assessment of the main drivers of the black sea ecosystem functioning mnemiopsis leidyi and beroe...
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Assessment of the main drivers of the Black Sea ecosystem functioning Mnemiopsis leidyi and Beroe ovata impact on
the Black Sea ecosystem. Modeling approach.
Tamara Shiganova
P.P.Shirshov Institute of oceanology RAS, Moscow, RUSSIIA
Paul NivalLaboratoire d'Oceanographie de Villefranche Villefranche-sur-Mer, FRANCE
Distribution of M. leidyi in the native and invaded areas
Costello J. H., J. E. Purcell, K. M. Bahya, H. W. Mianzan & T. A. Shiganova, 2011
Native area
Invaded area
In the Black Sea, native gelatinous species belong to moderately cold-water species: the ctenophore Pleurobrachia pileus, scyphomedusa Aurelia aurita, and the pyrophyte alga Noctiluca scintillans. Two warm water invasive ctenophores arrived and established in the heated upper layer
Cold water species Warm water species
Subdivision of the gelatinous species in their relation to mean seasonal, annual and minimal winter SST: analyses of field data according to main component method )
Population genetic analyses supported its invasion from the Gulf of Mexico (e.g., Tampa Bay) into the Black Sea, then secondary into the Azov, northern Aegean and into the Caspian Sea and the Mediterranean (Ghaboolia, Shiganova et al., 2010)
1982
1988
1992
19901999
1993
2009
2005
2005
20092009
2006
2005
2006
2006
2007
2007
Dispersal of Mnemiopsis leidyi in the Eurasian seasDispersal of Mnemiopsis leidyi in the Eurasian seasMnemiopsis leidyi
19971997
1999
2004
2011
20052000
Dispersal of Beroe ovata in the Eurasian seasBeroe ovata
Black Sea
Sea of Azov
Aegean Sea
Caspian Sea
Baltic Sea
The dark areaCorresponds to theperiod of Mnemiopsis leidyi occurance(observations)
Environmental data and
M.leidyi invasion in the seas of Eurasia
After Shiganova et al., in press
Interannual variation of M.leidyi and B.ovata in the Black Sea
0
500
1000
1500
2000
2500
19841988
1989
1990
1991
1992
1993
1994
1995
1996
1997
1998
Years
Abun
danc
e, ind
.m-2
M.leidyi,ind.m2
0
500
1000
1500
2000
2500
1999
2000
2001
2002
2003
2004
2005
2006
2007
2008
0
200
400
600
800
1000
1200
M.leidyi,ind.m2
B.ovata,ind.m2
Zooplankton are the main food of M.leidyiChange of interannual variability of edible zooplankton after
appearance of invasive ctenophores in the inshore and offshore waters in August in the northeastern Black Sea
Zooplankton in the inshore waters,ind.m2
0
50000
100000
150000
200000
250000
300000
350000
400000
450000
500000
VIII.1989
VIII.1993
VIII.1994
VIII.1995
VII.1996
VIII.1998
VIII.1999
IX.2000
VIII.2001
VIII.2002
VIII.2003
VIII.2004
VIII.2005
VIII.2006
VIII.2007
VIII.2008
Years
Zoop
lank
ton,
ind.
m-2
Calanus euxinus Paracalanus parvusPseudocalanus elongatus Acarthia clausiOithona similis Pontella mediterraneaLarvae_Bivalvia Larvae_PolychaetaLarvae_Decapoda Larvae GastropodaCirripedia Larvae Parasagitta setosaCladocera without Penilia Penilia avirostrisCentropages ponticus Oikopleura dioicaHarpacticoida
Zooplankton in the offshore waters,ind.m2
0
50000
100000
150000
200000
250000
300000
350000
400000
450000
VIII.1989
VIII.1993
VIII.1994
VIII.1995
VII.1996
VIII.1998
VIII.1999
IX.2000
IX.2002
VIII.2003
VIII.2004
IX.2005
VIII.2006
VIII.2007
VIII.2008
zoop
lankto
n,ind
.m2
Calanus euxinus Paracalanus parvusPseudocalanus elongatus Acarthia clausiOithona similis Pontella mediterraniaOithona nana Oikopleura dioicaParasagitta setosa Larvae_BivalviaLarvae_Polychaeta Larvae_DecapodaLarvae Gastropoda Cirripedia LarvaeCladocera without Penilia Penilia avirostrisCentropages ponticus IsopodaOthers
Mnemiopsis leidyi seasonal cycle in cold (1993) and warm (1994) year before B.ovata arrival
Field data in the coastal area
r=0,6-08, p<0,001Cold yearCold year Warm year
Before the arrival of B.ovata, M.leidyi abundance was controlled by temperature and zooplankton prey
1994
0
1
2
3
4
5
6
7
8
9
I II III IV V VI VII VIII IX X XI XII
mn
emio
psi
s,г.
м-3
0
1
2
3
4
5
6
7
8
9
Zo
op
lan
kto
n,g
.m-3
Mnemiopsis Zooplankton
1992
0
1
2
3
4
5
6
7
8
9
I II III IV V VI VII VIII IX X XI XII
Mn
emio
psi
s,g
.m-3
0
1
2
3
4
5
6
7
8
9
Zo
op
lan
kto
n,m
g.m
-3
Mnemiopsis Zooplankton
2001
0
5
10
15
20
25
30
0 20 40 60 80 100 120 140 160 180 200 220 240 260 280 300 320 340 360
Time (days
Tem
pera
ture
, 0C
2003
0
5
10
15
20
25
30
0 20 40 60 80 100 120 140 160 180 200 220 240 260 280 300 320 340 360
Time (days)
Tem
pera
ture
, 0 C
2008
0
5
10
15
20
25
30
0 20 40 60 80 100 120 140 160 180 200 220 240 260 280 300 320 340 360
Time (days)
Tem
pera
ture
, 0 C
0
100
200
300
400
500
600
0 20 40 60 80 96 101 120 140 166 210 235 236 237 249 254 264 276 300 320 340 360
Zo
op
lan
kto
n, m
g.m
-3
0
100
200
300
400
500
600
20 40 60 80 100 120 148 160 180 200 220 240 250 260 266 270 280 300 320 340 350Z
oo
pla
nkt
on
,mg
.m-3
0
50
100
150
200
250
300
350
400
450
500
550
600
0 20 40 60 80 100 115 139 162 168 169 200 220 234 252 280 300 320 340 360
Zo
op
lan
kto
n,
mg
.m-3
0
20
40
60
80
100
120
140
160
180
200
220
240
260
0 20 40 60 80 100 120 140 160 180 200 220 240 260 280 300 320 340 360
Mn
emio
psi
s, i
nd
. m
-3
Mn.adult 1
Mn.Juv
Mn.eggs
Mn.larvae
0
20
40
60
80
100
120
140
160
180
200
220
240
260
0 20 40 60 80 100 120 140 160 180 200 220 240 260 280 300 320 340 360
Mn
emio
psi
s, i
nd
.m-3
0
20
40
60
80
100
120
140
160
180
200
220
240
260
0 20 40 60 80 100 120 140 160 180 200 220 240 260 280 300 320 340 360
Mn
emio
psi
s, i
nd
.m3
0
10
20
30
40
50
60
70
80
90
100
0 20 40 60 80 100 120 140 160 180 200 220 240 260 280 300 320 340 360Days
Ber
oe,
in
d.
m-3
0
10
20
30
40
50
60
70
80
90
100
0 20 40 60 80 100 120 140 160 180 200 220 240 260 280 300 320 340 360
Time (days)
Ber
oe,
ind
.m3
0
10
20
30
40
50
60
70
80
90
100
0 20 40 60 80 100 120 140 160 180 200 220 240 260 280 300 320 340 360Days
Ber
oe,
ind
.m-3
Ber.adult 1
Ber.Juv
Ber.eggs
Ber.larva
Temperature
Zooplankton
M.leidyi
B.ovata
A
JL
E
E
A
A
JL
Interannual variability of phenology of ctenophores M. leidyi and B.ovata in the coastal area of the NE Black Sea
Interannual variability of phenology of ctenophores M. leidyi and B.ovata in the coastal area of the NE Black Sea
Time of appearance of B.ovataTime of beginning reproduction of B.ovata
Time of appearance of M.leidyiTime of pick of reproduction of M.leidyi
120
140
160
180
200
220
240
260
280
300
320
340
360
1999 2000 2001 2002 2003 2004 2005 2006 2007 2008Year
Tim
e (d
ays)
appearance B.ovata Reproduction start of M.leidyiReproduction start of B.ovata Peak of M.leidyi reproduction Peak of B.ovata reproduction Disappearance M.leidyi larvaeDisappearance B.ovata larvae
--- M.leidyi
--- B.ovata
Life stages and forcing factors
for M.leidyi and B.ovataL J
E
A
food
Temperature
Individual based modeling approach was used to take into account life cycle and physiological features
E – eggsL-larvaJ-juvenaleA-adult
-mortality
0 50 100 150 200 250 300 350 4000
5
10
15
20
25
time (days)
biom
asse
zoo
pk (
mm
olN
.m-3
)
0 50 100 150 200 250 300 350 4005
10
15
20
25
30
temps days
Tem
pera
ture
(°C
)
Forcing functions
Zooplankton biomass
Temperature
Mortality rate
00.1
0.20.3
0.4
0
10
20
30
400
0.5
1
1.5
2
2.5
3
mmA1
0
0,5
1
1,5
2
2,5
0 5 10 15 20 25 30 35
Temperature (°C)
Ad
ult
mo
rta
lity
(d
ay
-1)
mm
A1
mortality
0
0,2
0,4
0,6
0,8
1
1,2
0 0,1 0,2 0,3 0,4
food concentration (mmol N.m-3)
mo
rtal
ity
rate
FoodTemperature
Mor
talit
y ra
te
Model
ZE
A
J
A
J
L
L
ETemperature
food
Prey – predator individual based model structure
M.leidyi
B.ovata
Age classes
L
A
J1
1
1
1
2
2
2
2
no
nL
nJ
nAprocess
ageing
Age classes
L
A
J1
1
1
1
2
2
2
2
no
nL
nJ
nAprocess
ageing
0% 50% 100%
fast growing
0
5
10
15
20
25
0 10 20 30 40 50
age (days)
length (
unit
s)
slow growing
egg
egg
larva juvenile
juvenilelarva
Adult
Adult
fast
slow
Change in stage duration depending on physiology (food, temperature)
0 5 10 15 20 25 300
50
100
150
temps days
n in
divi
dus
per
unit
volu
me
eggs
larvae
juveniles
adults
old adults
Time (days)
Num
ber
indi
vidu
als
per
unit
of
volu
me
first larva
first egg
first adult
first juvenile
Model
Ontogenetic cycle of M.leidyi and B.ovata development
5 10 15 20 25 300
0.1
0.2
0.3
0.4
0.5
0.6
0.7
0.8
0.9
1
temperature (°C)
spaw
ning
rat
e co
ef.
(d-1
)
Reproduction rate
0
100
200
300
400
500
600
700
20 21 22 23 24 25 26 27 28
Temperature, 0C
WW
,g,e
gg-
2.d
ay-1
0
20
40
60
80
100
120
140
160
180
20 21 22 23 24 25 26 27 28
Temperature, C
egg.
gWW
-1,d
ay
Experiments Model
M.leidyi
B.ovata
0 0.05 0.1 0.15 0.2 0.25 0.30
0.1
0.2
0.3
0.4
0.5
0.6
0.7
0.8
0.9
1
available food concentration
inge
stio
n ra
te c
oef.
(d-
1)
Ingestion rate coefficient (0 – 1)
C2= (Food- minFood)/ (Kf+ (Food-minFood))
C2 = 0 when Food < minFood
FoodminFood
0
0,5
1
1,5
2
2,5
3
3,5
4
0 2000 4000 6000 8000 10000 12000
zooplankton,ind.m-3
Ratio
n,DW
.day
-1
experiment Model
Ingestion rate
0
0.05
0.1
0.15
0.2 5 10 15 20 25 30 35
0
5
10
15
20
Stage duration
age to 50% transfer
Temperature (°C)
Ingestion
Sta
ge d
urat
ion
(day
s
optimum conditions
0 50 100 150 200 250 300 350 4000
50
100
150
200
250
300
350
400
temps (days)
num
bers
(n
m-3
)
First stage: M.leidyi present
B.ovata absent
A
L
J
E
0 50 100 150 200 250 300 350 4000
50
100
150
200
250
300
350
400
temps (days)
num
bers
(n
m-3
)
0 50 100 150 200 250 300 350 4000
50
100
150
200
250
time (days)
num
be
rs (
n m
-3)
M.leidyi
B.ovata
A
L
A
L
Second stage
B.ovata appears in surface water at time 200th day
Simulation of input of B.ovata in surface water at time 200th day.M.leidyi develops a bloom, which is grazed by B.ovataPredation on larvae, juveniles and adult on M.leidyi makes them disappear
Model
E
JE
J
0 50 100 150 200 250 300 350 4000
50
100
150
200
250
300
350
400
temps (days)
num
bers
(n
m-3
)
0 50 100 150 200 250 300 350 4000
50
100
150
200
250
time (days)
nu
mb
ers
(n
m-3
)
M.leidyi
B.ovata
A
L
A
L
0
20
40
60
80
100
120
140
160
180
200
220
240
260
0 20 40 60 80 100 120 140 160 180 200 220 240 260 280 300 320 340 360
Time days
Mne
mio
psis
, ind
. m-3
Mn.adult 1 Mn.Juv Mn.ova Mn.larvae
0
10
20
30
40
50
60
70
80
90
100
0 20 40 60 80 100 120 140 160 180 200 220 240 260 280 300 320 340 360Time days
Ber
oe,
ind
.m-3
Ber.adult 1 Ber.Juv Ber.ova Ber.larva
Model Field observations
M.leidyi
B.ovata B.ovata
L
J
A
L
E
A
JE
Conclusions
In present Black Sea ecosystem there is a bottom up control from zooplankton to its consumer Mnemiopsis leidyi and finally to its predator Beroe ovata.
Annual changes in temperature and food availability are considered as the main factors that control these predators’ dynamics and their impact on pelagic ecosystem of the Black Sea.Both field data analyses and individual based modeling confirmed that, with appearance B. ovata that controlled M. leidyi population, a recovering shift of the ecosystem appeared but was controlled by climate forcing.
Now in any case it is another ecosystem with two ctenophores that affected ecosystem but the time of high effect of M.leidyi is much shorter.
Acknowledgement
The research was performed in framework of project GK-0422
Thank you for attention