Journal of Nannoplankton Research, 22, I , 2000.

L. Cros: Variety of exothecal coccoliths of Syracosphaera, p. 41 - 51.

VARIETY OF EXOTHECAL COCCOLITHS OF SYRACOSPHAERA
Llui"sa Cros
Institut de Ciencies del Mar (CS/C), Passeig l oan de Borb6 sin, 08039, Barcelona, Spainfy
Abstract: Species of the genu s Syracosphaera produce an outer coccolith layer, the exo theca. The
coccoliths of thi s layer differ considerably from species to species. This study examines the diversity ofthe exothecal coccoliths of Syracosphaera species found in NW Mediterranean waters , presents !I newrecords of exothecal coccoliths, and proposes a tentative classifi cation of the exothecal coccoliths of
Sy racosphaera. Within the groups that emerge from this c lass ification, it is poss ibl e to recogni secommon characteristics of the coccospheres that could be relevant for understanding the phylogeny of
the genus.

futroduction
Syracosphaera is a diverse genus with a high number of
extant species (Kleijne, 1993; Jordan & Green, 1994) but
few fossil representatives (Perch-Nielsen, 1985). Due tothe small size of the coccoliths of most Syracosphaera
species and their low preservation potential (Young, 1998),
the development of their systematic study has depended
upon extant nannoplankton work. u sing electron
microscopy. The morphological complexity of this group,
especially with regard to the presence or absence of an
exotheca, has resulted in very different views regarding
the characterisation of the genus (Gaarder & Heimdal, 1977;
Okada & Mclntyre, 1977; Jordan & Young, 1990). Moreover,representatives of Syracosphaera have also been observed
as hetero-holococcolithophorid combination coccospheres(Cros et al., 2000).
Most Syracosphaera species produce dithecate coccospheres. The inner layer, or endotheca, is composed ofmurolith-shaped coccoliths with characteristic structuralfeatures, normally termed caneoliths (Jordan et al., 1995;
Young et al., 1997). All caneoliths have a central area flooredby radiating laths, and the rim is rather variable, having
one to three flanges on the outer wall; nonetheless theendothecal coccoliths are very similar in many different
spec ies. By contrast, the exo thec al coccoliths of
Syracosphaera are much more variable in morphology. But,
in spite of their wide variety of shapes, all these exothecal
coccoliths are grouped together as ,cyrtoliths' (Jordan etal., 1995; Young et al., 1997). Arising from the Greek kyrt6s,
curved, the term cyrtolith (Braarud et al., 1955) has
undergone a complex evolution of meaning. It was usedlo
designate the heterococcoliths in a calotte-likearrangement, including the calyptroform coccoliths of thefamily Rhabdosphaeraceae (Heimdal, 1993), but nowadays
it only embraces the exothecal coccoliths of Syracosphaera
species (Kleijne, 1992; Jordan et al. , 1995; Young et al.,1997). Some authors have recognised differences in shapeand distribution patterns in these exothecal coccoliths(Heimdal & Gaarder, 1981 ; Jordan et al. , 1995; Young et al.,1997) but to date, there have not been any specific studies
on this subject.
This study examines the diversity of exothecal
coccoliths of Syracosphaera species found in the NW
Mediterranean and presents a propo sal for theirclassification, based on their morphology.

Material and methods
The material was collected by filtering NW Mediterranean
sea-water obtained during several cruises of the Institut
de Ciencies del Mar (CS IC) carried out in 1995 (MES0-95,
from 30th May to 16th June; FRONTS-95, from 17th to
23rd June), 1996 (MES0-96, from 18th June to 3rd July;FRONTS-96, from 16th to 21 st September; FANS-I, from
1st to 8th November), 1997 (FANS-2, from 4th to 14th
February) and 1999 (Hivem-99, from 20th to 28th February).
For each sample, about 200ml of sea-water were filtered ,using a vacuum pump, onto polycarbonate Nucleopore
filters of 0.8f.l.lll pore-size and 25mm diameter. Salt wasremoved by rinsing the filters with - 2ml of tap-water. An
aliquot of the filter was glued, with silver colloidal
suspension, onto a SEM stub and coated with gold before
examination of the specimens in an Hitachi S-57 SEM.
Tentative classification of the exothecal coccoliths of
Syracosphaera species
The exothecal coccoliths of the dithecate Syracosphaera
species found in the NW Mediterranean waters aregrouped by morphological affinities to outline the present
tentative classification. Some common Syracosphaera
species not found in the study, but for which the exothecal
coccoliths have been illustrated by other workers, are added
as ,other examples' to better illustrate the characteristics
of some groups.
Most of the Syracosphaera species have flattened,
exothecal coccoliths which exhibit a disc-like shapecompletely differentiated from the endothecal coccoliths.
Others present exothecal coccoliths with a distinctly
concavo-convex shape, and in some of these it is possible
to distinguish as exothecal coccoliths real caneoliths, whichsometimes are very similar to the endothecal ones.
Disc-like coccoliths: These coccoliths appear as planoliths; they are commonly flat and present a diversity ofoutlines and thickness. Usually, they are composed of threeparts: (1) a central structure with two plates or an
undetermined number of elements; (2) a radial cycle, withnarrow elements; and (3) an external cycle, the rim, of broad
elements.

Disc coccoliths: Large, circular, plate-like coccoliths witha central, conical protrusion. They cover the endothecal

41

Journal of Nannoplankton Research, 22, 1, 2000.

L Cros: Variety of exothecal coccoliths of Syracosphaera, p. 41 - 51.

caneoliths with overlapping disposition and, sometimes,
entirely conceal the endotheca. Example: Syracosphaera
anthos (Plate 1, Figures 3, 4).

coccoliths are distributed, in an overlapping pattern, onlyaround the apical pole of the cell. Due to their distribution,
they have been called deviating coccoliths (Heimdal &Gaarder, 1981; Heimdal, 1993; Jordan et al., 1995). Examples:
Syracosphaera molischii (Plate 5, Figure 1), S. ossa (Plate
5, Figure 2), S. marginaporata (Plate 5, Figure 3), and S.
sp.II cf. S. epigrosa ofKleijne (1993) (Plate 5, Figure 4).

Wheel-like coccoliths: Circular and flat coccoliths withcharacteristic long, narrow laths in the radial cycle and acentral-structure of two plates. They cover the endothecal
caneoliths with overlapping disposition and, sometimes,
entirely conceal the endotheca. Examples: Syracosphaera
nodosa (Plate 2, Figure 3) and S. sp. ,aff. S. nodosa' (S. cf.
S. nodosa ofHeimdal & Gaarder, 1981) (Plate 2, Figure4).

Vaulted coccoliths: Exothecal coccoliths with a rounded
dome-like structure have been designated as vaulted
cyrtoliths (Jordan et al., 1995) or dome-shaped cyrtoliths(Heimdal, 1993). They are distributed over the endotheca
in a non-overlapping pattern. Examples: Syraco~phaera
pulchra (Plate 1, Figure 1) and S. histrica (Plate 1, Figure 2).
N.B. Studies on Syracosphaera pulchra havedemonstrated that S. pulchra exothecal coccoliths show
basically the same structure as the endothecal coccoliths(lnouye & Pienaar, 1988). So, this group of vaulted ·exothecal coccoliths might be considered as inverted
muroliths and close to the grouping of the caneoliths (seebelow).

Thin, (sub )circular coccoliths: Circular or subcircular
sheet-like coccoliths which are very thin and flat. Examples:
Syracosphaera lamina (Plate 2, Figure 1), S. tumularis(Plate 2, Figure 2), S. sp. type L ofKleijne (1993) (Plate 2,
Figures 5, 7).
Oval coccoliths: Oval to elliptical coccoliths withlongitudinal symmetry. They can be flat or have the central
elements slightly raised. Examples: Syracosphaera nana
(Plate 2, Figures 6, 8) and S. sp. ,aff. S. nana, very small'(Plate 3, Figures 5, 6).

Caneoliths: True exothecal caneoliths with a murolithmorphology, a wall that can have laterally extending flanges
and a central area floored by radiating laths. (See originaldescriptions in Braarud et al., 1955; Heimdal, 1993; Jordan
et al., 1995.) In the studied samples, three different types
were observed:

Asymmetrical coccoliths: Asymmetrical, subcircular to
subelliptical coccoliths with a rim which varies in widtharound the coccolith. Often, they appear grouped in an
imbricate arrangement forming a ribbon of coccoliths (seeHalldal & Markali, 1955), which can surround theendotheca. Examples: Syracosphaera delicata (Plate 3,
Figures 1, 2), S. bannockii (Plate 3, Figures 3, 4), S. sp. ,aff.
S. orbiculus, ovoid' (Plate 4, Figures 1, 3) and S. sp. ,aff. S.
orbiculus, spherical ' (Plate 4, Figures 2, 4 ).

Elliptical caneoliths with flanges: Elliptical caneoliths witha proximal flange, an almost imperceptible mid-wall flange
and a distal flange expanded outwards. These exothecal
caneoliths have a delicate appearance and are larger, thinner,with higher walls and weaker central structures than theirrespective endothecal body caneoliths, with which they
can be easily confused. They appear more concentrated
near the apical pole. Examples: Syracosphaera cf. S.
dilatata (Plate 6, figures 1, 2) and S. sp. typeD ofKleijne(1993) (Plate 6, Figures 3, 4).

Stratified coccoliths: Thick, multilayered coccoliths,
apparently with a construction of stratified laminae. This
stratification can be seen in distal view. They present anoverlapping pattern around the endotheca. Example:
Syracosphaera sp. , with stratified coccoliths' (Plate 1,
Figures 5, 6). Other example: Syracosphaera sp. type J of
Kleijne (1993).

Simple undulating coccoliths: Symmetrical and of simple
elliptical shape. Usually very few coccoliths, only on one
side of the coccosphere. Example: Syracosphaera sp.I cf.
S. epigro~a ofKleijne (1993) (Plate 5, Figures 5, 6). Other
example: S. borealis (see Winter & Siesser, 1994, p.135,
fig.l05, from C. Samtleben).

(Sub )circular caneoliths with nodes: Rounded caneoliths
with a beaded proximal flange, short wall and a very narrow,incipient distal flange. They do not posses a mid-wallflange. These exothecal caneoliths can be found all aroundthe endotheca, but they are usually more frequent near the
apical pole. Examples: Syracosphaera prolongata sensu
Throndsen (1972) (Plate 7, Figures 3, 4) and S. prolongata
sensu Heimdal & Gaarder (1981) (Plate 7, Figures 5-7). Other
example: S. pirus.
N. B. Gaarder & Heimdal ( 1977) suggested, by analogy
with Syracosphaera pulchra, that these exothecal
coccoliths were disposed with their convex side outwards,
i.e. that they were inverted relative to the endothecal
caneoliths. However, careful examination of many
specimens reveals that they are usually Jlisposed with theconvex side inwards, although they ctrft. be found in both
positions, convex side outwards and inwards.

Complex, undulating coccoliths or deviating coccoliths:
Asymmetrical and omamented with lateral extensions andwing-like protrusions. These complex, undulating

Elliptical caneoliths with nodes: Elliptical caneoliths
omamented by a characteristically beaded proximal flange.
They have neither mid-wall flange nor distal flange. In

Undulating coccoliths: These are rather irregular, discshaped coccoliths, having a broadly convex distal sidewith an elliptical central depression. There is a distinct
division into rim and central area. Individual elements areusually obscure. The central area usually displays
parenthesis-shaped slits at the ends of the coccolith; theseslits are more visible in proximal view. Based on theirstructural complexity, two groups can be distinguished:42

L. Cros: Variety of exothecal coccoliths of Syracosphaera, p. 41 - 51.

Journal of Nannoplankton Research, 22, 1, 2000.

proximal view, they appear delicate and more ornamented
than their respective endothecal body caneoliths, due tothe more separated laths and the presence of the beads.
They can be found all around the endotheca in a nonoverlapping pa~tern . Examples: Syracosphaera noroitica(Plate 6, Figures¥5, 6) and S. sp. type Gin Kleijne (1993)(Plate 7, Figures 1, 2).

possess exothecal elliptical caneoliths with a beaded
proximal flange display a high varimorphism in theirendothecal caneoliths.
Discussion and conclusions
This article describes, for the first time, 11 instances of
exothecal coccoliths not previously recognised:
Syracosphaera sp.I cf. S. epigrosa of Kleijne (1993), S.
marginaporata, S. sp.II cf. S. epigrosa ofKleijne (1993), S.
tumularis, S. sp. (aff. S. orbiculus, ovoid), S. sp. (aff. S.
orbiculus, spherical), S. sp. (aff. S. nana, very small), S. cf.
S. dilatata, S. sp. typeD ofKleijne (1993), S. noroitica andS. sp. type G of Kleijne (1993) . These observations,
together with published studies (Gaarder & Heirndal, 1977;
Okada & Mclntyre, 1977; Winter et al., 1979; Heimdal &Gaarder, 1981; Kleijne, 1993), indicate that exothecal
coccoliths in the genus Syracosphaera show a greatmorphological diversity. These exothecal coccoliths areoften disc-like, with a planolith morphology, but real
caneoliths, with a m urolith morphology, can be also found.
Other exothecal coccoliths appear highly curved, such as
the undulating (simple and complex) and the vaulted
coccoliths, which can be modified planoliths and inverted
muroliths, respectively. Among the disc-like exothecal
coccoliths, we can distinguish the following kinds: disc,wheel-like, thin (sub )circular, oval, asymmetrical andstratified. Among the caneoliths, we can distinguish thefollowing groups: elliptical with flanges, (sub )circular withnodes and elliptical with nodes. The exothecal coccolithsof S. rotula are difficult to classify within this scheme
because they present intermediate features between thevaulted, wheel-like and the (sub)circular caneoliths.
There is a strong relationship between exothecal
coccolith type, distribution pattern around the endotheca,
general shape of the coccosphere, and characteristics ofthe endothecal caneoliths (e.g. number and type of flanges,
possession and shape of circum-flagellar coccoliths).
Because of these connections, the study and classificationof exothecal coccoliths may be a useful tool to clarify the

Syracosphaera rotula
Syracosphaera rotula (Plate 4, Figures 5, 6) has circular
exothecal coccoliths which superficially look very like theexothecal coccoliths of S. nodosa and S. sp. , aff. S.
nodosa'. They have a wide, radial cycle formed of longlaths with sinistral obliquity, and a plate-like central area
composed of two tiles. However, in S. rotula exothecal
coccoliths, the outer part of the rim is bent through 90°,
forming a real wall, and resembling the exothecal subcircularcaneoliths . Nevertheless , the disposition on the
coccosphere, with the convex side outwards, is similar tothe disposition of the vaulted coccoliths. It can besuggested that S. rotula exothecal coccoliths could be alink between disc-like coccoliths and caneoliths.
Relationship between exothecal coccoliths and body
caneoliths
The dithecate Syracosphaera species display relationships between the exothecal coccolith type and morphologic characters of the endothecal coccoliths (see Table
1 ). The species which possess vaulted exothecal
coccoliths, S. pulchra and S. histrica, both exhibit circumflagellar caneoliths with spines that have bifurcate endings.
Species which have complex undulating coccoliths(deviating coccoliths) exhibit circum-flagellar caneoliths
with spines, which divide into four at the tips; often, these
species, also have an antapical spine-bearing caneolith.
Species which have (sub)circular exothecal caneoliths withnodes exhibit circum-flagellar caneoliths with spines thathave bifurcate endings and have a well-known tendency
to form very long coccospheres . The species which

Body caneoHth flanges

Syracosphnera s~les

Exothecal coccolith type

S. anthosS. nodosa • S. sp. 'aff. S. nodosa 'S. lamina , S. tumularis , S. sp. type L of K.leijne
S. nana , S. sp. 'aff. S. nana . very small'S. delicata , S. bannockii , otherS. spp.
S. sp. (with stratified cyrtoliths)

DISC-LIKE COCCOLITHS
disc coccoliths
wheel-like coccoliths
thin, (sub)circular coccoliths
oval coccoliths
asymmetrical coccoliths
stratified coccoliths

S. sp.I cf. S. epigrosa of K.leijne
S. molischii . S. ossa , S. marginaporata, S. sp.

UNDULATING COCCOLITHS
simple, undulating coccoliths
proximal and distal
complex, undulating coccoliths
proximal and distal
VAULTED COCCOLITHS

S. pulchra , S. histrica

proximalproximalproximalproximalproximalproximalproximal, mid-wall and distal

Differentiated apical caneoliths

yes. with robust spines
yes, with robust spines
noyes. with small spines
yes, with spines
yes, with spinesnoyes, with four-ended spines
yes. with bifurcated spines

S. cf. S. dilatata , S. sp. typeD of Kleijne
S. prowngata (two different types)
S. noroitica • S. sp. type G K.leijne

CANEOLITHS
proximal, mid-wall and distal
elliptical caneoliths with flanges(sub)circular caneoliths with nodes proximal, mid-wall and distal
elliptical caneoliths with nodes
proximalyes, with robust spines
yes, with bifurcated spines
yes, with bi-ended long spines

S. rotula

TRANSITIONAL FORM?
wheel-like coccoliths 'with wall'no- proximal and distal

Table 1: Relation ship between exothecal coccolith type and endothecal caneolith characters.

43

Journal of Nannoplankton Research, 22, 1, 2000.

L. Cros: Variety of exothecal coccoliths of Syracosphaera, p. 41 - 51.

taxonomic and phylogenetic relationships within thecomplex Syracosphaera genus.

class Prymnesiophyceae) from the Indian Ocean, Red Sea,
Mediterranean Sea and North Atlantic Ocean. Scripta
Geologica, 100:1-63.
Kleijne, A. 1993. Morphology, taxonomy and distribution ofextant
coccolithophorids (calcareous nannoplankton). Unpublished
PhD thesis, FEBO, Enschede, The Netherlands, Vrije
University, Amsterdam: 321pp.
Okada, H. & Mclntyre, A. 1977. Modern coccolithophores of
the Pacific and North Atlantic Oceans. Micropaleontology,
23(1): 1-55.
Perch-Nielsen, K. 1985. Cenozoic calcareous nannofossils. In:
H.M. Bolli, J.B. Saunders & K. Perch-Nielsen (Ec!,.s). Plankton
Stratigraphy. Cambridge University Press: 427-554.
Throndsen, J. 1972. Coccolithophorids from the Caribbean Sea.
Norwegian Journal of Botany, 19: 51-60:
Winter, A. & Siesser, W.G. 1994. Atlas ofliving coccolithophores.
In: A. Winter & W.G. Siesser (Eds). Coccolithophores.
Cambridge University Press.
Winter, A., Reiss, Z. & Luz, B. 1979. Distribution of living
coccolithophore assemblages in the Gulf of Eilat (Aqaba).
Marine Micropaleontology, 4: 197-223.
Young, J.R. 1998. Neogene. In: P.R. Bown (Ed.). Calcareous
Nannofossil Biostratigraphy. British MicropalaeontologicalSociety Publications Series/Chapman & Hall/Kluwer
Academic Publishers: 225-265.
Young, J.R., Bergen, J.A., Bown, P.R., Burnett, J.A., Fiorentino,A., Jordan, R.W., Kleijne, A., van Niel, B.E., Romein, A.J.T.
& von Salis, K. 1997. Guidelines for coccolith and calcareous
nannofossil terminology. Palaeontology, 40(4): 875-912.

Acknowledgements
I thank R. Margalef and M. Estrada for their direction andassistance. I thank also L. Arin for collecting the samples;
J.M. Fortuiio for the microphotography and assistancewith the SEM; J. Biosca for technical assistance with thephotography and D. Blasco for valuable criticisms. The
manuscript was highly improved by the comments of thereviewers P.R. Bown and J.R. Young. The research was
supported by the CSIC and the projects CODENET (EUFMRX-Cf97-0113),GRQ93-8041(CIRIT)andMAR98-0932(CICYT). Samples were provided, too, by the projectMAS3-Cf95-0037.
References
Braarud, T., Deflandre, G., Halldal, P. & Kamptner, E. 1955.
Terminology, nomenclature, and systematics of the
Coccolithophoridae. Micropaleontology, 1(2): 157-159.
Cros, L., Kleijne, A., Zeltner, A., Billard, C. & Young, J.R. 2000.
New examples ofholococcolith-heterococcolith combination
coccospheres and their implications for coccolithophorid
biology. Marine Micropaleontology, 39: 1-34.
Gaarder, K.R. & Heimdal, B.R. 1977. A revision of the genus
Syracosphaera Lohmann (Coccolithineae). ,Meteor"
Forschungs-Ergebnisse, Reihe D, 24: 54-71.
Halldal, P. & Markali, J. 1955. Electron Microscope Studies on
Coccolithophorids from the Norwegian Sea, the Gulf Stream
and the Mediterranean. Av. Ut. Av. Det. Norske VidenskapsAkademi Oslo, No.l: 5-28.
Heimdal, B.R. 1993. Modem Coccolithophorids. In: C.R. Tomas(Ed.). Marine Phytoplankton. A Guide to Naked Flagellates
and Coccolithophorids. Academic Press Limited, London:147-250.
Heimdal, B.R. & Gaarder, K.R. 1981. Coccolithophorids fromthe northern part of the eastern central Atlantic. II.
Heterococcolithophorids. ,Meteor" Forschungs-Ergebnisse,
Reihe D, 33: 37-69.
Inouye, I. & Pienaar, R.N. 1988. Light and Electron Microscope
Observations of the Type Species of Syracosphaera, S.
pulchra (Prymnesiophyceae). British Phycological Journal,
23: 205-217.
Jordan, R.W. & Green, J.C. 1994. A check-list of the extant
Haptophyta of the world. Journal of the Marine Biological
Association UK, 74: 149-174.
Jordan, R.W., Kleijne, A., Heimdal, B.R. & Green, J.C. 1995. A
glossary of the extant haptophyta of the world. Journal of
the Marine Biological Association UK, 75: 769-814.
Jordan, R.W. & Young, J.R. 1990. Proposed changes to theclassification system of living coccolithophorids. INA
Newsletter, 12(1): 15-18.
Kleijne, A. 1992. Extant Rhabdosphaeraceae (coccolithophorids,

44

Taxononricappendix
Syracosphaera anthos (Lohmann, 1912) Janin, 1987
Syracosphaera bannockii (Borsetti & Cati, 1976) Cros etal., 2000 (aff. S. sp. type K of Kleijne, 1993)
Syracosphaera borealis Okada & Mclntyre, 1977
Syracosphaera delicata Cros et al., 2000
Syracosphaera dilatata Jordan et al., 1993
Syracosphaera histrica Kamptner, 1941
Syracosphaera lamina Lecal-Schlauder, 1951
Syracosphaera marginaporata Knappertsbusch, 1993
( S. sp. type H of Kleijne, 1993)
Syracosphaera molischii Schiller, 1925
Syracosphaera nancr (Kamptner, 1941) Okada & Mclntyre,1977 ( S. sp. type A of Kleijne, 1993)
Syracosphaera nodosa Kamptner, 1941
Syracosphaera noroitica Knappertsbusch, 1993 (orthog. emend.
Jordan & Green, 1994) (S. sp. type E of Kleijne, 1993)
Syracosphaera orbiculus Okada & Mclntyre, 1977
Syracosphaera ossa (Lecal, 1966) Loeblich & Tappan, 1968
Syracosphaera pirus Halldal & Markali, 1955
Syracosphaera prolongata Gran ex Lohmann, 1913
Syracosphaera pulchra Lohmann, 1902
Syracosphaera rotula Okada & Mclntyre, 1977
Syracosphaera tumularis Sanchez-Smirez, 1990 (S. sp.
type C of Kleijne, 1993)

Journal of45

annoplanl1on R earch, 22, I, 2000.

Journal of47

annoplankton Research, 22, I, 2000.

48

Journal of

L. Cros: Vruiel)' f ex thecal coccoliths of Syraco plwera, p. 41 - 51 .

annoplankton Res r h, 22. l , 2000.

m pie undulating coccoliths deviating
te antapical pine-49

Journal ofannoplankton Research, 22, I, 2000.

Plate 6: yraco phaera e othecal caneoliths. cale bars= 21ffil. Figs 1-4: Elliptical caneolith with tlang : Fig.l: Cocco phereof S. f. S. dilatata with ex.othecal caneolith in the apical area and in deta hed po ition (top of figure, ee arrow ). MES -96, t. 12
41 ° 1 'N,2 O'E), m. Fi .2: DetailofFig. l with thee othecalcaneolith all pright. oti lhelargere th alcancoliths( eearrow ) in front of the endolhccal on ( ttom let corn r). Fig.3: Cocco phcre of racosphaera p. typeD of Kleijne (1993)
exhibiting th ex.othecal caneoliths ( ee arrow ) around it ( ome of them in deta hed po ition . Hivem-9 , t. 25[40°49' , 2°44 •E ,60m. Fig.4: Ex. thecal can lith of S ra osplwera p. type 0 of Kleijne (1993 . FRONT -96, t. 01 (41 ° 1 , 3°51 'E), 66m .
Fi 5, 6: Elliptical caneoliths with nod ·: Fig.S: Coccosph re of S. noroitica c hibiting ome deta hed ex. thecal can olith withnodes forming a pro jmaJ flange ee arrow ). MESO- 6 l. E3-4 (41 °23' 3°1O'E), 70m. Fig.6: D tail of S. noroitica caneolith .
lice (top right) ome e othecal caneolith exhibiting nod in pia e of the pro imal flange of the end thecal caneolith . At bottom
I fl corner there i an endothecal caneolith, in ide-view, exhibiting the u ual pro imal flange. MES0-96, St. I (40°37' , 2°55 ' E),

70m.

50

Journal f

annoplankton Re

rch, 22, I, 2000.