K. v. Sa/is: Some fossil "limg coooolilhophorid"..., p. I 09- 114.

Journal ofNannoplanldon Research, 16, 3, 1994.

SOME FOSSIL "LIVING COCCOLITHOPHORID" SPECIES
Katharina v. Sa/is, Geological Institute ETHZ, CH-8092 ZUrich, Switzerland
Abstract: Fossil representatives of the following living coccolithophorids never or rarely reported from the fossil
record are illustrated: Coronosphaera binodata, Helicosphaera pavimentum, Poricalyptra gaarderae, Poricalyptra
lsselii, Syraco/ithus confusus, Syracosphaera anthos, Syracosphaera pirus, Tetralithoides quadrilaminata,
Umbilicosphaera angustiforamen. This significantly increases the number of living species with an illustrated
ancestor. In addition, one new species, Pontosphaera decora/a is described, from the Pliocen

Introduction
Winter & Siesser ( 1994) provide us with a useful atlas ofthe living coccolithophorid species. For the specialist
interested in fossil coccolithophorids they provide a long
wished-for overview of what the result of over 200 million
years of calcareous nannoplankton evolution looks like inthe form of living species.
In the atlas the authors also make statements about
the known geological ranges of some 150 of the nearly 200
presently known, living species. According to this information only 2 5, (i.e. 17%) of the living species have a fossil
record (Fig. 1). The authors warn, that " this information
is presented with caution, as comparatively little is yet
known about the range/distribution of many species'' .
Species
living

20

40

60

80

100

120 110[t}:;:~;W:?-\c Heterococcoliths :·\·[ .-:' Holococcoliths . :·/·:

Pleist_?~Pliocell!L_

f- 10

Q)

c

Q)

my

g

1- 20

~
Q)

1- 30

~

g
.Q>

0

1- 40

2!

§
f-

so

LU

<..i

f- 60

0

Q)

n;
0..

Cretaceous

The geological ranges ofliving species according
to WINTER & SIESSER, 1994.
NOTE: These authors only givedata on some 1 SO of the total
ea. zoo living species and haveoverlooked the fossil rep resentatives of some living
species.

While we have known for a long time that not allliving species survive transport to the seafloor (Honjo,
1976) and so are not found in the underlying sediments,
the non-fossilising percentage of only 17% comes assomething of a surprise. Partial explanations may be thatnearly40%ofthecoccoliths in Winter & Siesser's (1994)
atlas are holococcoliths which have a very low fossilisation potential, and that the authors were not speciallyinterested in providing ancestors for the living
coccolithophorids. But there also must be a lack of interest
from the "palaeontological community" , to make aneffort to find and report ancestors of living species in thesediment. None of the authors who have presented
overviews ofthe evolution of the calcareous nannoplankton
have addressed this problem in earnest or attempted aconcentrated search for such species (Haq, 1983; Roth,1987; Perch-Nielsen, 1986; Bown et al., 1993).
This is an highly unsatisfactory state of affairs

when we want to investigate rates of species appearancesand extinctions, or the longevity of species and comparethem with other groups ofphyto- and zooplankton. Triggered by the realisation, that more must be known than has
been reported, I had another look at my TEM and SEM
pictures collected over the past quarter of a century. This
yielded the Neogene forms discussed in this paper. They
are illustrated here in order to illuminate the ranges given
by Winter & Siesser ( 1994 ). In addition, a f~st attempt tosearch the literature al!¥> furnished a fe~ ~Palaeogene
species that may be ancestors of living forms ,I hope this presentation will initiate f~jendly cooperation among colleagues who also have such ' ~orpses "in their EM-collections of calcareous nannofosSils, andlead to a better understanding of nannoplankton evolution.
Material
The following samples yielded the forms illustrated on
Plate 1:- KvS 645 from the "Mio-Pliocene" of Rio Gurabo, St.
Domingo (Saunders et al. , 1986). The assemblage includesReticu/ofenestrapseudoumbilicus, Helicosphaera
carteri, Spheno/ithus abies, Scapholithusfossilis and thespecies shown here. The presence of Reticulofenestra
pseudoumbi licus indicates the presence ofNN 15 or below(early Pliocene or older). Since discoasters are very rare
and no ceratoliths were found, the absence or presence ofthe marker species of this interval can not be used to allow
for a more precise age assignment. Other details of thesamples are given_by Saunders et al. (1986).
- DSDP Site 119-26; (Perch-Nielsen, 1971)- DSDP Site 354- 2cc; (Perch-Nielsen, 1977)- DSDP Site 355-2-4, 70cm; (Perch-Nielsen, 1977)- DSDP Site 357-5-4, 135cm; (Pen;h-Nielsen, 1977)
Systematic descriptions
CLASS PRYMNESIOPHYCEAE HIBBERD, 1976
Family Calyptrosphaeraceae Boudreaux & Hay, 1969·
(Holococcoliths)
Poricalyptra isselii (Borsetti & Cati, 1976) Kleijne,
1991
PI. 1, Fig. 2
Remarks: This holococcolithfeatures usually four, sometimes also more or fewer pores and som~times a central
horn, like the Pliocene form illustrated here.
Occurrence: living (Winter & Siesser, 1994); Early
Pliocene, Rio Gurabo, Dominican Republic.

===========================109==========================

Joumal ofNannoplankton Research , 16, 3, 1994.

Poricalyptra gaarderae (Borsetti & Cati, 1972)
Kleijne, 199 1
PI. I, Fig. I
Remarks: The characteristics of this species are the twopores, one on either side of the central bridge with orwithout a central process. The fossil form illustrated has
the same characteristics.
Occurrence: living (Winter & Siesser, 1994); Early
Pliocene, Rio Gurabo, Dominican Republic.
Sy raco/ithus confusus Kleijne 1991
PI. I, Figs 4, 5
OR Calyptro/ithina multipora (Gaarder, 1980) Norris
1985
OR Hel/adosphaera dalmatica (Kamptner, 1927)
Okada & Mclntyre, 1977 (in Steinmetz, 1991 ).
Remarks : S. confusus is characterised by 5-8 cavities
according to the original description. Kleijne (1991)
however, illustrates a paratype (pl. Vl, fig. 4) with morethan the " allowed " 8 cavities, since there are already 78 visible on half a coccolith. In the specimens illustrated
here from the Pliocene, there are from 10 to more than 20
cavities.Thus although the two Pliocene specimens illustrated here are similar to recent forms a precise assignment
is difficult. Nevertheless if we assume, that these specimens belong to one oft he possible candidates this increases
the number of living species with an illustrated ancestor
by another species.
Note the different states of preservation shown inthe detail PI. I, Fig. 5. Here the small size of the crystals
in the well preserved wall of the specimen can be observed
next to somewhat larger units in the distal, dome-shapedpart of the coccolith. The wall includes, however, alsoheavily overgrown zones where the originally discrete
crystals fuse into larger blocks, as is usually the case inolder holococcoliths.
Similar but larger specimens from the upper
Palaeocene, which were described by Perch- Ni~lsen ( 1971)
as Semihololithus biskayae also have more than 5-8
cavities (PI. 1, Fig. 6).
Occurrence: Living (Winter & Siesser, 1994 ); Early
Pliocene, Rio Gurabo, Dominican Republic; probably also
in the Late Palaeocene of the Bay of Biscay (PerchNielsen, 1971).
"In search ofa descendant??"
PI. 1, Figs 7,8
Remarks: The two holococcoliths illustrated here have no
obvious descendant among the living holococcoliths. The
form shown in PI. 1, Fig. 7 is polygonal and features acentral cavity, while Fig. 8 has an elliptical outline. The
latter might correspond to the proximal view of several
species, while there seems to be no correspondent living
form to the former.
Occurrence: Early Pliocene, Rio Gurabo, Dominican
Republic.

K. v.Sa/is: Somefossil '1img~ ..., p. l09-114.

Family Coccolithaceae Poche, 1913
Gemini lithe/la rotula Kamptner ( 1956) Back man 1980
PI. 1, Fig. 14.
This is included as a prime example of a common, but
relatively poorly known, fossil coccolith whose descendants are unclear.
Occurrence: Early Miocene to Late Pliocene.
Umbilicosphaera angustiforamen Okada & Mclntyre,
1977
PI. 1, Fig. 15
Remarks: The two shields are typically equal in size inthis genus and species, a feature that can also be observedin the distal view illustrated here. The angular plates
described from the holotype are not preserved in the fossil
specimen illustrated here. Another living species ofUmbilicosphaera with a wide open central area, U.
maceria, is smaller and has only some 30 elements, while
U. angustiforamen has more than 40 elements.
Occurrence: living (Winter & Siesser, 1994); Early
Pliocene, Rio Gurabo, Dominican Republic.
Family Helicosphaeraceae Black, 1971
Helicosphaera pavimentum Okada & Mclntyre 1977
PI. 1, Fig. 12
Remarks: Like most living species, H. pavimentum isusually illustrated in distal view. This shows a form witha central area that is formed of a pavement of minute laths
arranged± parallel to the long axis of the coccolith which,at around 5Jlm long, is considerably smaller than the morecommon H. carterilkamptneril wallichii, which is wellknown from the fossil record. The proximal view illustrated here shows a broken specimen featuring a suture-line±along the long axis of the coccolith.
Occurrence: Living-- LatePleistocene (Winter & Siesser,
1994 ); Early Pliocene, Rio Gurabo, Dominican Republic;
probably widespread but not generally reported.
Family Pleurochrysidaceae Fresnel & Billard, 1991
Tetralithoides quadrilaminata (Okada ·& Mclntyre,1977) Jordan et al. ( 1993)
PI. 1, Figs 9, 10
Remarks: T. quadrilaminata is characterised by a verythin rim and a very wide central area covered by four thinplates~ In the forms illustrated here the thin rim and wide
central area are well developed, but the exact structure of
the central area in the form ofa~~late, plates or oJ}ler is notclearly visible due to overgrowth and other particles
overlying the central area. This species is listed by Winter& Siesser (1994) as Cricosphaeraquadrilaminata, with=================================== 110 =================================~

K. v. Salis: Some folisiJ "limg coccol~d" ..., p. 109- 114.

no known fossil record. Subsequent work has revealed its
identity with Tetralithoides symeondesii (Theodoridis
1984) and so given it a lengthy fossil range (Jordan et al.
1993).
Occurrence: Living (Winter & Siesser, 1994); DSDP Site
372 in the western Mediterranean, the Atlantic and Indian
Oceans in the Miocene (NN 2- NN 11) according to
Theodoridis (1984); Quaternary of the Philippine Sea(Matsuoka & Okada, 1989) and DSDP Site 354 on Ceara
Rise north of Brazil, samples from NN 16 & NN 19.

Journal ofNamoplankton Research, 16,3, 1994.

is considered to have become extinct at the CretaceousTertiary boundary, with the extinction oftheMaastrichtian
species ofRote lapi/Ius, R. munilus, some 65 my ago. Someof the forms assigned to the recent family
Syracosphaeraceae however, might well be descendants offorms that would be assigned to the Stephanolithiaceae in
the fossil realm.

?, Coronosphaera binodata {Kamptner 1937) Gaarder& Heimdal, 1977 or? Coronosphaera medilerranea (Lohmann 1902)
Gaarder, 1977
PI. 1, Figs 17, 18

Family Pontosphaeraceae Lemmermann, 1908
Pontosphaera decorala n.sp.
PI. 1, Fig. 13

Holotype: PI. 1, Fig. 13. Negative No. 613412-5; deposited at the Museum of Natural History, Basel.
Locus typicus: Rio Gurabo, Dominican Republic.
Stratum typicum: NN 15 or below, Early Pliocene orolder.
Diagnosis: Flat species of Pontosphaera with a narrow
wall surrounding a distal area composed of± tangentiallyoriented laths pierced by very small holes/depressions.
More than four small blocks about 1~ni high are arranged
around the central area pierced by the holes/depressions.

Remarks: C. binodata as illustrated in Steinrnetz (1991)
has a wall of imbricated elements and a central area
featuring a floor of± radially arranged elements topped by
two thick nodes. C. mediterranea, on the other hand,features a central, elongated boss along the longaxis
surmounted or not by a central process. Coccoliths withsuch a wall are also found in the fossil record. The central
area in the forms illustrated here is empty. Forms withnodes have, however, been observed in several samples
with the light microscope. In the Cretaceous, such simple
rings as those illustrated here would be assigned to theCretaceous species Loxolithus armilla, of which no specimen with a filled central area has been illustrated to date.
Occurrence: living (Winter & Siesser, 1994); Early
Pliocene, Rio Gurabo, Dominican Republic.

Remarks: No other flat species of Pontosphaera has
small blocks arranged around the pierced part of thecentral area.

Syracosphaera anthos (Lohmann 1912) Janin, 1987
S. variabilis Halldal & Markali, 1955
Deutsch/andia gaarderae Perch-Nielsen,1980

Range: Forms similar to P. decorata have not beenreported from living or fossil assemblages so far. NN 15
or below.

Remarks: Perch-Nielsen (1980) described D. gaarderae
from the middle Pliocene. It has since been suggested thatthe differences used by Perch-Nielsen(1980) to distinguish the fossil specimens from the recent ones -- thenearly double size and the higher central cone in the fossil
form -- are insufficient to characterise a new species
different from the living one.

Family Syracosphaeraceae (Lohmann)
Lemmermann, 1903
This family includes coccoliths of widely varying construction. What the specialists ofliving coccolithophorids
call ''caneoliths'' would, by some authors specialised infossil forms be called coccoliths with a zeugoid wall and± open central area. In the fossil realm, such forms are
assigned to the recent families Pontosphaeraceae,
Calciosoleniaceae and the Syracosphaeraceae and thefollowing fossil families:-the Cbiastozygaceae which feature wall(s) ofvertical or
oblique elements and a central area spanned by a X or moreelaborate central arrangement of elongated elements.
Such forms disappear with the extinction of Jsthmolithus
recurvus in the Oligocene, some 32 million years ago.
-the Stephanolithiaceae which include forms of varying
outline, a wall of vertical elements and a central areaspanned by a great variation of usually± radially oriented
elements topped or not with a central process. The family

Occurrence: living (Winter & Siesser, 1994) ; Early
Pliocene (NN 15) of DSDP Site 357 (Perch-Nielsen,1980).
Syracosphaera ?plrus Halldal & Markali, 1955
PI. 1, Fig. 16

Remarks: S. pirus as illustrated in Steinrnetz (1991) ischaracterised by its lenticular shape, its wall of two cycles
of vertically standing elements surrounding a central area
bridged by radially oriented laths that bear a central
process of vertically standing, long parallel laths. In thespecimen illustrated here the wall also consists of verticalelements. It is, however, not clear beyond.doubtthat there.,,
are two cycles of elements, since overgrowth has affected
the fossil coccolith. The central stem is shorter than in theliving form, a feature probably due to breakage. Its struc-==========================111

Journal ofNamoplanldon Research , 16, 3, 1994.

ture is not discernible. The radial laths of the floor arepresent.
Similar forms lacking any remains of the central
area are not uncommon in some samples. Their relationship with the Palaeocene species Nodosella? franzii fromthe Lo\\rer Palaeocene ofTunisia (Perch-Nielsen, 1981, pl.
1, figs 4,5, 7-9) will have to be investigated further.
Similar forms with a wall of± vertical elements are also
known from the Mesozoic family Stephanolithiaceae (seeabove).
The identification is given asS ?pirus since mostillustrations of S.pirus and S.prolongata (a closely associated form), show three distinct flanges around the coccolith(see e.g. figs in Winter & Siesser, 1993 or INA Newsletter
14/1:40), so it is not certain that the form illustrated by
Steinmetz was correctly identified.
Occurrence: living (Winter & Siesser, 1994); NN 5
(middle Miocene) DSDP Site 355-2-4,70 (Brazil Basin).
Conclusions

The additional fossil representatives of living
coccolithophorids presented here -- actually also including G/adiolithus jlabellatus which has been found in
Holocene sediments with the light microscope -- add
another ten species to the 25 noted by Winter & Siesser( 1994) as species with a fossil record. It will need a major
effort from the calcareous nannofossil community, to gaina more realistic picture of the history of evolution of theancestors of the living species of coccolithophorids thanthe one illustrating the presently acknowledged state (Fig.
1). It will also need co-operation from the authors illustrating living coccolithophorids. From the published pictures
it is often impossible to see the structure of the single·
coccoliths. Thus I have a feeling that some small species
may have larger ancestorsin the fossil record but that we
do not realise this, because we do not have any illustrationof those species from which the fine structure is clearlyvisible. On the other hand species of Syracosphaera lookrather similar to some forms of the Mesozoic
Stephanolithiaceae.
Below is an attempt at assigning some fossil
Palaeogene and Jurassic species to living species based on
EM-illustrations

K. v. Sa/is: Some bsil "livWIB ooooolahophorid"•., p. 109- 114.

Homozygosphaera schilleri

Discolith. macroporusDef:JaOOre,
1954 (in Varol, 1989)
Pontosphaera syracusana
Pontosphaera crucifera Roth,1970
Syracolithus confusus
Semihololithus biskayeae P-N,
1971
Syracolithus quadripeiforatus Zygosphaera brytika Roth ,1970
Syracosphaera ossa

Living species=
Calyptrolithina multipara
Calyptrolithina divergens
Syracolithus bicorium

Coccolithus joensuui in Hay et
al. , 1967

Jurassic fossil species?
Anfractus variabilis Medil, 1979
Anfractus harrisonii Medd,
1979'' Neococcolithes dub ius''
Medd, 1979

lfwe want to make more educated guesses as to longevity
and changes in the evolutionary rates over the past 20 mythan has up to now been done, we have several ways to do
this:- one person works in splendid isolation for 10 years.
-several people pool their work and embark on intelligent,coordinated investigation of those sections that promise
results. These will be sections with exceptionally wellpreserved assemblages and, at the same time, a good age
control.
- everybody works on her /his own, thus probably duplicating a lot of work.

As INA approaches 20 years of " friendly cooperation" I
hope to be able to entice INA members to join me in theeffort to make a giant step forwards in the " acquisition"of ancestors of living coccolithophorid species. The goal
could be to present INA with a " jubilee-gift" for its first
20 years in 1997.

Palaeogene fossil species?

Acknowledgements
I would like to thank those who gave me the samples which
formed the basis of this study: John Saunders, Basel,
supplied the samples from the Dominican Republic andthe DSDP those from the Atlantic Ocean. Jeremy Young(London) and Ric Jordan (Yamagata) suggested welcome
improvements.

Cruciplacolithus tarquinius inRoth, 1970
Cr. primus P-N 1977 a.o.
Cyrtosphaera cucullata
Cepekiella e/ongata Roth, 1970
Homozygosphaera wensteinii Discolithus solidus Deflandre,
1954 (in Varol, 1989)

REFERENCES
Taxonomic references that are included in· Perch-Nielsen( 1985 - nannofossils) or Jordan & Kleijne ( 1994 - living
nannoplankton) are not listed below. They can also be found in
the Index and Bibliography of Calcareous Nannoplankton -Living species
Coccolithus neohelis

Magnifications
Figs 1: 7.500x; 2: 15.000x; 3: 15.000x; 4: lO.OOOx; 5: 9.000x; 6: 13.000x; 7: lO.OOOx; 8: 13.000x; 9: ???; 10: ???; 11 : lO.OOOx;
12: 6.500x; 13: 7.800x; 14: 7.500x; 16: 9.000x; 17: 7.500x; 18: lO.OOOx.
Localities
DominicanRepublic,RioGurabo5ection;"' NN 15, EarlyPiiocene: Figs 1, 2, 3, 4, 5, 7, 8, 11, 12, 13, 14, 17, 18. DSDPSUe 11926, NP 9, Late Palaeocene: Fig. 6. DSDP Site 354-2cc, NN 19, Early Pleistocene: Fig. 9. DSDP Site 354-3-2,115cm, NN 16, Late
Pliocene: Fig. 10. DSDP Site 357-5-4, 135cm, ,.. NN 7, middle Miocene: Fig. 15. DSDP Site 355-2-4, 70cm, NN 5, Early Miocene:
Fig.16.

==========================112=========================

K. v. Sa/is: Some fussj) "limgOOCXIOIIhophorid"... ,p. 109-114.

Journal ofNarmoplanldon Research , 16, 3, 1994.

Poricalyptra gaarderae

Poricalyptra isselii Syracolithus confusus

Syracolithus confusus

Semihol. biscayae Pal.

Holococcolith

Syracolithus confusus

Holococcolith~;.

==========================113========================~

K v. Sa/is: Some fOssil "limg coocolilhophorid''..., p. I 09- 114.

Journal ofNannoplankton Research, 16, 3, 1994.

Loeblich & J:appan ( 1966 to 1973) and subsequently in INA
Newsletters 1-15 (1979-1993).
Bown, P.R., Bumett, J.A & Gallagher, L.T. , 1993: Calcareous
nannoplanlctonevolution.Mem. di ScienzeGeol., Padova,
43,1-17.
Haq, B.U. 1983: Calcareous nannoplankton. Benchmark Papers
in Geology, v. 79, Hutchinson Ross Publishing Co.,
Stroudsburg, Pa., 338 p.
Honjo, S., 1976: Coccoliths: production, transportation, andsedimentation. Mar. Micropal., 1, 65-79.
Janin, M.-C .,l987: Micropaleontologie de concretions
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Jordan, R.W., Kleijne, A & Heimdal, B., 1993: Proposedchanges to the classification system ofcoccolithophorids.
ill. INA Newsletter, 15/1, 18-22.
Jordan, R.W., & Kleijne, A, 1994: A classification system forliving coccolithophores. In: Winter, A & Siesser, W.G.
(eds) " Coccolithophores' ', Cambridge University Press,
p. 83-105.
Kleijne, A , 1991 : Holococcolithophorids from the Indian
Ocean, Red Sea, Mediterranean Sea and the North
Atlantic Ocean. Marine Micropal. 17, 1-76.
Loeblich Jr., AR. and Tappan, H., 1966-73: Annotated index
and bibliography of the calcareous nannoplankton. I.
Phycologia, S, 81-216. II: 1968, J. Paleontol., 42, 584598; ill: 1969: J. Paleontol. , 43, 568-688; IV: 1970, J.
Paleonto/., 44, 668-674; V: 1970 Phycologia, 9, 167174; VI: 1971. Phycologia,10, 316-339; VII: 1973, J.
Paleontol., 47, 715-769. [Index of all nannoplanlcton
taxa with comments about their non-validity under
Botanical or Zoological Codes if not described properly.
Bibliography up to 1972].
Matsuoka, H. & Okada, H., 1989: Quantitative analysis of
Quaternary nannoplankton in the subtropical Northwestern Pacific Ocean. Marine Micropal., 14, 97-118.

Okada, H. & Mclntyre, A , 1977:. Modem coccolithophores of
the Pacific and North Atlantic Oceans .
Micropaleontology, 23/1 ,1-55.
Perch-Nielsen, K., 1971 :. Einige neue Coccolithen a us dem
Paleozlln der Bucht von Biskaya. Bull. geol. soc. Denmark, 20/4, 347-361.
Perch-Nielsen, K. 1977a Albian to Pleistocene calcareous
nannofossils from the Western South Atlantic, DSDP
Leg 39. JRDSDP, 39, 699-823
Perch-Nielsen, K., 1980: New Tertiary calcareous nannofossils
from the South Atlantic. Eclogae geol. Helv. 73/1 , 1-7.
Perch-Nielsen, K. 1985 . Mesozoic calcareous nannofossils;
Cenozoic calcareous nannofossils. In: Bolli, H.M.,
Saunders, J.B. & Perch-Nielsen, K. (eds.): " Plankton
Stratigraphy", Cambridge University Press, p. 329426;427-554; Index, p. 1006-1022.
Perch-Nielsen, K., 1986: Geologic events and the distribution of
calcareous nannofossils - some speculations. Bull.
Centre Rech. Explor.-Prod. Elf-Aquitaine, 10, 422-432.
Roth, P.H. 1987 Mesozoic calcareous nannofossil evolution:
relationtopaleoceanographicevents.Pa/eoceanography,
2, 601-611.
Saunders, J.B., Jung, P. & Bijou-Duval, B., 1986: Neogene
Paleontology in ·the Northern Dominican Republic.! .
Field surveys, Lithology, Environment, and age. Bull. of
American Paleontology, 8913, 231-279.
Steinmetz, J.C., 1991 : Calcareous nannoplankton biocoenosis:
sediment trap studies in the Equatorial Atlantic, Central
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Theodoridis, S., 1984: Calcareous nannofossil biostratigraphyof the Miocene and revision of the helicoliths anddiscoasters. Utrecht Micropal. Bull. 32, 1-271.
Varol, 0 ., 1989: Eocene calcareous nannofossils from Sile,(Northwest Turkey). Revista Esp. de Micropal. 21/2,
273-320.
Winter, A & Siesser, W .G ., 1994 : Atlas of living
coccolithophores. In: Winter, A & Siesser, W.G. (eds)" Coccolithophores", Cambridge University Press, p.
107-159.

•=========================114=========================