Journal of Sea Research 58 (2007) 283 290

www.elsevier.com/locate/seares

Unusual diatoms linked to climatic events in the

northeastern English Channel
Fernando Gmez , Sami Souissi
Station Marine de Wimereux, Universit des Sciences et Technologies de Lille-Lille1,
FRE 2816 ELICO CNRS, 28 Avenue Foch, F-62930 Wimereux, France
Received 26 February 2007; accepted 2 August 2007
Available online 21 August 2007

Abstract
The composition of the micro-phytoplankton was investigated at two fixed stations in the northeastern English Channel from
November 1997 to December 2005. In late summer-autumn 2005, proliferations of the centric diatoms Proboscia indica and
Rhizosolenia hebetata f. semispina were recorded for the first time in this area. The weather in 2005 was abnormal: a cold
winter and early arrival of summer conditions that extended to late autumn. This resulted in an unusually long, sustained calm and
warm period. We suggest that the reduction in mixing may have allowed diatoms of more-stratified waters to be competitive in the
normally highly mixed northeastern English Channel in summer-autumn 2005. In 2003 and in 2005, two warm-water diatom
species were recorded for the first time. A colony of Eucampia cornuta was observed after an exceptional heat wave in September
of 2003, and again in September 2005. Simultaneously at both fixed stations, Chaetoceros peruvianus was recorded in the warm
October of 2005. The climatic events are associated with the northward spreading of thermophilic diatoms such as E. cornuta and
Ch. peruvianus in the European Atlantic waters, and the proliferation of diatoms of stratified environments such as P. indica and
R. hebetata f. semispina in the usually highly turbulent waters of the northeastern English Channel.
2007 Elsevier B.V. All rights reserved.
Keywords: Diatoms; Thermophilic and warm-water phytoplankton; Time series; Climate change; Interannual variability

1. Introduction
It is generally accepted that we are in a climate phase
of accelerated warming coupled with increased high
frequency of temperature variation (Schr et al., 2004).
Meteorological or climatic factors affect the temperature, salinity, and physical structure of the ocean water
column. Climate change will influence the boundaries
of biogeographical regions and thus the ranges of
distribution of individual species. This hypothesis is
Corresponding author.
E-mail address: fernando.gomez@fitoplancton.com (F. Gmez).
1385-1101/$ - see front matter 2007 Elsevier B.V. All rights reserved.
doi:10.1016/j.seares.2007.08.002

widely supported today, and has been reinforced by

recent studies that show responses of marine organisms
to climatic features of the northeastern Atlantic (e.g.,
Alheit and Hagen, 1997; Beaugrand et al., 2000;
Hawkins et al., 2003).
The summer of 2003 was characterised by exceptionally hot weather in Europe, with the mean
temperature exceeding that of any summer for the past
500 years (Luterbacher et al., 2004). A heat wave
occurred from 1 to 15 August 2003, which caused
excessive human mortality throughout Europe, to a
catastrophic extent in France (Vandentorren et al.,
2004). The year 2005 was also anomalous in Europe,

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F. Gmez, S. Souissi / Journal of Sea Research 58 (2007) 283290

Fig. 1. Map of the English Channel. The arrow indicates the sampling stations (solid circles).

with a severe winter that caused snowfalls extending to

unusual places such as Algeria. In late spring-summer,
rainfall and snowmelt caused severe flooding of central
and eastern European rivers, and multi-month drought
conditions and forest fires also affected much of Western
Europe during July, August, and September (World
Meteorological Organization, 2005).
Long-term monitoring is essential for advances in
understanding the interannual variability and climatic
linkages of marine plankton. The French national
SOMLIT (Service d'Observation en Milieu LITtoral)
monitoring program was established off Boulogne-surMer (NE English Channel) in November 1997. The
present study describes qualitative changes in the
diatom composition since that time. Eucampia cornuta

(Cleve) Grunow was recorded for the first time in 2003,

and Chaetoceros peruvianus Brightwell in 2005. In late
summer and autumn 2005, unusual rhizosolenioid
diatoms reached high abundances. We describe and
illustrate these observations of unusual diatoms, which
reveal changes in the structure of the marine ecosystem
in European Atlantic waters.
2. Material and methods
Overall, 158 cruises were carried out on board RV
Sepia II from November 1997 to December 2005 off
Boulogne-sur-Mer in the northeastern English Channel
(Fig. 1). Two fixed stations were sampled during high
tides. One station was located 2 km offshore (5040

Fig. 2. Temporal distribution (19982005) of mean daily air temperature (C) at Boulogne-sur-Mer.

F. Gmez, S. Souissi / Journal of Sea Research 58 (2007) 283290

285

Fig. 3. Abundance (103 cells L 1) of Proboscia indica, Rhizosolenia hebetata f. semispina, and Neocalyptrella robusta in autumn 2005. Note the
different scale of Fig. 3d.

75N; 13117E; 21 m depth) under the influence of the

coastal flow (Brylinski et al., 1991). The other station,
8 km offshore (504075N; 12460E, 50 m depth), was
often separated by a tidal coastal front. The sampling
frequency was intended to be biweekly, but cruises were
sometimes cancelled or were restricted to the shore
station because of bad weather. Conductivity-temperature-depth (CTD) profiles were measured by a SeaBird
SBE 25 probe. However, in order to avoid erroneous
tendencies due to the undersampled periods or malfunctions leading to missing data during the eight years,
the meteorological variability was determined from the
meteorological station closest to the sampling stations.
Continuous records of air temperature were provided by
the French Meteorological Agency (Mto-France)
from a meteorological station located at Boulogne-surMer (5044N, 0136E, altitude 73 m).
Seawater samples were collected with a Niskin bottle
at the surface and at one metre above the bottom. Lugolfixed samples of 25 or 50 ml were settled in composite
settling chambers. The entire chamber was scanned at
200 magnification with an Olympus IX71 inverted
microscope, and specimens were photographed at 400
magnification. The sample analyses of the 8y time
series were carried out by the first author using the same
method.

a minimum of 4.56 C on 8 January 2003. In winter

1998 and during the winter of 2005, lower temperatures
were also recorded (Fig. 2). The years with the highest
number of days when mean air temperature was lower
than 0 C were 1998 (11 d), 2003 (9 d), and 2005 (9 d).
The fewest days with mean daily temperature b0 C
were recorded in 1999 (1 d), 2000 (2 d), and 2004 (2 d).
In winter 2005, mean air temperature fell below 0 C on
3 March, which was the latest temperature below 0 C in
the 8y time series (Fig. 2).
The highest mean daily air temperature of the 8y
time series occurred on 10 August 2003, 28.35 C. The
warmest years, based on the number of days with mean
daily air temperatures higher than 22 C, were 2003
(8 d) and 2005 (6 d). In contrast, 1998 and 2000 had
mean daily air temperatures higher than 22 C on only
two days. As usual, the highest temperatures were
recorded in July, August, and early September. In 2005,
the temperature reached daily means of up to 25.32 C
in early June. These data indicate that 2003 and 2005
were anomalous years. The winters were very cold in
both years, and extremely high temperatures were
recorded in the first half of August 2003. In 2005, the
low temperatures extended to March; high temperatures
were reached unusually early, in June, and continued to
late autumn (Fig. 2).

3. Results

3.2. General diatom assemblage characteristics

3.1. Air temperature at Boulogne-sur-Mer (November

1997December 2005)

The sampling stations were located near the Strait of

Dover, which constitutes the maritime corridor between
the English Channel and the North Sea. This region is
subject to strong tidal currents and winds (Salomon and
Breton, 1991; Prandle et al., 1993). Within this highly

The lowest mean daily air temperatures were

recorded from mid-December to February 2003, with

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F. Gmez, S. Souissi / Journal of Sea Research 58 (2007) 283290

Fig. 4. Photomicrographs of unusual diatoms off Boulogne-sur-Mer (NE English Channel), bright-field optics. (ad) Rhizosolenia hebetata
f. semispina. (ef) Proboscia alata. (gh) Proboscia indica. (i) Neocalyptrella robusta. (j) Corethron cf. hystrix. (kl) Chaetoceros peruvianus.
(mn) Eucampia cornuta. Dates of records: (n) 11/09/2003; (a, e) 08/08/2005; (b, c, f) 06/09/2005; (d, g, i, m) 19/09/2005; (h, j, k, l) 18/10/2005.
Scale bars represent 50 m.

turbulent environment, a few species of diatoms

dominate the phytoplankton assemblage, even in
summer. The spring diatom bloom is dominated by
Guinardia delicatula (Cleve) Hasle, Rhizosolenia

imbricata var. shrubsolei (Cleve) Schrder, Guinardia

striata (Stolterfoth) Hasle, and Guinardia flaccida
(Castracane) H. Peragallo. Although these diatoms
may also re-appear in late summer-early autumn

F. Gmez, S. Souissi / Journal of Sea Research 58 (2007) 283290

proliferations, other species are more characteristic of

this period, such as Pseudo-nitzschia spp. accompanied
by Ditylum brightwellii (West) Grunow, Thalassionema
nitzschioides (Grunow) Grunow ex Hustedt, Eucampia
zodiacus Ehrenberg, Meuniera membranacea (Cleve) P.
C. Silva, and, towards winter, the tychoplanktonic
species Paralia sulcata (Ehrenberg) Cleve and Raphoneis amphiceros (Ehrenberg) Ehrenberg. Rhizosolenia
setigera Brightwell was also a common component of
autumn assemblage, with an exceptionally high abundance in October 2003.
3.3. Unusual rhizosolenioid diatom assemblage in late
summer-autumn 2005
From August to November 2005, unusual members of
the family Rhizosoleniaceae Petit were observed. In
September, October, and November 2005, Proboscia
indica (H. Peragallo) Hernndez-Becerril (=Rhizosolenia
alata var. indica) was recorded for the first time in the 8y
time series. Rhizosolenia hebetata f. semispina (Hensen)
Gran was very rare before 2005. However, these two large
species reached high densities in autumn 2005. Also the
large diatom Neocalyptrella robusta (Norman) Hernndez-Becerril et Meave (=Rhizosolenia robusta), very
rarely observed previously, reached a remarkable abundance in autumn 2005. This assemblage of large rhizosolenioid diatoms in late summer and autumn 2005 is
highly unusual.
3.3.1. Rhizosolenia hebetata f. semispina (Fig. 3d,
Fig. 4ad)
Associated with Proboscia indica was Rhizosolenia
hebetata f. semispina. Whereas P. indica is morphologically distinctive, R. hebetata f. semispina may be
misidentified with congeneric species such as Rhizosolenia styliformis Brightwell, especially the variety
R. styliformis var. longispina Hustedt. R. hebetata f.
semispina is characterised by two columns of segments
and claspers, and a pointed otaria along the proximal
part of the process (Fig. 4ad). The specimens of
R. hebetata f. semispina reached up to 800 m in length.
No proliferation of R. hebetata f. semispina was
observed before autumn 2005. This taxon showed the
highest abundance at the beginning of October, reaching
20 000 cells L 1 at the subsurface depth of the onshore
station (Fig. 3d).
3.3.2. Proboscia indica (=Rhizosolenia alata var.
indica) (Fig. 4 gh)
Very few specimens of Proboscia alata (Brightwell)
Sndstrom (=Rhizosolenia alata var. alata) were re-

287

corded in late summer 2005 (Fig. 4e, f). However, the

dominant species of that genus was Proboscia indica
(=Rhizosolenia alata var. indica). This diatom differs
from other rhizosolenioid diatoms by the proboscis, an
elongated part of the valve with truncate tip and no
process. P. indica exhibits a number of characteristics
that distinguish it from P. alata and related taxa: its
frustule and valve dimensions are larger than in other
species, and as in P. alata the chain arrangement is
asymmetrical because of displaced claspers. The long
proboscis is more strongly sloped than in P. alata
(Fig. 4eh). No misidentification is possible because
of this distinctive morphology (Jordan et al., 1991;
Hernndez-Becerril, 1995), although the nomenclature
of the species is complicated because it has been often
considered a variety of R. alata. During this 8y period,
P. indica was only recorded in 2005, persisting between
August and late October. In September 2005, P. indica
reached an abundance of 10 000 cells L 1, becoming
one of the main components of the autumnal diatom
assemblage (Fig. 3).
3.3.3. Neocalyptrella robusta (=Rhizosolenia robusta)
(Fig. 4i)
This is a highly distinctive and very large species. A
single record in April 2002 and two records in February
2005 were the only observations before autumn 2005.
Neocalyptrella robusta reached a maximum abundance
of 150 cells L 1 in September and October 2005
(Fig. 3). Although its abundance was low compared
with the previous taxa, it should be taken into account
that the diatom is very large and thick, and contributes
considerably to the phytoplankton biomass. This species
completed the unusual diatom assemblage of large
rhizosolenioid diatoms observed in autumn 2005.
3.4. First records of tentative warmwater diatoms
The 2005 autumnal rhizosolenioid assemblage was
composed of diatoms that are common in surrounding
waters such as the North Sea and in the French Atlantic.
The diatoms appeared for the first time in our study area
and reached high abundances, coming to dominate the
assemblage. In 2003 and 2005, two diatoms, Eucampia
cornuta and Chaetoceros peruvianus, were for the first
time recorded in the 8y time series. In contrast to the
unusual rhizosolenioid diatoms, E. cornuta and
Ch. peruvianus were restricted to a few records and a
short period in later summer or early autumn. These
species are here considered to be biological indicators of
warmer conditions. Specimens of the genus Corethron
Castracane, tentatively identified as C. hystrix Hensen

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F. Gmez, S. Souissi / Journal of Sea Research 58 (2007) 283290

(Fig. 4j) appeared for the first time. The first records
coincided with Ch. peruvianus and re-appeared later in
mid December. The consideration of Corethron cf. hystrix as a thermophilic species requires further research.
3.4.1. Chaetoceros peruvianus (Fig. 4kl)
Ch. peruvianus is easily identified, and was recorded
simultaneously at both sampling stations and both
depths on 18 October 2005. The specimens appeared
as solitary cells (Fig. 4k), with the exception of one
twin-cell chain, probably recently divided (Fig. 4l).
Ch. peruvianus belongs to the Phaeoceros subgroup,
with thick setae that contain mobile chloroplasts. The
curved convex setae reached up to 400 m in length.
The maximum abundance was only 5 specimens per
sample in the offshore surface station (100 cells L 1).
The offshore seawater temperature was 16.8 C, and the
salinity was 34.9. Slightly lower values were observed
onshore (16.4 C and salinity of 34.6).
3.4.2. Eucampia cornuta (Fig. 4mn)
A colony of a diatom tentatively identified as Eucampia cornuta was recorded for the first time on 11
September 2003 (seawater temperature 18.5 C and
salinity 34.4) and reappeared on 19 September 2005
(seawater temperature 18.8 C and salinity 34.2). In both
years, E. cornuta appeared as single 4-cell chains in the
onshore station, at the surface in 2003, and at 21 m depth
in 2005. The cells were slightly curved in broad girdle
view, elevations long, narrow and apertures tall and
elliptical. The apical length of each cell was 12 m
and the pervalvar axis 50 m (Fig. 4m, n). Another
congeneric species, Eucampia zodiacus Ehrenberg, also
has a concave valve face. It is quite common in autumn
but may occasionally appear throughout the year. The
ribbed bands of E. cornuta were not observed due to the
optical limitations. The identification is based on the
slightly curved shape of the colony (Fig. 4n) and the tall
and elliptical apertures of E. cornuta (Fig. 4m, n) when
compared with E. zodiacus.
4. Discussion
Climate and meteorological events are well-known
regulators of phytoplankton growth, i.e., seasonal wind
regulation of the annual spring bloom, interannual
variations in this event and in mini-blooms, and species
successions (Robinson and Hunt, 1986). According to
Dickson et al. (1988), the timing of stratification of the
water column and the resulting spring diatom outbreak
is crucial in determining the extent of primary
productivity during a particular year.

In the North Atlantic, sea-surface temperature,

precipitation, and wind and weather patterns are
strongly influenced by atmospheric circulation patterns
indexed as the North Atlantic Oscillation (NAO). High
(positive) winter (December to February) NAO index
values imply increased westerly wind strength and
mixing, increased precipitation, and milder winters.
Stronger mixing (increased winds) and nutrient levels
(increased river runoff) should favour diatoms rather
than flagellates. Low (negative) NAO index values are
associated with southerly or south-easterly winds and
with colder winters. Highly negative monthly NAO
values in summer are associated with increased seasonal
stratification, shallower mixed depths, and rising
temperatures (Pingree, 2005).
The winter NAO index values from 1998 to 2005
were highly positive in 1999 (1.70) and 2000 (2.80), but
there was a single highly negative value in 2001
(1.89). The values were 0.20 in 2003,0.07 in 2004,
and 0.12 in 2005. Consequently these NAO winter
index values close to 0 cannot be associated with the
highly positive or negative values. Negative values
could be expected, taking into account the cold winters
in 2003 and 2005. The relationship between values of
the NAO index and the meteorology in our study area is
not obvious. Consequently, the relationship between the
NAO index values and the unusual qualitative changes
of the phytoplankton composition first observed in 2003
is also not immediately apparent.
4.1. Autumnal rhizosolenioid diatom assemblage
The occurrence, persistence, and high abundance of
two species, Rhizosolenia hebetata f. semispina and
Proboscia indica, in late summer-autumn are unusual
when compared with the previous 8y time series.
These species are cosmopolitan and associated with the
summer stratification in northern European waters
(Nehring, 1998) and along the French Atlantic coast
(Paulmier, 1997). Establishing which factors caused the
appearance of these species in 2005 is not easy. For
unknown reasons, these species were very rare at our
sampling stations, whereas they were more common in
surrounding regions. The main distinctive characteristic
of our study region is its high turbulence level. A
reduction of mixing in summer-autumn 2005 could have
favoured the development of species that were more
common in stratified waters of surrounding regions. Our
monitoring stations are located near the Strait of Dover,
which is governed by a northerly tidal flux (amplitude)
of 1.06 106 m3 s 1 from the English Channel to the
North Sea, enhanced by frequent southwest winds, with

F. Gmez, S. Souissi / Journal of Sea Research 58 (2007) 283290

a high tidal amplitude (8 m maximum tidal range)

(Salomon and Breton, 1991; Prandle et al., 1993). The
tides are periodic astronomical-forced phenomena that
have little influence on the interannual variability of the
phytoplankton.
A factor that affects the reduction of water column
mixing is thermal stratification (Pingree et al., 1978).
The high temperatures in summer 2003 were exceptional (Luterbacher et al., 2004). However, this was a
short two-week heat wave that did not extend to autumn.
For example, the monthly mean air temperature in
October 2003 was 9.73 C, the lowest values in any
October of the 8y time series. In contrast, the October
of 2001 and of 2005 showed the highest mean
temperatures in the 8y time series (15.23 C and
15.00 C, respectively). The summer conditions began
unusually early in 2005, and the high temperatures
continued to late October. Consequently the summerautumn 2005 was an exceptionally long period of
sustained high temperatures, which may have favoured
deeper thermal stratification of the water column.
We hypothesise that the long periods of sustained
high temperatures resulted in an unusual period of
stability of the water column. In 2005, species of later
stages of the phytoplankton succession, such as large
Rhizosolenia diatoms, proliferated (Margalef, 1978).
However, in the northeastern English Channel during the
normal years, the diatom species succession was
interrupted by the frequent mixing events. The unusually
mild conditions in 2005 may have enabled the diatoms
with larger vacuoles, which accumulate nutrient reserves
during mixing periods, such as Proboscia indica and
Rhizosolenia hebetata f. semispina, to outcompete and
displace the normal local diatom species. It is uncertain
whether the change in the water column stratification
was the only factor that could have triggered and maintained the unusual diatom assemblage.
4.2. Northward spreading of warm-water diatoms
The current climate period is characterised by more
rapid warming to greater maxima than during any other
period of the twentieth century (Luterbacher et al.,
2004). Climate change will influence boundaries of
biogeographic regions, and in consequence the northward displacement of the limit between the borealLusitanian (southern Europe) boundaries. Two diatoms,
Eucampia cornuta and Chaetoceros peruvianus, increased their abundances to levels high enough to be
detected. Both species are considered in the literature to
be warm-water species (Hasle and Syvertsen, 1997), and
to the best of our knowledge are unknown from the

289

North and Norwegian Seas, some of the best-investigated seas in the world. Both of these species were
recorded in late summer and early autumn, coinciding
with the warmer seawater conditions.
E. cornuta was not included in the atlas of diatoms
found along the French Atlantic coasts by Paulmier
(1997). This author illustrated two species in the genus
Eucampia, the common E. zodiacus and the borealarctic species E. groenlandica Cleve. E. cornuta has the
valve face concave in broad girdle view, whereas in
E. groenlandica the valve face is convex or flat (Hasle
and Syvertsen, 1997). E. cornuta is largely cited in the
Mediterranean Sea and known from the subtropical
northeast Atlantic (Margalef, 1973) and Portugal (Moita
and Vilarinho, 1999).
Although E. cornuta could be misidentified with
E. zodiacus during routine microscopical observations,
Ch. peruvianus is a highly distinctive species and its
misidentification is most improbable. In this study,
Ch. peruvianus appeared simultaneously at the inshore
and offshore stations, indicating a wide intrusion of this
warm-water species. Ch. peruvianus is known from the
south of the British Isles (Hendey, 1974; Hartley, 1986).
In the southern English Channel, Bygrave (1911)
remarked that the inflow of Atlantic water into the
English Channel was associated with the species
Ch. peruvianus and R. hebetata f. semispina. Paulmier
(1997) did not report Ch. peruvianus along the French
Atlantic coasts. However, Paulmier (1997, p. 188)
illustrated the closely related species Chaetoceros
pendulus Karsten, which in some cases shows a
distinctive short tubular central process. Hasle and
Syvertsen (1997) considered that Ch. pendulus may be a
synonym of Ch. aequatorialis Cleve. These two species
are morphologically close to Ch. peruvianus. In any
case, these three species are warm-water species that to
the best of our knowledge are unknown from the North
and Norwegian Seas or higher latitudes.
The results of this study indicate a link between the
spread of warm-water species such as Ch. peruvianus
and E. cornuta and climatic warm episodes in the
northeast Atlantic. So far these species have shown low
abundance, and it can be assumed that they have had
little influence on the structure of the pelagic food web.
In contrast, the proliferation of an unusual assemblage
of large rhizosolenioid species, Rhizosolenia hebetata
f. semispina and Proboscia indica, could have more
consequences for the local pelagic food web, which
needs to be investigated. This indicates that climate and
meteorological changes can influence niche structure,
pre-empted by a more-competitive species (Smayda,
2002). Alterations in the species composition, as

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F. Gmez, S. Souissi / Journal of Sea Research 58 (2007) 283290

reported in this study, may ultimately be responses to

climatic fluctuations.
Acknowledgements
Samples were collected within the context of the
SOMLIT program on board RV Sepia II (INSUCNRS). We thank N. Degros, E. Lecuyer, D. Devreker
and G. Flamme for their help in sample collection, J.W.
Reid for assistance with the English, and two anonymous J. Sea Res. reviewers for their valuable comments.
This is a contribution to the French IFB Biodiversit et
Changement Global and PNEC Programme dEnvironnement Ctier programs.
References
Alheit, J., Hagen, E., 1997. Long-term climate forcing of European
herring and sardine populations. Fish. Oceanogr. 6, 130139.
Beaugrand, G., Ibanez, F., Reid, P.C., 2000. Spatial, seasonal and longterm fluctuations of plankton in relation to hydroclimatic features
in the English Channel, Celtic Sea and Bay of Biscay. Mar. Ecol.,
Prog. Ser. 200, 93102.
Bygrave, W., 1911. Report on the plankton of the English Channel in
1906. Marine Biology Association International. Fish. Invest. 3rd
Report (Southern Area), 19061908, pp. 235267.
Brylinski, J.M., Lagadeuc, Y., Gentilhomme, V., Dupont, J.P., Lafite, R.,
Dupeuple, P.A., Huault, M.F., Auger, Y., Puskaric, E., Wartel, M.,
Cabioch, L., 1991. Le fleuve ctier un phnomne hydrologique
important en Manche Orientale. Exemple du Pas de Calais. Oceanol.
Acta 11, 197203.
Dickson, R.R., Kelly, P.M., Colebrook, J.M., Wooster, W.S., Cushing, D.H.,
1988. North winds and production in the eastern North Atlantic.
J. Plankton Res. 10, 151169.
Hartley, B., 1986. A check-list of the freshwater, brackish and marine
diatoms of the British Isles and adjoining coastal waters. J. Mar.
Biol. Assoc. UK 66, 531610.
Hasle, G.R., Syvertsen, E.E., 1997. Marine diatoms. In: Tomas, C.R.
(Ed.), Identifying Marine Phytoplankton. Academic Press, San
Diego, pp. 5385.
Hawkins, S.J., Southward, A.J., Genner, M.J., 2003. Detection of
environmental change in a marine ecosystem-evidence from the
western English Channel. Sci. Total Environ. 310, 245256.
Hendey, N.I., 1974. A revised check-list of the British marine diatoms.
J. Mar. Biol. Assoc. UK 54, 277300.

Hernndez-Becerril, D.U., 1995. Planktonic diatoms from the Gulf of

California and coasts off Baja California: the genera Rhizosolenia,
Proboscia, Pseudosolenia, and former Rhizosolenia species.
Diatom Res. 10, 251267.
Jordan, R.W., Ligowski, R., Nthig, E.-M., Priddle, J., 1991. The
diatom genus Proboscia in Antarctic waters. Diatom Res. 6, 6378.
Luterbacher, J., Dietrich, D., Xoplaki, E., Grosjean, M., Wanner, H.,
2004. European seasonal and annual temperature variability,
trends, and extremes since 1500. Science 303, 14991503.
Margalef, R., 1973. Fitoplancton marino de la regin de afloramiento
del NW de frica. Res. Exp. Cient. B/O Cornide 2, 6594.
Margalef, R., 1978. Life-forms of phytoplankton as survival alternatives in an unstable environment. Oceanol. Acta 1, 493509.
Moita, M.T., Vilarinho, M.G., 1999. Checklist of phytoplankton
species off Portugal: 70 years (19291998) of studies. Port. Acta
Biol., Sr. B : Sist. Ecol. Biogeogr. Paleontol. 18, 550.
Nehring, S., 1998. Establishment of thermophilic phytoplankton
species in the North Sea: biological indicators of climatic changes?
ICES J. Mar. Sci. 55, 818823.
Paulmier, G., 1997. Atlas des diatomophyces des ctes franaises et
des aires ocaniques adjacentes. Direction des Ressources
Vivantes de lIFREMER. RI-DRV 97-14, RH/L'Houmeau.
Pingree, R., 2005. North Atlantic and North Sea climate change: curl
up, shut down, NAO and ocean colour. J. Mar. Biol. Assoc. UK 85,
13011315.
Pingree, R.D., Holligan, P.M., Mardell, G.T., 1978. The effects of
vertical stability on phytoplankton distributions in the summer on
the northwest European shelf. Deep-Sea Res. 25, 10111028.
Prandle, D., Loch, S.G., Player, R., 1993. Tidal flow through the
Straits of Dover. J. Phys. Oceanogr. 23, 2337.
Robinson, G.A., Hunt, H.G., 1986. Continuous plankton records:
annual fluctuations of the plankton in the western English Channel,
195883. J. Mar. Biol. Assoc. UK 66, 791802.
Salomon, J.C., Breton, M.M., 1991. Courants de mare et courants
rsiduels dans la Manche. Oceanol. Acta 11, 4753.
Schr, C., Vidale, P.L., Lthi, D., Frei, C., Hberli, C., Liniger, M.A.,
Appenzeller, C., 2004. The role of increasing temperature
variability in European summer heatwaves. Nature 427, 332336.
Smayda, T.J., 2002. Turbulence, watermass stratification and harmful
algal blooms: an alternative view and frontal zones as pelagic
seed banks. Harmful Algae 1, 95112.
Vandentorren, S., Suzan, F., Medina, S., Pascal, M., Maulpoix, A.,
Cohen, J.C., Ledrans, M., 2004. Mortality in 13 French cities
during August 2003 heat wave. Am. J. Public Health 94,
15181520.
World Meteorological Organization, 2005. WMO statement on the
status of the global climate in 2005. Geneva, Switzerland. WMONo. 998, http://www.wmo.int.