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Exceptionally well preserved upper Eocene

to lower Oligocene calcareous nannofossils
(Prymnesiophyceae) from the Pande Formation
(Kilwa Group), Tanzania
a b c
Tom Dunkley Jones , Paul R. Bown & Paul N. Pearson
a
Department of Earth Sciences, University College London, Gower Street, London, WC1E
6BT, UK E-mail:
b
Department of Earth Sciences, University College London, Gower Street, London, WC1E
6BT, UK E-mail:
c
School of Earth, Ocean and Planetary Sciences, Cardiff University, Main Building, Park
Place, Cardiff, CF10 3YE, UK E-mail:
Published online: 09 Mar 2010.

To cite this article: Tom Dunkley Jones , Paul R. Bown & Paul N. Pearson (2009) Exceptionally well preserved upper
Eocene to lower Oligocene calcareous nannofossils (Prymnesiophyceae) from the Pande Formation (Kilwa Group),
Tanzania, Journal of Systematic Palaeontology, 7:4, 359-411, DOI: 10.1017/S1477201909990010

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 Journal of Systematic Palaeontology 7 (4): 359-411 Issued 24 November 2009
doi:10.1017/S1477201909990010 © The Natural History Museum

EXCEPTIONALLY WELL PRESERVED

UPPER EOCENE TO LOWER OLIGOCENE
CALCAREOUS n a n n o f o s s i l s
(PRYMNESIOPHYCEAE) FROM THE
PANDE FORMATION (KILWA GROUP),
TANZANIA
Tom Dunkley Jones†
Department of Earth Sciences, University College London, Gower Street, London, WC1E 6BT, UK
Downloaded by [University of Nebraska, Lincoln] at 23:04 13 July 2013

Paul R. Bown‡
Department of Earth Sciences, University College London, Gower Street, London, WC1E 6BT, UK

Paul N. Pearson§
School of Earth, Ocean and Planetary Sciences, Cardiff University, Main Building, Park Place, Cardiff CF10
3YE, UK

SYNOPSIS The most well preserved and diverse upper Eocene to lower Oligocene assemblage of
calcareous nannofossils (coccolithophores) known to date is described from the Pande Clays, Kilwa
Group, Tanzania. The assemblage is exceptionally diverse, with a total of 115 species described herein,
which significantly exceeds the current globally compiled nannofossil species diversity of 67 for the
latest Eocene (NP19/20). The enhanced diversity observed in these sections is concentrated in the
numerous Rhabdosphaeraceae (20 species), Syracosphaeraceae and holococcolith taxa (19 species)
that are unknown from any other contemporaneous location. Scanning electron microscope (SEM)
studies reveal exquisite preservation down to the sub-micron scale in many of these taxa, including
the architecture of < 3 μm holococcoliths and the details of grills and processes in other very small
fragile taxa - a size class which is rarely preserved even in Recent sediments. A distinct assemblage
of at least four specialist lower-photic zone taxa - three Gladiolithus species and Algirosphaera
fabaceus - is present in these sediments. The occurrence of these highly specialised coccoliths
in the Palaeogene sediments of Tanzania extends their previously known late Quaternary fossil
record by tens of millions of years. The controls on the exquisite preservation of such a diverse
nannofossil assemblage are difficult to determine but we speculate that a diverse open-ocean,
oligotrophic coccolithophore assemblage was being rapidly buried and sealed within a clay-rich facies
that is more characteristic of shelf-environments, a combination that, to date, makes the Palaeogene
sediments of Tanzania unique. One new genus, Pocillithus, is described, consisting of very small spine-
bearing muroliths that may be related to extant narrow-rimmed muroliths of uncertain affinity. Ten
new species are described: Pocillithus spinulifer, Reticulofenestra macmillanii, Calcidiscus parvus,
Syracosphaera monechiae, Syracosphaera raffiae, Blackites culter, Blackites shafikii, Acanthoica
backmanii, Orthozygus occultus and Orthozygus arcus.

KEY WORDS coccolithophores, taxonomy, nannoplankton, Eocene, Oligocene, preservation

Contents
Introduction 363
Implications for the Palaeogene history of coccolithophores 364
Diversity 364
Holococcoliths 365
Lower photic zone assemblage 366

† Email: tom.dunkleyjones@ucl.ac.uk
‡ Email: p.bown@ucl.ac.uk
§ Email: pearsonp@Cardiff.ac.uk
 360 T O M D U N K L E Y J O N E S ET AL.

Exceptional preservation 366

Materialand methods 367
Systematic framework 368
Systematic descriptions 369
Placolith coccoliths 371
Order Isochrysidales Pascher, 1910 371
Family Noelaerhabdaceae Jerkovic, 1970 emend. Young & Bown, 1997 371
Genus Cyclicargolithus Bukry, 1971 371
Cyclicargolithus floridanus (Roth & Hay in Hay et al., 1967)
Bukry, 1971 371
Genus Reticulofenestra Hay et al., 1966 371
Reticulofenestra bisecta (Hay et al., 1966) Roth, 1970 371
Reticulofenestra dictyoda (Deflandre in Deflandre & Fert, 1954)
Stradner in Stradner & Edwards, 1968 371
Reticulofenestra umbilicus (Levin, 1965) Martini & Ritzkowski,
1968 373
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Reticulofenestra macmillanii sp. nov. 373

Reticulofenestra minuta Roth, 1970 373
Reticulofenestra filewiczii Wise & Wiegand in Wise, 1983
stat. nov. 373
Reticulofenestra cf. R. filewicziiWise & Wiegand in Wise,
1983 stat. nov. 373
Order Coccolithales Schwarz, 1932 emend. Edvardsen & Eikrem in
Edvardsen et al., 2000 373
Family Coccolithaceae Poche, 1913 emend. Young & Bown, 1997 374
Genus Coccolithus Schwarz, 1894 374
Coccolithus pelagicus (Wallich, 1877) Schiller, 1930 374
Coccolithus formosus (Kamptner, 1963) Wise, 1973 374
Coccolithus cachaoi Bown, 2005a 374
Genus Cruciplacolithus Hay & Mohler in Hay et al., 1967 374
Cruciplacolithus cruciformis (Hay & Towe, 1962) Roth, 1970 374
Cruciplacolithus cf. C. primus Perch-Nielsen, 1977 374
Genus Chiasmolithus Hay et al., 1966 376
Chiasmolithus nitidus Perch-Nielsen, 1971 376
Chiasmolithus titus Gartner, 1970 376
Genus Bramletteius Gartner, 1969 376
Bramletteius serraculoides Gartner, 1969 376
Genus Clausicoccus Prins, 1979 376
Clausicoccus subdistichus (Roth & Hay in Hay et al., 1967)
Prins, 1979 376
Clausicoccus fenestratus (Deflandre & Fert, 1954) Prins, 1979 376
Genus Coronocyclus Hay et al., 1966 376
Coronocyclus nitescens (Kamptner, 1963) Bramlette &
Wilcoxon, 1967 376
Coronocyclus cf. C. nitescens (Kamptner, 1963) Bramlette &
Wilcoxon, 1967 376
Family Calcidiscaceae Young & Bown, 1997 378
Genus Umbilicosphaera Lohmann, 1902 378
Umbilicosphaera bramlettei (Hay & Towe, 1962) Bown et al.,
2007 378
Umbilicosphaera jordanii Bown, 2005a 378
Genus Calcidiscus Kamptner, 1950 378
Calcidiscus protoannulus (Gartner, 1971) Loeblich & Tappan,
1978 378
Calcidiscus parvus sp. nov. 378
Calcidiscus? s p. 1 380
Calcidiscus? s p. 2 380
Calcidiscus s p. 3 380
 UPPER EOCENE TO LOWER OLIGOCÈNE CALCAREOUS NANNOFOSSILS FROM TANZANIA 361

Placolith coccoliths incertae sedis 380

Genus Tetralithoides Theodoridis, 1984 380
Tetralithoides symeonidesii Th eodo ridis, 1984 380
Genus Hayella Gartner, 1969 380
Hayella situliformis Gartner, 1969 380
Genus Pedinocyclus Bukry & Bramlette, 1971 380
Pedinocyclus larvalis Bukry & Bramlette, 1971 380
Murolith coccoliths 381
Order Zygodiscales Young & Bown, 1997 381
Family Helicosphaeraceae Black, 1971 381
Genus Helicosphaera Kamptner, 1954 381
Helicosphaera compacta Bramlette & Wilcoxon, 1967 381
Helicosphaera bramlettei (Muller, 1970) Jafar & Martini, 1975 381
Helicosphaera wilcoxonii (Gartner, 1971) Jafar & Martini, 1975 381
Helicosphaera gartneri Theodoridis, 1984 381
Helicosphaera euphratis Haq, 1966 381
Helicosphaera reticulata Bramlette & Wilcoxon, 1967 381
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Family Pontosphaeraceae Lemmermann, 1908 381

Genus Pontosphaera Lohmann, 1902 381
Pontosphaera plana (Bramlette & Sullivan, 1961) Haq, 1971 383
Pontosphaera versa (Bramlette & Sullivan, 1961) Sherwood,
1974 383
Pontosphaera formosa (Bukry & Bramlette, 1968) Romein, 1979 383
Pontosphaera pectinata (Bramlette & Sullivan, 1961)
Sherwood,1974 383
Pontosphaera multipora (Kamptner, 1948) Roth, 1970 383
Pontosphaera alta Roth, 1970 383
Genus Scyphosphaera Lohmann, 1902 383
Scyphosphaera columella Stradner, 1969 383
Scyphosphaera apsteinii Lohmann, 1902 383
Order Syracosphaerales Hay, 1977 emend. Young et al., 2003 385
Family Syracosphaeraceae Lemmermann, 1908 385
Genus Syracosphaera Lohmann, 1902 385
Syracosphaera tanzanensis Bown, 2005a 385
Syracosphaera monechiae sp. nov. 385
Syracosphaera raffiae s p. nov. 385
Genus incertae sedis Coronosphaera Gaarder in Gaarder &
Heimdal, 1977 385
Coronosphaera? sp. 386
Family Calciosoleniaceae Kamptner, 1927 386
Genus Calciosolenia (Gran, 1912) Young et al., 2003 386
Calciosolenia sp. 386
Family Rhabdosphaeraceae Haeckel, 1894 386
Genus Blackites Hay &Towe, 1962 386
Blackites? furvus Bown & Dunkley Jones, 2006 387
Blackites deflandrei (Perch-Nielsen, 1968) Bown, 2005a 387
Blackites clavus Bown, 2005a 387
Blackites cf. Cruxia mericii Varol, 1989 387
Blackites fustis Bown, 2005a 387
Blackites culter s p. n ov. 387
Blackites tenuis (Bramlette & Sullivan, 1961) Sherwood, 1974 389
Blackites spinosus (Deflandre & Fert, 1954) Hay & Towe, 1962 389
Blackites singulus Bown & Dunkley Jones, 2006 389
Blackites shafikii s p. n ov. 389
Blackites s p. 1 389
Genus Rhabdosphaera Haeckel, 1894 391
Rhabdosphaera vitrea (Deflandre in Deflandre & Fert, 1954)
Bramlette & Sullivan, 1961 391
 362 T O M D U N K L E Y JONES ET AL.

Rhabdosphaera gracilentus (Bown & Dunkley Jones, 2006)

comb. nov. 391
Rhabdosphaera xiphos (Deflandre & Fert, 1954) Norris, 1984 391
Rhabdosphaera type 1 391
Genus Algirosphaera Schlauder, 1945 emend. Norris, 1984 391
Algirosphaera fabaceus (Bown & Dunkley Jones, 2006)
Bown et al., 2009 391
Algirosphaera sp. 1 393
Algirosphaera? sp. 2 393
Algirosphaera? sp. 3 393
Genus Acanthoica Lohmann, 1903 emend. Schiller, 1913 and
Kleijne, 1992 393
Acanthoica backmanii sp. nov. 393
Holococcoliths 394
Genus Orthozygus Bramlette & Wilcoxon, 1967 394
Orthozygus occultus sp. nov. 394
Orthozygus aureus (Stradner, 1962) Bramlette & Wilcoxon, 1967 394
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Orthozygus brytika (Roth, 1970) Aubry, 1988 394

Orthozygus arcus sp. nov. 394
Orthozygus sudis Bown & Dunkley Jones, 2006 397
Genus Lanternithus Stradner, 1962 397
Lanternithus minutus Stradner, 1962 397
Lanternithus procerus Bown, 2005a 397
Genus Quadrilateris Varol, 1991 397
Quadrilateris imparidividuus Varol, 1991 397
Genus Varolia Bown, 2005a 397
Varolia cistula Bown, 2005a 397
Varolia sicca Bown & Dunkley Jones, 2006 397
Varolia macleodii Bown & Dunkley Jones, 2006 397
Varolia boomeri Bown & Dunkley Jones, 2006 397
Varolia spp. 397
Genus Corannulus Stradner, 1962 397
Corannulus germanicus Stradner, 1962 398
Genus Holodiscolithus Roth, 1970 398
Holodiscolithus macroporus (Deflandre in Deflandre & Fert,
1954) Roth, 1970 398
Holodiscolithus solidus (Deflandre in Deflandre & Fert, 1954)
Roth, 1970 398
Holodiscolithus geisenii Bown, 2005a 398
Holodiscolithus minolettii Bown, 2005a 398
Genus Zygrhablithus Deflandre, 1959 398
Zygrhablithus bijugatus subsp. bijugatus (Deflandre in
Deflandre & Fert, 1954) Deflandre, 1959 398
Zygrhablithus bijugatus subsp. cornutus Bown, 2005a 398
Genus Clathrolithus Deflandre in Deflandre & Fert, 1954 398
Clathrolithus joidesa (Bukry & Bramlette, 1968) Bown, 2005a 398
Nannoliths 399
Family Braarudosphaeraceae Deflandre, 1947 399
Genus Braarudosphaera Deflandre, 1947 399
Braarudosphaera bigelowii (Gran & Braarud, 1935)
Deflandre, 1947 399
Braarudosphaera cf. B. hockwoldensis Black, 1973 399
Braarudosphaera stylifera Troelsen & Quadros, 1971 399
Genus Micrantholithus Deflandre in Deflandre & Fert, 1954 399
Micrantholithus minimus Bown & Dunkley Jones, 2006 399
Micrantholithus excelsus Bown, 2005a 399
Micrantholithus hebecuspis Bown, 2005a 399
Micrantholithus triquetra (Bown & Dunkley Jones, 2006)
comb. nov. 399
 UPPER EOCENE TO LOWER OLIGOCÈNE CALCAREOUS NANNOFOSSILS FROM TANZANIA 363

Genus Pemma Klumpp, 1953 402

Pemma papillatum Martini, 1959 402
Family Discoasteraceae Tan, 1927 402
Genus Discoaster Tan, 1927 402
Discoastersaipanensis Bramlette & Riedel, 1954 402
Discoaster tanii Bra mlette & Riedel, 1954 402
Discoaster nodifer Bra mlette & Riedel, 1954 402
Discoaster ornatus (Bramlette & Wilcoxon, 1967) Bown, 2005a 402
Discoaster distinctus Martini, 1958 402
Discoaster deflandrei Bramlette & Riedel, 1954 403
Family Sphenolithaceae Deflandre, 1952 403
Genus Sphenolithus Deflandre, 1952 403
Sphenolithus moriformis (Bronnimann & Stradner, 1960)
Bramlette & Wilcoxon, 1967 403
Sphenolithus conicus Bukry, 1971 403
Sphenolithus radians Deflandre, 1952 403
Sphenolithus pseudoradians Bramlette & Wilcoxon, 1967 403
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Sphenolithus predistentus Bramlette & Wilcoxon, 1967 403

Sphenolithus akropodus de Kaenel & Villa, 1996 403
Sphenolithus tribulosus Roth, 1970 405
Nannoliths incertae sedis 405
Pemma?uncinata Bown & Dunkley Jones, 2006 405
Genus Trochoaster Klumpp, 1953 405
Trochoastersimplex Klumpp, 1953 405
Genus Pocillithus gen. nov. 405
Pocillithus spinulifer sp. nov. 405
Genus Gladiolithus Jordan & Chamberlain, 1993 405
Gladiolithus flabellatus (Halldal & Markali, 1955) Jordan &
Chamberlain, 1993 405
Gladiolithus brevis Bown et al., 2009 406
Gladiolithus contus Bown et al., 2009 406
References 408

INTRODUCTION
Fossil Konservat-Lagerstatte - sedimentary units that con-
tain extraordinarily well preserved fossil material (Briggs
2001) - are the only source of information for many taxo-
nomic groups that have limited preservation potential in
standard sedimentary environments. The most famous ex-
amples of Konservat-Largerstatte typically host exquisitely
preserved soft-bodied organisms (e.g. Conway Morris 1990;
Viohl 1990), which provide both a wealth of biological in-
formation and a record of ancient community diversity that
is unavailable from the 'normal' hard-part dominated fossil
record. The consistent presence and often great abundance of
calcareous microfossils in a variety of sedimentary environ-
ments would, at first glance, suggest that these, often robust,
skeletal remains are not subject to significant taphonomic
losses between living and fossil assemblages and hence do
not require exceptional sedimentary conditions for the pre-
servation of a representative flora or fauna. Although this
may be true to some extent for the relatively large and robust
calcareous foraminifera, it is becoming clear that the minute
size, typically < 10 μm, and often fragile nature of calcareous
nannoplankton introduces a significant preservational bias in
their typical fossil record (Young et al. 2005; Bown et al.
2008).
Calcareous nannofossils are principally the remains of
calcite-scale (coccolith) bearing haptophyte alga, known as
coccolithophores (Phylum Haptophyta Hibberd ex Cavalier-
Smith, 1986; Class Prymnesiophyceae Hibberd, 1976; Sub-
class Calcihaptophycidae De Vargas et al., 2007). The fossil
record of coccolithophores extends from Upper Triassic to
Recent marine sediments (Bown et al. 2004), often occur-
ring in rock-forming abundances in chalks and deep-sea cal-
careous oozes. In spite of this apparently complete, continu-
ous Mesozoic and Cenozoic fossil record, studies of living
coccolithophore communities coupled with sediment trap
and core top assemblages have revealed a significant tapho-
nomic loss of small, <2.5 μm, and fragile taxa (Andruleit
et al. 2004; Young et al. 2005). Recent work in the Arabian
Sea recorded a 45% loss of species diversity between liv-
ing coccolithophore communities and assemblages recovered
from supralysoclinal sediment traps at depth in the water
column due to the loss of fragile taxa (Andruleit et al. 2004).
On a global scale, comparisons of the total extant coccolitho-
phore species diversity and the diversity of the Holocene
fossil record indicates that preservation potential is highly
 364 TOM DUNKLEY J O N E S ET AL.
size dependent - more than 90% of living taxa with coc-
coliths > 3 μm are preserved in the fossil record, whereas
only 20% of taxa with coccoliths < 3 μm have a fossil re-
cord (Young et al. 2005). In this case the loss of species
diversity represented by the non-preservation of small taxa
is significant; if rare and exceptional fossil occurrences are
included, only 54% of living species have a Holocene fossil
record but only ~30% are preserved in typical sedimentary
environments (Young et al. 2005).
A large number of taxa within the major coccolitho-
phore clades produce distinct coccolith morphologies in the
two phases of their haplo-diplontic life cycle: robust het-
erococcoliths formed of interlocking calcite crystals in the
diploid phase and fragile holococcoliths formed of small
equidimensional crystallites in the haploid phase (Billard
1994; Young et al. 1999). This difference in construction
makes holococcoliths highly susceptible to dissolution and
greatly reduces their preservation potential in the fossil re-
cord; of the 90 living holococcolith taxa only one, slightly
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larger (3-4 μm) form, Syracolithus schilleri (often identi-
fied as Holodicolithus macroporus), is commonly preserved
in Neogene sediments (Young et al. 2003). This severe
under-representation of both holococcolith taxa and small
fragile heterococcoliths in the fossil record, when compared
to modern diversities, imposes a significant limitation on
our current understanding of ancient coccolithophore di-
versity, phylogeny and evolution. However, much of this
'diversity loss' is concentrated within relatively few ex-
tant groups, notably the Syracosphaeraceae, Rhabdosphaer-
aceae and the incertae sedis groups Alisphaeraceae and Pap-
posphaeraceae, which, although limiting our understanding
of the evolutionary history of these groups, may not af-
fect the remaining heterococcolith families to the same ex-
tent. Similarly, the significant undersampling of holococco-
lith taxa in the fossil record may not significantly impact
the long-term diversity record of coccolithophores, which
is likely to be adequately represented by the record of
the heterococcolith producing life-cycle phase alone. Nev-
ertheless, with the ongoing development of molecular ge-
netic phy logenies for a large proportion of living taxa (Saez
et al. 2004) an accurate long time-series of ancient cocco-
lithophore diversity, from locations with exceptional nanno-
fossil preservation, is highly desirable in order to test phylo-
genetic relationships, provide reliable independent estimates
of key divergence times and test the reliability of the 'stand-
ard' fossil record.
The ongoing research of the Tanzania Drilling Project
(TDP) (Pearson et al. 2004, 2006a; Nicholas et al. 2006),
which has concentrated on the study of exceptionally well
preserved calcareous microfossils from the Cretaceous and
Palaeogene sediments of the Kilwa Group, coastal Tanzania,
has already led to significant improvements in the interpreta-
tion of foraminiferal taxonomy (Pearson et al. 2006b; Wade
& Pearson 2008) and geochemical proxy data (Pearson et al.
2001, 2007, 2008; Lear et al. 2008). The extraordinary and
exquisite assemblages of Palaeogene foraminifera and cal-
careous nannofossils justify the status of the Kilwa Group
as a world-class Konservat-Lagerstatte deposit (Bown et al.
2008). Initial light microscope assessments of sediment cores
drilled between 2002 and 2005 led to the preliminary descrip-
tion of exceptionally diverse assemblages of Palaeogene cal-
careous nannofossils, with species diversities equalling or
exceeding existing estimates of global Palaeogene diversit-
ies (Bown 2005a; Bown & Dunkley Jones 2006; Bown et al.
2007, 2008). Electron microscope imaging of this material
has revealed a quality of preservation and morphological de-
tail that is comparable to well preserved specimens from the
modern plankton (Bown et al. 2008).
The purpose of this paper is to document the calcareous
nannofossil flora from three shallow boreholes, TDP Sites
11, 12 and 17, which recovered an upper Eocene to lower
Oligocene succession including complete sections through
the Eocene-Oligocene Boundary (EOB). This interval of
the Palaeogene marks the final decline in global coccolitho-
phore species diversity from a Cenozoic maximum in the
late Paleocene-early Eocene to a Cenozoic minimum in the
early Oligocene (Bown et al. 2004). This parallels general
trends of both deep-water cooling (Zachos et al. 2001) and
declining atmospheric pCO2 through the Palaeogene (Pear-
son & Palmer 2000; Pagani et al. 2005), which is suggestive
of a potential macroevolutionary coupling between measured
coccolithophore species diversity and global climate, but one
whose nature is as yet poorly understood (Bown et al. 2004;
Bown 2005b).
The long-term trends in middle to late Eocene global cli-
mate appear to accelerate across the Eocene-Oligocene trans-
ition coincident with the rapid formation of a continental-
scale Antarctic ice sheet and a major shift in the global carbon
cycle (Coxall et al. 2005; Zachos & Kump 2005; Lear et al.
2008). As one of the major phy toplankton groups, the record
of coccolithophores through the Eocene-Oligocene trans-
ition is vital for understanding these rapid changes in the
marine ecosystem and carbon cycle. This period is also
an ideal interval to study the effect of large and rapid cli-
matic change on the patterns and processes of coccolitho-
phore evolution. The detailed taxonomic data presented here
is designed to complement both the recent study of biotic
events through this interval (Pearson et al. 2008) and a set
of high-resolution nannofossil assemblage data, and asso-
ciated palaeoceanographic interpretations, through two of
these boreholes (TDP 12 & 17) (Dunkley Jones et al. 2008).
Together, those two papers provide an unprecedented snap-
shot of the taxonomic diversity and ecological responses of
a coccolithophore community through the EOB.
IMPLICATIONS FOR THE PALAEOGENE
HISTORY o f c o c c o l i t h o p h o r e s
Diversity
The distinct and exceptional nature of the Tanzanian cal-
careous nannofossil assemblages becomes apparent when
their species composition and overall diversity is compared
to contemporaneous open-ocean sites. Data for three stand-
ard 'moderately to well preserved' upper Eocene nannofossil
assemblages from recent Ocean Drilling Program sites in the
Pacific (Shatsky Rise Site 1209; Bralower 2005), Atlantic
(Iberia Abyssal Plain Hole 900A; Liu 1996) and Southern
Ocean (Agulhas Ridge Hole 1090B; Marino & Flores 2002)
are compared to the Tanzanian assemblage in Table 1. The
most striking feature is the difference in overall species di-
versities, with values of 16, 47 and 44, respectively, for the
open-ocean sites compared to the 115 species-equivalent
morphotypes described from Tanzania. The additional
diversity in the Tanzanian sections is dominated by
 UPPER EOCENE TO LOWER OLIGOCÈNE CALCAREOUS NANNOFOSSILS FROM TANZANIA
Table 1 Comparison of species diversity across taxonomic groups and between the Tanzanian
sections and open-ocean sites.
Shatsky Rise
Taxonomic Group (Site 1209)
Total species 16
Rhabdosphaeraceae 1 (6%)
Zygodiscales 0 (10%)
Holococcoliths 1 (6%)
Braarudosphaeraceae 0 (0%)
Number of species in each group with percentage contribution to total species diversity in brackets. Note that
data for open-ocean sections are compiled across the majority of the upper Eocene whereas the Tanzanian
sections span only a short stratigraphic interval across the Eocene-Oligocene Boundary (EOB) and so the observed
differential in total species diversity is conservative. See the text for references.
holococcoliths (19 species in Tanzania; maximum of 5 spe-
cies in open-ocean sites) and members of the Rhabdosphaer-
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aceae (20 species in Tanzania; maximum of 2 species in
open-ocean sites); typically members of both these groups
are fragile and highly susceptible to fragmentation and dis-
solution.
The apparent bias against the preservation of fragile taxa
in the open-ocean sites may partially explain the 'excess' spe-
cies diversity in the Tanzanian sections; however, when com-
pared to similar contemporaneous low-latitude shelf-slope
clay facies, with generally excellent preservation, such as the
Yazoo Clay of Mississippi, USA (Dockery et al. 1991) and
the Tegalsari Marls of Java (Lunt & Sugiatno in press), the
Tanzanian material still shows considerable additional spe-
cies diversity (species diversities of 54 for the Yazoo Clay and
40 for the Tegalsari Marls: pers. obs.). Again, this differential
is largely due to the preservation of a diverse holococcolith
and rhabdolith assemblage in Tanzania, as well as the pres-
ence of previously unknown Palaeogene deep-photic zone
coccolithophores (Gladiolithus, Algirosphaera).
The discovery of these new and extraordinarily diverse
assemblages significantly alters the results of recent com-
pilations of global nannofossil species diversity through the
Palaeogene (Bown et al. 2004), which for the upper Eocene
interval of concern here (NP19/20) documented a global nan-
nofossil diversity of 67 species. The description of ~70%
more species from this interval at one location indicates
both the significant taphonomic loss of species diversity in
standard marine environments and the exceptional nature of
the Tanzanian sediments. Although such unique occurrences
have the potential to significantly bias long-term records of
diversity and macroevolutionary patterns of extinction and
origination, the initial species richness data taken from Tan-
zanian sediments ranging from late Paleocene to early Oli-
gocene age produce similar temporal trends, but with higher
absolute diversities, to previously compiled global species
richness data (Bown et al. 2008).
Holococcoliths
Holococcolith production during the haploid phase of the
coccolithophore life-cycle, and heterococcoliths during the
diploid phase, has only recently been established (Billard
1994; Young et al. 1999) and understanding the environ-
mental trigger and adaptive function of the holococcolith-
heterococcolith (haploid-diploid) transition in the cocco-
365
Iberia Abyssal Plain Agulhas Ridge Tanzania
(Hole 900A) (Hole 1090B) (TDP11,12,17)
1
47 44 5
2 (4%) 1 (2%) 20 (17%)
8 (17%) 3 (7%) 12 (10%)
5 (17%) 2 (5%) 19 (17%)
2 (4%) 1 (2%) 8 (7%)
lithophores is at an early stage (Cros et al. 2000; Noel
et al. 2004). Plankton sampling of the modern oceans has
established that holococcoliths are generally more abundant
in surface waters and in oligotrophic environments (Kleijne
1991; Cros et al. 2000). For example, in the Western Medi-
terranean there is a clear peak in abundance of the holococ-
colith phase of Helicosphaera carteri (formerly known as
Syracolithus catilliferus) in near surface waters, while the
heterococcolith phase reaches peak abundances at depths of
40-70 m (Cros et al. 2000). Recent modeling suggests that
the layered crystallites characteristic of many holococcolith
taxa are an adaptation to the near-surface environment as
they are more efficient at backscattering harmful ultra-violet
radiation away from the cell whilst allowing the transmission
of wavelengths within the photosynthetically active band of
400-700 nm (Quintero-Torres et al. 2006).
In spite of the potential environmental sensitivity of
holococcoliths, their use in palaeoceanographic reconstruc-
tions has been minimal due to their generally low preser-
vation potential (Ziveri et al. 2000; Young et al. 2005).
The Tanzanian assemblages presented here are exceptional
in the preservation of holococcolith morphologies with ori-
ginal crystallites down to the 0.1 μm scale, as well as in the
diversity of holococcolith morphologies, with a total of 19
species recorded through a relatively short interval, and in the
total abundance of holococcoliths, which constitute ~17% of
the late Eocene nannofossil assemblage. Based on our cur-
rent understanding of holococcolith ecology, combined with
the abundance of lower photic zone taxa, it appears that the
Tanzanian sediments are sampling a nannofossil assemblage
from a relatively oligotrophic, well-stratified surface water
column with high primary holococcolith diversity and abund-
ance. Detailed nannofossil assemblage data through the EOB
indicates a marked fall in holococcolith abundance into the
earliest Oligocene, which may be related to increased mix-
ing and a relative eutrophication of surface waters (Dunkley
Jones et al. 2008). The large shift in holococcolith abund-
ances through this interval of major oceanographic change
indicates the potential for holococcolith abundances to be
used as a powerful palaeoceanographic proxy in other sec-
tions with well-preserved holococcolith assemblages. The
holococcolith taxonomy presented in this paper will also
form the basis for detailed comparisons with other time slices
from the Palaeogene record of Tanzania to produce a long-
term macroevolutionary record of holococcolith diversity,
morphology and evolution.
 366 T O M D U N K L E Y J O N E S ET AL.
Lower photic zone assemblage
The scanning electron microscope (SEM) examination of
rock-chip surfaces in this study and in studies of lower Pa-
leocene and middle Eocene sediments has documented an ex-
traordinary assemblage of previously unknown Palaeogene
lower photic zone (LPZ) coccolithophores (Bown et al. 2008,
2009). This Palaeogene LPZ assemblage was first noticed
when highly modified, tubular, coccoliths where observed in
SEM studies which were indistinguishable from those of the
extant LPZ coccolithophore Gladiolithus flabellatus. Sub-
sequent extensive examination of sediments from late Paleo-
cene (nannofossil zone NP6) to early Oligocene (zone NP21)
age revealed a diverse LPZ assemblage including four spe-
cies of Gladiolithus and representatives of the modern LPZ
genera Solisphaera and Algirosphaera (Bown et al. 2009).
In the late Eocene-early Oligocene assemblages described
here, the distinctive LPZ assemblage consists of three Gladi-
olithus species (G. flabellatus, G. brevis and G. contus), and
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the earliest known occurrence of the genus Algirosphaera.
The occurrence of these taxa in the Palaeogene of Tanzania
extends their fossil record, which was previously limited
to sporadic occurrences in the late Quaternary (Okada &
Matsuoka 1996; Lars Legge et al. 2006), by between 35 and
55 Ma and is, to our knowledge, the first reported occurrence
of the fragile basal coccoliths (lepidoliths) of Gladiolithus in
the fossil record. The presence of this LPZ flora, which in
the modern oceans is found in water depths of 100-200 m
in clear, blue-water environments (Okada & Honjo 1973;
Okada 1983,1992) supports existing palaeo-water-depth es-
timates of 300-500 m for the Tanzanian sections (Pearson
et al. 2004, 2006a; Nicholas et al. 2006).
These new Palaeogene occurrences of extant LPZ coc-
colithophore taxa have three main implications: (1) they con-
strain the origin of a specialised LPZ flora to at least the early
Palaeogene; (2) they imply long Neogene ghost ranges for
these taxa and the possibility that Neogene deposits with
similar exceptional preservation may preserve some of this
'missing' diversity; (3) they document the highly conservat-
ive nature of many coccolithophore morphologies over tens
of millions of years, such as Gladiolithus flabellatus, which
indicates the presence of a strong stabilising selection on
these morphologies and a high degree of coccolith function-
ality; presumably, in the case of Gladiolithus this represents a
long-term and successful adaptation to the LPZ environment.
Exceptional preservation
The controls on exceptional calcareous microfossil preser-
vation in the Tanzanian material remain only partially un-
derstood. The Kilwa Group is dominated by a relatively
impermeable clay facies that would have severely restricted
post-burial fluid flow and thus facilitated a rapid equilibration
of carbonate ion concentrations between buried calcareous
microfossils and the surrounding pore fluids. Associated or-
ganic biomarkers from these sediments are also exceptionally
well preserved (van Dongen et al. 2006) suggesting rapid but
not deep burial. The relatively low sediment organic carbon
contents may also be a prerequisite for excellent preservation
but these factors alone do not appear sufficient to explain
the unique nature of these assemblages (Dunkley Jones &
Bown 2007; Bown et al. 2008). It is striking that sediment
trap samples of settling particulate matter within the water
column of the modern Arabian Sea, a location with generally
good to excellent coccolith preservation in sediments, con-
tains only limited holococcolith material and considerably
altered laths of Gladiolithus flabellatus without the associ-
ated basal lepidoliths (Andruleit et al. 2004). This is in dir-
ect contrast with the preservation of a diverse and abundant
holococcolith assemblage as well as numerous occurrences
of near complete collapsed Gladiolithus coccospheres, with
abundant associated lepidoliths (see below), throughout the
Palaeogene sediments of the Kilwa Group.
From the Arabian Sea data and similar sets of water
column sampling it appears that there are two primary loca-
tions for coccolith and coccosphere dissolution and breakage:
first during biological cycling in the surface ocean and second
in the early stages of sediment burial and diagenesis, with re-
latively little alteration occurring once coccolith material has
been efficiently 'packaged' into fast-sinking faecal pellets or
marine snow aggregates. In contrast to the results of Andruleit
et al. (2004), coccolith material contained within zooplank-
ton faecal pellets collected at water depths of 1000-3000 m
have been shown to contain very well preserved coccolith
material including holococcoliths and coccospheres compar-
able to the Tanzanian assemblages (Honjo 1976; Sprengel
et al. 2000). During detailed SEM observations of rock-chip
surfaces of the Tanzanian material it was qualitatively noted
that the best preservation is found in relatively large patches
or aggregates - most probably the remnants of zooplankton
faecal pellets - of several tens to hundreds of coccoliths,
whereas the preservation of isolated liths throughout the sed-
iment matrix is more variable, ranging between good and
moderate preservation.
In a number of other clay-rich sections (e.g. late Eocene
Yazoo Clay of the US Gulf Coast; middle Eocene Hamp-
den Formation of New Zealand) where we have undertaken
similar SEM examinations of rock-chip surfaces there is a
comparable exceptional preservation of fine structure, such
as sub-micron grills, in many common placoliths but without
the same extraordinary diversities of holococcoliths, mem-
bers of the Syracosphaerales and the LPZ flora seen in Tan-
zania. It should be noted that much of the diversity that is lost
between the plankton assemblages and settling assemblages
of Andruleit et al. (2004) and between the modern plankton
and the Holocene fossil record of Young et al. (2005) is con-
centrated within taxa that are both rare and fragile. It is the
export and preservation of a greater portion of this rare and
fragile diversity in the Tanzanian sediments that is contrib-
uting to these unique Palaeogene assemblages. From these
available lines of evidence we tentatively suggest that the
exceptional nannofossil assemblages of Tanzania are due to
an unusual combination of an open-ocean, oligotrophic and
hence diverse coccolithophore assemblage being preserved
within a clay-rich facies with a relatively rapid sediment-
ation rate. Normally clay-rich facies with good nannofossil
preservation are found in shelf environments that tend to have
low primary coccolithophore diversity. This is in contrast to
open-ocean assemblages with high primary coccolithophore
diversities but which are subject to significant nannofossil
dissolution and diagenesis in carbonate-rich sediments. We
therefore suggest that the Tanzanian upper slope palaeoen-
vironment, adjacent to a relatively narrow shelf, brought to-
gether a diverse 'blue-water' coccolithophore assemblage
with the rapid deposition of clay-rich sediments necessary
for its preservation.
 UPPER EOCENE TO LOWER OLIGOCÈNE CALCAREOUS NANNOFOSSILS FROM TANZANIA 367

u
d
o
U N
ö
u No
eu
TDP Cores

KENYA

Pemba Is.

J\ Zanzibar Is.
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Mafia TDP11, 12, 17

Figure 1 Stratigraphic extent of the Palaeogene TDP boreholes recovered between 2002 and 2005 (left); nannofossil zonation (NPNf Zones)
from Martini (1971) and planktonicforaminifer zonation (PF Zones) from Berggren & Pearson (2005); correlation of the two plankton zonation
schemes and time-scale are from the latter. Location of the Eocene-Oligocene boundary boreholes, Pande Pennisula, used in this study (right).

MATERIAL AND METHODS
The three boreholes that recovered EOB sections, TDP Site
11 (drilled October 2004; UTM 37L 560250 8983211; total
depth 102.8 m), TDP Site 12 (drilled October 2004; UTM
37L 560222 8981309; total depth 147.4 m) and TDP Site 17
(drilled August 2005; UTM 37L 560539 8984483; total depth
125.9 m), are all located along a 3 km stretch of the 'Pande
Road' between the villages of Mkazambo and Pande, Lindi
District, southern coastal Tanzania (Fig. 1). The drilling tech-
niques, sampling strategy, geological setting and lithostrati-
graphy are described in Pearson et al. (2004, 2006a) and
Nicholas et al. (2006). All cores are archived at the Tanzania
Petroleum Development Corporation in Dar-es-Salaam,
Tanzania.
All three boreholes recovered a succession of dark
greenish-grey clays with occasional limestone interbeds, typ-
ical of the middle Eocene to lower Oligocene Pande Form-
ation (Nicholas et al. 2006). The sections span the EOB,
nannofossil zones NP19/20 - undifferentiated due to the
unreliability of the marker species Sphenolithus pseudora-
dians (Perch-Nielsen 1985) - to NP21 (Martini 1971) and
planktonic foraminifera zones E15/16 to O1 (Berggren &
Pearson 2005). The in situ benthic foraminiferal assemblage
of the Pande clays recovered from these boreholes consists
of deep-water agglutinated and calcareous forms, domin-
ated by Bathysiphon, which is typical of outer shelf to up-
per slope environments (300-500m water depth: Nicholas
et al. 2006). Abundant allocthonous microscopic shell debris,
transported from the innermost shelf, is present throughout
the succession and often dominates benthic foraminiferal
assemblages. The recent identification of the deep-photic
zone, calcareous phytoplankton genus Gladiolithus (see Pl.
16, figs 2-6) throughout the TDP12 and TDP17 successions
 368 T O M D U N K L E Y J O N E S ET AL.
constrains the setting to a clear-water, open ocean envir-
onment (Jordan & Chamberlain 1997; Takahashi & Okada
2000).
Samples were prepared as smear-slides (Bown & Young
1998) and analysed using a Zeiss Axiophot microscope at
x 1000 magnification in cross-polarised (XPL) and phase-
contrast light (PC). Assemblages from TDP11, 12 and 17
were first logged semi-quantitatively, with slides observed
for at least 45 minutes; assemblages from sites TDP12 and
TDP17 were counted at ~1 m intervals; the method and res-
ults are presented in Dunkley Jones et al. (2008). Light mi-
croscope images were captured using Scion Image software
and macros written by Dr Jeremy Young (Bown & Young
1998). SEM studies were conducted on broken rock-surfaces
using a JEOL Digital JSM-6480LV SEM (Lees et al. 2004) -
simple smear and strew preparations were tried but these yiel-
ded poor results due to the abundance of fine clay particles,
which tend to coat nannofossils during these preparations.
All light microscope images were captured at the same mag-
Downloaded by [University of Nebraska, Lincoln] at 23:04 13 July 2013
nification (×2180); SEM images are scaled to appropriate
magnifications with 2 μm scale-bars shown on each plate.
Systematic framework
The detailed systematic descriptions, from order to spe-
cies level are based on recent advances in coccolithophore
systematics and higher taxonomy (Young & Bown 1997;
Edvardsen et al. 2000; Young et al. 2003; Jordan et al. 2004;
Saez´ et al. 2004). The long-used informal grouping of 'coc-
colithophores' to denote the calcite-scale producing hapto-
phyte algae, has recently been formalised by the definition
of the subclass Calcihaptophycidae, which forms a mono-
phyletic group stemming from the last common ancestor of
all calcifying haptophytes. This group is defined by a num-
ber of ultrastructural features of the flagellar/haptonematal
complex and includes some taxa that do not calcify or only
calcify in one phase of their life cycle (De Vargas et al.
2007). The taxonomy of the coccolithophores is based largely
upon the structure of the calcite scales, or coccoliths, pro-
duced by these single-celled algae. This contrasts with the
taxonomy of other extant unmineralised haptophyte algae,
which is based upon cell cytology, biology and, increasingly,
molecular genetics, with organic scale morphology not be-
ing used in classifications above generic level (Saez´ et al.
2004). However, the specific nature of calcite biomineral-
isation - involving the controlled growth of an anisotropic
crystalline medium - requires the operation of complex bio-
chemical processes, which appear to be highly conservative
and hence closely correspond with coccolithophore phylo-
geny (Saez´ et al. 2004). Indeed the taxonomy of coccolitho-
phores, which has largely been the work of palaeontologists,
has generally been proved robust by subsequent cytological
and molecular genetic studies on extant taxa (Saez´ et al.
2004).
The classification of coccolithophores, based upon the
complex structure of their coccoliths, depends upon the reli-
able recognition of homologous structural elements. It is now
understood that there is an underlying deep homology in coc-
colith structure rooted in the fundamental biomineralisation
mechanism common to all calcifying haptophytes (Young
et al. 1992, 2004). In all cases of heterococcolith formation
observed to date, biomineralisation begins with the nucle-
ation of a ring of calcite crystals - the proto-coccolith ring -
upon an organic base-plate scale within an intra-cellular ves-
icle. The proto-coccolith ring is formed of regularly spaced
calcite crystals with alternating sub-vertical and sub-radial
c-axis orientations (Young et al. 1992, 2004). These have
been termed V- and R-units respectively, and their subsequent
growth into complex coccolith structures forms the basis for
coccolithophore taxonomy (Fig. 2). One of the fundamental
advances of this 'V/R model' of coccolith formation is the
identification of characters useful for higher taxonomy -
those stemming from initial growth from the proto-coccolith
ring, notably rim/shield structure and crystallography - and
those formed late in the process of coccolith biomineral-
isation, which exhibit a high degree of plasticity and are
only useful for species level taxonomy, such as central area
structures and process/spine morphologies. The character-
istic coccolith morphology and V/R structure of five extant
(and Palaeogene) families are shown in Fig. 3 (after Young
et al. 2004).
The V/R model has proved to be a powerful tool for coc-
colithophore taxonomy and systematics, forming the basis of
recent revisions to the classification of Cenozoic nannoplank-
ton (Young & Bown 1997; Young et al. 2003). This purely
coccolith-structure-based classification is now being gradu-
ally integrated with new phylogenetic data from the molecu-
lar genetic analysis of extant haptophyte algae (Edvardsen
et al. 2000; Saez´ et al. 2004). Where molecular genetic data
is available it has become clear that the clades identified
by coccolith-structure analysis are robust. Figure 4 presents
a summary of extant haptophyte phylogeny based on mo-
lecular genetics integrated with coccolith-structure analysis
(after Young et al. 2005; see also Jordan et al. 2004). The
main discrepancy between these two approaches is in the
assignment of the appropriate levels of higher classification
and the relationships between families within the 'holococco-
lithophore producing clade' identified in Fig. 4. On the basis
of molecular genetics, Saez´ et al. (2004) recognise this entire
group as an order level clade, the 'Coccolithales', consisting
of the well-supported sub-clade of the Coccolithaceae, Cal-
cidiscaceae, Hymenomonadaceae and Pleurochrysidaceae,
and a number of other groups with poorly resolved relation-
ships. Herein, following Jordan et al. (2004) and Young et al.
(2005), the order Coccolithales has a more restricted use
denoting the sub-clade Coccolithaceae, Calcidiscaceae, Hy-
menomonadaceae and Pleurochrysidaceae, whilst the orders
Zygodiscales and Syracosphaerales are retained on the basis
of the homologies of coccolith structure - this is not in-
consistent with the available molecular genetic data. Recent
analyses of single cell small subunit rDNA has confirmed the
placement of the enigmatic genus Braarudosphaera within
the coccolithophore clade but its affinities within this group
are still uncertain (Takano et al. 2006).
This study could represent a further step in the integ-
ration of palaeontological/coccolith-structure data with the
molecular genetic approach to coccolithophore phylogeny
and classification. The Tanzanian assemblages provide the
most detailed view on Palaeogene coccolithophore diversity
available to date, both in terms of species diversities and also
in the presence of the majority of extant coccolithophore
families (Fig. 4). The description of such well-preserved ma-
terial will both aid and, hopefully, encourage future detailed
taxonomic, palaeobiological and evolutionary studies of Pa-
laeogene coccolithophores.
 UPPER EOCENE TO LOWER OLIGOCÈNE CALCAREOUS NANNOFOSSILS FROM TANZANIA 369

distal shield element lower tube

element

complete
coccolith

incomplete
coccolith

upper & lower upper tube

proto-coccolith ring V-unit nuclei proximal shield elements element
R-unit nuclei
Downloaded by [University of Nebraska, Lincoln] at 23:04 13 July 2013

Figure 2 V/R model of coccolith growth (after Young et al. 2004). Growth pattern of Coccolithus pelagicus from proto-coccolith ring of
alternating V- and R-units (left) and final coccolith structure (right).

SYSTEMATIC DESCRIPTIONS clarifications and is not a verbatim statement of the original

diagnosis or description of the taxon concerned. The term
Descriptive terminology follows the guidelines of Young diagnosis is also used where new species-level taxa are be-
etal.(1 997). The layout of the taxonomy and the higher taxo- ing defined. Where available, light microscope images are
nomic rationale follows recent classification schemes for ex- selected as holotypes as the microscope slides these were
tant coccolithophores (Young et al. 2003; Jordan et al. 2004) derived from can be preserved and archived. Abundance and
combined with the review of Cenozoic coccolithophore tax- occurrence of species is noted using the terms 'very rare'
onomy by Young & Bown (1997). The term 'diagnosis' is (sporadic occurrences; ~ l - 2 per slide), 'rare' (<1 specimen
used for taxa above the species level to denote a summary per field of view (FOV)), 'frequent' (1 perFOV), 'common'
diagnosis of current taxonomic understanding. This is based (1-10perFOV) and 'abundant' (>10perFOV). Size classes,
on both the original descriptions and later modifications or based on long-axis length, are 'very small' (<3 μm), 'small'

Coccolithaceae (Coccolithus) Noelaerhabdaceae (Gephyrocapsa)

Helicosphaeraceae (Helicosphaera)
Calcidiscaceae (Calcidiscus)

T-units

Calcidiscaceae ( Umbilicosphaera) Syracosphaeraceae (Syracosphaera)

Figure 3 Coccolith structure of five extant (and Palaeogene) coccolithophore families - R-units are white, V-units dark grey, circles indicate
the location of the proto-coccolith ring. The Syracosphaeraceae have additional T-units, which form the radial lath cycle, and have a tangential
c-axis orientation (after Young et al. 2004).
 370 TOM DUNKLEY J O N E S ET AL.

Polymorphic coccospheres? N Y Y Y n Y Y
CALCIHAPTOPHYCIDAE

HOLOCOCCOLITHOPHORE PRODUCING CLADE

o Qn) Qn) 2n Qn) Qn) Qn) QnJ Qn; Qn; Qn; Qn;
? fn ,
(n) nn [n) fn V nn Cn , C n , vn 1 vn ;

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oliths
I il)
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B33B
i i 1 'S

cosphae
Cale iosoleni

Rhabbdospha

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Downloaded by [University of Nebraska, Lincoln] at 23:04 13 July 2013

Isochrysidales Syraco- Zygo- Cocco ithaales

1 sphaerales discales
1

Class PRYMNESIOPHYCEAE

Figure 4 Consensus phylogeny of extant Haptophyte families based on molecular genetics-dotted lines indicate inferred relationships
where genetic data are not available (after Young et al. 2005). Shaded circles indicate life-cycle phases that produce coccoliths (n, haploid
phase; 2n, diploid phase; no shading indicates a life-cycle phase that does not produce coccoliths). Top line indicates the presence of
polymorphic coccospheres (N, no polymorphism; n, variomorphism but no true polymorphism; y/Y, polymorphism rare/common in group).
Families highlighted in grey are represented in the Eocene-Oligocene sediments described herein.

(3-5 μm), 'medium' (5-8 μm), 'large' (8-12 μm) and 'very
large' (> 12 μm) (Young et al. 1997). Morphometric data are
given for all new taxa.
Range information, where available, is given first for
a taxon's stratigraphic distribution within the EOB sections
presented here (TDP Sites 11, 12 and 17), second for all
the Cenozoic Tanzania Drilling Project sites following Bown
(2005a) and Bown & Dunkley Jones (2006) but updated with
more recent information and,finally,for its previously known
occurrences within the literature. Some caution is required
when using this data. First, due to the short stratigraphic in-
terval covered by the EOB cores studied here, only a few
species have reliable first or last occurrences within these
sections. The range data presented across all of the TDP sites
is fundamentally limited by the intervals recovered to date
(Fig. 1), which are focused around the Paleocene-Eocene
boundary, the middle Eocene zone NP15 and the EOB. This
biases much of this range data - for example any taxa appear-
ing between NP11 (TDP Site 3) and NP14b/15a (TDP Site 2)
will all register lowest occurrences in NP14b/15a. Likewise
there can be no range data extending beyond the outer limits
of NP6 (TDP Site 19) and NP23 (TDP Sites 1 and 6). In
the literature there is an absence of recently compiled, reli-
able range data for Palaeogene calcareous nannofossils. The
last extensive and reliable review of Palaeogene nannofossil
biostratigraphy is that of Perch-Nielsen (1985), which we use
as a basis for the literature ranges quoted here. For ranges
that extend into the Neogene we use highest occurrences
from the biostratigraphic scheme of Young (1998). Several
groups have very poorly known species ranges - notably
those whose occurrence is highly dependent on preserva-
tion or are generally limited to shelf-facies (e.g. holococco-
liths, Pontosphaeraceae, Rhabdosphaeraceae). In these cases
occurrence or range data from the original species descrip-
tion is quoted. Finally, a large number of species discussed
here were first described from the Tanzanian sections (Bown
2005a; Bown & Dunkley Jones 2006) and, to date, have not
been reported from other locations. In this case the Tanzanian
range data, with the caveats outlined above, is the only data
available.
All calcareous nannofossil zones refer to the NP
biozones of Martini (1971). The abbreviations LO and HO
are used to denote lowest occurrence, i.e. oldest/lowest oc-
currence in the sections, and highest occurrence, i.e. young-
est/highest occurrence in the sections, respectively. Addi-
tional abbreviations used in the descriptions are: CW, coc-
colith width; D, diameter; Ds, distal shield; H, height; L,
length; LM, light microscope; PC, phase-contrast illumina-
tion; Pr, proximal shield; SW, spine width; W, width; XPL,
cross-polarised light. Type material and images are stored
in the Department of Earth Sciences, University College
London.
 UPPER EOCENE TO LOWER OLIGOCÈNE CALCAREOUS NANNOFOSSILS FROM TANZANIA 371

Placolith coccoliths occurrence of the large form, in these sections, is immedi-

ately below the EOB and is then consistently present from
Order ISOCHRYSIDALES Pascher, 1910 this point upwards into the lower Oligocene (NP21). The
DIAGNOSIS. Includes the extant placolith-forming Noelaer- large morphotype is, however, commonly observed in middle
habdaceae and the non-calcifying Isochrysidaceae as well as Eocene sediments from Tanzania (Bown 2005a). Cyclicar-
the extinct family Prinsiaceae. The grouping of the extant golithus floridanus is present in the Tanzanian sections from
families Noelaerhabdaceae and Isochrysidaceae is based on NP14b to NP23. Reported range in the literature is from
flagellar characteristics (haptonema vestigial), the production NP12 (Okada 1980) to NN7 (Young 1998).
of alkenones and molecular genetics (Edvardsen et al. 2000;
Fujiwara et al. 2001; Young et al. 2003; Saez´ et al. 2004).
Genus RETICULOFENESTRA Hay et al., 1966
The inclusion of the Prinsiaceae is based on a shared cocco-
lith structure with the Noelaerhabdaceae, namely a well de- DIAGNOSIS. Circular or elliptical Reticulofenestra-type pla-
veloped R-unit that forms the proximal shield-element, two- coliths lacking in distinctive features; central-area can be
tube elements with opposite senses of imbrication and usually open, partially closed by extensions of the inner tube-
a central-area element (Young & Bown 1997). Central-area elements or spanned by a net or plug.
structures are always conjunct and formed from either the
central-area element or the inner tube-element of the prox- Reticulofenestra bisecta (Hay et al., 1966) Roth, 1970
imal shield. (Pl. 1, figs 4 & 18; Pl. 2, fig. 19)
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Family NOELAERHABDACEAE Jerkovic, 1970 DESCRIPTION. Medium to large reticulofenestrid coccoliths

emend. Young & Bown, 1997 with a central plug that are <10μm (NB the holotype is
DIAGNOSIS. Placolith-bearing forms with Reticulofenestra- ~8 μm).
type placolith structure; i.e. V-unit virtually absent with R-
units forming the proximal and distal shields, the inner and REMARKS. Most specimens of the R. bisecta morphology
outer tube-cycles, grill and any central-area structures (see are within the 8-10 μm size range, although occasional spe-
Fig. 3). Strongly birefringent in XPL. cimens larger than 10 μm were rarely observed and classified
as R. stavensis (Levin & Joerger, 1967) Varol, 1989 following
Bown (2005b).
Genus CYCLICARGOLITHUS Bukry, 1971
DIAGNOSIS. Sub-circular to circular Reticulofenestra-type OCCURRENCE. Frequent to common throughout these EOB
placoliths with a narrow central opening. Often possess a sections. Ranges from NP17 to NP23 in the Tanzanian sec-
higher tube cycle than typical Reticulofenestra coccoliths. tions. Reported literature range is from NP17 (Perch-Nielsen
1985) to NN1 (Young 1998).
Cyclicargolithusfloridanus(Roth & Hay in Hay et al.
1967) Bukry, 1971 (Pl. 1, figs 1-3,14-15 & 20) Reticulofenestra dictyoda (Deflandre in Deflandre &
Fert, 1954) Stradner in Stradner & Edwards, 1968 (Pl.
1, figs 5,16-17 & 19)
DESCRIPTION. Small to large subcircular reticulofenestrid
coccoliths with a high tube-cycle and in light-microscopy a
narrow, apparently vacant, central-area. DESCRIPTION. Used here for all elliptical reticulofenestrid
coccoliths between 3 and 14 μm with a relatively wide and,
REMARKS. During SEM analysis a finely perforate net, or in LM, apparently vacant central area.
remnants of one, was consistently present within the central
areas of all observed specimens (Pl. 1, figs 14-15 & 20). REMARKS. AS with Cyclicargolithus floridanus, SEM ima-
In proximal views these have a similar reticulate stucture to ging reveals the consistent presence of a fine central-area net
other reticulofenestrids (compare C. floridanus Pl. 1, fig. 14 in all specimens observed (Pl. 1, figs 16-17 & 19). Specimens
with R. dictyoda Pl. 1, fig. 16 or R. macmillani Pl. 1, fig. smaller than 3 μm are included in Reticulofenestra minuta
21) but are distinctly convex, curving up into the tube rather and those larger than 14 μm in R. umbilicus. Following
than being flush with the proximal surface (Pl. 1, fig. 20) Bralower & Mutterlose (1995) the 'intermediate' reticulofen-
and on the distal side some of the elements extend upward estrid taxon R. samodurovii (Hay et al. 1966) Roth, 1970 has
forming a pattern of curving ridges, in contrast to the nets not been used to separate specimens in the 3-14 μm range.
of Reticulofenestra species, which appear to lie flatter on the It should be noted that the size ranges of the original de-
distal surface (e.g. Pl. 1, figs 16-17 & 21-22). The pres- scriptions of both species overlap; R. samodurovii holotype
ence of central area nets does not conflict with the original length is 5.9 μm; the size range of the original description
description of the genus (Bukry 1971), which makes no spe- of R. dictyoda is 5.6-6.6 μm. The majority of R. dictyoda
cific mention of central area structures. SEM images of the specimens in these sections are large (~10-14 μm) and prob-
distal shield clearly show a raised tube-cycle surrounding the ably intergrade with R. umbilicus; however without detailed
central area (Pl. 1, figs 15 & 20). Specimens were informally morphometric study, ideally from a number of locations, it
divided into small (<5 μm) and large (>5 μm) morphotypes; is difficult to separate these late Eocene forms reliably into
the diameter of the large morphotype is typically in the range meaningful size classes.
5-8 μm.
OCCURRENCE. Common in the upper Eocene (NP 19/20 and
OCCURRENCE. The small morphotype is common through- lower NP21) but becomes very rare in the lower Oligo-
out these EOB sections (NP19/20-21) but the first common cene (upper NP21) of these EOB sections. Ranges from
 372 TOM D U N K L E Y J O N E S ET AL.
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Plate 1 LM images of Cyclicargolithus and Reticulofenestra placoliths (XPL): Figs 1-3 Cyclcargolithus floridanus; 1, TDP17/15-1,10 cm; 2,
small (< 5 μm) morphotype TDP17/5-1,10 cm; 3, TDP17/5-1, 7 cm. Fig. 4 Reticulofenestra bisecta TDP12/10-1, 55 cm. Fig. 5 Reticulofenestra
dictyoda TDP11/23-1, 23 cm. Fig. 6,7 Reticulofenestra macmillani sp. nov.; 6, TDP11/23-1, 23 cm (holotype); 7, TDP12/10-1, 55 cm (paratype).
Fig. 8 Reticulofenestra minuta TDP12/47-1,61 cm. Fig. 9 Reticulofenestra umbilicus TDP11/23-1, 23 cm. Figs 10-12 Reticulofenestra filewiczii;
10, TDP11/23-1, 23 cm; 11, TDP12/38-1,90 cm; 12, TDP11/15-1, 24 cm. Fig. 13 Reticulofenestra cf. R. filewiczii TDP17 / 42-1, 72 cm. SEM images of
Cyclicargolithus and Reticulofenestra placoliths. Figs 14,15,20 Cyclicargolithus floridanus, TDP12/26-2,62 cm. Fig. 16,17,19 Reticulofenestra
dictyoda; 16, TDP12/23-2, 79 cm; 17,19, TDP12/26-2, 62 cm. Fig. 18 Reticulofenestra bisecta TDP12/26-2, 62 cm. Figs 21,22 Reticulofenestra
macmillanii sp. nov. TDP12/26-2, 62 cm; 21, proximal view; 22, distal view. Figs 23-25 Reticulofenestra minuta? TDP12/26-2,62 cm. All LMs and
SEMs above the white dividing line are to the same scale; SEMs below the white line are enlarged relative to the LMs (see scale bars).
 UPPER EOCENE TO LOWER OLIGOCÈNE CALCAREOUS NANNOFOSSILS FROM TANZANIA
NP14b to NP23 in the Tanzanian sections. Reported literature
range is from NP13 to NP16 (Perch-Nielsen 1985) although
this species was originally described from both Eocene and
Oligocene sediments (Deflandre & Fert 1954). The last oc-
currence of R. dictyoda is poorly constrained due to taxo-
nomic difficulties within the reticulofenestrid lineage.
Reticulofenestra umbilicus (Levin, 1965) Martini &
Ritzkowski, 1968 (Pl. 1, fig. 9)
DESCRIPTION. Very large (> 14 μm) elliptical reticulofenes-
trid coccoliths with a relatively wide and, in LM, apparently
vacant central area.
REMARKS. The use of a placolith length of 14 μm as the
lower boundary for R. umbilicus follows the morphometric
work of Backman & Hermelin (1986), which concluded that
this condition was the most precise for the biostratigraphic
use of the LO of R. umbilicus in the middle Eocene. The size
distribution of large reticulofenestrids varies through time in
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the late Eocene and early Oligocene (Backman & Hermelin
1986) and probably varies biogeographically as well, making
species concepts somewhat arbitrary without detailed, site-
specific morphometric data. Reticulofenestra hillae Bukry
& Percival, 1971 has been used for large late Eocene and
early Oligocene reticulofenestrids with a narrower central-
area opening of less than a third of the placolith width.
OCCURRENCE. Rare throughout these EOB sections. Ranges
from NP16 to NP21 in the Tanzanian sections. Reported
literature range is from NP16 to NP22 (Perch-Nielsen 1985).
Reticulofenestra macmillanii sp. nov. (Pl. 1, figs 6-7
& 21-22)
DERIVATION OF NAME. Named after Dr Ian McMillan, ex-
pert on the micropalaeontology of Southern Africa.
DIAGNOSIS. Small elliptical reticulofenestrids with a narrow
rim, narrowly separated proximal and distal shields and a very
wide central-area, approximately three times the width of the
rim, which is spanned by a weakly birefringent net the out-
er edge of which is less heavily calcified than the central part.
DIMENSIONS. L 4.7 μm, W 4.3 μm.
TYPE MATERIAL. Holotype, Pl. 1, fig. 6. Paratype, Pl. 1,
fig. 7. Type locality, TDP Site 11, Pande, Tanzania. Type
level, lower Oligocene, Sample TDP11/23-1, 23 cm (Sub-
zone NP21).
OCCURRENCE. Rare throughout these EOB sections;
NP19/20-21; TDP Site 11, 12, 17.
Reticulofenestra minuta Roth, 1970 (Pl. 1, figs 8 &
23-25)
DESCRIPTION. Very small (<3 μm) elliptical reticulofenes-
trid coccoliths with, in LM, an apparently vacant central-area.
REMARKS. Small (~3 μm), elliptical reticulofenestrids were
observed in SEM (Pl. 1, figs 23-25), which probably corres-
pond to R. minuta placoliths observed in LM. In SEM these
have a distinctly raised but narrow tube-cycle giving the ap-
pearance of a beaded rim around the central-area, which in
turn is spanned by a coarse grill.
373
OCCURRENCE. Common to abundant throughout these EOB
sections. Ranges from NP14b to NP23 in the Tanzanian sec-
tions. Reported literature range is from NP18 (Perch-Nielsen
1985) to NN21 (Young 1998).
Reticulofenestra filewicziiWise & Wiegand in Wise,
1983 stat. nov. (Pl. 1, figs 10-12)
BASIONYM. Reticulofenestra bisecta filewiczii Wise &
Wiegand in Wise, 1983: pl. 6, fig. 2.
DIAGNOSIS. Medium to large, elliptical reticulofenestrids
with a narrow, elongated central-area spanned by a weakly
birefringent central area net.
REMARKS. This upper Eocene subspecies of R. bisecta was
described by Wise & Wiegand, solely on the basis of SEM
images, as an R. bisecta type morphology with a central-area
that is only partially closed, unlike the fully closed central-
area in R. bisecta bisecta. LM images of a form described
as R. bisecta filewiczii, and similar to those figured here,
are shown in de Kaenel & Villa (1996). This morphology is
distinct from specimens of R. bisecta with its closed central-
area, and is common in upper Eocene and lower Oligocene
sediments; warrants species status.
DIMENSIONS. L 5.7-8.3 μm, W 4.7-7.3 μm.
OCCURRENCE. Frequent throughout the EOB section.
Ranges from NP19/20 to NP21 in the Tanzanian sections.
First reported in the literature from the upper Eocene and
Oligocene of the Falkland Plateau (Wise 1983).
Reticulofenestra cf. R. filewiczii Wise & Wiegand in
Wise, 1983 stat. nov. (Pl. 1, fig. 13)
DIAGNOSIS. Medium to large, elliptical reticulofenestrids
with a R. filewiczii form but with slightly wider central areas
that are spanned by a faint grill that has a ring of pores near
its outer margin.
REMARKS. The presence of a faint central-area grill perfor-
ated by a ring of pores at its margin is similar to descriptions
of both R. locken Müller, 1970 and R. daviesii (Haq, 1968)
Haq, 1971; however both of these species appear to have a
significantly wider central-area and in this section R. cf. R.
filewiczii intergrades with R. filewiczii.
OCCURRENCE. Rare throughout these EOB sections.
Order COCCOLITHALES Schwarz, 1932 emend.
Edvardsen & Eikrem in Edvardsen et al., 2000
DIAGNOSIS. Includes the placolith-forming families Coc-
colithaceae, Calcidiscaceae and Pleurochrysidaceae and the
murolith-forming Hymenomonadaceae (Young et al. 2003;
Jordan et al. 2004). The placolith-forming families share two
common structural features; first, growth occurs downward
as well as upwards from the proto-coccolith ring, which be-
comes embedded within the rim (Fig. 3) and second, the distal
shield and most of the tube is formed of V-units, whereas the
proximal shield and sometimes part of the tube is formed
of R-units, making the distal shield dark and the proximal
shield bright in XPL (Young et al. 2004).
 374 TOM D U N K L E Y J O N E S ET AL.
Family COCCOLITHACEAE Poche, 1913 emend.
Young & Bown, 1997
DIAGNOSIS. Produce placoliths with V-unit forming both the
distal shield and the proximal layer of the inner tube, while
the R-unit forms a bicyclic proximal shield and the distal
layer of the inner tube (Fig. 2) (Young et al. 1992).
Genus COCCOLITHUS Schwarz, 1894
DIAGNOSIS. Placoliths with a Coccolithaceae-type structure
and an open or closed central-area, sometimes spanned by a
disjunt bar or diminutive cross.
REMARKS. Although Cruciplacolithus and Chiasmolithus
are the two accepted Coccolithaceae genera with axial/non-
axial central-area crosses, the genus Coccolithus does include
a number of Eocene species with reasonably well-developed
central-area crosses (C. mutatus, C. staurion, C. crucis; see
Bown 2005a).
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Coccolithus pelagicus (Wallich, 1877) Schiller, 1930
(Pl. 2, figs 1,13 & 19)
DESCRIPTION. Small and medium-sized elliptical placoliths
with Coccolithus-type structure and an apparently open
central-area in LM.
REMARKS. There are two accepted extant sub-species of C.
pelagicus; a subarctic form, C. pelagicus pelagicus, and a
temperate form, C. pelagicus braarudii, which have been
consistently distinguished on the basis of ecology, heterococ-
colith size, the production of distinct holococcolith morpho-
logies in the haploid phase and molecular genetics (Geisen et
al. 2002; Saez´ et al. 2003). The estimated divergence time for
these two subspecies is between 1.6 and 2.7 Ma (Saez´ et al.
2003). There is significant size variation within Palaeogene
C. pelagicus, which combined with our knowledge of the
genetic and morphological variation within the modern spe-
cies, suggests the presence of multiple sub- or pseudo-cryptic
species within the Palaeogene C. pelagicus species-concept.
SEM imaging of C. pelagicus in upper Paleocene/lower Eo-
cene sediments of Tanzania have noted the presence of a C.
pelagicus morphotype with a gracile central-area cross that
is not visible in LM (Bown et al. 2008). The considerable ge-
netic variability within the modern C. pelagicus, combined
with the observed persistence of a relatively conservative
morphotype from the earliest Palaeogene to the Recent, in-
dicates the action of a strong stabilising selection on this
coccolith morphology (Saez´ et al. 2003).
OCCURRENCE. Common throughout these EOB sections.
Ranges from NP9 to NP23 in the Tanzanian sections. Re-
ported literature range is from the early Paleocene to the
modern (Perch-Nielsen 1985).
Coccolithus formosus (Kamptner, 1963) Wise, 1973
(Pl. 2, fig. 2)
DESCRIPTION. Medium-sized circular to sub-circular
Coccolithus-type coccoliths with a circular and open
central-area.
OCCURRENCE. Rare to frequently throughout these EOB
sections. Ranges from NP9 to NP21 in the Tanzanian sec-
tions. Reported literature range is from the early Eocene to
the top of zone NP21 (Perch-Nielsen 1985).
Coccolithus cachaoi Bown 2005a (Pl. 2, fig. 3)
DESCRIPTION. Medium to large elliptical Coccolithus-type
coccoliths with a narrow central-area filled by a broad trans-
verse disjunct bar.
OCCURRENCE. Rare throughout these EOB sections. Ranges
from NP14b to NP23 in the Tanzanian sections.
Genus CRUCIPLACOLITHUS Hay & Mohler in Hay
etal., 1967
DIAGNOSIS. Elliptical Coccolithaceae-type placoliths with a
central-area spanned by an axial or near-axial cross.
Cruciplacolithus cruciformis (Hay & Towe, 1962)
Roth, 1970 (Pl. 2, fig. 4)
DESCRIPTION. Large elliptical placoliths with narrow
shields and a wide, open central-area spanned by an axial
to diagonal cross.
REMARKS. Shows considerable variation in the angle of the
cross bars, with early forms having a more axial cross, which
justifies its inclusion in Cruciplacolithus.
OCCURRENCE. Very rare throughout these EOB sections.
Ranges from NP14b to NP21 in the Tanzanian sections. First
described from the lower Eocene of France (Hay & Towe
1962).
Cruciplacolithus cf. G primus Perch-Nielsen, 1977
(Pl. 2, figs 5 & 14)
DESCRIPTION. Small to medium-sized placoliths with a nar-
row elliptical central-area that is almost filled by an axial
cross.
REMARKS. It has been suggested that C. primus is restric-
ted to Paleocene sediments (Perch-Nielsen 1985), however
this closely related form is consistently present through-
out the Eocene and into the lower Oligocene of Tanzania
(Bown 2005a). This long-ranging morphotype may be an-
cestral to the Neogene and extant C. neohelis, which, on
the basis of recent molecular genetic data, appears to have an
early Paleocene divergence from the genus Coccolithus (Saez
et al. 2004). The absence of the C. primus morphotype from
many deep-sea Palaeogene sequences may be due to either a
low preservation potential and/or a long history of restriction
to neritic/coastal environments, as observed for the extant C.
neohelis (Young et al. 2003).
OCCURRENCE. Rare throughout these EOB sections. Ranges
from NP9 to NP23 in the Tanzanian sections. Reported liter-
ature range for C. primus s.s is NP1 to NP8 (Perch-Nielsen
1985).
 UPPER EOCENE TO LOWER OLIGOCÈNE CALCAREOUS NANNOFOSSILS FROM TANZANIA 375
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Plate 2 LM images of Coccolithus, Cruciplacolithus, Bramletteius, Chiasmolithus, Clausicoccus and Tetralithoides placoliths (XPL unless
stated otherwise). Fig. 1 Coccolithuspelagicus TDP17/42-1, 72 cm. Fig. 2 Coccolithus formosus TDP11/23-1, 23 cm. Fig. 3 Coccolithus cachaoi
TDP12/17-2, 56 cm. Fig. 4 Cruciplacolithus cruciformis TDP12/26-1, 56 cm; 4a (XPL); 4b (PC). Fig. 5 Cruciplacolithus cf. Cr. primus TDP12/10-1,
55 cm; 5a (XPL); 5b (PC). Fig. 6 Chiasmolithus nitidus TDP12/40-3,97 cm. Fig. 7 Chiasmolithus titus TDP12/26-1, 56cm; 7a (XPL); 7b (PC). Figs 8,
9 Bramletteius serraculoides; 8, TDP11/15-1, 24cm; 9, TDP12/46-1,40cm. Fig. 10 Clausicoccus subdistichus TDP12/38-1,90cm. Fig. 11
Clausicoccus fenestratus TDP11/23-1, 23 cm. Fig. 12 Tetralithoides symeonidesii TDP12/26-2,62 cm. SEM images of Coccolithus,
Cruciplacolithus, Bramletteius, Reticulofenestra, Cyclicargolithus and Clausicoccus placoliths. Fig. 13 Coccolithus pelagicus, TDP12/47-2, 64 cm.
Fig. 14 Cruciplacolithus cf. Cr. primus TDP12/26-2, 62 cm. Figs 15-18 Bramletteius serraculoides; 15,17,18, TDP12/26-2,62 cm; 16, TDP12/47-2,
64 cm. Fig. 19 Coccolithus pelagicus (A), Reticulofenestra bisecta (B), Cyclicargolithus floridanus (C), Clausicoccus subdistichus (D) TDP12/26-2,
62 cm. All LMs are to the same scale; all SEMs are enlarged relative to the LMs (see scale bars), except for Fig. 19 (same scale as LMs).
 376 T O M D U N K L E Y J O N E S ET AL.

Genus CHIASMOLITHUS Hay et al., 1966
DIAGNOSIS. Elliptical Coccolithaceae-type placoliths with a
diagonal, usually offset, central-area cross.
REMARKS. The genus Chiasmolithus has previously been
subdivided into Chiasmolithus and Sullivania on the basis
of bar morphology and the presence/absence of a distinct
collar formed by the centro-distal cycle (Varol 1992). Here,
we retain the genus Chiasmolithus for all these forms as the
most practicable taxonomic grouping pending an improved
understanding of Chiasmolithus phylogeny.
Chiasmolithus nitidus Perch-Nielsen, 1971 (Pl. 2,
fig. 6)
DESCRIPTION. Medium to small Chiasmolithus species with
a simple oblique central-area cross.
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tions. Reported literature range is NP15 to NP22 (Perch-
Nielsen 1985).
Genus CLAUSICOCCUS Prins, 1979
DIAGNOSIS. Placoliths with typical Coccolithaceae-type rim
and a wide central-area spanned by a disjunct perforate plate.
Clausicoccus subdistichus (Roth & Hayin Hay et al.
1967) Prins, 1979 (Pl. 2, figs 10 & 19)
DESCRIPTION. Small elliptical Clausicoccus species with a
narrow central-area spanned by a disjunct plate with perfor-
ations that are only clearly visible in SEM (Pl. 2, fig. 19).
OCCURRENCE. Rare to frequent throughout these EOB sec-
tions. Ranges fromNP15b to NP21 in the Tanzanian sections.
Reported literature range from the late Eocene to early Oli-
gocene (Perch-Nielsen 1985).

OCCURRENCE. Rare in the upper Eocene becoming very rare

in the lower Oligocene of these EOB sections. Ranges from
Clausicoccus fenestratus (Deflandre & Fert, 1954)
NP9 to NP21 in the Tanzanian sections. Reported literature
Prins, 1979 (Pl. 2, fig. 11)
range is NP15 to NP16 (Perch-Nielsen 1985).

Chiasmolithus titus Gartner, 1970 (Pl. 2, fig. 7)
DESCRIPTION. Medium-sized Chiasmolithus species with
an oblique central-area cross with one axis composed of
simple arms and the other composed of kinked arms due to
additional distal segments angled obliquely to the axis of the
main cross, producing a swastika-like appearance.
OCCURRENCE. Rare in the upper Eocene becoming very rare
in the lower Oligocene of these EOB sections. Ranges from
NP15 to NP21 in the Tanzanian sections. Reported literature
range is NP15 to NP21 (Perch-Nielsen 1985).
Genus BRAMLETTEIUS Gartner, 1969
DIAGNOSIS. Cruciplacolithus-like base with a very large
monocrystalline 'paddle' or spine.
DESCRIPTION. Medium-sized elliptical Clausicoccus spe-
cies with a wide central-area spanned by a disjunct plate
with perforations visible in LM.
OCCURRENCE. Rare to frequent throughout these EOB sec-
tions. Ranges from NP14b to NP23 in the Tanzanian sec-
tions. Reported literature range from the middle Eocene to
early Oligocene (Perch-Nielsen 1985).
Genus CORONOCYCLUS Hay et al., 1966
DIAGNOSIS. Open ring-like coccolith without distinct
shields, elements of rim complexly intergrown, apparently
with outer V-unit and inner R-unit.
Coronocyclus nitescens (Kamptner, 1963) Bramlette
& Wilcoxon, 1967 (Pl. 3, figs 1 & 19)

Bramletteius serraculoides Gartner, 1969 (Pl. 2, figs DESCRIPTION. Medium to large ring-shaped coccoliths, bi-
8-9 & 15-18) cyclic in XPL. The birefringent cycle dominates the XPL
image and is crossed by extinction lines that are non-axial.
Outer edge appears serrated in LM due to a large number of
DESCRIPTION. Small placoliths with a Cruciplacolithus-like small radiating nodes developed in concentric circles around
rim and an open central-area spanned by an axial cross that the lower and outer surfaces of the ring (Pl. 3, fig. 19).
bears a monocrystalline paddle-like process (Pl. 2, figs 8,15
& 18) along its long-axis. There is a considerable variation OCCURRENCE. Rare throughout these EOB sections. Ranges
in the birefringence of the inner cycle. from NP15b to NP23 in the Tanzanian sections. Reported
literature range from the late Eocene (Perch-Nielsen 1985)
REMARKS. The c-axis of the monocrystalline paddle is to NN6 (Young 1998).
aligned at ~45° to the paddle long-axis and lies at a high
angle (Pl. 2, fig. 18). At present there is no evidence for di- Coronocyclus cf. C. nitescens (Kamptner, 1963)
morphism with paddle- and non-paddle-bearing coccoliths Bramlette & Wilcoxon, 1967 (Pl. 3, figs 20, 21)
occurring on the same coccosphere although there is con-
siderable variation in the length and width of the observed
paddles. The axial cross consists of multiple, lath-like ele- DESCRIPTIONS. TWO SEM images of what appear to be
ments (Pl. 2, fig. 16) and on the distal side is surrounded by Coronocyclus-type coccospheres (Pl. 2, figs 20, 21) show
a ring of elliptical elements (Pl. 2, fig. 17). placoliths with a proximal shield slightly wider than the
distal shield and an irregular grill-like structure spanning the
OCCURRENCE. Frequent to common throughout these EOB wide central area. One of these coccospheres is completely
sections. Ranges from NP15 to NP21 in the Tanzanian sec- contained within a larger Calcidiscus-type coccosphere
 UPPER EOCENE TO LOWER OLIGOCÈNE CALCAREOUS NANNOFOSSILS FROM TANZANIA 377
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Plate 3 LM images of Coronocyclus, Umbilicosphaera, Calcidiscus, Hayella and Pedinocyclus placoliths (XPL unless stated otherwise). Fig. 1
Coronocyclus nitescens TDP17/42-1, 72 cm. Fig. 2 Umbilicosphaera bramletteii TDP12/40-3,97 cm. Figs 3,4 Umbilicosphaera jordanii; 3,
TDP17/38-1,10cm; 4, TDP12/10-1, 55 cm. Fig. 5 Calcidiscusprotoannulus TDP17/42-1, 72 cm. Figs 6, 7 Calcidiscus?sp. 1; 6, TDP17/42-1, 72 cm;
7, TDP12/10-1, 55 cm; Fig. 7a (XPL); Fig. 7b (PC). Figs 8 , 9 Hayella situliformis TDP12/10-1, 55 cm. Fig. 10 Pedinocyclus larvalis, TDP12/10-1,
55 cm. SEM images of Calcidiscus, Umbilicosphaera, Coronocyclus and Pedinocyclus placoliths. Figs 11-16 Calcidiscus protoannulus; 11,12,15,
distal views TDP12/26-2,62 cm; 13, distal view TDP12/36-1,44 cm; 14, distal view TDP12/47-2,64 cm; 16, proximal view TDP12/30-1, 35 cm. Figs
17,18 Umbilicosphaera bramlettei; 17, distal view TDP12/47-2,64 cm; 18, proximal view TDP12/26-2, 62 cm. Fig. 19 Coronocyclus nitescens
TDP12/30-1, 35 cm. Figs 20,21 Coronocyclus cf. C. nitescens TDP12/47-2,64 cm; 20, coccosphere; 21, part-coccosphere within a
Calcidiscus-type coccosphere. Fig. 22 Calcidiscus? sp. 1TDP12/26-2, 62 cm. Fig. 23 Pedinocyclus larvalis TDP12/26-2,62 cm. All LMs are to the
same scale; all SEMs are enlarged relative to the LMs (see scale bars).
 378 TOM D U N K LE Y J O N E S ET AL.

(Pl. 2, fig. 21). This may be a chance occurrence, represent
a transitional life-cycle stage, or be a combination of endo-
and exothecal coccoliths on the same coccosphere; similar
complex associations are observed in the extant species Ce-
ratolithus cristatus where planoliths often occur inside coc-
cospheres of hoop coccoliths (Cros et al. 2000; Young et al.
2003).
OCCURRENCE. Rarely observed in SEM observations of
these EOB sections. Similar forms have been observed in
Tanzanian sediments of middle Eocene zone NP15.
Family CALCIDISCACEAE Young & Bown, 1997
DIAGNOSIS. Form elliptical, subcircular and circular placo-
liths where V-unit forms the (non-birefringent) distal shield
and tube, extending to the proximal surface, and R-unit forms
the proximal shield (Calcidiscus-typerim)(Fig. 3).
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OCCURRENCE. Frequent in the upper Eocene becoming
common in the lower Oligocene of these EOB sections.
Ranges from NP9 to NP23 in the Tanzanian sections.
Genus CALCIDISCUS Kamptner, 1950
DIAGNOSIS. Sub-circular to circular placoliths with typic-
ally monocyclic distal shields with straight sutures between
elements; proximal shield either mono- or bicyclic.
REMARKS. The placolith morphologies of the genera Cal-
cidiscus and Umbilicosphaera are similar, with Umbil-
icosphaera being distinguished by its kinked distal-shield
sutures and wide, open central-area. This generic distinction
is supported by molecular genetic analyses of extant taxa
(Saez´ et al. 2003).
Calcidiscus protoannulus (Gartner, 1971) Loeblich &
Tappan, 1978 (Pl. 3, figs 5 & 11-16)

Genus UMBILICOSPHAERA Lohmann, 1902

DIAGNOSIS. Circular or elliptical placoliths with wide, open DIAGNOSIS. Small to medium-sized (3.5-8.0 μm) ring-
central-area, birefringent tube-cycle and complex kinked su- shaped coccoliths that are distinctly bicyclic in XPL; the
tures between distal shield elements. Proximal shield mono- bright inner cycle is narrower than the outer cycle and crossed
or bicyclic. by extinction lines that are near axial.

REMARKS. SEM images show that distal shield sutures are

Umbilicosphaera bramlettei (Hay & Towe, 1962) straight and near radial (Pl. 3, figs 11-15). At the inner mar-
Bown et al., 2007 (Pl. 3, figs 2,17 & 18) gin of the distal shield there may be a raised tube-cycle with
strong clockwise imbrication (Pl. 3, figs 11 & 12); this is vari-
ably developed between specimens, with some possessing
DESCRIPTION. Small to medium-sized (3.5-7.5 μm), nar-
smooth inner margins to the distal shield (Pl. 3, figs 13 & 14).
row, ring-shaped coccoliths, distinctly bicyclic in XPL with a
This form is similar to the extant Umbilicosphaera anulus,
wide, open central-area. The two cycles are similar in width;
which also has a wide central-area (characteristic of Umbil-
the bright inner cycle is crossed by non-axial extinction lines.
icosphaera) and straight sutures between elements (charac-
REMARKS. SEM images of U. bramlettei clearly show teristic of Calcidiscus). Molecular genetic analysis supports
its morphological similarities to modern Umbilicosphaera, the division between Calcidiscus-type species with straight
namely the presence of kinked sutures with anticlockwise sutures and closed central-areas (C. leptoporus, C. quad-
obliquity on the distal shield (Pl. 3, fig. 17) and a proximal riperforatus, Oolithotus fragilis) and Umbilicosphaera spe-
shield with a strong clockwise obliquity and a narrow, slightly cies with open central-areas and kinked sutures (U sibogae,
raised collar at its inner edge (Pl. 3, fig. 18). This distal shield U.foliosa) (Saez´ et al. 2003). Data is not yet available for U.
element morphology is very similar to the extant U. sibogae anulus, so the generic placement of forms with open central-
(Bown et al., 2007), supporting its reassignment to the genus areas and straight sutures is still uncertain.
Umbilicosphaera.
OCCURRENCE. Common in the upper Eocene becoming rare
OCCURRENCE. Rare throughout these EOB sections. Ranges to very rare in the lower Oligocene of these EOB sections.
from NP9 to NP21 in the Tanzanian sections. First described Ranges from NP11 to NP23 in the Tanzanian sections. Re-
from the lower Eocene of France (Hay & Towe 1962). ported literature range from middle to late Eocene (Perch-
Nielsen 1985).

Umbilicosphaera jordanii Bown, 2005a (Pl. 3, figs 3

Calcidiscus parvus sp. nov. (Pl. 4, figs 1-3)
&4)

DIAGNOSIS. Medium-sized (3.5-6.0 μm) circular ring-
shaped placoliths with a non-birefringent distal shield, a
narrow bright tube-cycle crossed by non-axial extinction
lines, and vacant central-area that is slightly wider than the
shields.
REMARKS. AS with U. bramlettei, recent SEM studies of
Palaeogene U. jordanii from Tanzania have shown that this
species has a distal shield element morphology similar to
the extant U sibogae (Bown et al., 2007) confirming its
assignment to the genus Umbilicosphaera.
DERIVATION OF NAME. From parvus meaning small, refer-
ring to the size of this placolith.
DIAGNOSIS. Small (3-4 μm) broadly elliptical to circular
Calcidiscus-type placoliths with a closed central-area; prox-
imal shield smaller than the distal shield and closed by an
inner set of irregular elements (Pl. 4,fig.1).
DIMENSIONS. L 3.3 μm, W 3.1 μm.
TYPE MATERIAL. Holotype, Pl. 4, fig. 1 (arrowed). Paratype,
Pl. 4, fig. 2. Type locality, TDP Site 12, Pande, Tanzania.
 UPPER EOCENE TO LOWER OLIGOCÈNE CALCAREOUS NANNOFOSSILS FROM TANZANIA 379
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Plate 4 SEM images of Calcidiscus-type placoliths. Figs 1-3 Calcidiscus parvus sp. nov.; 1 (holotype) & 3, TDP12/26-2, 62 cm; 2, TDP12/30-1,
35 cm (paratype). Fig. 4 Calcidiscus sp. 2 TDP12/26-2,62 cm. Figs 5-9 Calcidiscus sp. 3; 5, 6 & 9, TDP12/26-2, 62 cm; 7 , 8 , distal views
TDP12/47-2,64cm. All SEMs are to the same scale.
 380 TOM DUNKLEY J O N E S ET AL.
Type level, upper Eocene, Sample TDP12/26-2,62cm (Zone
NP19/20).
REMARKS. The inclusion of broadly elliptical and circular
forms in one species is based on the association of the two
morphologies on collapsed coccospheres (see Pl. 4, fig. 1).
These small placoliths have not been reliably identified in
LM.
OCCURRENCE. Rarely observed in SEM studies of TDP Site
12 within nannofossil zone NP19/20 undifferentiated.
Calcidiscus? sp. 1 (Pl. 3, figs 6, 7 & 22)
DIAGNOSIS. Small to medium-sized elliptical placolith ob-
served in SEM with a Calcidiscus-type distal shield and a
moderately wide central-area completely closed by a plate
formed of multiple elements (Pl. 3, fig. 22). Similar sized el-
liptical placoliths with a moderately wide, dark outer cycle,
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very narrow bright inner-cycle and a wide, apparently vacant,
central-area are frequently observed in LM (Pl. 3, figs 6 &
7) and may correspond to the placoliths observed in SEM.
REMARKS. Specimens tentatively assigned to the genus Cal-
cidiscus and paired with placoliths observed in LM. Similar
forms have been observed in SEM studies of late Paleocene-
early Eocene sediments of Tanzania (pers. obs.) and further
investigation of these forms is being undertaken before they
are assigned to either an existing or new genus within the
Calcidiscaceae.
OCCURRENCE. Rare to frequent throughout these EOB sec-
tions. Similar forms are rarely observed in SEM studies of
Tanzanian sediments from the late Paleocene (NP9), early
and middle Eocene.
Calcidiscus? sp. 2 (Pl. 4, fig. 4)
DIAGNOSIS. Small (4-5 μm) circular placoliths with a
closed central-area. Proximal shield smaller than distal; distal
shield elements show slight clockwise obliquity, whilst prox-
imal shield elements show strong anticlockwise obliquity (Pl.
4, fig. 4).
REMARKS. This is a very unusual form, with the strong an-
ticlockwise obliquity of distal shield elements being atypical
for the Calcidiscaceae, making the generic assignment un-
certain. The general appearance of the distal (larger) shield is
similar to the extant Oolithotus fragilis, but the coccoliths of
this modern genus have a characteristic asymmetry (Young
et al. 2003). These small placoliths have not been reliably
identified in LM. As with Calcidiscus sp. 1, further investig-
ation of these forms is being undertaken before they are as-
signed to a new or existing genus within the Calcidiscaceae.
OCCURRENCE. Rarely observed in SEM studies of these
EOB sections.
Calcidiscus sp. 3 (Pl. 4, figs 5-9)
DIAGNOSIS. Small (3-5 μm) circular-polygonal
Calcidiscus-type placoliths usually with a narrow,
open central-area but occasionally this is closed by a set
of irregular petaloid elements (Pl. 4, figs 6-8); proximal
shield smaller than distal and with clockwise imbrication of
elements.
REMARKS. The polygonal outline and small number of ele-
ments in the distal shield are similar to those of the modern
Hay aster, but this form does not have the distinctive diminut-
ive proximal shield of Hayaster. These small placoliths have
not been reliably identified in LM and as with Calcidiscus
sp. 1 and sp. 2, further investigation of these forms is be-
ing undertaken before they are assigned to a new or existing
genus within the Calcidiscaceae.
OCCURRENCE. Rarely observed in SEM studies of these
EOB sections. Similar forms have been observed in Tan-
zanian sediments from middle Eocene zone NP15.
Placolith coccoliths incertae sedis
Genus TETRALITHOIDESTheodoridis, 1984
DIAGNOSIS. Elliptical narrow-rimmed placoliths with wide
central-areas filled by four plates.
Tetralithoides symeonidesiiTheodoridis, 1984 (Pl. 2,
fig. 12)
DIAGNOSIS. Small, dark, low-birefringence placoliths with
wide central-area spanned by four distinct plates.
OCCURRENCE. Very rare in LM studies of these EOB sec-
tions. Ranges from NP15 to NP21 in the Tanzanian sections.
This is a significant extension to the reported literature range
of early Miocene zone NN2 to late Miocene/early Pliocene
zone NN12 (Young 1998).
Genus HAYELLA Gartner, 1969
DIAGNOSIS. Modified placolith(?) coccoliths formed of a
single cycle of subvertical crystal units and a tube cycle with
two flanges (Young & Bown 1997).
Hayella situliformis Gartner, 1969 (Pl. 3, figs 8 & 9)
DESCRIPTION. Medium-sized circular placolith coccoliths
with a wide, open central-area and highly birefringent tube-
cycle. Shield elements are distinct and show obliquity.
OCCURRENCE. Rare throughout these EOB sections. Ranges
from NP17 to NP21 in the Tanzanian sections. Reported lit-
erature range from the late Eocene to early Oligocene (Perch-
Nielsen 1985).
Genus PEDINOCYCLUS Bukry & Bramlette, 1971
DIAGNOSIS. LOW birefringence circular coccoliths.
Pedinocyclus larval is Bukry & Bramlette, 1971 (Pl. 3,
figs 10 & 23)
DESCRIPTION. Medium to large circular coccoliths with a
large thin flat distal shield of distinct, slightly inclined ele-
ments and relatively small open central-area. These forms
may be placoliths but if so the proximal shield has not been
observed to date and may be greatly reduced.
 UPPER EOCENE TO LOWER OLIGOCÈNE CALCAREOUS NANNOFOSSILS FROM TANZANIA 381

OCCURRENCE. Rare throughout these EOB sections. Ranges
from NP11 to NP23 in the Tanzanian sections. Reported
literature range from late Eocene to early Oligocene (Perch-
Nielsen 1985).
M urolith coccoliths
Order ZYGODISCALES Young & Bown, 1997
DIAGNOSIS. Muroliths and modified descendants with an
outer rim-cycle of V-units showing anticlockwise imbrica-
tion and an inner rim-cycle of R-units showing clockwise
imbrication.
Family HELICOSPHAERACEAE Black, 1971
DIAGNOSIS. V-units of outer rim modified to form a helical
flange ending in a wing or spike. R-units form the baseplate
and extend to form a blanket of small elements.
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REMARKS. Similar to H. bramlettei but with a distinct spur.
OCCURRENCE. Rare throughout these EOB sections. Ranges
from NP16 to NP23 in the Tanzanian sections. Reported
literature range from NP18 to NP24 (Perch-Nielsen 1985).
Helicosphaera gartneriTheodoridis, 1984 (Pl. 5, figs
7-9)
DESCRIPTION. Medium-sized and elongated, symmetrically
elliptical helicosphaerids with an open central-area spanned
by a disjunct bar.
REMARKS. The original description ofH. gartneri noted that
a fine 'grid' often fills the central-area; in SEM studies of the
Tanzanian specimens a fine grill on the proximal side was
observed spanning the central-area (Pl. 5, fig. 9).
OCCURRENCE. Rare throughout these EOB sections

(NP19/20 to NP21). Not reliably observed in other Tanzanian

Genus HELICOSPHAERA Kamptner, 1954
sites. First described from the Oligocene of Egypt (Theodor-
DIAGNOSIS. See family diagnosis. idis 1984).
Helicosphaera compacta Bramlette & Wilcoxon, 1967
(Pl. 5, figs 1 & 2) Helicosphaera euphratis Haq, 1966 (Pl. 5, figs 10 &
11)
DESCRIPTION. Oviform coccolith with a well-defined bi-
refringent blanket in XPL and narrow central-area spanned
DESCRIPTION. Elliptical with a narrow central area almost
by a conjunct bar.
completely filled by a broad oblique disjunct bar.
OCCURRENCE. Frequent throughout these EOB sections.
Ranges from NP16 to NP23 in the Tanzanian sections. Re- OCCURRENCE. Rare throughout these EOB sections. Ranges
ported literature range from NP17 to NP24 (Perch-Nielsen from NP19/20 to NP21 in the Tanzanian sections. Reported
1985). literature range from NP18 to NN6 (Perch-Nielsen 1985).

Helicosphaera bramlettei (Muller, 1970) Jafar & Helicosphaera reticulata Bramlette & Wilcoxon, 1967
Martini, 1975 (Pl. 5, figs 3-5) (Pl. 5, figs 12 & 13)

DESCRIPTION. Asymmetrically elliptical with a distinctly DESCRIPTION. Broadly rhombohedral in outline with a mod-
pointed anterior wing and a relatively wide central-area erately well-developed spur in some specimens (Pl. 5, fig.
spanned by a broad, diabolo-shaped disjunct bar. 13), and a broad oblique disjunct bar. Entire distal surface
REMARKS. Here we follow Theodoridis (1984) in including appears to be covered by a blanket with multiple perforations.
in H. bramlettei forms with oblique to sub-horizontal bars. OCCURRENCE. Rare to frequent in the upper Eocene becom-
DIFFERENTIATION. Helicosphaera bramlettei is distin- ing frequent to common in the lower Oligocene of these EOB
guished from the middle Eocene H. seminulum by the lat- sections. Ranges from NP15b to NP21 in the Tanzanian sec-
ter's rounded outline and large rectangular central open- tions. Reported literature range from NP17 to NP22 (Perch-
ing spanned by a sub-horizontal bar. Following Theodoridis Nielsen 1985).
(1984) we include smaller, relatively elongated and symmet-
rically elliptical forms in H. gartneri. Helicosphaera wilcox- Family PONTOSPHAERACEAE Lemmermann, 1908
onii has a similar morphology to H. bramlettei but has an
DIAGNOSIS. Murolith coccoliths with V-units forming a nar-
anterior wing that ends in a distinct spur.
row outer rim-cycle with anticlockwise imbrication and R-
OCCURRENCE. Rare to frequent throughout these EOB sec- units forming the inner rim with clockwise imbrication, the
tions. Ranges from NP14b/15a to NP23 in the Tanzanian sec- base-plate and blanket.
tions. Reported literature range from NP15 to NP25 (Perch-
Nielsen 1985).
Genus PONTOSPHAERA Lohmann, 1902
DIAGNOSIS. Central-area solid or with a variable number of
Helicosphaera wilcoxonii (Gartner, 1971) Jafar &
Martini, 1975 (Pl. 5, fig. 6) pores.
REMARKS. Extant members of the genus are distinguished
DESCRIPTION. Elliptical with relatively wide central-area from Scyphosphaera by their monomorphic coccospheres -
spanned by a broad, transverse disjunct bar and a wing that Scyphosphaera coccospheres are dimorphic with elevated
ends in a distinct spur. equatorial coccoliths (Young et al. 2003).
 382 T O M D U N K L E Y J O N E S ET AL.
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Plate 5 LM (XPL) and SEM images of Helicosphaera, Scyphosphaera, Pontosphaera and Syracosphaera muroliths. Figs 1,2 Helicosphaera
compacta; 1, TDP12/10-1, 55 cm; 2, TDP12/36-2, 70cm. Figs 3-5 Helicosphaera bramlettei; 3, TDP12/40-3,97 cm; 4, TDP12/47-2,64cm (double
scale of LM images); 5, TDP12/38-1, 90 cm. Fig. 6 Helicosphaera wilcoxonii TDP12/10-1, 55 cm. Figs 7-9 Helicosphaera gartneri; 7, TDP12/15-1,
12 cm; 8, TDP12/40-3,97 cm; 9, TDP12/26-2,62 cm (double scale of LM images). Figs 10,11 Helicosphaera euphratis; 10, TDP12/46-1, 40 cm; 11,
TDP11/23-1, 23 cm. Figs 12,13 Helicosphaera reticulata TDP12/26-1, 56cm. Fig. 14 Scyphosphaera columella TDP12/26-1, 56cm. Fig. 15
Scyphosphaera apsteinii TDP12/26-1, 56 cm. Fig. 16 Pontosphaera plana TDP12/26-2, 62 cm. Figs 17,18 Pontosphaera versa; 17, TDP12/40-1,
56 cm; 18, TDP12/40-3,97 cm. Fig. 19 Pontosphaera formosa TDP17/15-1,10 cm. Fig. 20 Pontosphaera cf. P. formosa TDP12/10-1, 55 cm. Figs 21,
22 Pontosphaera pectinata; 21, TDP11/10-1, 29 cm; 22, TDP11/15-1, 24cm. Figs 23,24 Pontosphaera multipora; 23, TDP12/10-1, 55 cm; 24,
TDP12/38-1, 90 cm. Figs 25,26 Pontosphaera alta TDP12/10-1, 55 cm. Figs 27,28 Syracosphaera tanzanensis; 27, TDP12/10-1, 55 cm; 28,
TDP12/47-4, 35 cm. All LMs are to the same scale; SEMs (Figs 4 & 9) are double this magnification.
 UPPER EOCENE TO LOWER OLIGOCÈNE CALCAREOUS NANNOFOSSILS FROM TANZANIA
Pontosphaera plana (Bramlette & Sullivan, 1961)
Haq, 1971 (Pl. 5, fig. 16)
DESCRIPTION. Simple unadorned plate, typically with two
very narrow longitudinal slits running parallel to the long-
axis.
OCCURRENCE. Rare to frequent throughout these EOB sec-
tions. Ranges from NP9 to NP23 in the Tanzanian sec-
tions. First described from the lower Eocene of California
(Bramlette & Sullivan 1961).
Pontosphaera versa (Bramlette & Sullivan, 1961)
Sherwood, 1974 (Pl. 5, figs 17 & 18; Pl. 6, figs 4 & 5)
DESCRIPTION. Simple unadorned plate with a broad raised
margin/rim. Many of the larger late Eocene Tanzanian spe-
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cimens have a single ring of perforations around the margin
of the plate (Pl. 5, fig. 18; Pl. 6, figs 4 & 5).
OCCURRENCE. Rare throughout these EOB sections. Ranges
from NP10 to NP23 in the Tanzanian sections. First described
from the lower Eocene of California (Bramlette & Sullivan
1961).
Pontosphaera formosa (Bukry & Bramlette, 1968)
Romein, 1979 (Pl. 5, figs 19 & 20)
DESCRIPTION. Large (9-17 μm) with a high flaring rim that
displays discernible elements in LM. The plate is thin and
indistinct in XPL although in some specimens multiple per-
forations are clearly visible (Pl. 5, fig 20).
OCCURRENCE. Rare throughout these EOB sections. Ranges
from NP14b/15a to NP21 in the Tanzanian sections. First
described from the middle Eocene of Mexico (Bukry &
Bramlette 1968).
Pontosphaera pectinata (Bramlette & Sullivan, 1961)
Sherwood, 1974 (Pl. 5, figs 21 & 22)
DESCRIPTION. Pontosphaera with a plate that is scalloped
towards its outer edge as defined by narrow radial ridges and
furrows that run inwards towards the centre where the plate
is perforated by multiple small holes.
REMARKS. Shows significant variability in the size and de-
velopment of radial ridges and the number of central perfor-
ations. May intergrade with forms of P. multipora.
OCCURRENCE. Frequent throughout these EOB sections.
Ranges from NP10 to NP21 in the Tanzanian sections. First
described from the lower Eocene of California (Bramlette &
Sullivan 1961).
Pontosphaera multipora (Kamptner, 1948) Roth,
1970 (Pl. 5, figs 23 & 24; Pl. 6, figs 1 & 3)
DESCRIPTION. Simple unadorned plate with a narrow indis-
tinct rim and up to three concentric cycles of small perfora-
tions.
383
REMARKS. AS with modern forms (Young et al. 2003), the
width of the outer rim is highly variable, as is the number
and size of perforations (Pl. 6, fig. 3).
OCCURRENCE. Frequent throughout these EOB sections.
Ranges from NP14b/15a to NP23 in the Tanzanian sections.
Reported literature range from the Palaeogene to modern
(Young 1998); first described from the Tortonian (Kamptner
1948).
Pontosphaera alta Roth, 1970 (Pl. 5, figs 25 & 26;
Pl. 6, fig. 2)
DESCRIPTION. Pontosphaera with an elevated, birefringent
rim and a thin, low birefringence plate with a large number
of small perforations.
REMARKS. Similar to P. versa but with a thinner and/or lower
rim and less birefringent central plate. This species was first
described by Roth (1970) from the lower Oligocene Red
Bluff Formation of Alabama, USA. The SEM images of the
original description are near identical to Pl. 6, fig. 2 and LM
examination of specimens from the Red Bluff Formation
confirmed the LM identification of P. alta presented here
(pers. obs.).
OCCURRENCE. Rare throughout these EOB sections but
more common than P. versa. Ranges from NP19/20 to NP21
in the Tanzanian sections. First described from the lower
Oligocene of the US Gulf Coast (Roth 1970).
Genus SCYPHOSPHAERA Lohmann, 1902
DIAGNOSIS. Extant forms are similar to Pontosphaera but
with dimorphic coccospheres consisting of elevated equat-
orial coccoliths, known as lopadoliths, and Pontosphaera-
type muroliths.
REMARKS. Fossil coccoliths with a lopadolith morpho-
logy are included within this genus. The (presumably)
Pontosphaera-like coccoliths that would be expected to oc-
cur on 'fossil' Scyphosphaera coccospheres would not be
distinguishable from true Pontosphaera coccoliths (unless
preserved on a fossil coccosphere), and so are almost cer-
tainly included within the genus Pontosphaera.
Scyphosphaera columella Stradner, 1969 (Pl. 5, fig.
14)
DESCRIPTION. Tall narrow murolith with near-straight walls
that run parallel or flare slightly to a maximum width at the
distal end.
OCCURRENCE. Very rare throughout these EOB sections.
Ranges from NP14a/15b to NP21 in the Tanzanian sections.
First described from the lower Eocene of Austria (Stradner
1969).
Scyphosphaera apsteinii Lohmann, 1902 (Pl. 5, fig.
15)
DESCRIPTION. Large elevated murolith with variable mor-
phology but typically broad with curved, gently flaring walls
that may narrow distally.
 384 T O M D U N K L E Y J O N E S ET AL.
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Plate 6 SEM images of Pontosphaera, Calciosolenia and Syracosphaera muroliths. Figs 1,3 Pontosphaera multipora TDP12/26-2,62 cm.
Fig. 2 Pontosphaera alta TDP12/26-2, 62 cm. Figs 4,5 Pontosphaera versa; 4, distal view TDP12/23-2, 79 cm; 5, proximal view TDP12/26-2,
62 cm. Figs 6, 7 Calciosolenia sp.; 6, TDP12/26-2,62 cm; 7, TDP12/10-1, 55 cm. Fig. 8 Coronosphaera?sp. TDP12/47-2,64cm. Figs 9-11
Syracosphaera monechiae; 9, TDP12/47-2,64cm; 10 (paratype) & 11 (holotype), TDP12/26-2,62 cm. Figs 12,13 Syracosphaera tanzanensis; 12,
TDP12/47-2,64 cm; 13, TDP12/26-2,62 cm. Fig. 14 Syracosphaera cf. S. tanzanensis TDP12/23-2, 79 cm. Figs 15,16 Syracosphaera raffiae; 15
(holotype) & 16 (paratype), TDP12/26-2,62 cm. All SEMs above the white dividing line are to the same scale; all SEMs below the dividing line are
enlarged relative to those above (see scale bars).
 UPPER EOCENE TO LOWER OLIGOCÈNE CALCAREOUS NANNOFOSSILS FROM TANZANIA
OCCURRENCE. Very rare throughout these EOB sections.
Ranges from NP14b/15a to NP21 in the Tanzanian sections.
Reported literature range from the Palaeogene to the modern
(Young 1998).
Order SYRACOSPHAERALES Hay, 1977 emend.
Young et al., 2003
DIAGNOSIS. Includes coccolithophores that produce com-
plex coccoliths typically consisting of three components: (1)
a rim with normal V/R structure but with a proto-coccolith
ring embedded within it; (2) a radial lath cycle, formed of
crystallites with sub-tangential c-axis orientation (‘T-units')
that interdigitate with rim elements; (3) an axial structure
in the centre of the coccolith that may be formed from ra-
dial lath elements and/or from additional, disjunct elements
(Fig. 3) (Young et al. 2003).
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Family SYRACOSPHAERACEAE Lemmermann, 1908
DIAGNOSIS. Extant forms often develop distinct inner and
outer layers of coccoliths around the cell (dithecatism). En-
dothecal (inner layer) coccoliths are relatively conservative
in form, normally muroliths with a well-developed central-
area lath-cycle and variable inner central-area (Fig. 3), whilst
exothecal (outer layer) coccoliths are more variable in form
including planolith, murolith and dome-shaped forms. In-
dividual coccospheres can show a significant variation in
coccolith size and shape dependent on their position on
the coccosphere (varimorphism) and may include a number
of distinct coccolith morphologies (polymorphism) includ-
ing spine-bearing circum-flagellar coccoliths. These features
make the taxonomy of fossil species problematic as a num-
ber of distinct coccolith morphologies, and hence fossil taxa,
may be produced by the same biological species.
Genus SYRACOSPHAERA Lohmann, 1902
DIAGNOSIS. Coccospheres usually dithecate with highly
variable exothecal coccoliths and murolith endothecal coc-
coliths with one, two or three flanges.
Syracosphaera tanzanensis Bown, 2005a (Pl. 5, figs
27 & 28; Pl. 6, figs 12 & 13)
DESCRIPTION. Medium-sized muroliths with narrow bicyc-
lic rim (bright inner cycle, dark outer cycle) and wide central-
area.
REMARKS. This species was tentatively assigned to the
genus Syracosphaera on the basis of its appearance in LM
(Bown 2005a). This is confirmed by SEM images presented
here (Pl. 6, figs 12 & 13), which show muroliths with a typ-
ical Syracosphaera structure, with a high rim, narrow radial
lath cycle, broad central plate formed of fused elements and
a central spine. The relatively thick and high rim produces
the distinct bright cycle in XPL.
OCCURRENCE. Frequent to common throughout these EOB
sections. Ranges from NP16 to NP23 in the Tanzanian sec-
tions.
385
Syracosphaera monechiae sp. nov. (Pl. 6, figs 9-11)
DERIVATION OF NAME. Named after Dr Simonetta Monechi
(Universita di Firenze, Italy), nannopalaeontologist.
DIAGNOSIS. Small (3-4 μm) simple elliptical murolith coc-
colith with a narrow rim formed of sub-vertical elements, a
distinct proximal flange, a wide central-area spanned by a
well-developed fragile lath cycle and a low central spine (Pl.
6,fig.11).
DIFFERENTIATION. The larger and more robust S. tanzanen-
sis has a basal plate formed of fused lath elements whereas in
S. monechiae the radial laths remain separated up to the base
of the low central spine. Differentiated from the smaller S.
raffiae, which has a lower and thickerrimand an elongated
central boss rather than the low spine of S. monechiae. Coro-
nosphaera? sp. differs from S. monechiae in having multiple
radial lath elements to each rim element and in having im-
bricatedrimelements.
DIMENSIONS. L 2.6 μm, W 1.7 μm, H 0.6 μm.
TYPE MATERIAL. Holotype: Pl. 6, fig. 11. Paratype: Pl. 6,
fig. 10. Type locality, TDP Site 12, Pande, Tanzania. Type
level, upper Eocene, Sample TDP12/26-2, 62 cm (Subzone
NP19/20).
OCCURRENCE. Rare throughout these EOB sections;
NP19/20-21; TDP Sites 11, 12, 17.
Syracosphaera raffiae sp. nov. (Pl. 6, figs 15 & 16)
DERIVATION OF NAME. Named after Dr Isabella Raffi (Uni-
versita' "G. d'Annunzio", Italy), nannopalaeontologist.
DIAGNOSIS. Very small (~2 μm) elliptical murolith cocco-
lith with a thick rim formed of elements showing a dextral
obliquity, a distinct radial lath cycle and central boss.
DIFFERENTIATION. Differentiated from S. monechiae by its
smaller size, lower and thicker rim and elongated central
boss.
DIMENSIONS. L 2.1 μm, W 1.6 μm.
TYPE MATERIAL. Holotype: Pl. 6, fig. 15. Paratype: Pl. 6, fig.
16. Type locality, TDP Site 12, Pande, Tanzania. Type level,
upper Eocene, Sample TDP12/26-2,62 cm (Zone NP19/20).
OCCURRENCE. Very rare in SEM observations of TDP 12
sediments from zone NP19/20.
Genus incertae sedis CORONOSPHAERA Gaarder
in Gaarder & Heimdal, 1977
DIAGNOSIS. Murolith coccoliths with an outer cycle show-
ing strong anticlockwise imbrication, a vertical inner cycle
and two radial laths to eachrimelement.
REMARKS. Coronosphaera is often placed within the Syra-
cosphaeraceae; here we follow Young et al. (2003) who
place the genera as an incertae sedis tax on within the Syra-
cosphaerales on the basis of its anomalous imbricate outer
rim cycle and higher number of radial laths per rim cycle
element.
 386 T O M D U N K L E Y J O N E S ET AL.
Table 2 Morphological terms applied to rhabdolith basal coccoliths
Kleijne (1992) Aubry (1999)
Outer rim cycle ri
Inner rim cycle M2
Radial cycle CARC ( c e n t r a l - a r e a r a d i a l cycle)Radial(lath)cycleRadial(lath)
Intermediate central-area cycle CAIC (intermediate central-area cycle)
(only Discosphaera)
Lamellar cycle CA4 (central-area cycle 4)
Cuneate cycle CACC (central-area cuneate cycle)
For Palaeogene rhabdoliths (principally Blackites) Aubry (1999) identifies only one rim cycle (M1), called the 'radial cycle' M2, and recognises four central-area cycles
(CA1-4), with CA4 forming the 'stem or cupola' and cycles CA1-3 variably developed in different 'groups'. For Blackites, we recognise the same rim and radial cycles
as for the extant genera, along with up to four additional, discrete central-area/lamellar cycles.
Coronosphaera? sp. (Pl. 6, fig. 8)
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DIAGNOSIS. Small (~3 μm) elliptical murolith coccolith
with an outer rim cycle that appears to show anticlockwise
imbrication and a high number of radial lath cycles.
OCCURRENCE. Very rare in SEM studies of these EOB sec-
tions. Not yet observed from other Tanzanian sites.
Family CALCIOSOLENIACEAE Kamptner, 1927
DIAGNOSIS. Includes coccolithophores that produce muro-
lith coccoliths without flanges, usually termed scapholiths.
Rim predominantly formed of V-units with small R-units
at the base/inner margin (Young et al. 2003). Central-area
structure formed of a single lath-cycle, with pairs of laths
from opposite sides of the coccolith rim forming a series of
transverse bars.
Genus CALCIOSOLENIA (Gran, 1912)
Young et al., 2003
DIAGNOSIS. Includes coccolithophores that form rhombic
muroliths (scapholiths).
Calciosolenia sp. (Pl. 6, figs 6 & 7)
DESCRIPTION. Isolated rhombic muroliths (scapholiths)
were frequently observed during SEM investigations of rock
chip surfaces and rarely observed in LM. These have not
been given a species assignment but in SEM are remarkably
similar in morphology to the living C. brasiliensis (Young
et al. 2003).
OCCURRENCE. Rare throughout these EOB sections. Similar
small rhombic scapholiths range from the Lower Cretaceous
to the Recent (Bown et al. 1998).
Family RHABDOSPHAERACEAE Haeckel, 1894
DIAGNOSIS. Includes coccolithophores that produce typic-
ally disc-shaped coccoliths (planoliths) formed of three com-
ponents: (1) a narrow, slightly elevated rim formed of two
cycles of elements - an upper/outer rim cycle of simple non-
imbricate V-unit elements and a lower/inner rim cycle with
strong obliquity and uncertain crystallography; (2) a cycle
of radial laths joining rim to lamellar cycle (although absent
Young et al. (2003) This study
Upper/outer rim cycle (V unit) Upper/outer rim cycle and
lower/inner rim cycle -
together labelled 'Rm' (rim)
Lower/inner rim cycle (R? unit)
Radial (lath) cycle Radial (lath) cycle 'RC
l-area cycle Lamellar cycle(s) 'LC
'Cuneate cycle' Cuneate cycle 'C
in Rhabdosphaera); (3) lamellar cycle(s) formed of lamel-
lar elements showing clockwise imbrication, often multiple
cycles with inner cycles more elongate, inclined and in helical
arrangement forming a spine or protrusion. Coccoliths of the
Rhabdosphaeraceae are often termed 'rhabdoliths' (Kleijne
1992).
REMARKS. The morphological terminology that has been
applied to Palaeogene members of the Rhabdosphaeraceae
can be confusing. Table 2 summarises previous classification
schemes and indicates a potential synthesis of terminology
that is coherent with the established V/R model of coccolith
morphogenesis and the structural homologies that define the
family Rhabdosphaeraceae.
Genus BLACKITES Hay & Towe, 1962
DIAGNOSIS. Spine- or process-bearing coccoliths with a
base formed of three components: (1) a bicyclic outerrim; (2)
a radial lath cycle; (3) multiple lamellar cycles with clock-
wise imbrication. Simple to highly complex spines or pro-
cesses can be formed by the multiple tiered lamellar cycles
extending up from the coccolith base.
REMARKS . Typically more than one radial lath cycle element
to each rim element. Proximal views often show the bicyclic
rim characteristic of the Rhabdosphaeraceae but this can be
obscured by overgrowth or the growth of the inner lamellar
cycles across the proximal side of the basal coccolith.
The differentiation of the extant genus Rhabdosphaera
and extinct Palaeogene genus Blackites has been problem-
atic. The genus Rhabdosphaera has been used by some au-
thors to include Palaeogene rhabdoliths with unknown basal
structure (Perch-Nielsen 1985) and its use for Palaeogene
forms persists in the literature (e.g. Varol 1998). The type
species of the two genera are the extant Rhabdosphaera clav-
igera (Haeckel, 1894) and the Eocene/Oligocene Blackites
spinosus (Hay & Towe, 1962). The original description of
the genus Blackites noted the characteristic radial cycle in
these coccoliths, which are 'characterized by a single circular
shield with a perforate ring spanned by interlocking ribs near
the periphery' (Hay & Towe 1962: 505). This characteristic
was later used as the distinguishing feature between Blackites
and Rhabdosphaera, with the latter 'compressed elliptically
and is lacking the cycle of interlocking ribs characteristic of
Blackites' (Stradner in Stradner & Edwards 1968: 29). As
 UPPER EOCENE TO LOWER OLIGOCÈNE CALCAREOUS NANNOFOSSILS FROM TANZANIA 387

with other coccolith groups (Young et al. 1992), rim struc-
ture is the most robust basis for generic taxonomy in the
Rhabdosphaeraceae and can be used to distinguish between
Blackites, which has a radial lath cycle, and Rhabdosphaera,
which lacks this radial cycle. The radial cycle in some Black-
ites species may be greatly reduced but is still present as a
series of residual radial elements between the outer cycle and
the lamellar cycle.
Morphological analyses of spines and processes have
been used to define a number of additional Palaeogene rhab-
dolith genera (Aubry 1999; Shafik 1989). At present we have
not attempted to subdivide the genus Blackites, which, based
on the definition above and the available SEM images ap-
pears to include the majority of Palaeogene rhabdoliths.
Blackites? furvus Bown & Dunkley Jones, 2006 (Pl. 7,
fig. 1)
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Blackites cf. Cruxia mericii Varol, 1989 (Pl. 7, fig. 5)
DESCRIPTION. Small rhabdoliths with relatively low, hollow
sacculiform spines. The spine broadens from the base and is
capped by a curving upper surface formed of one element,
which differs from Cr. mericii, which has a two-part upper
surface. The spine is slightly wider than the coccolith. The
coccolith rim appears to be narrow and is formed of two bright
cycles, whereas only one cycle is observed in Cr. mericii.
OCCURRENCE. Occurs very rarely throughout these EOB
sections. Also found in the middle Eocene sub-zone NP15b
of Tanzania.
Blackites fustis Bown, 2005a (Pl. 7, fig. 6)
DESCRIPTION. Rhabdolith with a relatively wide, hollow
ampulliform spine that is initially near parallel-sided, ex-

DESCRIPTION. Medium-sized, subcircular-circular cocco- pands distally, reaching a width similar to that of the cocco-
liths with broad, low-birefringence shields crossed by lith, and then terminates sharply in a domed top. The rim is
strongly curving extinction lines. The central-area is narrow relatively broad and the outer cycle does not lie distally over
or closed. the inner cycle.
REMARKS. Subcircular, spineless Rhabdosphaera-type pla- OCCURRENCE. Occurs very rarely throughout these EOB
noliths have been observed in SEM (see Pl. 8,fig.8), which sections. Also found in the middle Eocene sub-zones NP15b-
may correspond to B. furvus identified in LM. At present 15c of Tanzania.
there is not sufficient data to confidently correlate LM and
SEM images, hence the retention of the original tentative Blackites culter sp. nov. (Pl. 7, figs 7, 8 & 18-20)
generic assignment.

OCCURRENCE. Frequent throughout these EOB sections. DERIVATION OF NAME. From culter meaning knife or spear-
Ranges from NP19/20 to NP21 in the Tanzanian sections. point, referring to the appearance of this rhabdolith in LM.
DIAGNOSIS.Rhabdolith with a spine that has concave de-
Blackites deflandrei (Perch-Nielsen, 1968) Bown, pressions running along its length, producing either a three
2005a (Pl. 7, fig. 3) (Y-shaped), or possibly four-fold (X-shaped), symmetry
across the axis of the spine (Pl. 7, figs 18-20). In LM the
spine is moderately dark in XPL and has a flat blade-like
DESCRIPTION. Rhabdolith with a relatively low, wide, hol-
appearance. The spine broadens away from the coccolith to
low sacculiform spine constructed from at least two distinct
a maximum width approximately two-thirds of the way up
structural units when viewed in XPL; there may be a small,
the spine and then tapers to a point with an overall height ap-
terminal papilla. The spine broadens slightly to a maximum
proximately four times the width of the coccolith. Maximum
in the lower cycle, before tapering to a point; the spine is
spine-width is slightly less than the width of the basal coc-
always slightly narrower than the coccolith with a height
colith. The rim is made up of two cycles lying side-by-side
similar to or slightly greater than the coccolith width.
typical of the B. morionum group.
OCCURRENCE. Occurs very rarely throughout these EOB REMARKS. Relatively large Blackites bases observed in
sections. Ranges from NP14a/15b to NP21 in the Tanzanian SEM (see Pl. 8, figs 3 & 9) are most likely the bases of
sections. First described from the upper Eocene of Denmark Blackites culter (compare with the base of the specimen in
(Perch-Nielsen 1968). Pl. 7,fig.20).

Blackites clavus Bown, 2005a (Pl. 7, fig. 4) DIFFERENTIATION. Gross morphology similar to B. flam-
meus Bown & Dunkley Jones, 2006 in LM but B. flammeus
possesses a hollow, slightly asymmetric spine whereas B.
DESCRIPTION. Rhabdolith coccoliths with a short spine that culter has a flat, blade-like spine.
tapers rapidly to a point, giving the overall appearance of
DIMENSIONS. H 6.7 μm, CW 3.3 μm, SW 1.7 μm.
a nail or pin. Typically the height of the spine is similar to
the width of the coccolith. The rim is relatively broad with TYPE MATERIAL. Holotype: Pl. 7, fig. 7. Paratype: Pl. 7,
a small outer cycle that does not lie distally over the inner fig. 8. Type locality, TDP Site 12, Pande, Tanzania. Type
cycle. level, upper Eocene, Sample TDP12/36-2, 70 cm (Subzone
NP19/20).
OCCURRENCE. Occurs very rarely throughout these EOB
sections. Ranges from NP9 to NP21 in the Tanzanian sec- OCCURRENCE. Rare throughout these EOB sections;
tions. NP19/20-21; TDP Sites 11, 12, 17.
 388 T O M D U N K L E Y J O N E S ET AL.
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Plate 7 LM (XPL unless stated otherwise) and SEM images of Blackites and Rhabdosphaera coccoliths. Fig. 1 Blackites? furvus TDP11/23-1,
23 cm; 1a (XPL); 1b (PC). Fig. 2 Blackites base TDP17/15-1,10 cm. Fig. 3 Blackites deflandrei TDP12/38-1,90cm. Fig. 4 Blackites clavus
TDP11/10-1, 29 cm. Fig. 5 Blackites cf. Cruxia mericii TDP12/38-1,90 cm. Fig. 6 Blackites fustis TDP12/10-1, 55 cm. Figs 7 , 8 Blackites culter sp.
nov.; 7, TDP12/36-2, 70 cm (holotype); 8, TDP12/26-1, 56cm (paratype). Fig. 9 Rhabdosphaera vitrea TDP12/10-1, 55 cm. Fig. 10 Blackites tenuis
TDP12/46-1, 40 cm. Figs 11,12 Blackites spinosus; 11, TDP17/15-1,10 cm; 12, TDP12/46-1,40 cm. Figs 13,14 Rhabdospharea gracilentus; 13,
TDP12/46-1, 40 cm; 14, TDP12/40-3,97 cm. Fig. 15 Blackites singulus TDP12/47-4, 35 cm. Fig. 16 RhabdosphaeraxiphosTDP12/10-1, 55 cm; 16a
(XPL); 16b (PC). Fig. 17 Blackites tenuis TDP12/36-1, 44 cm. Figs 18-20 Blackites culterTDP12/26-2,62 cm. Fig. 21 Blackites spinosus
TDP12/26-2,62 cm. Figs 22-24 Rhabdosphaera vitrea; 22, TDP12/26-2,62 cm; 23, 24, TDP12/47-2,64 cm. Fig. 25 Rhabdosphaera gracilentus
TDP12/26-2,62 cm. Fig. 26 Rhabdosphaera spine TDP12/26-2, 62 cm. All LMs are to the same scale; all SEMs are enlarged relative to the LMs
(see scale bars).
 UPPER EOCENE TO LOWER OLIGOCÈNE CALCAREOUS NANNOFOSSILS FROM TANZANIA 389

Blackites tenuis (Bramlette & Sullivan, 1961) DIAGNOSIS. Small, sub-circular to elliptical rhabdoliths with
Sherwood, 1974 (Pl. 7, figs 10,17) a low domed process, formed of a continuous whorl of one
lamellar cycle, and a distinct collar formed from an outwardly
directed lamellar cycle.
DESCRIPTION. Rhabdolith with a tall, narrow styliform spine
that is narrow at its base, broadens slightly and then tapers to REMARKS. Clusters of associated rhabdoliths, presumably
a point but is never as broad as the coccolith rim. The rim is from the same coccosphere, show distinct varimorphism in
relatively narrow with a bright inner cycle. the degree of process development from domal to distinctly
conical (Pl. 8,fig.2). This is similar to many extant members
REMARKS. Isolated Blackites bases, with an outer dark cycle
of the Rhabdosphaeraceae, which have more pronounced
and inner bright central cycle in XPL, crossed by curving processes in apical or antapical regions (cf. Acanthoica, Cyr-
but roughly north-south trending extinction lines (Pl. 7, tosphaera, Algirosphaera, see Young et al. 2003). Blackites
fig. 2), are relatively common throughout these EOB sec- shafikii rhabdoliths are similar to those described within the
tions. Similar forms have been described as Blackites amplus novel genera Amitha and Notiocyrtolithus from the upper
Roth & Hay in Hay et al., (1967) but, based on their size and Eocene-lower Oligocene of the Otway Basin, SE Australia
structure, are very likely to be the isolated bases of Blackites (Shafik 1989), which all possess a distinct collar and low
tenuis (Aubry, 1999). domed process. The principal difference observed in the SE
DIFFERENTIATION. Differs from B. spinosus in having a nar-
Australian forms is the presence of a broadly spaced more
radially orientated lamellar cycle, with clear slits between
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row base to the spine whereas B. spinosus has a wide base

adjacent elements, located between the collar and the lamel-
to the spine, from which the spine tapers continuously to a
lar cycle that forms the process. A similar, but less well-
point.
developed, more radially orientated lamellar cycle is visible
OCCURRENCE. Rare throughout these EOB sections. Ranges in some of the Tanzanian specimens (such as lamellar cycle
from NP15 to NP23 in the Tanzanian sections. First de- LC3 in Pl. 8, fig. 12). We prefer to retain these forms within
scribed from the middle and upper Eocene of California, the genus Blackites on the basis of their structural similarities
USA (Bramlette & Sullivan 1961). with other members of the genus (Pl. 8, figs 9-12). All Black-
ites rhabdoliths have the characteristic (bicyclic) rim and ra-
dial lath cycle and then a variable number of lamellar cycles
Blackites spinosus (Deflandre & Fert, 1954) Hay & forming a spine or process. In Plate 8, figs 9-12 the lamellar
Towe, 1962 (Pl. 7, figs 11,12 & 21) cycles at the base of the process are labelled as LC1, LC2,
etc., the last of which forms the bulk of the process or spine.
DESCRIPTION. Rhabdolith with a tall, narrow styliform spine Using this scheme it becomes clear that the distinct collar
that tapers gradually to a point. The width of the spine is observed in Blackites shafikii - formed of an outward direc-
2
around half that of the coccolith base but considerably taller ted, radially orientated lamellar cycle (LC ) - is analogous to
(~6x width of the base). The rim is relatively broad and the vertically directed 3
lamellar cycle that forms the collar in
distinctly convex. Blackites culter (LC ) and the horizontal, radially orientated
lamellar cycle in Blackites amplus (LC2). This demonstrates
OCCURRENCE. Rare throughout these EOB sections. Ranges both the plasticity of the lamellar cycle orientation in Black-
from NP14b/15a to NP21 in the Tanzanian sections. First ites but also the common structural elements between these
described from the Lutetian of France and Oligocene of New species. Retaining all of these, and related species, within
Zealand (Deflandre & Fert 1954). the genus Blackites prioritises the similarities of rim and ra-
dial lath cycle construction over more subtle distinctions of
lamellar cycle orientation and process morphology.
Blackites singulus Bown & Dunkley Jones, 2006 (Pl.
7, fig.15)
DIMENSIONS. L 3.6 μm, W 2.9 μm, H ~2-3 μm.

DESCRIPTION. Long, slender spine that tapers gradually TYPE MATERIAL. Holotype: Pl. 8, fig. 1 (arrowed), Paratype:
from a slightly thickened broader end. The spine is birefrin- Pl. 8,fig.2 (arrowed). TDP Site 12, Pande, Tanzania. Upper
gent when parallel with the polarizing directions and has a Eocene, Sample TDP12/26-2, 62 cm (Subzone NP19/20).
narrow, axial canal.
OCCURRENCE. Rare in SEM studies of TDP Site 12 sedi-
REMARKS. The spine has never been observed associated ments from zone NP19/20.
with a basal coccolith.
OCCURRENCE. Frequent throughout these EOB sections Blackites sp. 1 (Pl. 8, figs 4 & 5)
becoming common in some intervals of the upper Eo-
cene (NP19/20-21). Ranges from NP19/20 to NP21 in the
Tanzanian sections. DESCRIPTION. Small dome-shaped rhabdolith, similar to
Blackites shafikii but with the first lamellar cycle forming
Blackites shafikii sp. nov. (Pl. 8, figs 1, 2 & 10) the lower part of the process rather than a distinct collar. The
top of the process ends in a distinct and more widely spaced
lamellar cycle (Pl. 8,fig.4).
DERIVATION OF NAME. Named after Dr Samir Shafik, mi-
cropalaeontologist. OCCURRENCE. Rare in SEM studies of these EOB sections.
 390 T O M D U N K LE Y J O N E S ET AL.
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Plate 8 SEM images of Blackites and Rhabdosphaera coccoliths; Figs 9-12 show main structural elements: rim cycle (Rm), radial cycle (RC)
and lamellar cycles numbered in sequence (LC). Figs 1,2 & 10 Blackites shafikii sp. nov.; 1, (holotype), 2 (paratype) & 10, TDP12/26-2,62 cm.
Figs 3,9 Blackites base TDP12/26-2,62 cm. Figs 4,5 Blackites sp. 1TDP12/26-2, 62 cm. Figs 6,7 Rhabdosphaera xiphos; 6, TDP12/47-2, 64 cm;
7, TDP12/47-2,64cm. Fig. 8 Rhabdosphaera type 1TDP12/23-2, 79 cm. Fig. 11 Blackites culterTDP12/26-2,62 cm. Fig. 12 Blackites cf. B. shafikii
sp. nov. TDP12/47-2, 64 cm. All SEMs above the white dividing line are to the same scale; all SEMs below the dividing line are enlarged relative
to those above (see scale bars).
 UPPER EOCENE TO LOWER OLIGOCÈNE CALCAREOUS NANNOFOSSILS FROM TANZANIA
Genus RHABDOSPHAERA Haeckel, 1894
DIAGNOSIS. Includes coccolithophores that produce both
spine and non-spine bearing rhabdoliths with no radial lath
cycle. Lamellar cycle fills central-area and forms spine where
present.
REMARKS. Based on the above discussion of the genus
Blackites, we restrict the use of the genus Rhabdosphaera
in fossil material to spine and non-spine bearing rhabdol-
iths that lack a radial cycle. The spine of Rhabdosphaera
is formed of one continuous lamellar cycle rather than the
multiple, tiered lamellar cycles characteristic of Blackites.
Rhabdosphaera vitrea (Deflandre in Deflandre &
Fert, 1954) Bramlette & Sullivan, 1961 (Pl. 7, figs 9 &
22-24)
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DESCRIPTION. Rhabdosphaera vitrea rhabdoliths are de-
scribed as having a very narrow styliform spine that typically
has a prominent basal collar. The basal coccolith is broad and
dominated by the inner cycle, with the outer cycle forming
only the outermost edge.
REMARKS. SEM image (Pl. 7, figs 22-24) shows the
buttress-like structures at the base of the spine that appear
as a 'collar' in LM. It also appears that R. vitrea lacks the
radial lath cycle characteristic of Blackites (Pl. 7, figs 23 &
24) hence should be assigned to the genus Rhabdosphaera
following Bramlette & Sullivan (1961).
OCCURRENCE. Rare throughout these EOB sections. Ranges
from NP11 to NP23 in the Tanzanian sections. First described
from the middle Eocene of France (Deflandre & Fert 1954)
and reported to range into the middle Oligocene (Perch-
Nielsen 1985). Genus ALGIROSPHAERA Schlauder, 1945 emend.
DIAGNOSIS. Rhabdosphaeraceae rim with a radial cycle but
Rhabdosphaera gracilentus (Bown & Dunkley Jones, with a domal or double-lipped protrusion formed of rod-like
2006) comb. nov. (Pl. 7, figs 13,14 & 25) elements and often covered with plate-like laths.
BASIONYM. Blackites gracilentus Bown & Dunkley Jones,
2006: 28, pl. 3, fig. 21.
DESCRIPTION. Rhabdolith coccoliths which, in LM, have a
thin, narrow coccolith base and a narrow spine with an axial
canal.
REMARKS. In SEM the spines of R. gracilentus appear sim-
ilar in morphology to the extant R. clavigera Murray &
Blackman, 1898 but are formed of less rapidly spiralling
elements (Pl. 7, fig. 25). The rhabdolith base lacks a ra-
dial lath-cycle, hence its reassignment to Rhabdosphaera.
The original description of B. gracilentus (Bown & Dunkley
Jones 2006) figured a specimen from the lower Oligocene
of TDP12 (pl. 3, fig. 20 in Bown & Dunkley Jones 2006;
reproduced here as Pl. 7, fig. 16a), which is now, on the basis
of SEM studies, reassigned to R. xiphos described below.
OCCURRENCE. Rare to frequent throughout these EOB sec-
tions. Ranges from NP14b/15a to NP21 in the Tanzanian
sections.
391
Rhabdosphaera xiphos (Deflandre & Fert, 1954)
Norris, 1984 (Pl. 7, fig. 16; Pl. 8, figs 6 & 7)
DESCRIPTION. Dimorphic rhabdoliths: spinose rhabdoliths
have a very narrow, gracile spine which is typically two to
three times longer than the width of the coccolith with a nar-
row collar at the base (Pl. 8, fig. 6); non-spinose rhabdoliths
are very small, broadly elliptical to subcircular with a nar-
row rim, no radial cycle, a lamellar cycle filling central-area
and approximately 13 imbricate plates of the lamellar cycle
form a distinctive circular whorl on the distal surface (Pl. 8,
fig. 7).
DIFFERENTIATION. Spinose forms similar to Rhab-
dosphaera gracilentus in LM but with a much finer, needle-
like spine. In SEM, differentiated from R. gracilentus by the
presence of a narrow collar at the base of the spine, which
is needle-like (Pl. 8, fig. 6), whereas the spine of R. graci-
lentus expands distally producing a club-shaped appearance
(Pl. 7, fig. 25). Appears indistinguishable from spine-bearing
coccoliths of extant R. xiphos (Young et al. 2003).
OCCURRENCE. Rare throughout these EOB sections. Ranges
from NP19/20 to 21 in the Tanzanian sections. The extant R.
xiphos had no previously known fossil record.
Rhabdosphaera type 1 (Pl. 8, fig. 8)
DIAGNOSIS. Small broadly elliptical to subcircular, slightly
domed, Rhabdosphaera-type planolith dominated by the
central lamellar cycle.
OCCURRENCE. Very rare in SEM studies of these EOB sec-
tions.
Norris, 1984
REMARKS. Differs from other members of the Rhab-
dosphaeraceae, which have processes formed from multiple
sets of lamellar cycles (Probert et al. 2007). Algirosphaera
was previously only known from the modern plankton and
rare occurrences in Pleistocene sediments (Aubry 1999;
Okada & Matsuoka 1996).
Algirosphaera fabaceus (Bown & Dunkley Jones,
2006) Bown et al., 2009 (Pl. 9, figs 1-7)
DIAGNOSIS. Coccolith with typical Rhabdosphaeraceae rim
structure but with a low two-part sacculiform spine (a wide
dome with narrower base) constructed from a series of ver-
tical and oblique lath-like elements and with distal portion
covered with multiple plate-like laths.
REMARKS. First described from LM observations of the
distal view of the coccolith (holotype figure is reproduced
here as Pl. 9, fig. 3), which have now been recognised as
strikingly similar to LM distal views of extant Algirosphaera
(J. Young, pers. comm.) and linked to side views in LM (Pl. 9,
figs 1 & 2) and a number of SEM images (Pl. 9, figs 4-7).
 392 T O M D U N K L E Y J O N E S ET AL.

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la.
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A Igirosphaera fabaceus

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s
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5. Algirosphaera fabaceus

CL j ^ ^ VL
LC ^

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' Rm
8. Algirosphaera sp. 1
^ A
6., ilgirosphaera fabaceus

non-spinous 2//m
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1
I "à Jw 1
;
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spinous
\f (paratype)
F
13. Afe/r.? sp. 3
\
holotype
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10. Acanthoica hackmanii

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11.
12. Algirosphaera? sp. 2
\4.Algir.? sp. 3

Plate 9 LM (XPL unless stated otherwise) and SEM images of Algirosphaera and Acanthoica coccoliths. Figs 1-3 Algirosphaera fabaceus
TDP11/15-1, 24 cm; 1a (XPL); 1b (PC); 2, TDP12/38-1,90 cm; 3, distal view TDP11/15-1, 24 cm. Figs 4-7 Algirosphaera fabaceusTDP12/26-2,
62 cm; rim cycle (Rm), radial cycle (RC), lamellar cycle (LC), vertical laths (VL), oblique laths (OL), cover laths (CL); 4, distal and side views; 5,
proximal and distal views; 6, side view, dotted line highlights diminutive cycles of oblique laths between vertical laths; 7, proximal view. Figs 8,
9 Algirosphaera sp. 1; 8, proximal view TDP12/23-2, 79 cm; 9, distal view TDP12/47-2,64 cm. Figs 10,11 Acanthoica backmanii sp. nov.
TDP12/26-2,62 cm; 10, showing a number of body coccoliths without processes and one spine-bearing (apical?) coccolith, holotype arrowed; 11,
(paratype) distal view. Fig. 12 Algirosphaera?sp. 2TDP12/26-2, 62 cm. Figs 13,14 Algirosphaera?sp. 3; 13, TDP12/47-2,64 cm; 14, TDP12/26-2,
62 cm. All LMs to the same scale; all SEMs above the white dividing line are to the same scale (enlarged relative to LMs); all SEMs below the
dividing line are enlarged relative to SEMs above (see scale bars).
 UPPER EOCENE TO LOWER OLIGOCÈNE CALCAREOUS NANNOFOSSILS FROM TANZANIA
The two-part process of A. fabaceus appears to be made up of
three structural elements directly comparable to those in A.
robusta (Probert et al., 2007), namely: (1) a thin outer layer
of cover laths (CL), which cover the distal, domal portion of
the process (Pl. 9, figs 4 & 6); (2) multiple sets of slightly ob-
lique laths (OL), which appear to form the bulk of the distal,
domal part of the process and are organised into two dis-
crete sets, along each side of the hood, with a central cavity
running between them (Pl. 9, figs 4 & 5); (3) a set of ver-
tical laths (VL) that extend vertically from the lamellar cycle
within the coccolith base, up to around a third of the total
process-height and form the exterior of the lower, narrower
part of the process (Pl. 9, fig. 6). There appear to be four
or five regular cycles of shorter oblique laths inter-digitated
with the vertical laths within the basal section of the process,
which potentially extend inwards to form the internal struc-
ture of the process (Pl. 9, fig. 6). The structure of the process
in A. fabaceus differs from A. robusta in three key ways: (1)
the vertical and oblique rods of A. robusta are broader in A.
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fabaceus (hence termed laths rather than rods) and are organ-
ised into distinct cycles, whereas in A. robusta they are less
regularly distributed in a roughly helical arrangement; (2) A.
fabaceus only has cover laths over the more distal, wider,
part of the spine, whereas the whole process is covered in A.
robusta; (3) the oblique laths in A. fabaceus are at a much
higher angle, near the vertical, whereas they are ~45° in A.
robusta. The regularity of the vertical laths in A. fabaceus, ex-
tending up from the characteristic Rhabdosphaeraceae basal
lamellar cycle, are intermediate in form between the mul-
tiple lamellar cycles of Palaeogene Blackites species and the
more irregular construction of the vertical and oblique rods of
the modern A. robusta, which may make A. fabaceus a trans-
itional form between these two genera. In LM A. fabaceus has
a distinctly bipartite low sacculiform spine in XPL, with the
lower-spine unit broadening away from the coccolith-base to
a maximum width approximately equal to the basal cocco-
lith, and an upper-spine of a two part flattened dome, which
is bright in XPL (Pl. 9, figs 1 & 2). This is consistent with
LM observations of A. robusta, where the hood is promin-
ent and formed of crystals with vertically orientated c-axes -
probably the oblique rods of the process. This correlates with
A. fabaceus, where the bright (in XPL) distal dome is dom-
inated by oblique laths and the lower, dark section of the
process is dominated by vertical laths and suggests that the
vertical and oblique laths have distinct c-axis orientations.
The rim is indistinct in XPL but appears to show a brighter
inner cycle (radial cycle?) and a faint, dark outer cycle (rim?).
Typically the spine height is slightly greater than the width
of the coccolith.
DIFFERENTIATION. In LM gross morphology is similar to
Cruxia mericii Varol, 1989 but with less distinct elements
making up the basal coccolith and lower-spine, and bright
distinct domal upper-spine.
OCCURRENCE. Rare throughout these EOB sections, zones
NP19/20 and NP21, and not yet observed in any of the other
Tanzanian sites.
Algirosphaera sp. 1 (Pl. 9, figs 8 & 9)
DIAGNOSIS. Small broadly elliptical coccolith with typical
Rhabdosphaeraceae base (rim/radial cycle/lamellar cycle)
393
and a wide, open central-area. Long lath-like elements pro-
trude vertically upwards from the lamellar cycle, similar to
the vertical rods in A. fabaceus.
REMARKS. Presence of vertical rods suggest placement
within Algirosphaera as this is the only Rhabdosphaeraceae
genus that has vertical rods, rather than multiple helical
lamellar cycles, forming part of the process (Probert et al.
2007).
OCCURRENCE. Very rare in SEM studies of these EOB sec-
tions; not observed in other Tanzanian sites.
Algirosphaera? sp. 2 (Pl. 9, fig. 12)
DIAGNOSIS. Very small (~2 μm) elliptical coccolith with
apparently Rhabdosphaeraceae-type rim (only the rim and
radial cycle are visible) and a sacculiform process formed of
multiple, helically arranged elements.
REMARKS. Tentatively placed in genus Algirosphaera on the
basis of the multiple elements forming the outer covering (?)
of the process.
OCCURRENCE. One individual observed in upper Eocene
zone NP19/20; not observed in other Tanzanian sites.
Algirosphaera? sp. 3 (Pl. 9, figs 13 & 14)
DIAGNOSIS. Very small (~2 μm) long-elliptical coccolith
with apparently Rhabdosphaeraceae-type rim (only the rim
and radial cycle are visible) and low, elongate process formed
of multiple elements.
REMARKS. Very tentatively placed in genus Algirosphaera
on the basis of the multiple elements forming the process, but
may be correctly placed elsewhere in Rhabdosphaeraceae.
OCCURRENCE. Very rare in SEM studies of these EOB sec-
tions; not observed in other Tanzanian sites.
Genus ACANTHOICA Lohmann, 1903 emend.
Schiller, 1913 and Kleijne, 1992
DIAGNOSIS. Polymorphic coccospheres with well-
developed spines on apical and antapical coccoliths; body
coccoliths have a well-developed radial lath cycle and a
lamellar cycle that forms a variably developed protrusion.
REMARKS. AS in the genus Rhabdosphaera, the spine/
process of Acanthoica is formed of one continuous lamellar
cycle rather than the multiple, tiered lamellar cycles of Black-
ites. Acanthoica coccoliths are distinguished from those of
Rhabdosphaera by the presence of a distinct radial cycle in
the former.
Acanthoica backmanii sp. nov. (Pl. 9, figs 10 & 11)
DERIVATION OF NAME. Named after Professor Jan Backman
(Stockholm University), micropalaeontologist and palaeo-
ceanographer.
DIAGNOSIS. Small (2-3 μm), broadly elliptical body cocco-
liths with well-developed rim, radial lath and lamellar cycles.
 394 T O M D U N K LE Y J O N E S ET AL.

Lamellar cycle forms very low domed protrusion or elongate
spine (Pl. 9, fig. 10).
REMARKS. Body-coccolith morphology is very similar
to extant Acanthoica species (cf. A. quattrospina-A.
acanthifera) (Young et al. 2003) and the observation of a
collapsed coccosphere with only one spine bearing coccolith
(Pl. 9, fig. 10) is consistent with the restriction of spine-
bearing coccoliths to the apical-antapical regions of modern
Acanthoica.
DIMENSIONS. L 2.4 μm, W 1.8 μm.
TYPE MATERIAL. Holotype, Pl. 9, fig. 11. Paratype, Pl. 9, fig.
10 (arrowed). TDP Site 12, Pande, Tanzania. Upper Eocene,
Sample TDP12/26-2, 62 cm (Subzone NP19/20).
OCCURRENCE. Rare in SEM; NP19/20; TDP Site 12.
Holococcoliths
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(in XPL) inner cycle. SEM images (Pl. 11, figs 1-5) show
that they are constructed of multiple, small (<0.1 μm) crys-
tallites, with the main oval/polygonal rim observed in LM
consisting of a thin, flat layer of crystallites surrounding a
central, flat-topped dome with a perforate cover on the distal
side that corresponds to the 'hole' seen in LM (Pl. 11, figs
1-3).
DIMENSIONS. L 5.0 μm, W 4.7 μm.
TYPE MATERIAL. Holotype, Pl. 10, fig. 1a (fig. 1b same spe-
cimen). Paratype, Pl. 10, fig. 2. TDP Site 12, Pande, Tanzania,
lower Oligocene, TDP12/10-1, 55 cm (Zone NP21).
OCCURRENCE. Rare throughout these EOB sections;
NP19/20-21; TDP Sites 11, 12, 17. Not observed in other
Tanzanian sites.
Orthozygus aureus (Stradner, 1962) Bramlette &
Wilcoxon, 1967 (Pl. 10, figs 3 & 4; Pl. 11, fig. 9)

(Formerly included in the family Calyptrosphaeraceae

Boudreaux & Hay, 1969)
All modern holococcoliths were traditionally included DESCRIPTION. Elliptical, low-birefringence holococcolith
within the single family Calyptrosphaeraceae. However, the comprising a narrow rim spanned by dome-shaped fluted
higher taxonomy of holococcoliths is problematical because: protrusion. The protrusion is broad and pierced by a central
(1) it appears likely that most, if not all, extant holococco- hole surrounded by a number of smaller perforations. Dis-
lith taxa will prove to have hetercoccolith equivalents; (2) tinguishable in both plan and side views.
the type species of this family, Calyptrosphaera oblonga, OCCURRENCE. Rare throughout these EOB sections. Ranges
has now been shown to be part of the life-cycle, and ju- from NP16 to NP23 in the Tanzanian sections. Reported
nior synonym, of Syracosphaera pulchra, which makes the literature range from the upper Eocene to middle Oligocene
Calyptrosphaeraceae a junior synonym of the family Syra- (Gartner & Bukry 1969).
cosphaeraceae Lohmann, 1902; and (3) the rather generalised
holococcolith morphologies make taxonomically meaning-
Orthozygus brytika (Roth, 1970) Aubry, 1988 (Pl. 10,
ful homologies difficult to identify (see Young et al. 2003
for further discussion). More fundamentally, the family Ca- fig. 5)
lyptrosphaeraceae is clearly polyphyletic, making its formal
use impossible (Jordan et al. 2004). However, the consider- DESCRIPTION. Small, narrowly-elliptical non-birefringent
ation of fossil holococcoliths as a single informal grouping holococcolith comprising a narrow rim spanned by diagonal
is, at present, the only feasible approach to their taxonomy. cross-bars.
The identification of associated fossil hetero-/holococcolith
life-cycle phases by, for example, observing fossil holo- OCCURRENCE. Rare throughout these EOB sections. Ranges
heterococcolith combination coccospheres is unlikely. The from NP 19/20 to NP23 in the Tanzanian sections. First de-
following taxonomy is largely consistent with previous taxo- scribed from the early Oligocene of the US Gulf Coast (Roth
nomic monographs (e.g. Perch-Nielsen 1985; Aubry 1988; 1970).
Bown 2005a), and applies established generic names where
possible but, for the most part, infers limited phylogenetic Orthozygus arcus sp. nov. (Pl. 10, figs 6-8)
information.
DERIVATION OF NAME. From arcus, meaning a 'bow' or
Genus ORTHOZYGUS Bramlette & Wilcoxon, 1967 'arch', referring to the characteristic arched appearance of
DIAGNOSIS. Basin-shaped holococcoliths with a central the bridge of this holococcolith.
bridge. Formed of crystallites with c-axes vertical, hence DIAGNOSIS. A holococcolith that is usually seen in ob-
appears non-birefringent in plan view. lique view and is comprised of a low basal coccolith and
a low arched bridge; both coccolith and process are weakly-
Orthozygus occultus sp. nov. (Pl. 10, figs 1 & 2; Pl. 11,birefringent.
figs 1-5)
DIFFERENTIATION. The spine is broader and lower than that
of O. aureus and does not have the distinct perforations seen
DERIVATION OF NAME. From occultus, meaning hidden, re- on the distal surface of O. aureus. Crystallographic properties
ferring to the dark appearance of this holococcolith in XPL. similar to those of O. sudis but with a low and smoothly
arching bridge instead of pointed spine. Differentiated from
DIAGNOSIS. Medium-sized irregularly elliptical to poly- O. occultus by its visible birefringence in XPL and distinct
gonal holococcoliths, which are dark in XPL but have dis- low arched bridge.
tinctly high relief in PC. Appear to have a central hole in
LM, which sometimes contains a poorly developed but bright DIMENSIONS. H 3.2 μm, W 4.6 μm.
 UPPER EOCENE TO LOWER OLIGOCÈNE CALCAREOUS NANNOFOSSILS FROM TANZANIA 395
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Plate 10 LM (XPL unless stated otherwise) images of various holococcoliths. Figs 1,2 Orthozygus occultus sp. nov. TDP12/10-1,49 cm; 1a
holotype (PC); 1b (XPL); 2, paratype (PC) TDP12/46-1, 40cm. Figs 3 , 4 Orthozygus aureus; 3, TDP12/38-1, 90 cm; 4, TDP12/10-1,49 cm. Fig. 5
Orthozygus brytika TDP12/10-1,49 cm; 5a (XPL); 5b (PC). Figs 6 - 8 Orthozygus arcus sp. nov.; 6, TDP17/15-1,10cm; 7, TDP12/40-3, 97 cm
(holotype); 8, TDP17/42-1, 72 cm (paratype). Figs 9,10 Orthozygus sudis TDP12/10-1,49 cm. Figs 11,12 Lanternithus minutus; 11, TDP12/38-1,
90cm; 12, TDP12/10-1,49 cm. Fig. 13 Lanternithusprocerus TDP11/15-1, 24cm. Fig. 14 Quadrilateris imparidividuus TDP12/26-1, 56cm. Fig. 15
Varolia cistula TDP12/10-1, 55 cm. Fig. 16 Varolia sicca TDP12/46-1, 40 cm. Figs 17-18 Varolia macleodii; 17, TDP12/10-1, 55 cm; 18, TDP12/46-1,
40cm. Figs 19,20 Varolia boomeri; 19, TDP12/46-1,40cm; 20, TDP12/10-1, 55 cm. Fig. 21 Corannulusgermanicus TDP17/42-1; 21a (XPL); 21b
(PC). Fig. 22 Holodiscolithus macroporus TDP12/40-3,97 cm. Fig. 23 Holodiscolithus solidus TDP12/10-1, 55 cm. Fig. 24 Holodiscolithus geisenii
TDP12/36-2, 70cm. Fig. 25 Holodiscolithus minolettiiTDP12/46-1, 40 cm; 25a (PC); 25b (XPL). Fig. 26 Zygrhablithus bijugatus subsp. cornutus
TDP12/38-1,90 cm. Figs 27,28 Zygrhablithus bijugatus subsp. bijugatus; 27, TDP12/40-3,97 cm; 28, TDP12/38-1, 90 cm. Fig. 29 Trochoaster
simplex TDP12/46-1, 40cm (PC). Fig. 30 Clathrolithus joidesa TDP12/10-1, 55 cm; 30a (XPL); 30b (PC). All LMs are to the same scale.
 396 T O M D U N K L E Y J O N E S ET AL.
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Plate 11 SEM images of various holococcoliths. Figs 1-5 Orthozygus occultus sp. nov.; 1&4, TDP12/47-2, 64 cm; 2, TDP12/38-3, 93 cm; 3,
TDP12/42-3,86 cm; 5, TDP12/23-2, 79 cm. Figs 6,7 Zygrhablithus bijugatus subsp. cornutus TDP12/38-3, 93 cm. Fig. 8 Zygrhablithus bijugatus
subsp. bijugatus TDP12/23-2, 79 cm. Fig. 9 Orthozygus aureus TDP12/38-2, 99 cm. Figs 10-13 Lanternithus minutus TDP12/47-2,64cm.
Figs 14-18 Varolia cistula; 14,16,17, TDP12/26-2,62 cm; 15, TDP12/23-2, 79 cm; 18, TDP12/47-2, 64 cm. Figs 19-21 Varolia?spp.; 19,
TDP12/23-2, 79 cm; 20, TDP12/26-2,62 cm; 21, TDP12/38-3,93 cm. Fig. 22 Holodiscolithus solidus TDP12/23-2, 79 cm. All SEMs above the
white dividing line are to the same scale; all SEMs below the dividing line are enlarged relative to those above (see scale bars).
 UPPER EOCENE TO LOWER OLIGOCÈNE CALCAREOUS NANNOFOSSILS FROM TANZANIA 397

TYPE MATERIAL. Holotype, Pl. 10, fig. 7. Paratype, Pl. 10,
fig. 8. TDP Site 12, Pande, Tanzania, upper Eocene, Sample
TDP12/40-3, 97 cm (Zone NP19-20).
OCCURRENCE. Rare throughout the section; NP19/20-21;
TDP Sites 11,12, 17. Not observed in other Tanzanian sites.
Orthozygus sudis Bown & Dunkley Jones, 2006 (Pl.
10, figs 9 & 10)
DESCRIPTION. A holococcolith that, in side view, comprises
a low basal coccolith and a relatively tall, hollow spine that
tapers to a point; both coccolith and spine are birefringent in
side view and crystallographically continuous in XPL.
OCCURRENCE. Rare throughout these EOB sections. Ranges
from NP19/20 to NP21 in the Tanzanian sections.
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Genus VAROLIA Bown, 2005a
DIAGNOSIS. Cavate, thin-walled, box-like holococcoliths
usually seen in side view.
Varolia cistula Bown, 2005a (Pl. 10, fig. 15; Pl. 11, figs
14-18)
DESCRIPTION. A small to medium-sized, thin-walled cavate
holococcolith with a gently curved distal-cover. The structure
broadens distally from the basal plate and is typically wider
than it is high. Formed of tiny (~0.1 μm) crystallites (Pl. 11,
figs 14-18).
OCCURRENCE. Frequent to common in the upper Eocene be-
coming rare in the lower Oligocene of these sections. Ranges
from NP 19/20 to NP23 in the Tanzanian sections.
Varolia sicca Bown & Dunkley Jones, 2006 (Pl. 10, fig.

Genus LANTERNITHUS Stradner, 1962 16)

DIAGNOSIS. Cavate sub-hexagonal holococcoliths with a
series of protruding flanges - birefringent in plan view. DESCRIPTION. A small, elliptical coccolith with a narrow,
birefringent rim. When orientated at 0• the rim is crossed by
Lanternithus minutus Stradner, 1962 (Pl. 10, figs 11 & narrow N-S and E-W extinction lines.
12; Pl. 11, figs 10-13)
OCCURRENCE. Rare throughout these EOB sections. Ranges
from NP15b to NP21 in the Tanzanian sections.
DESCRIPTION. Slightly rectangular holococcolith in plan
view, with distinct lateral and end rim blocks and a large Varolia macleodii Bown & Dunkley Jones, 2006 (Pl.
central pore. In side view it has a box-like appearance that 10, figs 17 & 18)
tapers slightly to one (the distal?) end where it is capped by
a distal cover.
DESCRIPTION. A thin-walled, cavate coccolith with a domed
OCCURRENCE. Frequent to common in the upper Eocene distal-cover. The cavate central-area space is spanned by at
becoming rare in the lower Oligocene of these EOB sections. least two vertical septa.
Ranges from NP14b/15a to NP23 in the Tanzanian sections.
Reported literature range from the middle Eocene to early OCCURRENCE. Rare throughout these EOB sections. Not ob-
Oligocene (Gartner & Bukry 1969). served in other Tanzanian sites.

Lanternithus procerus Bown, 2005a (Pl. 10, fig. 13) Varolia boomeri Bown & Dunkley Jones, 2006 (Pl. 10,
figs 19 & 20)

DESCRIPTION. Broadly similar to L. minutus but with elong-

ated sides, forming a taller 'process' when observed in side
view.
OCCURRENCE. Rare throughout these EOB sections. Ranges
from NP15b to NP23 in the Tanzanian sections.
Genus QUADRILATERISVarol, 1991
DIAGNOSIS. Quadrilateral birefringent holococcolith
formed of four blocks.
Quadrilateris imparidividuus Varol, 1991 (Pl. 10,
fig.14)
DESCRIPTION. A small, narrowly rectangular coccolith
formed from four thin blocks that are birefringent in XPL.
OCCURRENCE. Rare throughout these EOB sections. Not
observed in other Tanzanian sites. First described from the
middle Eocene of Pakistan (Varol 1991).
DESCRIPTION. A thin-walled, cavate coccolith with a poin-
ted distal-cover. At least two distinct septa are present within
the cavity, running sub-parallel to and just beneath the distal-
cover.
OCCURRENCE. Rare throughout these EOB sections. Not ob-
served in other Tanzanian sites.
Varolia? spp. (Pl. 11, figs 19-21)
A number of <3 μm, simple cavate holococcolith morpho-
logies without distal covers (calicalith-type) were observed
during SEM studies. These are tentatively assigned to the
genus Varolia pending further studies of Paleocene to middle
Eocene sediments of Tanzania, which contain similar holo-
coccolith morphologies.
Genus CORANNULUS Stradner, 1962
DIAGNOSIS. Non-birefringent in plan view, discoidal holo-
coccolith with large central opening and marginal perfora-
tions or indentations.
 398 TOM D U N K LE Y J O N E S ET AL.

Corannulusgermanicus Stradner, 1962 (Pl. 10, fig. 21) Holodiscolithus minolettii Bown, 2005a (Pl. 10,
fig. 25)
DESCRIPTION. Ring-shaped, broadly elliptical, non-
birefringent holococcolith with multiple (~7-ll) thorn-like DESCRIPTION. A small, elliptical coccolith with a narrow
projections around the outer edge. Morphology of holo- dark rim visible in PC (Pl. 10, fig. 25a) and a complexly-
coccolith clearest in PC where outer rim shows high constructed, birefringent central-area plate with bright blocks
relief. towards each end.

OCCURRENCE. Very rare in these EOB sections. Ranges
from NP17 to NP21 in the Tanzanian sections. Reported
literature range is from the middle to the late Eocene (Siesser
1983; Perch-Nielsen 1985).
Genus HOLODISCOLITHUS Roth, 1970
DIAGNOSIS. Elliptical non-birefringent holococcoliths with
perforations.
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OCCURRENCE. Rare throughout these EOB sections. Ranges
from NP15b to NP21 in the Tanzanian sections.
Genus ZYGRHABLITHUS Deflandre, 1959
DIAGNOSIS. Spinose holococcoliths that are birefringent in
side view. In plan view the coccoliths are non-birefringent
with a distinct figure-of-eight shaped central structure.
Zygrhablithus bijugatus subsp. bijugatus (Deflandre

in Deflandre & Fert, 1954) Deflandre, 1959 (Pl. 10, figs

REMARKS. Following Bown (2005a) and Bown & Dunkley 27 & 28; Pl. 11, fig. 8)
Jones (2006), we have employed this genus more widely
to include small elliptical holococcoliths with low or no bi-
refringence in XPL. Medium to large spinose holococcolith com-
DESCRIPTION.
prising a rim and central process; the spine has a narrow
axial canal and tapers uniformly to a distal point or blunt
Holodiscolithus macroporus (Deflandre in Deflandre termination.
& Fert, 1954) Roth, 1970 (Pl. 10, fig. 22) OCCURRENCE. Rare to frequent in the upper Eocene becom-
ing rare to very rare in the lower Oligocene of these EOB
DESCRIPTION. Elliptical non-birefringent holococcoliths sections. Ranges from NP9 to NP23 in the Tanzanian sec-
that appear as a dark plate pierced by a variable number tions. Reported literature range is from the early Paleocene
of pores. (Perch-Nielsen 1985) to NN2 (Young 1998).

OCCURRENCE. Rare throughout these EOB sections. Ranges
from NP9b to NP23 in the Tanzanian sections. Reported
literature range is from the Palaeogene to NN18 (Young
1998).
Holodiscolithus solidus (Deflandre in Deflandre &
Fert, 1954) Roth, 1970 (Pl. 10, fig. 23; Pl. 11, fig. 22)
DESCRIPTION. Elliptical holococcoliths comprising a rim
and six central-area bars, which are typically weakly- or
non-birefringent. The bars merge centrally to form a ridge
that bifurcates at either end.
OCCURRENCE. Rare throughout these EOB sections. Ranges
from NP9 to NP23 in the Tanzanian sections. Reported lit-
erature range is from the middle Eocene (Siesser 1983) to
middle Oligocene (Roth 1970).
Holodiscolithus geisenii Bown, 2005a (Pl. 10, fig. 24)
Zygrhablithus bijugatus subsp. cornutus Bown,
2005a (Pl. 10, fig. 26; Pl. 11, figs 6 & 7)
DESCRIPTION. Medium-sized spinose holococcolith com-
prising a rim and central process; the spine is relatively long,
has a narrow axial canal and short lateral protrusions or horns
at its distal end (two in side view).
OCCURRENCE. Rare to frequent in the upper Eocene becom-
ing rare to very rare in the lower Oligocene of these EOB
sections. Ranges from NP14b/15a to NP23 in the Tanzanian
sections.
Genus CLATHROLITHUS Deflandre in Deflandre &
Fert, 1954
DIAGNOSIS. Applied here to unusual holococcoliths com-
prised of a domed, perforate framework.
Clathrolithus joidesa (Bukry & Bramlette, 1968)
Bown, 2005a (Pl. 10, fig. 30)

DESCRIPTION. Large, non-birefringent, C-shaped holococ-

DESCRIPTION. A small, broadly elliptical coccolith with a
narrow dark rim and a weakly birefringent central-area plate,
which is crossed by gently curving axial sutures. Shows low
relief in PC.
OCCURRENCE. Rare in the upper Eocene of these EOB sec-
tions. Ranges from NP11 to NP23 in the Tanzanian sections.
colith only observed in side view. Inner margin of coccolith
usually shows higher birefringence in XPL. No visible pores.
OCCURRENCE. Rare throughout these EOB sections. Ranges
from NP19/20 to NP23 in the Tanzanian sections. First de-
scribed from upper Eocene into the Oligocene (Bukry &
Bramlette 1968).
 UPPER EOCENE TO LOWER OLIGOCÈNE CALCAREOUS NANNOFOSSILS FROM TANZANIA 399

Nannoliths Genus MICRANTHOLITHUS Deflandre in Deflandre

Family BRAARUDOSPHAERACEAE Deflandre, 1947
DIAGNOSIS. The extant species Braarudosphaera bigelowii
has a cell-wall covering of 12 pen taliths forming an imper-
forate dodecahedron. Recent molecular genetic analysis has
confirmed the placement of B. bigelowii within the Hapto-
phyta (Takano et al. 2006). Extinct members of the family
Braarudosphaeraceae include all pentalith nannoliths. Pent-
aliths have a laminar ultrastructure with c-axes tangential to
pentalith outline.
Genus BRAARUDOSPHAERA Deflandre, 1947
DIAGNOSIS. Pentagonal nannoliths formed of five, non-
perforated trapezoidal segments; sutures between segments
go to the edges of the pentagon.
Braarudosphaera bigelowii (Gran & Braarud, 1935)
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& Fert, 1954
DIAGNOSIS. Pentagonal nannoliths formed of five trapezoi-
dal elements; sutures between segments go to the vertices of
the pentagon.
Micrantholithus minimus Bown & Dunkley Jones,
2006 (Pl. 12, figs 7, 8, 20 & 21)
DESCRIPTION. A small (typically <4 μm) Micrantholithus
with raised ridges along the line of the segment sutures
that give the appearance of a deeply indented pentalith. In
LM, many specimens appear to have a thin calcite laminar
between the raised, birefringent areas, which is confirmed by
SEM observations (Pl. 12, figs 20 & 21). This form has a
similar morphology to M. excelsus but is considerably smal-
ler.

Deflandre, 1947 (Pl. 12, figs 1-3,15 & 16) OCCURRENCE. Frequent to common throughout these EOB
sections. Ranges from NP16 to NP21 in the Tanzanian sec-
DESCRIPTION. Regular pentagonal nannoliths with a tions.
smooth, flat or slightly concave surface. Type species of the
genera Braarudosphaera. This species often exhibits consid- Micrantholithus excelsus Bown, 2005a (Pl. 12, fig. 9)
erable size variability; in this section pentaliths range in size
between 3.3 and 11.6 μm with larger forms tending to have
DESCRIPTION. Large, elevated Micrantholithus with deeply
more rounded apices.
indented sides and gracile, long rays.
REMARKS. Recent genetic and morphological studies of OCCURRENCE. Rare throughout these EOB sections. Ranges
modern B. bigelowii populations strongly suggest that they from NP11 to NP23 in the Tanzanian sections.
consist of several pseudo-cryptic species (Takano et al.
2006).
Micrantholithus hebecuspis Bown, 2005a (Pl. 12,
OCCURRENCE. Frequent to common throughout these EOB figs 10 & 11)
sections. Ranges from NP9 to NP23 in the Tanzanian sec-
tions. Reported literature range is from the Aptian to the
DESCRIPTION. Large Micrantholithus with deeply indented
modern (Young 1998).
sides and blunt,flat-endedapices.

Braarudosphaera cf. B. hockwoldensis Black, 1973 OCCURRENCE. Rare throughout these EOB sections. Ranges
(Pl. 12, fig. 4) from NP10 to NP21 in the Tanzanian sections.

Micrantholithus triquetra (Bown & Dunkley Jones,

DESCRIPTION. Braarudosphaera with protruding pentalith
2006) comb. nov. (Pl. 12, figs 13,14, 22 & 23; Pl. 13,
segments.
fig. 1)
OCCURRENCE. Rare throughout these EOB sections. Ranges
from NP10 to NP23 in the Tanzanian sections. First described BASIONYM. Pemma? triquetra Bown & Dunkley Jones,
from the Lower Cretaceous (Black 1973). 2006: 34, pl. 9, fig. 7.

Braarudosphaera stylifera Troelsen & Quadros, 1971 DESCRIPTION. Relatively large (typically >5 μm),
(Pl. 12, figs 5 & 6) triangular-shaped nannoliths that thicken along the longer
sides and exhibit low birefringence in XPL. Appear similar
to disaggregated Pemma or Micrantholithus segments in
DESCRIPTION. Tall pentalith, which in side view tapers up- LM but with a more acute angle between the longer edges
wards from a wide base. In plan view the pentaliths show an (~50-60°, compared to 72• for typical pentaliths).
indistinct, lower birefringence, outer margin and increased
birefringence towards the centre. The standard pentalith mor- REMARKS. Segments have been very rarely observed lying
phology of the base becomes visible by altering the depth of with long-edges adjacent (Pl. 12, fig. 13) similar to disag-
focus. gregated pentaliths but this may be a chance occurrence. The
c-axis orientation is similar to other pentaliths, i.e. tangential,
OCCURRENCE. Rare throughout these EOB sections. Ranges which was the basis for the tentative generic designation. In
from NP15b to NP23 in the Tanzanian sections. First de- SEM, segments are observed to consist of a series of fine cal-
scribed from the middle Eocene of Brazil (Troelsen & cite laminae similar to other members of the Braarudosphaer-
Quadros 1971). aceae (Pl. 12, figs 22 & 23). A simple explanation of the more
 400 TOM D U N K L E Y J O N E S ET AL.
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Plate 12 LM (XPL unless stated otherwise) and SEM images of Braarudosphaera, Micrantholithus and Pemma nannoliths. Figs 1-3
Braarudosphaera bigelowii; 1, TDP17/15-1,10 cm; 2-3, TDP12/40-3,9 cm. Fig. 4 Braarudosphaera cf. B. hockwoldensis TDP12/47-1, 61 cm.
Figs 5,6 Braarudosphaera stylifera; 5, TDP12/40-3,9 cm; 6, TDP12/26-1, 56 cm. Figs 7,8 Micrantholithus minimus; 7, TDP12/46-1,40 cm; 8,
TDP12/10-1, 55 cm. Fig. 9 Micrantholithus excelsus TDP11/23-1, 23 cm. Figs 10,11 Micrantholithus hebecuspis TDP12/46-1,40 cm. Fig. 12 Pemma
papillatum TDP12/46-1,40cm; 12a (XPL); 12b (PC). Figs 13,14 Micrantholithus triquetra; 13, TDP12/38-1,90 cm; 14, TDP12/10-1, 55 cm. Figs 15,
16 Braarudosphaera bigelowii TDP12/47-1,61 cm. Figs 17,18 Pemma papillatum; 17, TDP12/42-3, 86 cm; 18, TDP12/47-2,64 cm. Fig. 19
Micrantholithus sphere TDP12/23-2, 79 cm. Figs 20,21 Micrantholithus minimus TDP12/23-2, 79 cm. Figs 22,23 Micrantholithus triquetra
TDP12/38-3,93 cm. All LMs and SEMs above the white dividing line are to the same scale; all SEMs below the dividing line are enlarged relative
to those above (see scale bars).
 UPPER EOCENE TO LOWER OLIGOCÈNE CALCAREOUS NANNOFOSSILS FROM TANZANIA 401
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Plate 13 SEM image of Micrantholithus triquetra and LM (all PC) images of Discoaster nannoliths. Fig. 1 Micrantholithus triquetra
TDP12/38-3,93 cm. Figs 2,3 Discoastersaipanensis; 2, TDP12/40-2,90 cm; 3, TDP12/46-1,40cm. Fig. 4 Discoaster tanii TDP12/10-1, 55 cm.
Figs 5,6 Discoaster nodiferTDP12/14-1, 21cm. Figs 7,8 Discoast er ornat us; 7, TDP11/23-1, 23 cm; 8, TDP12/26-1, 56 cm. Figs 9,10 Discoaster
distinctus; 9, TDP12/38-1, 90 cm; 10, TDP12/14-3, 29 cm. Figs 11-14 Discoaster deflandrei; 11, TDP12/46-1, 40 cm; 12, TDP12/10-1, 55 cm; 13-14,
TDP12/36-2, 70 cm. All LM images are to the same scale; SEM image is enlarged relative to LMs (see scale bars).
 402 T O M D U N K L E Y JONES ET AL.
acute opening angle and higher c-axis orientation is that these
are segments of a Micrantholithus-type pentalith with a con-
cave base (cf. Pl. 12, fig. 19). This evidence, along with the
absence of any distinct perforation, warrants the reassign-
ment to the genus Micrantholithus.
OCCURRENCE. Frequent to common throughout these EOB
sections (NP19/20 to NP21). Not observed in other Tanzanian
sections.
Genus PEMMA Klumpp, 1953
DIAGNOSIS. Pentaliths with a depression in each segment;
often producing elevated, robust structures that are com-
monly observed in side view.
Pemmapapillatum Martini, 1959 (Pl. 12, figs 12 &
17-18)
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DESCRIPTION. Large pentaliths with distinct depression near
the apex of each pentalith and characteristic multiple nodes
along the outer margin of each pentalith.
OCCURRENCE. Common in the upper Eocene of the section
(zones NP19/20-21) but, in this section, it has a distinct last
occurrence within zone NP21 of TDP12, ~7 m below the HO
Hantkeninidae, which characterises the Eocene-Oligocene
boundary. Varol (1998) reports a similar highest occurrence
just below the EOB in the North Sea. Ranges from NP15b to
NP21 in the Tanzanian sections. Reported first occurrence of
Pemma is in the middle Eocene (Perch-Nielsen 1985).
Family DISCOASTERACEAE Tan, 1927
DIAGNOSIS. Discoidal nannoliths with 3–40 elements radiat-
ing from a common centre or axis. c-Axes are vertical hence,
in plan view, are non-birefringent in XPL. Discoasters are
rare in the TDP material and not especially well preserved;
no new species are described or possible coccospheres ob-
served.
Genus DISCOASTER Tan, 1927
DIAGNOSIS. Star or rosette-shaped nannofossils with low
birefringence in XPL.
Discoastersaipanensis Bramlette & Riedel, 1954
(Pl. 13, figs 2 & 3)
DESCRIPTION. Discoasters with normally six or seven
straight or curved rays joined through half their length and
which then taper to a point. A central boss and distinct radial
sutural ridges and depressions are visible in some specimens
(Pl. 13, fig. 2).
OCCURRENCE. Rare through the upper Eocene zone
NP19/20 of these EOB sections. Ranges from NP14b/15a
to NP19/20 in the Tanzanian sections. Reported literature
range is from NP15 to NP19/20 (Perch-Nielsen 1985); HO
defines the top of NP20 (Martini 1971).
Discoaster tanii Bramlette & Riedel, 1954 (Pl. 13,
fig. 4)
DESCRIPTION. Typically five-rayed discoaster with rays of
uniform width terminating in a flat or slightly notched ray-
end. May show poorly developed unpaired lateral nodes on
some rays but does not have the regular paired ray-nodes of
Discoaster nodifer. This form has close morphological affin-
ities to and may intergrade with D. nodifer and D. ornatus,
which are frequently classified as subspecies or closely re-
lated species of D. tanii.
OCCURRENCE. Rare throughout these EOB sections. Ranges
from NP14b/15a to NP21 in the Tanzanian sections. Reported
literature range is from NP17 to NP23 (Perch-Nielsen 1985).
Discoaster nodifer Bramlette & Riedel, 1954 (Pl. 13,
figs 5 & 6)
DESCRIPTION. Potentially six to nine but typically six-rayed
discoaster with rays of uniform width and blunt or notched
ray-ends. Differentiated from Discoaster tanii by the con-
sistent presence of well-developed paired ray-nodes on all
rays.
OCCURRENCE. Rare throughout these EOB sections. Ranges
from NP14b/15a to NP23 in the Tanzanian sections. Reported
literature range is from NP15 to NP23 (Perch-Nielsen 1985).
Discoaster ornatus (Bramlette & Wilcoxon, 1967)
Bown, 2005a (Pl. 13, figs 7 & 8)
DESCRIPTION. Five-rayed discoaster, rays of uniform width
with well-developed, slightly bifurcating, terminal ray-end
notches and paired lateral ray-nodes. Discoaster ornatus
tends to have thicker rays, a more conspicuous central boss
and better developed ray-end notches than D. tanii or D.
nodifer.
OCCURRENCE. Rare throughout these EOB sections. Ranges
from NP19/20 to NP23 in the Tanzanian sections. Reported
literature range is from NP21 to NP23 (Perch-Nielsen 1985).
Discoaster distinctus Martini, 1958 (Pl. 13, figs 9 &
10)
DESCRIPTION. Five or six-rayed stellate discoasters with
prominent ray-tip bifurcations that resemble spanners. Ap-
pears similar to some forms of Discoaster deflandrei but
has rays of uniform width terminated by distinct ray-tip bi-
furcations whereas D. deflandrei often has rays that slightly
increase in width towards the ray-tip.
OCCURRENCE. Rare throughout these EOB sections. Ranges
from NP14b/15a to NP23 in the Tanzanian sections. Reported
literature range is from NP12 to NP14 (Perch-Nielsen 1985).
 UPPER EOCENE TO LOWER OLIGOCÈNE CALCAREOUS NANNOFOSSILS FROM TANZANIA
Discoaster deflandrei Bramlette & Riedel, 1954
(Pl. 13, figs 11-14)
DESCRIPTION. Normally six-rayed, but includes five- to
seven-rayed, discoasters with a terminal bifurcation but
lacking a well-developed central boss. The general ray-
morphology can be highly variable between specimens with
some showing a moderate to extreme widening of rays to-
wards the ray-ends (Pl. 13, fig. 14).
OCCURRENCE. Rare throughout these EOB sections. Ranges
from NP14b/15a to NP23 in the Tanzanian sections. Reported
literature range is from NP11 (Perch-Nielsen 1985) to NN7
(Young 1998).
Family SPHENOLITHACEAE Deflandre, 1952
DIAGNOSIS. Conical-shaped nannoliths consisting of several
superimposed cycles of elements all radiating from a com-
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mon point of origin; c-axes of the elements run along their
length.
Genus SPHENOLITHUS Deflandre, 1952
DIAGNOSIS. See family description.
Sphenolithus moriformis (Bronnimann & Stradner,
1960) Bramlette & Wilcoxon, 1967 (Pl. 14, fig. 1)
DESCRIPTION. Small to medium-sized (H 4-7 μm) non-
spined sphenolith with a domed upper surface.
OCCURRENCE. Frequent to common throughout these EOB
sections; becomes more abundant in the lower Oligocene
(NP21). Ranges from NP9 to NP23 in the Tanzanian sections.
Reported literature range is from NP12 (Perch-Nielsen 1985)
to NN10 (Young 1998).
Sphenolithus conicus Bukry, 1971 (Pl. 14, figs 2 & 3)
DESCRIPTION. Medium to large sphenolith with short, blunt
spine and broad robust base. Lower quadrants are large and
the spine is dark at 0•.
OCCURRENCE. Rare in the lower Oligocene (zone NP21)
of these EOB sections. Ranges from NP19/20 to NP23 in
the Tanzanian sections. Reported literature range is from the
Oligocene to NN3 (Young 1998).
Sphenolithus radians Deflandre, 1952 (Pl. 14, fig. 4)
DESCRIPTION. Medium to large sphenolith with a tall com-
pound spine and square-shaped base with equidimensional
quadrants. The spine is dark but visible at 0• and charac-
terised by a median suture line; at 45• the spine is brightest.
Sometimes difficult to reliably distinguish from Sphenolithus
pseudoradians; in these sections S. radians is used for forms
with a smoother spine and a basal-width that is not signific-
antly greater than the base of the apical spine.
OCCURRENCE. Rare throughout these EOB sections. Ranges
from NP11 to NP23 in the Tanzanian sections. Reported
literature range is from NP11 to NP19 (Perch-Nielsen 1985).
403
Sphenolithus pseudoradians Bramlette & Wilcoxon,
1967 (Pl. 14, fig. 5)
DESCRIPTION. Medium to large sphenolith with a tall com-
pound spine. Distinguished from Sphenolithus radians by its
more irregular outline and apical cycles that extend laterally
beyond the base of the apical spine (Pl. 14, fig. 5b).
REMARKS. The first occurrence of S. pseudoradians marks
the base of zone NP20 (Martini 1971), however the applic-
ation of this event is problematic due to the difficulties in
reliably distinguishing between S. radians and S. pseudora-
dians.
OCCURRENCE. Rare throughout these EOB sections. Ranges
from NP17 to NP23 in the Tanzanian sections. Reported
literature range is from NP15 to NP24 (Perch-Nielsen 1985).
Sphenolithus predistentus Bramlette & Wilcoxon,
1967 (Pl. 14, figs 6 & 7)
DESCRIPTION. Small to medium-sized sphenolith (H 3.4-
7.5 μm excluding bifurcations) with tall, narrow bifurcating
spine and very short, lower quadrants.
OCCURRENCE. Frequent to common throughout these sec-
tions; becomes more abundant in the lower Oligocene
(NP21). Ranges from NP16 to NP23 in the Tanzanian sec-
tions. Reported literature range is from NP17 to NP24 (Perch-
Nielsen 1985).
Sphenolithus akropodus de Kaenel & Villa, 1996 (Pl.
14, fig. 8)
DESCRIPTION. Medium to large sphenolith with a long taper-
ing apical spine, sometimes bifurcated, and short laterally
extending proximal elements forming two pointed feet. In
XPL the spine is bright at 45• and weakly birefringent at 0•.
REMARKS. The separation of the early Oligocene spheno-
lith lineage S. predistentus - S. akropodus - S. distentus -
S. ciperoensis into consistent taxonomic entities is notori-
ously difficult. Here we follow de Kaenel & Villa (1996)
in differentiating S. akropodus from S. predistentus by the
appearance of the basal elements in XPL; at 45• to crossed
nicols the basal elements of S. akropodus form two bright
pointed feet that appear imbedded into the spine (Pl. 14, fig.
8a); in S. predistentus the basal elements form two smaller
feet that appear distinct from the main spine of the sphenolith
(Pl. 14, figs 6 & 7). Sphenolithus akropodus is distinguished
from S. distentus by its larger size and more massive apical
spine.
OCCURRENCE. Rare in the lower Oligocene zone NP21 of
TDP Site 17. Previously reported from lower Oligocene
zones NP22 and NP23 of ODP Site 900, Iberia Abyssal
Plain.
 ZJOZJ T O M D U N K L E Y J O N E S ET AL.
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Plate 14 LM (XPL unless stated otherwise) and SEM images of Sphenolithus and incertae sedis nannolith Pemma? uncinata. Fig. 1
Sphenolithus moriformis TDP12/10-1, 55 cm. Figs 2,3 Sphenolithus conicus; 2, TDP11/23-1, 23 cm; 3, TDP12/10-1, 55 cm. Fig. 4 Sphenolithus
radians TDP11/23-1, 23 cm. Fig. 5 Sphenolithus pseudoradians TDP12/13-1,10 cm. Figs 6,7 Sphenolithus predistentus; 6, TDP12/10-1, 55 cm; 7,
TDP11/10-1, 29 cm. Fig. 8 Sphenolithus akropodus TDP17/11-1,10cm. Figs 9,10 Sphenolithus tribulosus TDP11/23-1, 23 cm. Figs 11-17 Pemma?
uncinata; 11, TDP11/23-1, 23 cm; 12, TDP12/40-3, 9 cm; 13, TDP12/14-3, 29 cm; 14, TDP12/47-4, 35 cm; 15, TDP12/26-1, 56 cm; 15a (XPL); 15b
(PC); 16, TDP12/38-3,93 cm; 17, TDP12/26-2,62 cm. All LMs are to the same scale; SEMs are enlarged relative to LMs (see scale bars).
 UPPER EOCENE TO LOWER OLIGOCÈNE CALCAREOUS NANNOFOSSILS FROM TANZANIA 405

Sphenolithus tribulosus Roth, 1970 (Pl. 14, figs 9 & Pocillithus spinulifer sp. nov. (Pl. 15, figs 1-5)
10)

DERIVATION OF NAME. From spinula, meaning ' small spine'

DESCRIPTION. Like S. predistentus but the basal part of the and referring to the tall spines of these coccoliths.
sphenolith is considerably broader than the spine, giving an
inverted T-shaped outline. DIAGNOSIS. Miniscule to very small murolith (protolith)
coccoliths with high, narrow walls and a central-area spanned
OCCURRENCE. Rare throughout these EOB sections. Not ob- by an axial cross bearing tall, hollow, near-parallel-sided
served in other Tanzanian sections. Reported literature range spines. The coccoliths have not been unequivocally observed
is from NP21 to NP23 (Perch-Nielsen 1985). in the light microscope but are often abundant on rock sur-
faces observed in SEM.
Nannoliths incertae sedis DIFFERENTIATION. Rather similar to the simple murolith
Pemma? uncinata Bown & Dunkley Jones, 2006 coccolith of the early Mesozoic but also shares some char-
(Pl. 14, figs 11-17) acters with extant narrow-rimmed muroliths with affinities
to the Papposphaeraceae (Young et al. 2003), i.e. a small,
murolith coccolith with a hollow, square spine, protruding
DESCRIPTION. Large (typically >7 μm), triangular-, from a central-area cross formed of four simple crystal units.
rectangular- or rhomb-shaped nannoliths, which are No evidence of coccolith dimorphism.
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thickened along two sides ('base' and 'side') to give a

hook-shaped appearance, with a faintly birefringent 'back' ;
c-axis orientation is parallel with the long axis. SEM
imaging shows they are formed of layered rods/lamellae,
which thicken to produce the higher birefringence base and
side. Generic designation is probably incorrect, however it
is premature to propose a new generic name for this form
until its nature is better understood.
OCCURRENCE. Rare to frequent throughout these EOB sec-
tions. Not observed in other Tanzanian sections.
DIMENSIONS. Murolith base L 1.1 μm, W 0.9 μm; spine
length 4.0-6.0 μm.
TYPE MATERIAL. Holotype, Pl. 15, fig. 2. Paratype, Pl. 15,
fig. 3. TDP Site 12, Pande, Tanzania, upper Eocene, Sample
TDP12/23-2, 79 cm (Zone NP21).
OCCURRENCE. Commonly observed in SEM studies of these
EOB sections; has also been observed in sediments from TDP
Sites 2, 12, 13, 14, 16b (NP9-21).

Genus GLADIOLITHUS Jordan & Chamberlain, 1993

Genus TROCHOASTER Klumpp, 1953
DIAGNOSIS. Dimorphic deep photic zone extant coccolitho-
DIAGNOSIS. Non-birefringent, hexaradiate nannoliths with
phore with basal elliptical plate-like coccoliths formed of
thin radiating rays and multiple pores. two elements (lepidoliths) and long hexagonal-section tube
coccoliths.
Trochoastersimplex Klumpp, 1953 (Pl. 10, fig. 29)
Gladiolithus flabellatus (Halldal & Markali, 1955)
DESCRIPTION. Hexagonal non-birefringent nannolith. Six Jordan & Chamberlain, 1993 (Pl. 16, figs 1-6)
thin rays extend a short distance beyond the margin of the
hexagonal disc with numerous pores. Secondary shorter rays DESCRIPTION. Gladiolithus with oval lepidoliths formed
are present between each of the longer rays, which terminate from two equidimensional elements joined along a transverse
at the disc edge. suture. The tube coccoliths are narrow, elongate and free of
external ornament. The lepidoliths are very small (<2 μm),
REMARKS. This form may be a holococcolith but, as yet,
smooth on both sides, often observed separating into their
there is no clear microstructural evidence for this.
constituent elements, and are indistinguishable from those of
OCCURRENCE. Very rare in the upper Eocene of these EOB the living species, G. flabellatus (see Pl. 16, fig. 1). The tube
sections. Ranges from NP9b to NP21 in the Tanzanian sec- coccoliths are long, narrow and gently tapering, and formed
tions. First described from the upper Eocene of Germany from five or six elongate elements. The elongate elements
(Klumpp 1953). gently taper, and are narrowest at the proximal end where
they terminate bluntly, and widest at the distal end where
they have triangular terminations. They are smooth on the
Genus POCILLITHUS gen. nov. inner and outer surfaces but typically have fine serrations
TYPE SPECIES. Pocillithus spinulifer sp. nov. along one of the long edges. The lith morphologies, dimen-
sions and coccosphere shape do not differ significantly from
DERIVATION OF NAME. Frompocillum, meaning 'little cup', modern specimens of G. flabellatus.
referring to murolith morphology of these coccoliths.
OCCURRENCE. Rare to frequent, in SEM studies of rock
DIAGNOSIS. Miniscule to small murolith coccoliths that chip surfaces from the late Eocene of TDP12 (NP19/20-21).
have tall narrow spines in the type species Poncillithus spin- Ranges from NP6 to NP23 in the Tanzanian sections (Bown
ulifer that extend from a small basal disc. The spines have a et al. 2009). Reported literature range is from late Quaternary
narrow axial canal and are usually seen in side view. to Recent (Okada & Matsuoka 1996; Lars Legge et al. 2006).
 406 T O M D U N K L E Y J O N E S ET AL.
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Plate 15 SEM images of incertae sedis nannolith Pocillithus spinulifer sp. nov. Figs 1,5 TDP12/26-2,62 cm, Fig. 2 (holotype arrowed)
TDP12/23-2, 79 cm, Figs 3 (paratype) &4TDP12/47-2,64cm. All images are to the same scale.

Gladiolithus brevis Bown et al., 2009 (Pl. 16, figs 3 & but also observed in middle Eocene sediments of Tanzania
6) (NP15).

DESCRIPTION. Gladiolithus with short, unornamented tube-
coccoliths with blunt distal terminations. This species is de-
scribed in Bown et al. (2009).
OCCURRENCE. Rare to frequent, in SEM studies of rock
chip surfaces from the late Eocene of TDP12 (NP19/20-21)
Gladiolithus contus Bown et al., 2009 (Pl. 16, figs 5 &
6)
DESCRIPTION. Gladiolithus with very long, narrow tube-
coccoliths that appear to taper at both ends. Edges of tube
 UPPER EOCENE TO LOWER OLIGOCÈNE CALCAREOUS NANNOFOSSILS FROM TANZANIA 407
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Plate 16 SEM images of incertae sedis nannolith Gladiolithus. Fig. 1 Gladiolithus flabellatus extant, plankton sample from Hawaii Ocean Time
Series station at 180 m, May 2005, courtesy of Jeremy Young. Figs 2,4 Gladiolithus flabellatus TDP12/26-2,62 cm. Fig. 3 Gladiolithus flabellatus
and G. brevis TDP12/26-2,62 cm. Fig. 5 Gladiolithus flabellatus and G. contus TDP12/26-2,62 cm. Fig. 6 Gladiolithus flabellatus, G. brevis and G.
contus TDP12/26-2,62 cm. All images are to the same scale.
 408 T O M D U N K L E Y J O N E S ET AL.
elements may show prominent serrations. This species is de-
scribed in Bown et al. (2009).
OCCURRENCE. Rare to frequent, in SEM studies of rock
chip surfaces from the late Eocene of TDP12 (NP19/20-21)
but also observed in upper Paleocene sediments of Tanzania
(NP9).
REFERENCES
Andruleit,H. A., Rogalla, U. & Stäger, S. 2004. From living communit-
ies to fossil assemblages: origin and fate of coccolithophores in the
northern Arabian Sea. Pp. 5-21 in M. Triantaphylou (ed.) Advances
in the Biology, Ecology and Taxonomy of Extant Calcareous Nan-
noplankton. Micropaleontology Press, American Museum of Natural
History, New York.
Aubry, M. P. 1988. Handbook of Cenozoic Calcareous Nannoplankton,
Book 2: Ortholithae (Catinasters, Ceratoliths, Rhabdoliths). Micropa-
Downloaded by [University of Nebraska, Lincoln] at 23:04 13 July 2013
leontology Press, American Museum of Natural History, New York.
— 1999. Handbook of Cenozoic Calcareous Nannoplankton, Book 5: He-
liolithae (Zygoliths and Rhabdoliths). Micropaleontology Press, Amer-
ican Museum of Natural History, New York.
Backman, J. & Hermelin, J. O. R. 1986. Morphometry of the Eocene
nannofossil Reticulofenestra umbilicus lineage and its biochronolo-
gical consequences. Palaeogeography Palaeoclimatology Palaeoeco-
logy 57: 103-116.
Berggren, W. A. & Pearson, P. N. 2005. A revised tropical to subtropical
Paleogene planktonic foraminiferal zonation. Journal of Foraminiferal
Research 35: 279-298.
Billard, C. 1994. Life cycles. Pp. 167-186 in J. C. Green & B. S. C.
Leadbeater (eds) The Haptophyte Algae. Clarendon Press, Oxford.
Black, M. 1971. The systematics of coccoliths in relation to the palaeon-
tological record. Pp. 611-624 in B. M. Funnel & W. R. Riedel (eds)
The Micropaleontology of the Oceans. Cambridge University Press,
Cambridge.
— 1973. British Lower Cretaceous Coccoliths. I - Gault Clay (Part 2).
Palaeontological Society of London (Monograph) 127: 49-112.
Boudreaux, J. E. & Hay, W. W. 1969. Calcareous nannoplankton and
biostratigraphy of the late Pliocene-Pleistocene-Recent sediments in
the Submarex cores. Revista Española de Micropaleontología 1: 249-
292.
Bown, P. R. 2005a. Paleogene calcareous nannofossils from the Kilwa
and Lindi areas of coastal Tanzania (Tanzania Drilling Project 2003-
4). Journal of Nannoplankton Research 27: 21-95.
— 2005b. Calcareous nannoplankton evolution: a tale of two oceans.
Micropaleontology 51(4): 299-308.
— & Dunkley Jones, T. 2006. New Paleogene calcareous nannofossil
taxa from coastal Tanzania: Tanzania Drilling Project Sites 11 to 14.
Journal of Nannoplankton Research 28: 17-34.
— & Young, J. R. 1998. Techniques. Pp. 16-28 in P. R. Bown (ed.)
Calcareous Nannofossil Biostratigraphy. Kluwer Academic, London.
—, Rutledge, D. C., Crux, J. A. & Gallagher, L. T. 1998. Lower
Cretaceous. Pp. 86-131 in P. R. Bown (ed.) Calcareous Nannofossil
Biostratigraphy. Kluwer Academic, London.
—, Lees, J. A. & Young, J. R. 2004. Calcareous nannoplankton evolution
and diversity through time. Pp. 481-508 in H. R. Thierstein & J. R.
Young (eds) Coccolithophores: From Molecular Processes to Global
Impact. Springer, Berlin.
—, Dunkley Jones, T. & Young, J. R. 2007. Umbilicosphaera jordanii
Bown, 2005 from the Paleogene of Tanzania: confirmation of generic
assignment and a Palaeocene origination for the family Calcidiscaceae.
Journal of Nannoplankton Research 29: 25-30.
—, —, Lees, J. A., Randell, R., Mizzi, J., Pearson, P. N., Coxall, H. K.,
Young, J. R., Nicholas, C. J., Karega, A., Singano, J & Wade, B. S.
2008. The Paleogene Kilwa Group of coastal Tanzania: a calcareous
microfossil Konservat-Lagerstätte. Geological Society of America Bul-
letin 120: 3-12.
—, —, Young, J. R. & Randell, R. 2009. A Paleogene record of extant
lower photic zone calcareous nannoplankton. Palaeontology 52: 457-
469.
Bralower, T. J. 2005. Paleocene-early Oligocene calcareous nan-
nofossil biostratigraphy, ODP Leg 198 Sites 1209, 1210, and
1211 (Shatsky Rise, Pacific Ocean). Proceedings of the Ocean
Drilling Program, Scientific Results 198. Available at: http://www-
odp.tamu.edu/publications/198_SR/115/1 15.htm.
— & Mutterlose, J. 1995. Calcareous nannofossil biostratigraphy of Site
865, Allison Guyot, Central Pacific Ocean: a tropical Paleogene ref-
erence section. Proceedings of the Ocean Drilling Program, Scientific
Results 143: 31-74.
Bramlette, M. N. & Riedel, W. R. 1954. Stratigraphic value of discoast-
ers and some other microfossils related to Recent coccolithophores.
Journal of Paleontology 28: 385-403.
— & Sullivan, F. R. 1961. Coccolithophorids and related nannoplank-
ton of the early Tertiary in California. Micropaleontology 7: 129-
188.
— & Wilcoxon, J. A. 1967. Middle Tertiary calcareous nannoplankton
of the Cipero section, Trinidad, W.I. Tulane Studies in Geology and
Paleontology 5: 93-131.
Briggs, D. E. G. 2001. Exceptionally preserved fossils. Pp. 328-332 in
D. E. G. Briggs & P. R. Crowther (eds) Palaeobiology. Blackwell
Publishing, Oxford.
Brönnimann, P. & Stradner, H. 1960. Die Foraminiferen und Discoaster-
idenzonen von Kuba und ihre interkontinentale Korrelation. Erdoel-Z
76: 364-369.
Bukry, D. 1971. Cenozoic calcareous nannofossils from the Pacific
Ocean. San Diego Society of Natural History, Transactions 16: 303-
327.
— & Bramlette, M. N. 1968. Stratigraphic significance of two genera
of Tertiary calcareous nannofossils. Tulane Studies in Geology and
Paleontology 6: 149-155.
— & — 1971. Validation of Pedinocyclus and Guinquerhabdus new cal-
careous nannoplankton genera. Tulane Studies in Geology and Pale-
ontology 8: 122.
— & Percival, S. F. 1971. New Tertiary calcareous nannofossils. Tulane
Studies in Geology and Paleontology 8: 123-146.
Cavalier-Smith, T. 1986. The Kingdom Chromista: origins and system-
atics. Progress in Phycological Research 4: 309-344.
Conway Morris, S. 1990. Burgess Shale. Pp. 270-274 in D. E. G. Briggs
& P. R. Crowther (eds) Palaeobiology. Blackwell, Oxford.
Coxall, H. K., Wilson, P. A., Palike, H., Lear, C. H. & Backman, J.
2005. Rapid stepwise onset of Antarctic glaciation and deeper calcite
compensation in the Pacific Ocean. Nature 433: 53-57.
Cros, L., Kleijne, A., Zeltner, A., Billard, C. & Young, J. R. 2000.
New examples of holococcolith-heterococcolith combination cocco-
spheres and their implications for coccolithophorid biology. Marine
Micropaleontology 39: 1-34.
Deflandre, G. 1947. Braarudosphaera nov. gen., type d'une famille nou-
velle de Coccolithophoridés actuels à éléments composites. Comptes
Rendus Hebdomadaires des Séances de l'Académie des Sciences, Paris
225: 439-441.
— 1952. Classe des Coccolithopridés (Coccolithophoridae Lohmann,
1902). Pp. 439-470 in P-P. Grassé (ed.) Traité de Zoologie. Ana-
tomie, Systématique, Biologie, 1, part 1, Phylogenie. Protozoaires:
Généralités. Flagellés. Masson and Cie, Paris.
— 1959. Sur les nannofossiles calcaires et leur systématique. Revue de
Micropaleontologie 2: 127-152.
— & Fert, C. 1954. Observations sur les Coccolithophoridés actuels
et fossiles en microscopie ordinaire et éléctronique.´ Annales de
Paléontologie 40: 115-176.
de Kaenel, E. & Villa, G. 1996. Oligocene-Miocene calcareous nanno-
fossil biostratigraphy and paleoecology from the Iberia Abyssal Plain.
Proceedings of the Ocean Drilling Program, Scientific Results 149:
79-145.
De Vargas, C., Aubry, M. P., Probert, I. & Young, J. R. 2007. Origin and
evolution of coccolithophores: from coastal hunters to oceanic farmers.
Pp. 251-287 in P. G. Falkowski & A. H. Knoll (eds) Evolution of
 UPPER EOCENE TO LOWER OLIGOCÈNE CALCAREOUS NANNOFOSSILS FROM TANZANIA
Primary Producers in the Sea. Elsevier Academic Press, Amsterdam,
Boston.
Dockery III, D. T., Stover, C. W., Weathersby, P., Stover Jr, W. C.
& Ingram, S. L. 1991. A continuous core through the undifferenti-
ated Yazoo Clay (late Eocene, Jackson Group) of Central Mississippi.
Mississippi Geology 12: 23-27.
Dunkley Jones, T. & Bown, P. R. 2007. Post-sampling dissolution and
the consistency of nannofossil diversity measures: a case study from
freshly cored sediments of coastal Tanzania. Marine Micropaleonto-
logy 62: 254-268.
—, —, Pearson, P. N., Wade, B. S., Coxall, H. K. & Lear, C. H.
2008. Major shifts in calcareous phytoplankton assemblages through
the Eocene-Oligocene transition of Tanzania and their implications
for low-latitude primary production, Paleoceanography, 23 PA4204.
doi:10.1029/2008PA001640.
Edvardsen, B., Eikrem, W., Green, J. C., Andersen, R. A., Yeo
Moon-Van Der Staay, S. & Medlin, L. K. 2000. Phylogenetic re-
constructions of the Haptophyta inferred from 18S ribosomal DNA
sequences and available morphological data. Phycologia 39: 19-
35.
Downloaded by [University of Nebraska, Lincoln] at 23:04 13 July 2013
Fujiwara, S., Tsuzuki, M., Kawachi, M., Minaka, N. & Inouye, I.
2001. Molecular phylogeny of the haptophyta based on the rbcL gene
and sequence variation in the spacer region of the RUBISCO operon.
Journal of Phycology 37: 121-129.
Gaarder, K. R. & Heimdal, B. R. 1977. A revision of the genus Syra-
cosphaera Lohmann (Coccolithineae). "Meteor" Forschungsergebn-
isse. Reihe D, Biologie 24: 54-71.
Gartner, S. J. 1969. Two new calcareous nannofossils from the Gulf
Coast Eocene. Micropaleontology 15: 31-34.
— 1970. Phylogenetic lineages in the lower Tertiary coccolith genus Chi-
asmolithus. North American Paleontology Convention Sept. 1969, Pro-
ceedings G: 930-957.
— 1971. Calcareous nannofossils from the JOIDES Blake Plateau cores
and revision of the Paleogene nannofossil zonation. Tulane Studies in
Geology and Paleontology 8: 101-121.
— & Bukry, D. 1969. Tertiary holococcoliths. Journal of Paleontology
43: 1213-1221.
Geisen, M., Billard, C., Broerse, A. T. C., Cros, L., Probert, I. & Young,
J. R. 2002. Life-cycle associations involving pairs of holococcolitho-
phorid species: intraspecific variation or cryptic speciation? European
Journal of Phycology 37: 531-550.
Gran, H. H. 1912. Pelagic plant life. Pp. 307-386 in J. Murray & J. Hjort
(eds) The Depths of the Ocean. Macmillan, London.
— & Braarud, T. 1935. A quantitative study of the phytoplankton in
the Bay of Fundy and the Gulf of Maine (including observations on
hydrography, chemistry and turbidity). Journal of the Biological Board
of Canada 1: 279-467.
Haeckel, E. 1894. Systematische Phylogenie der Protisten und Pflanzen.
Reimer, Berlin, 400 pp.
Halldal, P. & Markali, J. 1955. Electron microscope studies on cocco-
lithophorids from the Norwegian Sea, the Gulf Stream and the Mediter-
ranean. Avhandlinger utgitt av det Norske Videnskapsakademi i Oslo.
Mat. - Naturvid. Klasse 1: 1-30.
Haq, B. U. 1966. Electron microscope studies on some upper Eocene cal-
careous nannoplankton from Syria. Stockholm Contributions in Geo-
logy 15: 23-37.
— 1968. Studies on upper Eocene calcareous nannoplankton from
NW Germany. Stockholm Contributions in Geology 18: 13-
74.
— 1971. Paleogene calcareous nannoflora. Parts I-IV. Stockholm Contri-
butions in Geology 25: 1-158.
Hay, W. W. 1977. Calcareous nannofossils. Pp. 1055-1200 in A. T.
S. Ramsay (ed.) Oceanic Micropalaeontology 2. Academic Press,
London.
— & Towe, K. M. 1962. Electron microscope examination of some cocco-
liths from Donzacq (France). Eclogae geologicae Helvetiae 55: 497-
517.
—, Mohler, H. & Wade, M. E. 1966. Calcareous nannofossils from
Nal'chik. Eclogae geologicae Helvetiae 59: 379-399.
409
—, —, Roth, P. H., Schmidt, R. R. & Boudreaux, J. E. 1967. Cal-
careous nannoplankton zonation of the Cenozoic of the Gulf Coast and
Caribbean-Antillean area, and transoceanic correlation. Transactions
of the Gulf Coast Association of Geological Societies 17: 428-480.
Hibberd, D. J. 1976. The ultrastructure and taxonomy of the Chryso-
phyceae and Prymnesiophyceae (Haptophyceae): a survey with some
new observations on the ultrastructure of the Chrysophyceae. Botan-
ical Journal of the Linnaen Society 72: 55-80.
Honjo, S. 1976. Coccoliths: production, transportation and sedimentation.
Marine Micropaleontology 1: 65-79.
Jafar, S. A. & Martini, E. 1975. On the validity of the calcareous nan-
noplankton genus Helicosphaera. Senckenbergiana Lethaea 56: 381-
397.
Jerkovic, L. 1970. Noelaerhabdus nov. gen. type d'une nouvelle famille de
Coccolithophorides fossiles. Noelaerhabdaceae du Miocène supérieur
de Yougoslavie. Comptes Rendus (Hebdomadaires des Séances) de
l'Académie des Sciences, Paris 270: 468-470.
Jordan, R. W. & Chamberlain, A. H. L. 1993. Vexillarius cancellifer
gen. et sp. nov. and its possible affinities with other living coccolitho-
phorids. Pp. 305-325 in B. Hamrsmid & J. R. Young (eds) INA 1991.
Nannoplankton Research, Biostratigraphy and Paleobiology. Quatern-
ary Coccoliths. MND Hodonin, Hodonin.
— & — 1997. Biodiversity among haptophyte algae. Biodiversity and
Conservation 6: 131-152.
—, Cros, L. & Young, J. R. 2004. A revised classification scheme for
living haptophytes. Micropaleontology 50: 55-79.
Kamptner, E. 1927. Beitrag zur Kenntnis adriatischer Coccolithophor-
iden. Archiv für Protistenkunde 58: 173-184.
— 1948. Coccolithen aus dem Torton des Inneralpinen Wiener Beckens.
Sitzungberichte der Osterreichischen Akademie der Wissenschaften.
Mathematisch-Naturwissenschaftliche Klasse 87: 152-158.
— 1950. Uber den submikroskopischen Aufbau der Cocco-
lithen. Akademie der Wissenschaften in Wien, Mathematisch-
Naturwissenschaftliche Klasse 87: 152-158.
— 1954. Untersuchungen uber den Feinbau der Coccolithen. Archiv für
Protistenkunde 100: 1-90.
— 1963. Coccolithineen-Skelettreste aus Tiefseeablagerungen des Pazi-
fischen Ozeans. Naturhistorischen Museum in Wien, Annalen 71:117-
198.
Kleijne, A. 1991. Holococcolithophorids from the Indian Ocean, Red Sea,
Mediterranean Sea and North Atlantic Ocean. Marine Micropaleonto-
logy 17: 1-76.
— 1992. Extant Rhabdosphaeraceae (coccolithophorids, class Prymne-
siophyceae) from the Indian Ocean, Red Sea, Mediterranean Sea and
North Atlantic Ocean. Scripta Geologica 100: 1-63.
Klumpp, B. 1953. Beitrag zur Kenntnis der Mikrofossilien des Mittleren
und Oberen Eozän. Palaeontographica 103A: 377-406.
Lars Legge, H., Mutterlose, J. & Arz, H. W. 2006. Climatic changes in
the northern Red Sea during the last 22,000 years as recorded by cal-
careous nannofossils. Paleoceanography doi:10.1029/2005PA001 142.
Lear, C. H., Bailey, T. R., Pearson, P. N., Coxall, H. K. & Rosenthal, Y.
2008. Cooling and ice growth across the Eocene-Oligocene transition.
Geology 36: 251-254.
Lees, J. A., Bown, P. R., Young, J. R. & Riding, J. B. 2004. Evidence
for annual records of phytoplankton productivity in the Kimmeridge
Clay Formation coccolith stone bands (Upper Jurassic, Dorset, UK).
Marine Micropaleontology 52: 29-49.
Lemmermann, E. 1908. Flagellatae, Chlorophyceae, Coccosphaerales
und Silicoflagellatae. Pp. 1-40 in K. Brandt & C. Apstein (eds) Nor-
disches Plankton. Botanischer Teil. Lipsius and Tischer, Kiel and
Leipzig.
Levin, H. L. 1965. Coccolithophoridae and related microfossils from the
Yazoo formation (Eocene) of Mississippi. Journal of Paleontology 39:
265-272.
— & Joerger, A. P. 1967. Calcareous nannoplankton from the Tertiary of
Alabama. Micropaleontology 13: 163-182.
Liu, L. 1996. Eocene calcareous nannofossils from the Iberia Abyssal
Plain. Proceedings of the Ocean Drilling Program, Scientific Results
149: 61-78.
 410 T O M D U N K LE Y J O N E S ET AL.
Loeblich, A. R. & Tappan, H. 1978. The coccolithophorid genus Cal-
cidiscus Kamptner and its synonyms. Journal of Paleontology 52:
1390-1392.
Lohmann, H. 1902. Die Coccolithophoridae, eine Monographie der Coc-
colithen bildenden Flagellaten, zugleich ein Beitrat zur Kenntnis des
Mittelmeerauftriebs. Archiv für Protistenkunde 1: 89-165.
— 1903. Neue Untersuchungen über den Reichthum des Meeres an Plank-
ton und über die Brauchbarkeit der verschiedenen Fangmethoden.
Zugleich auch ein Beitrag zur Kenntnis des Mittelmeerauftriebs. Wis-
senschaftliche Meeresuntersuchungen Abteilung Kiel 7: 1-87.
Lunt, P. & Sugiatno, H. in press. A review of the Eocene and Oligocene
in the Nanggulan area, south central Java. Bulletin of the Indonesian
Geological Survey, Bandung.
Marino, M. & Flores, J. A. 2002. Middle Eocene to early Oligocene
calcareous nannofossil stratigraphy at Leg 177 Site 1090. Marine Mi-
cropaleontology 45: 383-398.
Martini, E. 1958. Discoasteriden und verwandte Formen im NW-
deutschen Eozän (Coccolithophorida). 1. Taxonomische Untersuchun-
gen. Senckenbergiana Lethaea 39: 353-388.
— 1959. Pemma angulatum und Micrantholithus basquensis, zwei neue
Downloaded by [University of Nebraska, Lincoln] at 23:04 13 July 2013
Coccolithophoriden-Arten aus dem Eozän. Senckenbergiana Lethaea
40: 415-21.
— 1971. Standard Tertiary and Quaternary calcareous nannoplankton zon-
ation. Pp. 739-785 in A. Farinacci (ed.) Proceedings of the Second
Planktonic Conference, Roma, 1970. Edizioni Tecnoscienza, Rome.
— & Ritzkowski, S. 1968. Die Grenze Eozän/Oligozän in der Typus-
Region des Unteroligozäns (Helmstedt-Egeln-Latdorf). ¨Memoires du
Bureau de Recherches Geologiques et Minieres 69: 233-237.
Müller, C. 1970. Nannoplankton-Zonen der Unteren-Meeresmolasse
Bayerns. Geologica Bavarica 63: 107-118.
Murray, G. & Blackman, V. H. 1898. On the nature of the Coccospheres
and Rhabdospheres. Philosophical Transactions of the Royal Society
of London (B) 190: 427-441.
Nicholas, C. J., Pearson, P. N., Bown, P. R., Dunkley Jones, T., Huber,
B. T., Karega, A., Lees, J. A., McMillan, I. K., O'Halloran, A.,
Singano, J. M. & Wade, B. S. 2006. Stratigraphy and sedimentology
of the Upper Cretaceous to Paleogene Kilwa Group, southern coastal
Tanzania. Journal of African Earth Sciences 45: 431-466.
Noël, M.-H., Kawachi, M. & Inouye, I. 2004. Induced dimorphic life
cycle of a coccolithophorid, Calyptrosphaera sphaeroidea (Prymne-
siophyceae, Haptophyta). Journal of Phycology 40: 112-129.
Norris, R. E. 1984. Indian Ocean nannoplankton. I. Rhabdosphaeraceae
(Prymnesiophy ceae) with a review of extant taxa. Journal of Phycology
20:27-41.
Okada, H. 1980. Calcareous Nannofossils from Deep Sea Drilling Project
Sites 442 through 446, Philippine Sea. Initial Reports of the Deep Sea
Drilling Project 58: 549-565.
— 1983. Modern nannofossil assemblages in sediments of coastal and
marginal seas along the western Pacific Ocean. Utrecht Micropaleon-
tological Bulletins 30: 171-187.
— 1992. Biogeographic control on modern nannofossil assemblages in
surface sediments of Ise Bay, Mikawa Bay and Kumano-Nada, off
coast of central Japan. Memorie di Scienze Geologich XLIII: 431-
449.
— & Honjo, S. 1973. The distribution of oceanic coccolithophorids in the
Pacific. Deep-Sea Research 20: 355-374.
— & Matsuoka, M. 1996. Lower-photic nannoflora as an indicator of the
late Quaternary monsoonal paleo-record in the tropical Indian Ocean.
Pp. 231-245 in A. Moguilevsky & R. Whatley (eds) Microfossils
and Oceanic Environments. University of Wales, Aberystwyth-Press,
Aberystwyth.
Pagani, M., Zachos, J. C., Freeman, K. H., Tipple, B. & Bohaty, S.
2005. Marked decline in atmospheric carbon dioxide concentrations
during the Paleogene. Science 309: 600-603.
Pascher, A. 1910. Chrysomonaden aus dem Herschberger Grossteiche.
Monographien und Abhandlungen zur Internationale Revue der
gasamten Hydrobiologie und Hydrographie 1: 1-66.
Pearson, P. N. & Palmer, M. R. 2000. Atmospheric carbon dioxide
concentrations over the past 60 million years. Nature 406: 695-699.
—, Ditchfield, P. W., Singano, J., Harcourt-Brown, K. G., Nicholas,
C. J., Olsson, R. K., Shackleton, N. J. & Hall, M. A. 2001. Warm
tropical sea surface temperatures in the late Cretaceous and Eocene
epochs. Nature 413: 481-487.
—, Nicholas, C. J., Singano, J., Bown, P. R., Coxall, H. K., van Dongen,
B. E., Huber, B. T., Karega, A., Lees, J. A., Msaky, E., Pancost,
R. D., Pearson, M. & Roberts, A. P. 2004. Paleogene and Cretaceous
sediment cores from the Kilwa and Lindi areas of coastal Tanzania:
Tanzania Drilling Project Sites 1-5. Journal of African Earth Sciences
39: 25-62.
—, —, —, —, —, —, —, —, —, MacLeod, K., McMillan, I. K.,
Pancost, R. D., Pearson, M. & Msaky, E. 2006a. Further Paleo-
gene and Cretaceous sediment cores from the Kilwa area of coastal
Tanzania: Tanzania Drilling Project Sites 6-10. Journal of African
Earth Sciences 45: 279-317.
—, Olsson R. K., Huber B. T., Hemleben C. & Berggren W. A. (eds)
2006b. Atlas of Eocene Planktonic Foraminifera. Cushman Foundation
for Foraminiferal Research, Special Publication No. 41, 513 pp.
—, van Dongen, B. E., Nicholas, C. J., Pancost, R., Schouten, S.,
Singano, J. & Wade, B. S. 2007. Stable warm tropical climate through
the Eocene Epoch. Geology 35: 211-214.
—, McMillan, I. K., Wade, B. S., Dunkley Jones, T., Coxall, H. K.,
Bown, P. R. & Lear, C. H. 2008. Extinction and environmental change
across the Eocene-Oligocene boundary in Tanzania. Geology 36: 179-
182.
Perch-Nielsen, K. 1968. Der Feinbau und die klassifikation der Cocco-
lithen aus dem Maastrichtien von Dänemark. Det Kongelige Danske
Videnskabernes Selskab Biologiske Skrifter 16: 1-93.
— 1971. Elektronenmikroskopische Untersuchungen an Coccolithen und
verwandten Formen aus dem Eozan van Dänemark. Det Kongelige
Danske Videnskabernes Selskab Biologiske Skrifter 18: 1-76.
— 1977. Albian to Pleistocene calcareous nannofossils from the Western
South Atlantic, DSDP Leg 39. Initial Reports of the Deep Sea Drilling
Project 39: 699-823.
— 1985. Cenozoic Calcareous Nannofossils. Pp. 329-554 in H. M. Bolli,
J. B. Saunders & K. Perch-Nielsen (eds) Plankton Stratigraphy. Cam-
bridge University Press, Cambridge.
Poche, F. 1913. Das System der Protozoa. Archiv für Protistenkunde 30:
125-321.
Prins, B. 1979. Notes on nannology - 1. Clausicoccus, a new genus of
fossil coccolithophorids. INA Newsletter 1: N2-N4.
Probert, I., Fresnel, J., Billard, C., Geisen, M. & Young, J. R. 2007.
Light and electron microscope observations of Algirosphaera robusta
(Prymnesiophyceae). Journal of Phycology 43: 319-322.
Quintero-Torres, R., Aragón, J. L., Torres, M., Estrada, M. & Cros,
L. 2006. Strong far-field coherent scattering of untraviolet radiation
by holococcolithophores. Physical Review 74. doi: 10.1103/Phys-
RevE.74.032901.
Romein, A. J. T. 1979. Lineages in early Paleogene calcareous
nannoplankton. Utrecht Micropaleontological Bulletins 22: 1-
230.
Roth, P. H. 1970. Oligocene calcareous nannoplankton biostratigraphy.
Eclogae Geologicae Helvetiae 63: 799-881.
Sáez, A. G., Probert, I., Geisen, M., Quinn, P., Young, J. R. & Medlin,
L. K. 2003. Pseudo-cryptic speciation in coccolithophores. Proceed-
ings of the National Academy of Sciences of the United States of
America 100: 7163-7168.
—, —, Young, J. R., Edvardsen, B., Eikrem, W. & Medlin, L. K.
2004. A review of the phylogeny of the Haptophyta. Pp. 251-270 in
H. R. Thierstein & J. R. Young (eds) Coccolithophores: from Molecular
Processes to Global Impact. Springer, Berlin.
Schiller, J. 1913. Vorlaufige Ergebnisse der Phtyoplankton-
Untersuchungen auf den Fahrten SMS 'Najade' in der Adria
1911/12. I. Die Coccolithophoriden. Sitzungberichte der Akademie
der Wissenschaften in Wien, Mathematisch-Naturwissenschaftliche
Klasse 122: 597-617.
—1930. Coccolithineae. Pp. 89-267 in L. Rabenhorst (ed.) Kryptogamen-
Flora von Deutschland, Osterrich und der Schweiz 10. Band, 2 Abt.
Akademische Verlagsgesellschaft, Leipzig.
 UPPER EOCENE TO LOWER OLIGOCÈNE CALCAREOUS NANNOFOSSILS FROM TANZANIA
Schlauder, J. 1945. Recherches sur les flagellés calcaires de la Baie
d'Alger. Université d'Alger, Algiers.
Schwarz, E. H. L. 1894. Coccoliths. Annals and Magazine of Natural
History, series 6 14: 341-346.
Schwarz, E. 1932. Beitrage zur Entwicklungsgeschichte der Protophyten.
IX. Der Formwechsel von Ochrosphaera neapolitana. Archiv für Pro-
tistenkunde 77: 434-462.
Shafik, S. 1989. Some new calcareous nannofossils from upper Eocene
and lower Oligocene sediments in the Otway Basin, southeastern Aus-
tralia. Alcheringa 13: 69-83.
Sherwood, R. W. 1974. Calcareous nannofossil systematics, paleoeco-
logy, and biostratigraphy of the middle Eocene Weches Formation of
Texas. Tulane Studies in Geology and Paleontology 11: 1-79.
Siesser, W. G. 1983. Paleogene Calcareous Nannoplankton Biostrati-
graphy: Mississippi, Alabama and Tennessee. Bulletin of the Missis-
sippi Department of Natural Resources 125:1-61.
Sprengel, C., Baumann, K. H. & Neuer, S. 2000. Seasonal and inter-
annual variation of coccolithophore fluxes and species composition in
sediment traps north of Gran Canaria (29°N 15°W). Marine Micropa-
leontology 39: 157-178.
Downloaded by [University of Nebraska, Lincoln] at 23:04 13 July 2013
Stradner, H. 1962. Über neue und wenig bekannte Nannofossilien aus
Kreide und Alttertiär. Sonderabdruck aus den Verhandlungen der Geo-
logischen Bundesanstalt 2: 363-377.
— 1969. The nannofossils of the Eocene flysch in the Hagenbach Valley
(northern Vienna Woods), Austria. Rocznik Polskiego Towarzystwa
Geologicznego 39: 493-432.
— & Edwards, A. R. 1968. Electron microscope studies on upper Eocene
coccoliths from the Oamaru Diatomite, New Zealand. Jahrbuch der
Geologischen Bundesanstalt Wien, Special Volume 13: 1-66.
Takahashi, K. & Okada, H. 2000. Environmental control on the biogeo-
graphy of modern coccolithophores in the southeastern Indian Ocean
offshore of Western Australia. Marine Micropaleontology 39: 73-86.
Takano, Y., Hagino, K., Tanaka, Y., Horiguchi, T. & Okada,
H. 2006. Phylogenetic affinities of an enigmatic nannoplankton
Braarudosphaera bigelowii based on the SSU rDNA sequences. Mar-
ine Micropaleontology 60: 145-156.
Tan, S. H. 1927. Discoasteridae incertae sedis. Koninklijke Akademie van
Wetenschappen te Amsterdam Proceedings 30: 411-419.
Theodoridis, S. 1984. Calcareous nannofossil biostratigraphy of the Mio-
cene and revision of the helicoliths and discoasters. Utrecht Micropa-
leontological Bulletin 32: 1-271.
Troelsen, J. C. & Quadros, L. P. 1971. Distribuição biostratigráfîca
dos nanofósseis em sedimentos marinhos (Aptian-Mioceno) do Brasil.
Annaes da Academia Brasileira de Ciencias 43: 577-609.
van Dongen, B. E., Talbot, H. M., Schouten, S., Pearson, P. N. &
Pancost, R. D. 2006. Well preserved Palaeogene and Cretaceous bio-
markers from the Kilwa area, Tanzania. Organic Geochemistry 37:
539-557.
Varol, O. 1989. Eocene calcareous nannofossils from Sile (northwest
Turkey). Revista Española de Micropalaeontología 21: 273-320.
— 1991. New Cretaceous and Tertiary calcareous nannofossils. Neues
Jahrbuch für Geologie und Paläontologie. Abhandlungen 182: 211-
237.
411
— 1992. Sullivania a new genus of Paleogene coccoliths. Journal of
Micropaleontology 11: 141-150.
— 1998. Paleogene. Pp. 200-224 in P. R. Bown (ed.) Calcareous Nanno-
fossil Biostratigraphy. Kluwer Academic, London.
Viohl, G. 1990. Solnhofen Lithographic Limestones. Pp. 285-289 in D. E.
G. Briggs & P. R. Crowther (eds) Palaeobiology. Blackwell, Oxford.
Wade, B. S. & Pearson, P. N. 2008. Planktonic foraminiferal turnover,
diversity fluctuations and geochemical signals across the Eocene/
Oligocene boundary in Tanzania. Marine Micropaleontology. 68:244-
255.
Wallich, G. C. 1877. Observations on the coccosphere. Annals and
Magazine of Natural History 19: 342-350.
Wise, S. W. 1973. Calcareous nannofossils from cores recovered during
Leg 18, Deep Sea Drilling Project: biostratigraphy and observations
on diagenesis. Initial Reports of the Deep Sea Drilling Project 18:
569-615.
— 1983. Mesozoic and Cenozoic calcareous nannofossils recovered by
Deep Sea Drilling Project Leg 71 in the Falkland Plateau Region,
Southwest Atlantic Ocean. Initial Reports of the Deep Sea Drilling
Project 71: 481-550.
Young, J. R. 1998. Neogene. Pp. 225-265 in P. R. Bown (ed.) Calcareous
Nannofossil Biostratigraphy. Kluwer Academic, London.
— & Bown, P. R. 1997. Cenozoic calcareous nannoplankton classifica-
tion. Journal of Nannoplankton Research 19: 36-47.
—, Didymus, J. M., Bown, P. R., Prins, B. & Mann, S. 1992. Crystal
assembly and phylogenetic evolution in heterococcoliths. Nature 356:
516-518.
—, 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 termin-
ology. Palaeontology 40: 875-912.
—, Davis, S. A., Bown, P. R. & Mann, S. 1999. Coccolith ultrastruc-
ture and biomineralisation. Journal of Structural Biology 126: 195-
215.
—, Geisen, M., Cros, L., Kleijne, A., Sprengel, C., Probert, I. &
Ostergaard, K. K. 2003. A guide to extant coccolithophore taxonomy.
Journal of Nannoplankton Research, Special Issue 1: 1-125.
—, Henriksen, K. & Probert, I. 2004. Structure and morphogenesis of the
coccoliths of the CODENET species. Pp. 191-216 in H. R. Thierstein
& J. R. Young (eds) Coccolithophores: from molecular process to
global impact. Springer, Berlin.
—, Geisen, M. & Probert, I. 2005. A review of selected aspects of coc-
colithophore biology with implications for paleodiversity estimation.
Micropaleontology 51: 267-288.
Zachos, J. C. & Kump, L. R. 2005. Carbon cycle feedbacks and the
initiation of Antarctic glaciation in the earliest Oligocene. Global and
Planetary Change 47: 51-66.
—, Pagani, M., Sloan, L., Thomas, E. & Billups, K. 2001. Trends,
rhythms, and aberrations in global climate 65 Ma to present. Science
292: 686-693.
Ziveri, P., Broerse, A. T. C., van Hinte, J. E., Westbroek, P. & Honjo,
S. 2000. The fate of coccoliths at 48°N 21°W, Northeastern Atlantic.
Deep-Sea Research 47: 1853-1875.