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I. Siokou-Frangou et al.: Mediterranean plankton
Fig. 7. The top panel shows the deepening of the DCM (Z Chl Max) and the chl a dispersion (Chl Dispersion)
as an average of the discrete depth difference from water column average of chl a concentration, in percentage.
Dispersion values in the WMS are closer to 100% than in the EMS, demonstrating a higher vertical patchiness
in the latter. The bottom panel represents the west to east decrease for calculated chl a in pico-, nano- and
microplankton. Note the Longitude scale: the two data points to the left are outside the MS, while the Levantine
basin was not sampled. Modified with permission from Dolan et al. (2002).
80
Fig. 7. The top panel shows the deepening of the DCM (Z Chl
Max) and the chl a dispersion (Chl Dispersion) as an average of
the discrete depth difference from water column average of chl a
concentration, in percentage. Dispersion values in the WMS are
closer to 100% than in the EMS, demonstrating a higher vertical
patchiness in the latter. The bottom panel represents the west to
east decrease for calculated chl a in pico-, nano- and microplankton.
Note the Longitude scale: the two data points to the left are outside
the MS, while the Levantine basin was not sampled. Modified with
permission from Dolan et al. (2002).
the exception of the short period of late winter mixing. The
DCM progressively sinks across a west to east gradient from
30 m in the westernmost area (Dolan et al., 2002, Fig. 7), to
70 m in the South Adriatic Sea (Boldrin et al., 2002), down
to 120 m in the Levantine basin (Christaki et al., 2001; Dolan
et al., 2002). The eastward increase in DCM depth is prob-
ably related to lower productivity and hence higher seawater
transparency in the Levantine Sea, but the level of DCM may
vary considerably between cyclonic and anticyclonic areas
(Ediger and Yilmaz, 1996). In the westerm MS, the depth
of the DCM is strongly affected by the Atlantic water inflow
and the consequent physical dynamics along the vertical axis
(Raimbault et al., 1993).
The distribution of biomass is clearly reflected in pri-
mary production rates (Table 1). Satellite-based estimates
range from 130 to 198 g C m
−
2
y
−
1
over the years 1997–
2001 (Bricaud et al., 2002; Bosc et al., 2004), with values
for the EMS generally in the lower portion of the range. Es-
timates from in situ incubations in previous decades were
as low as 80–90 g C m
−
2
y
−
1
(Sournia, 1973). More recent
measurements get closer to satellite-based estimates, e.g., in
the Gulf of Lion (140–150 g C m
−
2
y
−
1
, Conan et al., 1998),
but remain consistently lower in other areas such as the Cre-
tan Sea (59 g C m
−
2
y
−
1
, Psarra et al., 2000). A clear east-
ward reduction in primary production was reported in the
Fig. 8. Integrated primary production (mg C m
−
2
day
−
1
) during the MINOS cruise (May-June 1996). Repro-
duced with permission from Moutin and Raimbault (2002).
81
Fig. 8. Integrated primary production (mg C m
−
2
day
−
1
) during
the MINOS cruise (May–June 1996). Reproduced with permission
from Moutin and Raimbault (2002).
results from a late-spring (May–June) trans-Mediterranean
cruise (Moutin and Raimbault, 2002), when maxima were
close to 1 g C m
−
2
d
−
1
in the south-western basin and min-
ima ranged between 150 and 250 mg C m
−
2
d
−
1
at several
stations of the Levantine Sea (Fig. 8). Interestingly, estimates
obtained in spring in other studies reflect the same spatial
pattern and are within the same ranges as those shown by
Moutin and Raimbault (2002) (Table 1). Comparably high
values (up to 1.7 g C m
−
2
d
−
1
) were reported in the Catalan
front area in March (Moran and Estrada, 2005), and in the
Alboran Sea in May–June (Lohrenz et al., 1988). At the DY-
FAMED station in the Ligurian Sea, primary production rates
were 240–716 mg C m
−
2
(over a 14 h incubation) (Vidussi
et al., 2001) but reached values as high as 1.8 g C m
−
2
d
−
1
in
April (Marty and Chiaverini, 2002). Measurements at other
sites of the EMS, namely in the South Adriatic Sea (Boldrin
et al., 2002) and in the North East Aegean Sea (Ignatiades
et al., 2002; Zervoudaki et al., 2007), also match the low val-
ues recorded by Moutin and Raimbault (2002).
Spatial and seasonal variability of primary production val-
ues can be high (Table 1), especially in very dynamic ar-
eas like the Alboran Sea (Mac´ıas et al., 2009) and the
Catalan Sea (Granata et al., 2004). Inter-annual variability
of primary production may also be high, mainly depend-
ing on the depth of the winter mixing (Estrada, 1996). In
spite of light limitation, the contribution of subsurface and
deep chlorophyll maxima can be significant, in some cases
reaching 30% of the total production of the water column
(Estrada et al., 1985).
While satellite-based estimates represent gross primary
production, values from in situ incubations are generally
closer to net production. In neither case, however, is an
estimate of the new versus recycled production provided,
and it is hence impossible to appreciate the actual amount
of biomass added to the system through phytoplankton ac-
tivity.
Indeed, values for new production, which can be
equated to the export production assuming a quasi-steady
state, are much lower, i.e. in the order of 4.5 mol C m
−
2
y
−
1
(1mol C≡12g C) and 1.5 mol C m
−
2
y
−
1
for the WMS and
EMS, respectively (e.g. Bethoux, 1989, but see also Minas
et al., 1988 for a discussion). These estimates correspond
Biogeosciences, 7, 1543–1586, 2010
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