J. Nannoplankton Res. 30 (1), 2008, pp.9-21 © 2008 International Nannoplankton Association, Inc.
ISSN 1210-8049 Printed by Cambridge University Press, UK

9Spatial distribution of living coccolithophores along an eastwest transect in the subtropical South Atlantic
Karl-Heinz Baumann*, Babette Boeckel, Martin Čepek
University of Bremen, Geoscience Department, PO Box 330440, D-28334 Bremen, Germany; *baumann@uni-bremen.de

Manuscript received 29th November, 2007; revised manuscript accepted 8th February, 2008

Abstract The upper-water-column distribution of living coccolithophores in the subtropical South Atlantic and
Benguela Upwelling regime was examined along an E-W transect of 11 phytoplankton stations. Coccolithophore
standing crops ranged from less than 10 000 cells/l in the nutrient-depleted, shallow-water stations of the western Subtropical Gyre to 396 300 cells/l in the medium-depth stations of the nutrient-enriched Benguela Current. Emiliania huxleyi was the most abundant species, with concentrations of up to 200 000 individuals/l. The present study reveals
hydrographic controls on the distribution of some of the major taxa. Assemblages in the Benguela Current were characterised by at least five species (E. huxleyi, Gephyrocapsa ericsonii, Rhabdosphaera xiphos, Umbellosphaera irregularis, Syracosphaera halldalii), which exhibit their highest cell numbers at the easternmost stations, reflecting higher
nutrient levels. The upper photic zone (0-80m water-depth) of the Subtropical Gyre generally contained holococcolithophores and the species Discosphaera tubifera, Umbellosphaera tenuis, Calciosolenia brasiliensis and Syracosphaera pulchra, which exhibit an affinity for lower nutrient-levels. The flora in the lower photic zone (>80m-100m
water-depth) was characterised by abundant Florisphaera profunda in the nutrient-depleted Subtropical Gyre, whereas
Calcidiscus leptoporus, Oolithotus fragilis and Umbilicosphaera sibogae occurred in the deep boundary-zone from the
Benguela Current to the Subtropical Gyre waters. Thus, the vertical distribution of all coccolithophore taxa, except thethree placolith-bearing species Gephyrocapsa oceanica, G. ericsonii and E. huxleyi, was probably controlled by upperphotic-zone water-temperature and stratification of the water-column.

Keywords South Atlantic, living, ecology, plankton
1. Introduction
Phytoplankton, at the base of the marine food-chain, serve
as a key to understanding both marine ecology and biogeochemistry. However, to integrate palaeobiology andmodelling of climatic changes into this study requires a detailed understanding of the ecology of these organisms.
Critical to such an understanding is knowledge on the extent of, and controls on, species-level biodiversity. Coccolithophores are the predominant group of calcifying
marine phytoplankton, have the best fossil record of all
phytoplankton, and play a unique role in the global carbon
cycle. They are characterised by calcareous scales (coccoliths), which surround the living cell to form an extracellular covering called a coccosphere (e.g. Winter &Siesser, 1994; Young, 1994). Classification of both living
and fossil coccolithophores is based exclusively on coccolith morphology and, at species level, on fine variations
in size and shape (Jordan & Green, 1994; Young et al.,2003).
General aspects of coccolithophore biogeography andhabitat are known from taxonomic surveys of the plankton
and of bottom-sediments (e.g. Knappertsbusch, 1990;
Samtleben & Schröder, 1992; Kleijne, 1993; Winter et al.,
1994; Samtleben et al., 1995; Baumann et al., 2000;
Hagino et al., 2000; Cros, 2001; Triantaphyllou et al.,2002). Individual species are mostly cosmopolitan, butwith more or less limited latitudinal distributions. Thus,
broad biogeographic coccolithophore zones are recognised(McIntyre & Bé, 1967; Okada & Honjo, 1973; Winter etal., 1994). In part, these latitudinal zones reflect tempera-ture tolerance but, even at this level, it is apparent that otherfactors are of major significance. For instance, the presenceof nannofloral zones associated with equatorial upwelling
and the subtropical oligotrophic gyres suggest control bynutrients/trophic level, rather than temperature alone. However, although much information is available on the oceanic
scale (e.g. McIntyre & Bé, 1967; Okada & Honjo, 1973;
Honjo & Okada, 1974; Kleijne, 1991, 1992, 1993; Hagino
et al., 2000), as well as of smaller areas (e.g. Winter et al.,
1979; Reid, 1980; Winter, 1985; Mitchell-Innes & Winter,
1987; Samtleben & Schröder, 1992; Ziveri et al., 1995a, b;
Baumann et al., 1999, 2000; Cros, 2001; Triantaphyllou etal., 2002, 2004), the environmental parameters that control
coccolithophore distribution are still poorly understood.
This reflects, in part, a shortage of suitable studies on natural populations. Much of the available data deal with relative abundances in coccolithophore assemblages, which
are likely biased. Little work has been done so far on absolute numbers or on absolute abundances of single species
relative to ecological parameters in modern communities(e.g. Venrick, 1982). In addition, in many studies, only surface-water samples (e.g. from 5m or 10m water-depth)
have been investigated.
Nevertheless, the eastern part of the study area is one of
the few regions where quite a number of coccolithophorestudies have been conducted during the last 15 years. Mostof these studies dealt with the Benguela Upwelling regime
off SW Africa. Distinct species assemblages have beenshown to be strictly associated with the various surface andsubsurface water-masses and related hydrological fronts

10

Baumann, Boeckel, Čepek

generated by the upwelling processes (e.g. Friedinger &Winter, 1987; Giraudeau et al., 1993; Giraudeau & Bailey,
1995). The temporal variations in coccolithophore populations also mirror the seasonal changes of the main hydrographic conditions off SW Africa (Giraudeau et al., 2000;
Romero et al., 2002). In turn, the highest coccolith fluxes
are inversely correlated with fluctuations in sea-surface
temperature and, thus, with the seasonal dynamics ofcoastal upwelling occurrence. Results obtained from surface-sediment studies also indicate that the coccolith distribution seems to be temperature- and nutrient-controlled,
covarying with the seaward extension of the upwelling filament zone in the Benguela region (Giraudeau, 1992; Baumann et al., 1999; Boeckel & Baumann, 2004). In contrast,
coccolithophores have been very scarcely investigated in
the oligotrophic South Atlantic since McIntyre & Bé
(1967), and data on living coccolithophores in this area are
extremely limited (Boeckel & Baumann, 2008). The
nannoplankton distribution to the south of Africa has beenstudied (Verbeek et al., 1989), but beyond that, only the occurrence and distribution of the species in surface-sediments has been investigated (Mostajo, 1984; Boeckel et al.,2006).
Therefore, the aims of this study were: 1) to investigate
the coccolithophore species composition in an E-W transect, from the nutrient-enriched waters of the Benguela
Current towards the nutrient-depleted waters of the Subtropical Gyre; (2) to examine and correlate the species occurrences to the means of phosphate and nitrate content, as
well as temperature, in order to add some precision to thehabitat preferences of the species; and finally (3) to study
the depth-distribution of the species in each of the oceanic
regimes.
2. Modern oceanography of the study
area
The oceanic upper-layer circulation of the eastern South
Atlantic has been summarised in a number of comprehensive reviews (Lutjeharms & Meeuwis, 1987; Peterson &Stramma, 1991; Reid, 1996; Shannon & Nelson, 1996), so
only a brief summary is given here.
An important feature of the South Atlantic circulation
system (Figure 1) is the cross-equatorial transport of warm
surface-waters from the Indian Ocean and from the anticyclonic South Atlantic Gyre into the North Atlantic Ocean,by means of the South Equatorial Current. In the easternSouth Atlantic off SW Africa, the surface-water circulation
is dominated by the northward-directed Benguela Coastal
Current, the coastal tongue of the Benguela Current (BC),and the warmer, southward-flowing Angola Current (Figure 1). The Benguela Coastal Current and Angola Current
converge between 14˚ and 16˚S, forming a marked front
on the shelf (Angola-Benguela Front), which is well-defined in terms of both temperature and salinity (Shannon& Nelson, 1996). In addition, the prevailing winds in thisregion in turn drive an offshore surface drift and cause
coastal upwelling of cold, nutrient-rich water, especially

Figure 1: Schematic map of the main circulation in the upper level of theSouth Atlantic Ocean (redrawn mainly from Peterson & Stramma, 1991).
Locations of the water-stations are indicated as black dots. AC – Agulhas
Current, BC – Benguela Current, SG – Subtropical Gyre

during austral winter. Upwelling occurs in a number of upwelling cells, south of about 18˚S, with a major, semi-permanent cell at 27˚S (Shannon & Nelson, 1996). This
upwelling leads to enhanced biological productivity off
Namibia. The typical westward extent of the upwelling is
between 150 and 250km off the coast.
Further offshore, the north-westward-flowing Benguela
Oceanic Current, which is the oceanic portion of the BC, ischaracteristic of the upper-layer waters. This flow feeds
into a broad, north-westward-flowing South Equatorial
Current, forming the eastern limb of the Subtropical Gyre
(SG). The South Equatorial Current itself consists of two
branches, a main stream flowing south of 10˚S and a tradewind-forced smaller, faster-flowing branch, between 2˚ and4˚S (Peterson & Stramma, 1991). At about 10˚S, off Brazil,
the South Equatorial Current splits into two branches, thesouthward-flowing Brazil Current and the northward-flowing North Brazil Current (Figure 1; Peterson & Stramma,1991). The Brazil Current transports relatively warm surface-water southwards, until it encounters the cold, northward-moving Malvinas Current. This generates the highly
dynamic Brazil-Malvinas Confluence that causes the eastward deflection of the Brazil Current into the South Atlantic Current. At the southern border of the Subtropical
Front, the warmer South Atlantic Current waters parallel
the cold and nutrient-rich Antarctic Circumpolar Current
in the south. The boundary between subtropical and subantarctic waters might extend regionally over a large area
between 30˚ and 45˚S (Smythe-Wright et al., 1998). Off
the south-western tip of Africa, the SAC meets the westward-directed Agulhas Current, which consists of warmand saline Indian Ocean water. At about 10˚E, the SAC deflects northward and merges into the BC. This southern endof the Benguela system, the retroflection of the Agulhas
Current, is responsible for a substantial amount of warmand salty water being transported from the Indian Ocean to
the South Atlantic.

Living coccolithophores...subtropical South Atlantic

The vertical distribution of temperature and nutrients(phosphate and nitrate) down to 200m water-depth are displayed along an E-W transect at 32.5˚S (Figure 2). Nutrient data were obtained using the World Ocean Atlas(Conkright et al., 1994). The data used in this study aremonthly means, so that the limited number of data-points
result only in a rather low resolution of nutrient and temperature in the presented transect. Monthly mean temperature data for February yields a rather consistent variation
throughout the investigated transect. A slight increase in
sea-surface temperatures, from 16-20˚C to 20-24˚C in theuppermost 100m of water-depth, is observed toward thewest. Phosphate concentrations of up to 0.3μmol/l at thesurface and >0.5μmol/l at a depth of >100m in the BC wereslightly higher than in the SG, where surface values ranged
from 0.1-0.2μmol/l. A similar distribution was observed fornitrate concentrations (Figure 2). In the BC, the nitrate content ranged from 2.0μmol/l at the surface to >7.0μmol/l at
a depth of >150m, whereas the values in the SG extended
from 0.5-1.5μmol/l at the surface (<50m water-depth) to
2.0-5.0μmol/l at a water-depth of >100m.
3. Material and methods
A total of 51 water-samples, from between the depths of
0m and 200m, were collected from 11 stations. The samples were taken during various R/V Meteor cruises in different years (see Table 1), forming an E-W transect
between SW Africa and SE South America (Figure 1).
Sample stations were located from about 29˚S to 37˚S andfrom 14˚E to 51˚W (Table 1). Although the gaps in sampling periods between the cruises may present difficulties
in directly comparing the coccolithophore assemblages, thedatasets were combined in order to gain a complete coverage of the various water-masses in the central South Atlantic.
Water-samples of generally 2l volume were taken byhydrocasts attached either to a multinet or to a rosette(Hydro-Bios, Kiel). Onboard, the sea-water was filtered
immediately through cellulose nitrate filters (SartoriusTM,
47mm or 25mm diameter, 0.45μm pore-size) and dried for
24 hours at 50˚C. The filters were then stored in plastic
Petri dishes and were shrink-wrapped with silica gel in film
to protect them from humidity.

11

In the laboratory at Bremen University, a piece of eachfilter was fixed, with double-sided adhesive tape, to an aluminium stub. These stubs were sputtered with gold and thefilters were examined quantitatively for coccolithophores
with a scanning electron microscope (SEM; Zeiss DSM
940A). During counting, each sample was examined at amagnification of 3000x in transects across the filter area.
Whole coccospheres, as well as coccoliths, were counted.
Identification and taxonomy of species follows that outlined by Kleijne (1993), Jordan & Kleijne (1994) and Winter & Siesser (1994). Thus, the taxonomy of the species
Emiliania huxleyi, and especially Calcidiscus leptoporus, isthe old, classical one – morphotypes and genotypes (e.g.
Young & Westbroek, 1991; Hagino et al., 2005; Hagino &Okada, 2006) have not been distinguished, since part of thedata stem from counts that were performed as part of a PhD
project over 10 years ago (Čepek, 1996).
Coccolith numbers were converted to coccosphere units
to obtain the number of individuals/l of sea-water. Therefore, numbers of coccoliths on a coccosphere of each of thedifferent species were counted. For each species, a mean
was then calculated. For species that were not observed as
coccospheres in the studied samples, the data was takenfrom the literature (Table 2). Coccolithophore cells/l of seawater were then calculated as follows:
Cell numbers per litre =

C*A
a*V

where C is the number of coccospheres, A is the entire areaof the filter, a is the counted area of the filter, and V is thevolume of water filtered. The data presented in this paperarearchived
inthe
PANGAEA
database(www.pangaea.de).

4. Results
4.1 Cell density and species diversity of
total coccolithophores
Coccoliths were found in all of the studied samples, andintact coccospheres were also present in most of the samples. In the upper water-column of the oligotrophic gyre
regime, the standing crop of coccolithophores was relatively low (<50*103 individuals/l), and the species diversity
was at its lowest. In contrast, in the vicinity of the temperate BC, high coccolithophore cell densities
of up to 396* 103 individuals/l were obCruise
Month/Year GeoB
Position
Sampling depths (m)Stationserved (Figure 3, Table 3), and the species_________________________________________________________________________ diversity was relatively high in comparison
to the SG. The highest species diversity was
M23-1
02/19932005
28˚53.7’S / 14˚09.4’E
0, 20, 40, 100
found at Stations GeoB 2008 (50m waterM23-1
02/19932006
29˚10.9’S / 13˚07.5’E
0, 20, 40, 60, 80, 100
M23-1
02/19932008
31˚06.1’S / 11˚42.9’E
25, 50, 100
depth) and GeoB 2012 (20m water-depth),M23-1
02/19932011
35˚36.6’S / 08˚15.8’E
20, 40, 60, 80, 100
which each yielded 34 coccolithophore taxa.
M23-1
02/19932012
33˚02.7’S / 03˚20.3’E
0, 20, 40, 60, 80, 100
The highest diversities generally occurred at
M23-1
02/1993201732˚50.6’S / 02˚23.4’W
0, 20, 40, 60, 80, 100
stations in the transition zone from the BC
M34-3
02/19963801
29˚30.0’S / 08˚17.6’W
20, 50, 150, 200
M34-3
02/19963808
30˚48.6’S / 14˚42.7’W
20, 50, 100, 150, 200
to the SG waters. In addition, the highest
M46-4
03/20006430
31˚27.6’S / 24˚30.9’W
5, 20, 50, 100, 150
numbers of total holococcolithophores (upM46-4
03/20006435
31˚14.7’S / 39˚20.0’W
5, 10, 50, 100, 150
to 31*103 individuals/l) also occurred in the
M46-2
12/19996229
37˚13.7’S / 51˚41.3’W
10, 20, 50
upper water-column of this area (Figure 3).

Table 1: Geographical positions and sampling depths of all GeoB stations

12

Baumann, Boeckel, Čepek

Figure 2: Maps of (a) temperature, (b) nitrate concentration and (c) phosphate concentration (all mean annual at 75m depth), as well as the corresponding transect at 32.5˚S (mean monthly data). Hydrographic data are available from the World Ocean Atlas (1994; see http://ingrid.ldgo.columbia.edu/SOURCES/.LEVITUS94/)

4.2 Distribution of coccolithophore
assemblages and species
A total of 70 species was identified. The assemblages comprised 51 heterococcoliths and 19 holococcoliths. However,
only 10 heterococcolithophore species, with the holococcolithophores, reached more than 10% of the nannoflora, in

at least one of the studied samples. Of these, Emiliania huxleyi was the most abundant species in many of the samples(generally 10-65%). From the floral composition and thespecies cell densities, three broadly-defined coccolithophore assemblages were identified, which can be correlated to the BC, the oligotrophic SG, and the lower photic

Living coccolithophores...subtropical South Atlantic
Species

Coccoliths per
Data source
coccosphere_________________________________________________________________
Acanthoica acanthifera
Calcidiscus leptoporus
Calciosolenia brasiliensis
Discosphaera tubifera
Emiliania huxleyi
Emiliania huxleyi
Florisphaera profunda
Gephyrocapsa ericsonii
Gephyrocapsa oceanica
Oolithotus fragilis
Rhabdosphaera clavigera
Rhabdosphaera xiphos
Syracosphaera halldalii
Syracosphaera pulchra
Umbellosphaera irregularis
Umbellosphaera tenuis
Umbilicosphaera sibogae

42
24
90
55
single-layered: 12
multi-layered: 44
65
16
16
37
42
45
44
26
20
22
80

Winter & Siesser (1994)
this study
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this study
this study
this study
Kleijne (1993)
this study
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this study

13

addition to the species mentioned above, E.
huxleyi also occurred in relatively high abundances (up to 40%) in this regime.
The nannoflora in the lower photic zone(>80-100m water-depth) is characterised byabundant Florisphaera profunda (up to 52%)in the nutrient-depleted western part of the SG(Figure 7), whereas common to minor occurrences of Calcidiscus leptoporus (up to 20%),
Oolithotus fragilis (up to 17%), and Umbilicosphaera sibogae (up to 10%) occurred in theboundary zone from the BC to the SG waters.
Unfortunately, sampling at the easternmost stations never exceeded 100m water-depths, sothat the depth-distribution in the BC region remains uncertain. E. huxleyi and Gephyrocapsa
oceanica, although exhibiting their highest cellnumbers in the middle photic BC region, wereconstrained to deeper waters in the warm, oligotrophic stations of the SG (Figure 4).

Table 2: Number of coccoliths per coccosphere of the most common species, as used inthis study

5. Discussion
5.1 Standing crop
Major changes in coccolithophore diversity, and especiallyin cell density, occur across the transition of the BC to theSG, which is associated with shifts in sea-surface temperature and nutrients. The coccolithophore cell numbers observed in the BC regime were significantly higher thanzone. A more distinct differentiation of the assemblages ishampered by the relatively low depth-resolution, as well as
the longitudinal resolution of the samples.
The predominant group in the BC (Figures 4, 5) consists of at least five species (E. huxleyi, Gephyrocapsa ericsonii,
Rhabdosphaera
xiphos,Umbellosphaera irregularis, Syracosphaera halldalii), all of which exhibit
their highest cell numbers at the easternmost stations (2005, 2006, 2008). In thisregime, the coccolithophore assemblage
was dominated by E. huxleyi (29-62%),
reaching its highest numbers in middepth waters (30-70m water-depth). In
addition, G. ericsonii (up to 32%), U. irregularis (up to 16%) and R. xiphos (upto 12%) were particularly common components of the nannofloras.
The SG group consists of fivespecies and species groups (Discosphaera tubifera, Umbellosphaera
tenuis, Calciosolenia brasiliensis, Syracosphaera pulchra, Acanthoica spp.), together with the total holococcolithophores. Although all of these taxa reach
their highest cell densities at stations in
the transition zone from the BC to the
SG waters, they all show high relative
abundances in the upper water-column(<80m water-depth) of the oligotrophic
SG (Figure 6). In particular, U. tenuis(up to 66%), D. tubifera (up to 18%),
and S. pulchra (up to 10%) were com- Figure 3: Vertical distribution of total standing crop of coccolithophores (above), and cell nummon components of the nannoflora. In bers of total holococcolithophores (below) along the studied transect. Hatched area represents
absence of taxa

14

Baumann, Boeckel, Čepek

Subtropical Gyre
Benguela Current
Cell density * 103 cells/lWaterTaxa
Cell density * 103 cells/lcolumn
min
max
mean
min
max
mean
______________________________________________________________________________

austral winter and summer. Therefore, it can be inferred that the goodcorrespondence of the present celldensities at the sites within the BC
entiretotal coccolithophores 7.3
302.4
109.1
77.6
396.3
170.6
with those previously reported fromE. huxleyi
0.6
129.9
31.8
34.4
199.7
76.7
the Namibian and South African
G. ericsonii
0.1
4.8
31.2
3.2
85.7
34.2
shelves reflects the influence of filaG. oceanica
0.1
2.1
0.3
0.1
2.3
0.8
ments which supply nutrients to the
UPZ
Acanthoica spp.
0.1
6.4
1.1
4.2
4.2
0.6
surface-waters, resulting in higher(0-<80m) C. brasiliensis
0.1
6.6
0.7
0.2
5.4
2.5
numbers of mainly placolith-bearing
D. tubifera
0.1
4.4
1.3
0.5
2.7
1.1
species.
R. xiphos
0.1
16.4
2.1
2.5
24.8
13.9
Placolith-bearing species occur
S. halldalii
0.1
16.7
1.9
0.1
17.9
5.2
S. pulchra
0.1
4.3
1.3
0.1
6.4
1.1
in much higher cell densities thanU. irregularis
0.1
27.9
1.8
0.2
36.6
9.6
umbelliform species, which typicallyU. tenuis
1.6
84.1
24.0
0.2
23.1
6.1
inhabit oligotrophic regimes (e.g.
holococcolithophores 0.8
30.6
8.6
1.4
10.7
4.5
Young, 1994; Haidar & Thierstein,
LPZ
C. leptoporus
2.3
30.5
6.0
2.3
17.9
2.3
2001; Hagino & Okada, 2006). Thus,(80-100m) F. profunda
0.5
34.6
7.3
2.2
12.7
5.4
a generally low standing stock was
Oolithotus spp.
0.7
41.0
12.1
0.1
0.7
0.3
observed, especially in the upper
U. sibogae
0.1
10.7
2.8
0.2
2.0
0.6
photic zone (<80m water-depth) of
the SG. Here, the coccolithophore
Table 3: Comparison of the most important nannofloral elements of the Subtropical Gyre andflora was rather similar to previBenguela Current. Minimum, maximum and mean absolute cell numbers within the upper and lower
ously-published assemblages fromphotic zone are displayedcomparable oceanic settings (e.g.
Hagino et al., 2000; Kinkel et al.,those in the oligotrophic gyre (Figure 3, Table 3). Althoughlocated far away from the actual upwelling, total cell den- 2000), in terms of cell density and the dominance of typisities are in the order of those previously reported from the cally non-placolith-bearing species. The extremely low
Namibian upwelling system (Giraudeau et al., 1993). Coc- coccolithophore cell densities in the upper photic zone
colithophore standing crops reach up to 466*103 cells/l, the (<50m water-depth) may be explained by the extremely
maximum cell density occurring at the shallow-water sta- low nitrate concentrations during the sampling period intions (Giraudeau et al., 1993). The
presented data is also in good
agreement with the results of Giraudeau & Bailey (1995), who examined coccolithophores in an
upwelling cell off Hondeclip Bay
and calculated total cell densities
of up to 278*103 cells/l. However,water-samples from a nearby region, investigated by MitchellInnes & Winter (1987), revealed
greater differences, compared toour data. The authors observed assemblages of up to 2340*103 cells/lin mature, upwelled water, adjacent to an upwelling cell off theCape Peninsula. The influence ofthe coastal upwelling cells on theoffshore waters, through offshore
migration of chlorophyll filaments,
is strongest during austral summer(Romero et al., 2002). During thistime of the year, the coccolith flux,
as observed in a mooring site close
to the Namibian Upwelling (atabout 29˚S, 13˚E), shows bimodal
Figure 4: Vertical distribution of Emiliania huxleyi and Gephyrocapsa oceanica (absolute cell numbers)seasonality, with major peaks in along the studied transect. Hatched area represents absence of taxa

Living coccolithophores...subtropical South Atlantic

15

austral summer.
Cell densities in the SG increased
below
80-100m
water-depth.
Florisphaera profunda dominated thecoccolithophore assemblage of the lower
photic zone and was largely responsible
for this increase in cell density. This
species is known to occur in relatively
high abundances, even under relatively
low-nutrient conditions (Haidar & Thierstein, 2001). In addition, the highest concentrations of coccolithophores wereobserved at the thermocline in well-stratified waters (e.g. Hagino et al., 2000).
The upper or lower vertical distribution
limits of many coccolithophore taxa coincided with the top of the thermocline.
However, species such as Emiliania huxleyi and Gephyrocapsa oceanica, which
exhibit their highest cell numbers in the
middle photic zone of the BC region,
also contributed to the lower-photic assemblage in the warm, oligotrophic stations of the SG (Figure 4).
5.2 Environmental controls
on coccolithophore species
distribution
The present study reveals hydrographic
controls on the distribution of some of
the major taxa. The nannofloras in the
nutrient-enriched stations of the BC are
characterised by high cell numbers ofEmiliania huxleyi and Gephyrocapsa ericsonii, while the other species occur inmuch lower absolute cell numbers.
Whereas E. huxleyi was often found to
dominate assemblages in temperate toarctic waters (e.g. Mitchell-Innes & Winter, 1987; Kleijne, 1993; Giraudeau &Bailey, 1995), another Gephyrocapsa
species, G. oceanica, is well known toalso prefer eutrophic waters (Kleijne,
1993; Andruleit et al., 2000). However,G. oceanica only occurs in extremely
low cell numbers in the studied samples,
whereas G. ericsonii is numerically Figure 5: Vertical distribution of cell numbers of the species of the Benguela Current assemmuch more abundant and exhibits a dis- blage (Gephyrocapsa ericsonii, Umbellosphaera irregularis, Rhabdosphaera xiphos, Syratribution pattern rather similar to that of cosphaera halldalii) along the studied transect. Hatched area represents absence of taxa
E. huxleyi. Nevertheless, not much is
known about the ecology of G. ericsonii. Winter et al. conditions. This is also supported by the correlation of cell(1979) reported G. ericsonii as a predominant nanno- numbers and nutrient concentrations in the present study
plankton species, along with E. huxleyi, in the Gulf of Elat, (Figure 8). In addition, a clear correlation of higher cellwith a preference for near-coastal currents with high nutri- numbers with decreasing temperatures can be seen, indient-contents. Work on sediment cores by Takahashi & cating a preference for eutrophic, upwelling conditions.
E. huxleyi is known to be the most ubiquitous of all livOkada (2000) and Okada & Wells (1997) from off westerning
coccolithophore species (e.g. McIntyre & Bé, 1967;
Australia also suggest that G. ericsonii prefers eutrophic

16

Baumann, Boeckel, Čepek

Figure 6: Vertical distribution of cell numbers of the upper-photic-zone species of the Subtropical
Gyre (Discosphaera tubifera, Umbellosphaera tenuis, Syracosphaera pulchra, Calciosolenia
brasiliensis, Acanthoica spp.) along the studied transect. Hatched area represents absence of taxa

Winter & Siesser 1994; Ziveri et al.,1995a; Hagino et al., 2000). It tolerates a temperature range from 1˚C to30˚C, although its highest abundances occur at water-temperaturesof <23˚C (Haidar & Thierstein,2001), and it is common in eutrophic
as well as oligotrophic conditions. In
the present study, the highest abundance of E. huxleyi appeared in the
BC at temperatures comparable tothose of previous investigations (e.g.
Haidar & Thierstein, 2001; Hagino& Okada, 2006). Nevertheless, theabundances at the other stations werehigh as well, so this species could becharacterised as a euryoecious
species, having a wide range of habitat tolerance. In addition, G. oceanica exhibits a rather similar
distribution to E. huxleyi. Although
cell numbers are extremely low, thespecies appeared with elevated cellnumbers in the upper photic layer inthe BC, whereas in the SG, G. oceanica was mainly observed below thethermocline. The species is also regarded as a eutrophic species (Kleijne et al., 1989; Kleijne, 1993), but it
is also adapted to higher sea-surface
temperatures (Hagino et al., 2000;
Hagino & Okada, 2006). As shownby Brand (1994), coccolithophores
such as E. huxleyi and G. oceanica
tend to live at greater depths in thecentral gyres because of the higher
nutrient availability at these depths.
This finding is supported by the observations of Hagino et al. (2000),
who found similar distribution patterns of these species in the Pacific.
The authors explained the distribution by invoking different morphotypes, some occurring in the photic
zone of the dynamic, eutrophic,
mixed-water regimes, whereas othermorphotypes occurred below thethermocline in the well-stratified stations.
The coccolithophore flora in
the surface layer of the well-stratified
SG could be distinguished from theBC nannoflora by the higher abundances of Discosphaera tubifera.
This species generally occurred outside the upwelling areas, and has
been referred to as part of the sub-

Living coccolithophores...subtropical South Atlantic

17

tropical zone flora (Jordan &Chamberlain, 1997), which isconsistent with our observations.
D. tubifera is reported to live atwater-depths down to 40-80m in
oligotrophic environments (e.g.
Honjo & Okada, 1974; Reid,
1980; Hagino et al., 2000; Malinverno et al., 2003), also rather
consistent with our observations(Figure 9). The same pattern isdisplayed by Syracosphaera pulchra, Calciosolenia brasiliensis
and Umbellosphaera tenuis, allof which are non-placoliths
known to prefer stable, stratified
waters (Okada & McIntyre,
1977; Hagino et al., 2000).
Umbilicosphaera irregularis
has also been reported as a typical oligotrophic species (e.g.
Young, 1994; Baumann et al.,
1999; Haidar & Thierstein,2001). Its occurrence in the BC
stations is surprising, and contradicts the usual notion. However, U. irregularis has beenfound in the Pacific Ocean, bothat stations with extremely hightemperatures (>30˚C) and highnutrient levels (Hagino et al.,2000), as well as in the usual
warm, oligotrophic environments. Hagino & Okada (2006)
also showed that the concentration of this species generally had
no relation to nutrient concentrations, except that it is absent
when the phosphate concentration is relatively high. And therefore, the threshold of >0.7μmol/l
was probably not reached in thestudied BC stations. It is notablethat the highest numbers of U.
tenuis occurred in the transitionzone from the BC to the SG. The
correlation of its cell numberswith nitrate concentration
clearly demonstrates that thisspecies is adapted to low nutri- Figure 7: Vertical distribution of cell numbers of the deep-dwelling species (Florisphaera profunda,
Oolithotus fragilis, Umbilicosphaera sibogae, Calcidiscus leptoporus) along the studied transect. Hatched
ent levels. Nevertheless, the tol- area represents absence of taxa
erance of some morphotypes ofU. tenuis (Kleijne, 1993; Hagino & Okada, 2006) for Baumann, 2008).
Calcidiscus leptoporus, Oolithotus fragilis and Umbilslightly nutrient-enriched, and possibly relatively stratified,
icosphaera
sibogae have been regarded as rather oligwaters cannot be excluded. So far, it is known that the SG
otrophic
speciesin previous investigations (Okada &assemblage within the upper 50m of the photic zone is
Honjo,
1975;
Okada& McIntyre, 1979; Mitchell-Innes &mainly composed of U. tenuis Types III and IV (Boeckel &18

Baumann, Boeckel, Čepek

well as the typical lower photic zone
(LPZ) assemblage (Figure 9). The
UPZ assemblage can generally berecognised until about 80m, to a maximum 100m, of water-depth, and ischaracterised by the highest coccolithophore cell numbers. Typical members of this group are species of the
Syracosphaerales, the Rhabdosphaeraceae (Discosphaera tubifera, Rhabdosphaera xiphos, Acanthoica spp.),
Syracosphaeraceae (Syracosphaera
halldalii, S. pulchra shown as examples), Umbellosphaeroideae (Umbellosphaera tenuis, U. irregularis), as
well as all of the holococcolithophore
species observed in this study. Thus,
the composition is rather similar to
those previously published as UPZ assemblages (e.g. Winter et al., 1994;
Jordan & Chamberlain, 1997; Hagino
et al., 2000; Malinverno et al., 2003).
Some of the species, particularly Calciosolenia brasiliensis, D.
tubifera, Acanthoica spp. and Gephyrocapsa ericsonii, do exhibit theirhighest cell numbers in the lowermost
part of this UPZ. Some of them are already known to occupy intermediate
Figure 8: Correlation of nitrate concentration, water temperature (from conductivity-temperaturewater-depths, and have been termed
depth [CTD] casts) and absolute cell numbers of the most common species. Monthly mean values
middle photic zone (MPZ) assemof nitrate at each station were obtained from the World Ocean Atlas (Conkright et al., 1994). The
blages, which are usually found bescale in the middle refers to the size of the bubbles (= absolute cell numbers)
tween 80-120m of water-depth
Winter, 1987; Giraudeau, 1992), which is supported by our (Okada & Honjo, 1973; Jordan & Chamberlain, 1997).
study. All of them showed their highest abundances below However, since all of these species are also present in thethe thermocline (80m-100m), in the transition zone be- upper photic layer, a further differentiation has not been
tween the BC and SG regimes. High abundances of U. si- made in this study. Unlike the LPZ nannoflora, there is nobogae in the upper photic zone have previously been hydrographic restriction for the MPZ nannoflora to moveobserved in the Gulf of Elat (Winter et al., 1979). Never- (or be moved) into shallower waters. In addition, the optitheless, these species are regarded as deep-living taxa, ba- mal depth-range of the species may vary due to changing
sically influenced by water-mass stratification and nutrient environmental conditions during different seasons of alevel.
year, when, for example, surface-water stratification may
Modern holococcolithophores have been intensively be more pronounced (Reid, 1980). For this area of coccolinvestigated with respect to their geographical distribution ithophore distribution, more study and detailed sampling(Kleijne, 1991). Their highest absolute abundances occur in across the thermocline is needed.
the Mediterranean Sea and in the NE Atlantic Ocean, andIn addition, most of the lower-photic dwellers are abunare associated with lower nutrient levels. This generally dant in tropical to subtropical regions, regardless of watercorresponds well with the results of the present study, with stratification, as has been commonly observed in previous
holococcolithophores reaching their highest abundances at studies (e.g. Hagino et al., 2000). In particular,
low nutrient concentrations.
Florisphaera profunda is well known to occupy this nichein the LPZ regime (Jordan & Chamberlain, 1997) and also
5.3 Coccolithophore depth-distribution
dominates the samples studied in the SG. Its cell density
Typical depth-related assemblages can be identified from drastically decreased eastwards, and is low in the area ofthe depth-distribution of the species, although the entire the BC. As reported previously (e.g. Okada & Honjo, 1973;
photic zone was not studied at high resolution. However, it Jordan & Chamberlain, 1997; Hagino et al., 2000; Malinis in fact possible to identify two main assemblages in the verno et al., 2003), other species, such as Oolithotus fragphotic zone, an upper photic zone assemblage (UPZ) as ilis, are mainly restricted to the LPZ. In addition,

Living coccolithophores...subtropical South Atlantic

Calcidiscus leptoporus and, especially, Umbilicosphaera
sibogae were observed as deep-living species (Figure 9).
Emiliania huxleyi, present in cell numbers similar to thoseof F. profunda, as well as G. ericsonii and G. oceanica, also
occurred in the LPZ of the SG.
Thus, the vertical distribution of all coccolithophoretaxa, except the three placolith-bearing species G. ericsonii,
G. oceanica and E. huxleyi, was probably controlled by
UPZ water-temperature and stratification of the water-column. The latter certainly has an influence on the presenceof nutrients in the study area.
6. Summary and conclusions
A total of 70 species (19 holococcolithophores, 51 heterococcolithophores) was observed in this study. Emiliania
huxleyi was numerically the most abundant species in the
majority of samples, but high cell numbers (>50*103
cells/l) of Gephyrocapsa ericsonii and Umbellosphaera irregularis were also observed.

191) Coccolithophores showed a difference in lateral distribution. This distribution was controlled by the relatively
cool, nutrient-enriched waters of the BC, as well as by thewarm, nutrient-depleted waters of the SG. Assemblages in
the BC were characterised by at least five species (E. huxleyi, G. ericsonii, Rhabdosphaera xiphos, U. irregularis,
Syracosphaera halldalii), which exhibit their highest cellnumbers at the easternmost stations, reflecting higher nutrient levels. The SG generally contained the holococcolithophores and the species Discosphaera tubifera,
Umbellosphaera tenuis, Calciosolenia brasiliensis andSyracosphaera pulchra, which exhibit an affinity for lower
nutrient levels.
2) A distinct vertical stratification in the coccolithophorid
assemblages was observed. Typical members of the UPZ
assemblage are D. tubifera, R. xiphos, Acanthoica spp., S.
halldalii, S. pulchra and, especially, U. tenuis and U. irregularis. The flora in the LPZ (>80-100m water-depth)
was characterised by abundant Florisphaera profunda in
the nutrient-depleted SG, whereas Calcidiscus leptoporus,
Oolithotus fragilis and Umbilicosphaera sibogae occurred in the lower-photic boundary
zone, from the BC to the SG waters.
3) Some species, such as E. huxleyi, G. ericsonii and G. oceanica, lived in a wide rangeof water-depths. In the BC, they appearedwith higher abundances in the mixed layer,
whereas they were more common at greater
depths in the SG.
4) In the study area, the vertical distributionof all coccolithophore taxa, except the threeplacolith-bearing species mentioned above,
was controlled by upper photic zone temperature and water stratification.
Acknowledgements
The authors would like to thank the officers andcrew of the R/V Meteor and the scientific crews of
the various Meteor cruises for their assistance. We
are grateful to the reviewers, Drs Maria Triantaphyllou and Ric Jordan, and the Editor, Jackie
Lees, for their constructive comments and suggestions on the manuscript. This research was
funded by the Deutsche Forschungsgemeinschaft(Research grant BA 1648/11).

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