in each area between time tstart and tend
262 then connectivity matrixes were made using R studio.

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263 To test the significance of differences, Analyses of Variance (ANOVA) were
264 calculated using SigmaPlot 12.5 software. If the overall ANOVA results were significant,
265 Bonferroni pair-wise comparisons (p < 0.05) were performed to test among combinations.

266 In order to link the in-situ dynamics of the zooplankton community and Mnemiopsis
267 leidyi with environmental data (biotic and abiotic), a principal component analysis (PCA) was
268 performed using the Primer software. The data matrix includes environmental data
269 (temperature, salinity, freshwater flows, chlorophyll concentration) and abundance data for
270 zooplankton and Mnemiopsis leidyi (adults, larvae and eggs). After transformation of the data
271 into log(n+1), the latter were normalized (Shapiro-Wilk test). The PCA was thus traced as
272 well as the circle of correlations allowing to identify the factors influencing the planktonic
273 dynamics of Berre Lagoon.

274

275 3. Results
276 3.1. Environmental parameters
277 3.1.1. Wind conditions
278 The Berre Lagoon is subject to mainly 3 wind regimes: the Mistral, the Ponant, and
279 the Marin (Supplementary figure 1). The Mistral, which dominates the region (59.7 ± 8.5 % of
280 the year with maximum speeds of 22 m s-1), is a cold and dry North-West wind (320° - 360°).
281 The Ponant, hot and dry, is in the West sector (270°) while the Marin, is a gentle and humid
282 wind, blowing from the South-East (120° - 165°).
283 The intensity of the Mistral shows strong seasonal variability (S1) with high speeds in
284 winter (19.4 ± 1.9 m s-1) and autumn (17.3 ± 1.3 m s-1) while in spring and summer the
285 Mistral only blows at 7.8 ± 0.5 m s-1 and 7.1 ± 0.8 m s-1 respectively.
286 The Marin and the Ponant represent respectively; 5.9 ± 2.1 % and 11.9 ± 1.5 % of the
287 annual winds. They are weaker winds: 3.3 ± 0.5 m s-1 and 3.1 ± 0.9 m s-1 respectively. The
288 Marin shows a higher intensity in January (10.4 ± 0.4 m s-1) and July (10.7 ± 1.9 m s-1) and
289 the Ponant in summer (5.5 ± 2.1 m s-1).
290

291 3.1.2. Temperature and salinity

292 The temperature of the Berre Lagoon, between 2010 and 2017, followed a classic
293 seasonal cycle of a temperate zone (Supplementary figure 2) with surface temperature often
294 below 6 °C in January-February, and as low as 0.3 °C on 13 February 2012 (surface layer
295 was frozen around the edge of the lagoon). Maximum temperatures usually above 24 °C
296 were found in July-August, reached up to 26 °C in August 2012. At SA2, no vertical

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297 stratification was observed (Figure 4). The vertical distribution of temperature over the six
298 coastal stations also did not show any pattern. The highest temperature was observed in
299 Rognac station (16.8 ± 7.2 °C) compared to the other stations (16.5 ± 6.6 °C) of the lagoon
300 (ANOVA, Bonferroni test, p < 0.05) was recorded.
301 The salinity at SA2 appeared very variable (Supplementary figure 3). Minimum
302 surface salinity was measured in February-March 2017 (S = 15.6) while highest value was
303 observed in September 2012 (S = 29.5). Large freshwater inflows signature from the rivers
304 (Arc and Touloubre) and the EDF powerplant were observed punctually as far as SA2 where
305 salinity reached ~ 15 all the way down to 6 m during winter 2010 but only impacted the top
306 meter in January-February between 2014 and 2017. This signature spread impacted also the
307 Istres station where surface salinities were significantly lower (ANOVA, Bonferroni test, p <
308 0.05) from February to April than during the rest of the year (Supplementary figure 3).
309 Bottom layer (7.3 m) salinity displayed a marked signature of marine entrances (S >
310 25 in bottom) throughout the year with highest values (S > 30) recorded between spring
311 (April-May) and late summer (September-October). No specific pattern in the vertical
312 distribution of the salinity was observed at the shallow stations. The Vaine sub-basin
313 appeared less affected by freshwater inputs than the Berre basin (ANOVA, Bonferroni test, p
314 < 0.05).
315 Mean winter surface values ranged from 17.3 ± 2.9 (February 2017) to 20.7 ± 3.5.
316 (January 2017) while a slight increase significantly occurred towards summer (24.0 ± 1.3 in
317 July 2017 to 28.2 in September 2017) in the Berre basin.
318

319 3.1.3. Chlorophyll a concentration

320 The chlorophyll a concentration was highly variable: from 0.19 µg Chla L-1 (January
321 2017) to 37.1 µg Chla L-1 (November 2016) (Supplementary figure 4). A phytoplankton bloom
322 (> 10 µg Chla L-1) was generally observed at the surface in autumn over the entire Berre
323 Lagoon. Although no significant difference was observed between the bottom and surface
324 layers (ANOVA, p > 0.05), they presented two distinct situations. At the surface, 2011 was
325 marked by a summer peak (up to 17.5 µg Chla L-1) and even larger autumn one (up to 25 µg
326 Chla L-1), whereas in 2016 only one maximum was reported in October-November (20 µg
327 Chla L-1). However, in autumn 2017 phytoplankton maximum was located at the bottom in
328 Istres (40 µg Chla L-1) and SA2 (15 µg Chla L-1).

329

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330 3.1.4. Planktonic biomass
331 The planktonic biomass comprises an assemblage of phytoplanktonic cells and
332 zooplanktonic organisms making up the seston (Figure 4). Silt and sediments (< 60 µm) were
333 removed after sieving the sample. The biomass ranged from 5.8 mgDW m-3 to 386.9 mgDW
334 m-3 in general, except for 2 samples with values well above 500 mgDW m-3 (658.0 mgDW m-3
335 in November 2011, 627.9 mgDW m-3 in January 2014). The planktonic biomass was highly
336 variable and did not show a trend (Figure 4A). Between January 2015 and September 2017,
337 planktonic biomass was dominated by middle size classes (200-500 µm and > 500 µm)
338 organisms (Figure 4B) representing 36.0 ± 16.6 % and 53.8 ± 20 % of the total biomass,
339 respectively. The 200-500 µm fraction dominated the planktonic community between
340 February (47.7 %) and June (63.6 %), while the > 500 µm fraction was mainly dominant
341 between August (42.2 %) and December (79.4 %). On average, the 80-200 µm fraction
342 contributed only 10.1 ± 8.0 % and ranges from 4.3 % (November) to 25.6 % (July).

343 Between 2010-2017, the estimated carbon plankton biomass at SA2 ranged from
344 12.4 µgC L-1 (January 2017) to 1866.4 µgC L-1 (November 2016) and was highly variable
345 (Supplementary figure 5). For the six coastal stations, values ranged from 1.0 µgC L-1 (3
346 frères station, August 2016) to 1 189.4 µgC L-1 (Anse de Berre station, September 2017)
347 (Supplementary figure 5). On average Istres stations presented the highest values (292.4 ±
348 205.5 µgC L-1 and 207.0 ± 196.6 µgC L-1 respectively). Phytoplankton carbon estimated from
349 the chlorophyll:carbon ratio of 50 contributed to 44 % to 99 % of the total planktonic carbon
350 biomass while zooplankton contributed from 1 % to 77 %.

351

352 3.2. Zooplankton dynamics

353 Zooplankton from SA2 showed a high temporal variability between 2010 and 2017
354 (Figure 5), with abundance ranging from 173 ind m-3 (December 2013) to 159 280 ind m-3
355 (July 2015). Zooplankton abundance was significantly (ANOVA, Bonferroni test, p < 0.05)
356 higher in spring (38 546 ± 21 338 ind m-3) and summer (23 017 ± 16 305 ind m-3) than in
357 autumn (13 407 ± 9 008 ind m-3) and winter (14 663 ± 8 015 ind m-3).

358 The zooplankton along the lagoon (6 stations) appeared relatively homogeneous
359 spatially (Figure 9) and showed the same temporal pattern than SA2: October-December :
360 4 073 ± 3 031 ind m-3; January-March: 6 483 ± 2 754 ind m-3; April-June: 4 635 ± 3 900 ind m-
3
361 ; July-September: 19 921 ± 7 718 ind m-3. In May 2016, an abundance seven times higher
362 than at the 6 other station (average 14 428 ± 6 661 ind m-3) was observed at the Arc station
363 (100 990 ind m-3), linked to an explosion in the Acartia tonsa population representing 95 % of
364 the zooplanktonic community (96 181 ind m-3).

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366 3.3 Mnemiopsis leidyi population dynamics

367 At SA2 Mnemiopsis leidyi adult’s population displayed a high temporal variability but
368 with no apparent seasonality (Figure 5). However, high abundances were observed in
369 September 2010 (20 ind m-3), July 2011 (11 ind m-3) and July 2017 (44 ind m-3). On the other
370 hand, Mnemiopsis was not collected at SA2 station at several occurrences between March
371 and May 2012, February-March 2013, June-July 2013, again July 2014 and May-July 2016
372 (Table 3). In 2016, even though Mnemiopsis was not reported at SA2, it was present at the
373 shore stations (Figure 6), where Mnemiopsis leidyi abundance was very variable. Rognac
374 showed the highest (ANOVA, Bonferroni test, p < 0.05) abundances observed during this
375 study with 940 ind m-3 in July 2017 and the highest annual average: 38 ± 177 ind m-3 with
376 Mnemiopsis reported at each sampling date. It is important to signal that in July-August
377 2016, Mnemiopsis were present at the Rognac station in very low abundance but were too
378 large (~ 15 cm total length) to be accurately sampled.
379 In the Berre basin, mean annual M. leidyi abundances at the different stations were
380 within the same range: Arc: 1 ± 2 ind m-3; EDF: 2 ± 7 ind m-3; Istres: 2 ± 7 ind m-3; 3 frères: 1
381 ± 3 ind m-3. Maximum abundances were observed in July 2017 with 35 ind m-3and 38 ind m-3
382 respectively at EDF and Istres.
383 Population dynamics of Mnemiopsis within the whole system showed that eggs were
384 mainly produced during autumn and winter, dominating then the population, while larvae
385 were more abundant in spring, and adults were present with high abundances during
386 summer and autumn.

387 The annual average abundance of eggs ranged from 114 ± 242 ind m-3 to 5 626 ± 29
388 107 ind m-3, and was significantly higher in 2016 (3 891 ± 18 947 ind m-3) than in 2017 (19 ±
389 41 ind m-3) (ANOVA, Bonferroni test, p < 0.05). In winter, highest eggs abundances (ANOVA,
390 Bonferroni test, p < 0.05) were observed in the Vaine sub-basin (140 ± 302 ind m-3) and near
391 freshwater sources (Arc: 105 ± 277 ind m-3; EDF: 113 ± 316 ind m-3) (Figure 6).

392 Larval stages (cydippid and transitional stage) overall abundances were comprised
393 between 37 ± 93 ind m-3 and 470 ± 1 291 ind m-3 and showed a spatial pattern (Figure 6). As
394 observed for eggs, in winter, larval abundance was highest in the Vaine sub-basin (97 ± 210
395 ind m-3), increasing throughout spring (357 ± 724 ind m-3). At that time population found near
396 freshwater sources was high too (Arc: 608 ± 680 ind m-3; EDF: 153 ± 249 ind m-3). High
397 levels of larvae were also reported in autumn, at Arc and EDF stations (33 ± 51 ind m-3 and
398 15 ± 8 ind m-3 respectively) while none or very low numbers were observed at the other

12
399 stations (Anse de Berre: 0 ind m-3; Istres: 1 ± 1 ind m-3; SA2: 6 ± 9 ind m-3; 3 frères: 8 ± 5 ind
400 m-3; and Rognac: 6 ± 5 ind m-3).

401 The PCA run on the data from 2016-2017, highlighted the seasonality of the
402 Mnemiopsis leidyi population (Figure 7), with the first axis explaining 41.5 % and the second
403 axe 23.4 % of the variability. Axis 1 shows a gradient between winter (high freshwater inflows
404 through the EDF powerplant, low salinity, and low Mnemiopsis abundances) and summer
405 conditions (hight salinity and high Mnemiopsis abundances). The second axis placed in
406 opposition spring-summer conditions 2016 with those of 2017. Effectively, 2017 was
407 characterized by a large bloom in spring-summer while in 2016 zooplankton abundances
408 were highest with low Mnemiopsis abundance.

409

410 3.4. Dispersion dynamics of Mnemiopsis leidyi: modelling approach with

411 the Ichthyop tool
412 3.4.1. Hydrodynamics of the Berre Lagoon
413 We considered here that the hydrodynamics in this small almost closed basin was
414 essentially subject to wind forcing. When the wind is blowing from the North-North-West
415 (Mistral regime: Figure 8), surface water is pushed towards the South section of the lagoon,
416 with the highest current speed being located along the eastern shore (between 0.25 and 0.5
417 m s-1). The surface water layer then dives when reaching the South shore of the main basin
418 initiating a bottom counter current. Finally, in the North, the circulation follows the rise of the
419 bottom depth. In the Vaine sub-basin, clockwise current was observed at the surface and a
420 deep anticlockwise current was created. Highest current values (0.45 m s-1) were recorded
421 under strong Mistral, while value up to only 0.25 m s-1 were observed under moderate
422 Mistral.

423 The light West-North-West wind regime (Figure 9) corresponds to wind shift period
424 with weak currents speed. Then, differences between surface and bottom circulation was
425 less clear. In the Vaine sub-basin, surface current was clockwise in contrast to the bottom
426 current direction which was opposed; both showed low current speed (< 0.1 m s-1).

427 Under moderate South-East wind conditions, the surface water mass is pushed
428 North-Westward along the West and East sides of the Berre lagoon where currents are the
429 strongest (~ 0.3 m s-1) of the lagoon and then plunges to depth (Figure 10). The water moves
430 then southward above the bottom, creating a counter-current in the middle of the Berre
431 basin. In the Vaine sub-basin, an anticlockwise surface current (0.1 m s-1) was observed.

432

13
433 3.4.2. Hydrodynamic circulation forcing the distribution of Mnemiopsis
434 leidyi
435 The release of particles from the 7 stations localised around the lagoon allowed to
436 follow their dispersion in detail over a 3 days period.

437

438 3.4.2.1. Particle transport dynamics under Mistral regime

439 Under Mistral conditions (Figures 11 and 12), particles released in the North of the
440 lagoon (Arc and EDF stations) followed, at the surface, the east coast of the Berre basin and
441 dive down to southward along a surface current.

442 Under moderate Mistral, particles released at Istres station at the surface followed the
443 coast to the South (16 %) but those released deeper were caught in an eddy and remained
444 in the same area slightly North of the station (84 %). Particles from the southern station (3
445 frères) whatever the depth of release were caught by the counter-current and reached the
446 Northern section of the lagoon (Area 1: 17 %; Area 2: 10 %; Area 3: 32 %) while 41 %
447 remain in their start area. All particles released from Area 2 (EDF and Arc) were carried
448 South East by the surface current (62 % in Area 4). A gyre located within the Vaine sub-
449 basin (Rognac station) was observed forcing 100% of the particles to be retained within.

450 In strong Mistral conditions (Figure 12) particles dispersion follows the same pattern
451 but over a wider area than under moderate mistral due to highest current speeds (Figure 14).
452 For Area 2, 51 % of the particles of the Arc/EDF stations reached SA2 station and covered
453 22 km, and 37 % reached to the South of the Berre basin (21 km covered). The distance
454 covered by particles from Area 2 under strong mistral regime was on average 12 km longer
455 than under moderate mistral regime (10 km). The same trend was observed for the other
456 stations. The momentum of the water dynamics in the Vaine sub-basin allowed 7 % of the
457 particles to overflow into the Berre lagoon to reach the 3 frères station. Area 1 (Istres station)
458 retained 71 % of particles due to eddy and 14 % reached the Berre basin South.

459

460 3.4.2.2. Particle transport dynamics under light West-North-West regime

461 Under the weak West-North-West wind regime considered here, dispersion was partly
462 similar to than under moderate mistral conditions but with a higher dispersion (Figure 13). Of
463 the 1000 particles released at Arc/EDF 53 % moved to the South (Area 4) with 3 %
464 overflowing into the Vaine sub-basin. Particles from Istres remained mainly within the Area 1
465 (80 %) with only 4 % drifting as far South as the Canal Caronte (Area 4). Particles from 3
466 frères stations showed higher dispersion over the entire Berre basin than over the other

14
467 stations (SA2, Arc, EDF) but did not spill over into the Vaine sub-basin. A total of 5 3% of
468 particles from 3 frères were retained in the Area 3.

469

470 3.4.2.3. Particulate matter transport dynamics under moderate South-

471 East regime
472 Particles released from the stations EDF/Arc moved to the North section of the Area 1
473 (74 % of particles). From SA2 (Area 3), 48 % of particles moved to the Area 2 North-
474 Westward at the surface with deeper layer particles following the bottom under-current
475 (Figure 14) and 23 % moving to the South (Area 4). Particles released at Istres showed a
476 very limited dispersion: 80 % of particles were retained and 6 % moving to Area 2 and 11 %
477 to Area 4. From 3 frères, 47 % of the particles were retained in the South of the Berre basin,
478 52 % moved to SA2 and just 1 % moved into the Vaine basin.

479 4. Discussion
480 High egg and larvae abundances near desalinated areas have already been
481 observed by Condon and Steinberg (2008), while for Lehtiniemi et al. (2007) low salinities
482 inhibit the expansion of the Mnemiopsis leidyi population. In the Berre basin, the different
483 areas near freshwater sources seemed to be favourable to the maintenance of M. leidyi.
484 During monthly monitoring between 2010 and 2015 at SA2, stations in the Berre basin
485 were characterized by low abundances or seasonal extinctions (reductions to levels below
486 detection limits) during very cold winters (T < 6 °C) as observed in 2012 and
487 2013. Mnemiopsis was absent from the SA2 samples in March-May 2012, February-March
488 and June-July 2013, and again in July 2014 and between May-July 2016 (Table 3).
489 Therefore, the absence of Mnemiopsis in samples during cold events and its reappearance
490 several months later suggested either a source outside the lagoon or the presence of a
491 refugia area within the whole lagoon system. The following criteria have been defined by
492 (Costello et al., 2006) for potential refugia area for Mnemiopsis overwintering:

493 • Hydrodynamics must be limited (no strong currents) and must play a role as a
494 concentrator of Mnemiopsis leidyi,
495 • The potential source area must have small variations in salinity,
496 • The potential source area must have enough zooplankton production to maintain M. leidyi
497 all year.
498 • The population of Mnemiopsis leidyi must survive (egg laying, growth) regardless of
499 seasonal cycles,
500

15
501 During our study, abundance of Mnemiopsis leidyi was significantly higher in the
502 Vaine sub-basin with all the other parameters in agreement to those defined for a refugia
503 area. In winter, the limitation of reproduction by low temperatures and high rates of water
504 turnover inevitably leaded to a decline in abundance to undetectable levels and/or extinction
505 of the population in the Berre basin.

506 Distribution and presence of M. leidyi appeared strongly linked with the wind patterns.
507 In fact as a planktonic species with very limited swimming capacity, organisms distribution
508 followed that of the water circulation, largely determined by winds and freshwater inputs
509 (Alekseenko et al., 2013a,b) characterized by high-speed reverse currents near the coasts.
510 In the Black Sea, Kubryakov et al. (2016) have already demonstrated the role of currents in
511 the distribution of this species, among others. The sudden arrival of adults in the Berre basin
512 in spring and summer suggests that the Vaine sub-basin, a refugia area for Mnemiopsis, was
513 a source of populations from which convective transport, particularly under conditions of
514 strong spring Mistral, provides the inoculum for the other regions of the Berre Lagoon. The
515 development of Mnemiopsis could be not possible according to environmental conditions
516 (temperature and food) in the Berre basin.

517 As observed in Narragansett Bay in winter (Pilson, 1985; Costello et al., 2006),
518 Mnemiopsis recovered in the Berre basin were ephemeral due to a limited reproductive
519 potential and high rates of water turnover (between 90 and 200 days).

520 Indeed, retention times most directly affected the winter persistence of the population
521 in an area and it was population persistence that primarily distinguishes the source habitat in
522 the Vaine sub-basin from the sink habitats in the Berre basin. If we assumed that Vaine sub-
523 basin was a source area for the lagoon, we looked at the conditions occurring before the re-
524 appearance at SA2. The Vaine sub-basin was used to reseed Mnemiopsis in the Berre
525 basin. From measurements made with Ichthyop and the percentages of connectivity between
526 the different areas of the Berre Lagoon, we were able to establish predictive maps of M.
527 leidyi reseeding in the lagoon. After identifying the periods of M. leidyi disappearance at SA2,
528 we looked at the wind conditions 3 days before the reappearance of M. leidyi to identify the
529 meteorological events allowing the return of Mnemiopsis to the Berre Lagoon
530 (Supplementary figure 6). For the seven periods of Mnemiopsis disappearance at SA2, we
531 identified wind conditions favourable to the reseeding of M. leidyi in the Berre basin. For each
532 period, strong Mistral or Marin winds conditions were identified, conditions that allowed the
533 export of M. leidyi from the Vaine sub-basin to the Berre basin. For example, before the
534 reappearance of Mnemiopsis at SA2 on May 10, 2012, Marin Wind conditions were observed
535 on May 8 and 9, 2012 allowing the export of Mnemiopsis leidyi from the Vaine sub-basin to

16
536 the Berre basin as identified in our measurements with Ichthyop. Following the
537 disappearance of M. leidyi in April 2013, we also identified strong Mistral conditions 3 days
538 (April 20, 2013) before the reappearance of Mnemiopsis at SA2 on April 23, 2013. Finally,
539 Mnemiopsis was absent from the samples on 5 July 2016 at SA2 while it was observed in the
540 Vaine sub-basin at the Rognac station on the same day. Following a Marin wind event on
541 July 17, 2016, Mnemiopsis was observed at SA2 on July 18, 2016. Thus, for each
542 disappearance of Mnemiopsis at SA2, we were able to show that in the 3 days before its
543 reappearance at SA2, strong Mistral and/or Marin winds conditions allowed the export of M.
544 leidyi from the Vaine sub-basin to the Berre basin. In fact, these observations illustrated that
545 the Vaine sub-basin was a real refugia area for Mnemiopsis leidyi.

546 Hydrographic features were likely the critical variables distinguishing source habitats
547 within M. leidyi metapopulations living in temperature regions characterized by cold winter
548 temperatures (< 8 °C). Temperature and food availability are likely to determine the role of
549 the Vaine sub-basin as a source region in the M. leidyi population of the Berre Lagoon.
550 Minimum winter temperatures (min: 3.5 °C; max: 14.8 °C; mean: 9.3 ± 3.4 °C) were
551 favourable for M. leidyi in the Vaine sub-basin, unlike the Berre basin stations (min: 3.0 °C;
552 max: 13.8 °C; mean: 8.9 ± 2.7 °C) which was 1°C warmer than Berre basin. Potential refugia
553 were highlighted, Areas 1 and 5, which acted as particle concentrators. Winter temperature
554 did not fall below 8.7 °C (15 February 2017) except for 1 day in 19 January 2017, T = 3.5
555 °C). Even during cold winters (2016-2017) with temperatures close to or equal to 3.5 °C (19
556 January 2017), M. leidyi was found in the Vaine sub-basin at all dates. In addition, the carbon
557 concentration available for M. leidyi showed less variability in Area 5 than in Area 1, provided
558 enough food to maintain Mnemiopsis. On the other hand, Marchessaux (2019) observed in
559 the laboratory the production of Mnemiopsis eggs at temperatures greater than or equal to 8
560 °C while Jaspers et al. (2014) and other references mentioned no reproduction below (12 °C
561 for Purcell et al., 2001a). Spring earlier warming in Vaine sub-basin allowed the population to
562 grow and reproduce weeks earlier than in the main basin. Once temperature control (19 °C)
563 was reached by spring warming, M. leidyi feeding (Reeve and Walter, 1978; Kremer, 1979)
564 and reproductive capacity (Kremer and Nixon, 1976; Marchessaux, 2019) allowed rapid
565 colonization and population expansion throughout the entire lagoon.

566 Reproduction and population persistence were supported by enough dietary carbon
567 available throughout the year (Jaspers et al., 2014). In fact, eutrophication maintained a high
568 production of phytoplankton with a significant contribution (up to 70 % to total planktonic
569 carbon biomass) and enough zooplankton biomass for M. leidyi to thrive (Deason, 1982;
570 Gavini et al., 2013; McNamara et al., 2014; Vansteenbrugge et al., 2016). In the Vaine sub-
571 basin, the available carbon concentration was always above 24 µgC L-1, then largely enough

17
572 for the physiological needs of Mnemiopsis even in winter. Kremer (1994) described the
573 relationship between temperature and food availability models and large-scale seasonality
574 models for M. leidyi. For example, it was only when temperatures exceeded the 8 °C
575 threshold for egg production (Marchessaux, 2019) that populations increased. The relative
576 benefits enjoyed by M. leidyi populations in the Vaine sub-basin were not limited in winter
577 and spring. Because M. leidyi is omnivorous, it can ingest ciliates, diatoms and
578 dinoflagellates (Deason and Smayda, 1982; Vansteenbrugge et al., 2016) as well as larger
579 prey. As well, reproduction decreased and overall abundances decreased at all stations in
580 the autumn, when zooplankton availability and temperature decreased. Nevertheless, It has
581 been demonstrated that in the lagoon, transitory larvae block their growth at temperatures
582 below 14 °C (Marchessaux, 2019), which could explain their dominance in the population
583 during winter (57 ± 39 %; T = 8.9 ± 2.8 °C) as observed by Reeve et al. (1989).

584 In fact, it can therefore be assumed that the Vaine sub-basin (Area 5) is a refugia
585 area for Mnemiopsis leidyi (Figure 15). This area has all the environmental (low temperature
586 variations: min: 3.5 °C; max: 27.1 °C; mean: 20.7 ± 5.6 °C; low salinity variations: min: 18.7;
587 max: 27.8; mean: 22.1 ± 2.6) and biological characteristics (enough carbon available: min:
588 2.01 µgC L-1; max: 854.5 µgC L-1; mean: 207.0 ± 196.58 µgC L-1) favourable to the
589 maintenance of M. leidyi, especially during the winter, as described by Costello et al. (2006).
590 By advective transport, an inoculum for the summer growth of M. leidyi in the Berre basin
591 was possible. Population growth could be rapid in these areas of the large basin under
592 favourable summer conditions but is then followed by seasonal growth. This pattern was
593 consistent with the seasonal distribution patterns described in studies in Narragansett Bay
594 and other areas of the world permanently occupied by M. leidyi (Table 4). The source
595 regions, particularly in temperate regions with cold winters, were characterized by seasonal
596 population growth in the coastal regions, i.e. New England (Kremer and Nixon, 1976),
597 Argentina (Mianzan and Sabatini, 1985) and the Black Sea (Shiganova et al., 2001). Studies
598 to delineate source and sink regions for M. leidyi have been limited in number because this
599 distinction requires multi-year sampling regimes with enough spatial variation to distinguish
600 between different habitat types within a metapopulation region.

601

602 5. Conclusion
603 To conclude, in the Vaine sub-basin, environmental conditions and trophic conditions well
604 above thresholds favourable for Mnemiopsis reproduction and predation were found in our
605 study between 2016-2017 The refugia area has a temperature and enough zooplankton
606 biomass to maintain Mnemiopsis leidyi year-round. The hydrodynamics of this basin show a

18
607 semi-permanent gyre that retains the M. leidyi under most wind regime. Thus, when
608 environmental conditions become favourable, this refugia area could allow M. leidyi to
609 expand into the rest of the lagoon in spring under favourable wind conditions, as observed in
610 the Chesapeake Bay.

611 The discovery of this refugia area in the Berre Lagoon therefore explains the
612 “reappearance” of M. leidyi in the entire lagoon after harsh winters and allows their
613 population to maintain itself in a sustainable way. In the future it will be important to study the
614 proportion of M. leidyi which are exported from the Berre Lagoon to the Mediterranean Sea
615 through the Caronte channel. It is not excluded that the Berre Lagoon represents a
616 population source for Mnemiopsis in French Mediterranean coasts.

617

618 Acknowledgements
619 We would like to thank Bruno Belloni and Justine Gadreaud for their help during in
620 situ sampling. We also thank the Microscopie et Imagerie Marine (MIM) and Service A la Mer
621 (SAM) platforms in MIO. We thank EDF for the temperature and salinity data. We also thank
622 three anonymous reviewers and the editor for their helpful comments.

623

624 Funding
625 The project leading to this publication has received funding from European FEDER
626 Fund under project 1166-39417. Guillaume Marchessaux was supported by a PhD fellowship
627 from the French Ministry of Higher Education and Research between 2015 to 2018.

628

629

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23
863 approach. Marine Biology 163, 1–17.
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865 zooplankton by the comb jelly Mnemiopsis leidyi and pelagic fishes in the Black Sea.
866 Oceanology of the Russian Academy of Sciences 35, 523–527.
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868 developments on the hydromorphology and ecology of the Durance channel and the
869 Etang de Berre, southeast France. Journal of environmental management 104, 35–
870 50.
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875
876
877
878
879
880
881

882 List of tables

883
884 Table 1. Characteristics (latitude, longitude and bottom depth) of the seven stations in the Berre
885 Lagoon.
886
887 Table 2. Meteorological scenarios established by Noveltis used in our study.
888
889 Table 3. Missing periods when of Mnemiopsis leidyi was not sampled in the Berre Lagoon (SA2
890 station) and corresponding environmental conditions.
891
892 Table 4. Patterns of Mnemiopsis leidyi distributions and population dynamics reported in literature.
893
894
895
896
897
898
899
900
901
902
903
904
905
906
907
908
909
910

24
911
912
913
914
915
916
917
918
919
920
921
922
923
924
925
926
927
928

929 List of figures

930

931 Figure 1. Study area and sampling stations (black circles). Black triangles: freshwater inputs
932 (Touloubre, Durance (EDF powerplant), Arc) and white triangle: connection to the sea (Caronte
933 Canal, South-West). Grey circles: long-term sampling stations from 2010-2015 (SA1, SA3). Blue lines:
934 rivers and Durance canal. White squares: cities. Med. Sea: the Mediterranean Sea.
935
936 Figure 2. Bathymetry and mesh description of the TELEMAC3D configuration on the Berre lagoon. Up:
937 bathymetry (colour) and horizontal mesh. Down: vertical mesh with the distribution of the 25 hybrid
938 plans.
939
940 Figure 3. Map of (A) the characteristic areas defined for the study of Lagrangian transport and (B)
941 characteristics of each areas. Black circles: sampling stations where simulations were started.
942
943 Figure 4. Temporal evolution of planktonic biomass (A) total biomass between 2010 and 2017, (B)
944 biomass by size classes between 2015-2017 at SA2.
945
946 Figure 5. Temporal evolution of the abundance of metazooplankton (ind m-3) and Mnemiopsis leidyi
947 (ind m-3) from 2010 to 2017 at SA2 station.
948
949 Figure 9. Different life stages (eggs, larvae, adults) Mnemiopsis leidyi abundance (log ind m-3) and
950 zooplankton abundance (ind m-3) between February 2016 and September 2017 for each station.

25
951 Figure 7. Principal component analyses (PCA) carried out on monthly average data according to
952 seasons and circle of correlations of measured variables (Spearman) between 2016 and 2017 on the
953 7 stations studied. Zpk: abundance of zooplankton (ind m-3); T: temperature (°C); Adults: abundance
954 of Mnemiopsis adults (ind m-3); Larvae: abundance of Mnemiopsis larvae (ind m-3); Eggs: abundance
955 of Mnemiopsis eggs (ind m-3); EDF : EDF powerplant flows (m3 s-1); Chlo : total chlorophyll
956 concentration (µg L-1); Arc : Arc river flows (m3 s-1); Wind spe: wind speed (m s-1); Wind dir: wind
957 direction (m s-1).
958
959 Figure 8. Currents of the Berre Lagoon under moderate and strong Mistral wind conditions (North-
960 West). Produced by Vincent Faure (GIPREB, 2019).
961
962 Figure 9. Current of the Berre Lagoon under a Ponant (West-North-West) regime. Produced by
963 Vincent Faure (GIPREB, 2019).
964 Figure 10. Current of the Berre Lagoon under a Marin (south-east) wind regime. Produced by Vincent
965 Faure (GIPREB, 2019).
966
967 Figure 11. Three-dimensional trajectories of particles over 72 hours under moderate Mistral regime
968 for the 7 stations studied. The black dot indicates the initial position of the particles. Yellow arrow:
969 wind direction.
970
971 Figure 12. Three-dimensional trajectories of particles over 72 hours under strong Mistral regime for
972 the 7 stations studied. The black dot indicates the initial position of the particles. Yellow arrow: wind
973 direction.
974
975 Figure 13. Three-dimensional trajectories of particles over 72 hours under a weak Ponant regime for
976 the 7 stations studied. The black dot indicates the initial position of the particles. Yellow arrow: wind
977 direction.
978
979 Figure 14. Three-dimensional trajectories of particles over 72 hours under moderate Marin regime
980 for the 7 stations studied. The black dot indicates the initial position of the particles. Yellow arrow:
981 wind direction.
982
983 Figure 15. Assessment of the conditions measured in the refugia area of M. leidyi in the Berre
984 Lagoon.
985

26
 986 List of supplementary figures
987
988 Supplementary figure 1. Wind speed (m s-1) as a function of direction (°) over time between 2010 and
989 2017.
990

991 Supplementary figure 2. Monthly evolution of temperature for SA2 station from 2010 to 2017 (2010-
992 2016: EDF CTD; 2016-2017: our CTD) and for other stations between May 2016 and September 2017.
993 X-axis: numbers between 1 to 12 represent months.
994
995 Supplementary figure 3. Monthly evolution of salinity for SA2 station from 2010 to 2017 (2010-2016:
996 EDF CTD; 2016-2017: our CTD) and for other stations between May 2016 and September 2017. X-
997 axis: numbers between 1 to 12 represent months.
998
999 Supplementary figure 4. Monthly evolution of chlorophyll a concentration for (A) SA2 station (surface
1000 and bottom) from 2010 to 2017 and (B) for other stations between May 2016 and September 2017.
1001 X-axis: numbers between 1 to 12 represent months.
1002

1003 Supplementary figure 5. Temporal evolution of estimated carbon concentrations (µgC L-1) between
1004 January 2010 and September 2017 for SA2 and between February 2016 and September 2017 for six
1005 coastal stations: carbon concentrations for phytoplankton and zooplankton.
1006
1007 Supplementary figure 6. Hypotheses of reseeding of Mnemiopsis leidyi in the Berre basin from
1008 refugia area (Vaine sub-basin). Estimates made on wind conditions 3 days before the reappearance
1009 of Mnemiopsis in the samples at SA2 station.
1010

1011

1012

1013

27
1
Stations Location Latitude Longitude Depth (m)
Istres Berre basin 43.4695 5.0206 6.5
EDF Berre basin 43.5243 5.0706 2.6
Arc Berre basin 43.5065 5.1113 2.6
SA2 Berre basin 43.4458 5.1139 8.2
3 frères Berre basin 43.4050 5.1157 5.0
Rognac Vaine basin 43.4644 5.2281 3.6
Anse de Between Berre and Vaine basins
43.4577 5.1610 1.5
Berre
2
3
 Localisation Speed (m s-1) Direction (°)

Strong Mistral (North-West) regime > 11 (> 39.6 km h-1) 300 – 345

Moderate Mistral (North-West) regime 8 – 11 (28.84 – 39.6 km h-1) 300 – 345

Ponant wind (Light West-North-West) regime 3 – 8 (10.8 – 28.84 km h-1) 255 – 300
Marin wind (Moderate South-East) regime 8 – 11 (28.84 – 39.6 km h-1) 120 – 165

5
1
Dates Temperature Salinity Chlorophyll Zooplankton
(°C) (µg L-1) abundance
(ind m-3)
12 March 2012 8.1 19.8 3.7 16 024
28 March 2012 11.8 22.3 0.9 38 380
23 April 2012 14.8 23.4 7.7 34 093
31 May 2012 19.6 24.5 1.3 56 707
7 March 2013 6.9 20.9 5.0 -
21 March 2013 8.1 21.0 8.3 6 933
20 June 2013 22.3 22.5 - -
11 July 2013 23.0 22.6 3.4 9 299
18 July 2013 24.3 23.2 2.3 1 728
16 July 2014 21.9 23.6 10.9 112 070
12 May 2016 15,9 22.9 4.2 78 677
29 June 2016 21.7 22.8 2.8 2 992
5 July 2016 22.0 23.5 3.8 1 718
2

3
 Region Seasonal variation pattern References
Narragansett Bay, Providence River regions characterized by longer Kremer and Nixon
USA seasonal presence and higher average abundances (1976), Costello et al.
than the mouth of the bay bordering the Atlantic (2006)
Ocean. Spring growth spread from inshore to
offshore stations.

Black Sea Spring and autumn reproduction peaks. High Shiganova et al. (2001)
abundances during summer. Shiganova et al. (2019)
and references included

Caspian Sea Reproduction all of the year and peak on abundance Shiganova et al. (2019)
during summer. and references included

Aral Sea Annual reintroduction necessary from adjoining Shiganova et al. (2001)
Black Sea due to yearly winter extinction in the
Aral Sea.

Sea of Marmosa Persistent year-round population provides a source Shiganova et al. (2001)
region for annual colonization of adjoining areas
via advective transport.

Bahia Bay, Argentina Winter persistence of M. leidyi in core region of bay Mianzan and Sabatini
occupied by a cluster of islands. Seasonal (1985)
population
expansion from this core region during summer
months followed by contraction to core region during
winter months over a 3-yr period.

Aegean Sea Reproduction period between spring and peak of Shiganova et al. (2019)
abundance and reproduction during summer. and references included

Levantine Sea Peak of abundances and reproduction during Shiganova et al. (2019)
autumn and winter. No-data for spring and summer. and references included

Ligurian Sea peak of abundances and reproduction during spring Shiganova et al. (2019)
and autumn. No-data for summer and winter. and references included

North Sea Winter an autumn reproduction peak and high Shiganova et al. (2019)
abundances during summer. and references included
1