A history and application of larger

foraminifera in Indonesian biostratigraphy,

calibrated to isotopic dating.

A summary of the subject for the 2004 GRDC workshop on

micropalaeontology
By
Peter Lunt & Tony Allan

Cite as:
Lunt, P., and T. Allan, 2004. Larger foraminifera in Indonesian biostratigraphy, calibrated to isotopic dating. GRDC Museum Workshop
on Micropalaeontology, June 2004. 109 pp.

CONTENTS
1. INTRODUCTION 1
2. HISTORICAL DEVELOPMENT 2
FIGURE 6: THE SUMMARY RANGE DIAGRAM OF THE LARGER FORAMS 10
3. MODERN SYNTHESIS 12
PRE-TERTIARY 12
THE PALEOCENE AND EARLY EOCENE (T.A1 & T.A2) 12
THE REGIONAL TRANSGRESSION IN T.A3 17
THE T.A3 TO T.B BOUNDARY 20
TERTIARY B 21
TERTIARY C 23
THE TRANSITION FROM T.C TO T.D 25
THE TOP OF T.D 27
SUBDIVISION OF T.E 27
ON THE UPPER T.E TO T.F BOUNDARY 27
TERTIARY F 29
FAUNAL DEVELOPMENT IN LOWER T.F (T.F1&2) 30
THE TOP OF T.F2 (FAUNAL TURNOVER) 31
LETTER STAGES AFTER T.F2 31
THE LATEST CENOZOIC RADIATION OF LARGER FORAMINIFERA 31
4. CAUSES OF FAUNAL TURNOVER 38
5. NOTES ON THE MAJOR TAXA (BY GENUS) 40
Image 1. The Late Cretaceous Barune Sandstone of eastern Papua 13
Image 2. The Jatibungkus Limestone, Paleocene C. Java 13
Image 3. The Jatibungkus Limestone, Paleocene C. Java 14
Image 4. Four images of “Nummulites nuttalli" and “N. thalicus”, Mankalihat, Kalimantan 15
Image 5. Tertiary a2? Limestone from Kali Sana, C. Java. Nummulites crasseornatus. 16
Image 6. Tertiary a3 Limestone from Halmahera 17
Image 7. Tertiary a3 Limestone with Planocamerinoides (=Assilina) 18
Image 8. Tertiary a3 Limestone with Alveolina 18
Image 9. Tertiary a3 Limestone from Karangsambung, C. Java, also N.javanus details 19
Image 10. Tertiary a3 Limestone with early Pellatispira 20
Image 11. Early part of Tertiary b Limestone from West Java. 21
Image 12. Tertiary b Limestone from Nanggulan. 22
Image 13. Tertiary b Limestone from Pasir E. Kalimantan. 23
Image 14. Tertiary c Limestone from SW Java 24
Image 15. Tertiary d Limestone from West Papua 24
Image 16. Tertiary e1 Limestone 25
Image 17. Tertiary e2-3 Limestone 26
Image 18. Tertiary e4 Limestone, Prupuh Lst, NE Java 28
Image 19. Tertiary e4 Limestone, Rajamandala Lst, C. Java 28
Image 20. Tertiary e5 (Upper T.e) Limestone, top of Prupuh Lst, type location 29
Image 21. Lower Tertiary f Limestone - T.f1 mid Early Miocene 30
Image 22. Lower Tertiary f Limestone - T.f2 base Middle Miocene, Geger, Madura 32
Image 23. Lower Tertiary f Limestone - T.f2 mid Middle Miocene, Jatiluhur, W. Java 33
Image 24. Upper Tertiary f Limestone - T.f3 intra Late Miocene 35
Image 25. Latest Miocene to Recent. Karren Lst. NE Java 36
Image 25. Latest Miocene to Recent. Un-named Lst near Semarang 37
1. INTRODUCTION
The biostratigraphic ranges of the larger foraminifera need to be brought into line with the standard Tertiary
time scale developed through the geo-magnetic polarity time scale (GPTS; see Cande & Kent, 1992, 1995) and the
planktonic zonal integration of Berggren et al. (1995). This has not yet been done because in their shallow carbonate
habitat there are few planktonic foraminifera or nannofossils to correlate with the open marine biostratigraphic zones,
and even if plankton is present, the indurated nature of the limestones makes such work very difficult. A secondary
problem is that shallow marine habitats are much more prone to hiatuses and periods of erosion from sea-level fall
and are usually an imperfect stratigraphic record. However, larger foraminifera remain an important fossil group for
the dating and environmental analysis of tropical limestone, and enough new data is now available to offer a zonal
calibration that extends the standard work of Adams (1970, 1984).
This review deals mostly with genera rather than species. However, even at generic level there are complications
that anyone using larger foraminifera must be aware of:
1. Larger foraminifera are a polyphyletic group with a common preference for tropical, shallow marine, photic, carbon-
ate environments. Such environments are made stable by a critically low nutrient level (i.e. they are oligotrophic).
Other environmental factors may fluctuate, but severe lack of nutrients is always the limiting condition. Hence the
environment, for the fauna and flora, is stable. This stability excerpts a strong evolutionary pressure for hetero-
morphic* modification and in particular what are classified as peramorphic descendants, that is; specialised adult
morphologies are naturally selected, with the result that ancestral morphologies – preserved as a growth stage in
the shells or tests of descendants – are progressively lost in a diminishing juvenile stage. This environmentally
driven evolutionary pressure produces very similar trends in distantly related lineages. If the juvenile stage of
growth is greatly diminished there may be no remaining clue as to the ancestor of a form. (e.g. the similar but
unrelated Discocyclina and Eulepidina)
As noted by many workers (cf. Haynes, 1984) there are five main characters that most larger foraminifera tend to
acquire;
a). A tendency to large size.
b). A tendency to increased numbers of chambers or chamberlets.
c). A tendency to acquire “lateral chamberlets” or cubiculae.
d). A tendency to radial symmetry.
e). A tendency to reduce the juvenile life / test stage, and to increase not only the proportion but the duration
of the adult stage.
2. The heterochronic nature of evolution produces orthogenetic [apparently directed] morphological trends lasting
many millions of years, so consequently larger forams have excellent examples of both gradual evolution, as well
as punctuated evolutionary events. Generally the punctuated morphological changes are used to define genera,
while the gradual trends may be used to define species (as in Miogypsinids and Nephrolepidina).
3. As with other types of foraminifera, there is repeated, or iterative, appearance of the same morphological forms or
features, which may be the “playing out” of the same genes at different geological times. Such iterative evolution
may cause the defining character of a genus to have more than one evolutionary record and consequently more
than one extinction. This is well documented in the genera Tansinhokella and Planostegina.
Biostratigraphy of Indo-Pacific larger forams: page 1

These evolutionary complications, plus the difficulty of identifying fossils in random thin section has resulted in a
taxonomy for the larger foraminifera that has over a century of prior work to consider, but which is still only at a basic
level in systematically naming the forms. It is hoped that soon we can identify a fairly complete set of evolutionary
characters, distinct from ecophenotypes, and such a catalogue will greatly assist the study of carbonate sediments by
assigning both an accurate age and environment of deposition.

* Heteromorphic evolution is the natural selection of characters related to the life stages of an organism. Natural selection may favour longer
juvenile or adult stages, or a change in the onset of reproductive maturity, or a combination of such changes.
2. HISTORY OF THE LETTER STAGES
(Most of the following historical summary first appeared in a 1998
IPA field guide)
The basic zones for larger foraminifera have long been
known as the Letter Stages, and while many have pointed
out that they are not true Stages but biozones, very few
have advocated replacing these popular and orderly-named
zones with cumbersome fossil-zonal names. For the most
part the application of these Letter Stages has been con-
stant over the last fifty years (with the exception of the
upper T.f). Also the Letter Stages appear to represent as-
semblages between faunal turnovers so it makes sense to
conserve them, and consider them as being of a similar
scale to “Eocene”, “Miocene” and “Pliocene’ etc., Epochs
first defined on fossil content but now used as names for
Series of Stages.
The Letter Stages were originally defined on asso-
ciations of certain microfossils; that is, the contents of
the biozones are defined. This contrasts with planktonic
microfossil zones that are defined by their boundaries,
which are single species evolution or extinction datums.
Ideally planktonic zonal datums should correlate well and
approximate time-lines. However in larger foraminiferal
carbonate sequences there is a good chance that Letter
Stage boundaries are delimited by facies changes, there-
fore correlation lines or the concept of “datum planes” are
considered inappropriate for the larger foram biozones.
THE EPOCHS OF LIFE
The term Tertiary first appeared in the eighteenth cen-
tury, in a four-fold stratigraphic subdivision comprising
Primary, Secondary, Tertiary, and Quaternary units. The
first referred to crystalline rocks and the second to hard
stratified rocks with some fossil content. The third, Terti-
ary, unit was recognised as hardly indurated sediments
that were obviously not part of young soils or alluvium.
The similar term Cainozoic (or Cenozoic; from the Greek
kainos = new and zoon = life) was defined in the mid nine-
teenth Century on broad biostratigraphic principles, being
the sediments (and their fossils) deposited in the period
after the end-Cretaceous mass extinction. It includes both
the Tertiary and Quaternary.
Just prior to the definition of the Cainozoic, Lyell
(1833) divided the Tertiary of Europe into the Eocene,
Miocene, Older Pliocene and Newer Pliocene (the latter
pair revised in 1839 into Pliocene and Pleistocene). As
with the Cainozoic these were biostratigraphy-based units,
as reflected in the Greek roots of the names, viz.: Eocene
= “dawn of the new”, Miocene = “less new”, Pliocene =
* from Palaeo + Eocene, = ancient Eocene. Combining these parts, begining and ending with vowels, gives "Pal-eocene" from "Palaeo-eocene", and not
"Pala-eocene". The similar word "Palaeogene" is a normal combination ["Palaeo-gene"], which involves no modification as there is no juxtapostion of
vowels. This follows the English convention that the Latin diphthong ai, from palaio, is transliterated to ae in palaeo, in much the same way that oe becomes
u in the Indonesian spelling of "Bandung", being the best representation of the pronunciation. American usage adopts "e" to represent the ai diphthong;
hence their "Pal-eocene" and "Paleo-gene". Choice depends on whether one pronounces oneself a "pal-ay-ontologist" or a "pal-ee-ontologist"
“more new”, Pleistocene = “most new”. Lyell defined his
Eocene as strata with less than 5% of mollusc fossils still
living, the Miocene as strata with about 17% extant forms,
and the Pliocene as strata with 35-50% extant forms. (The
Paleocene* and Oligocene were added in 1874 and 1854
respectively as workers discovered significant sections of
sediments at the base of the Eocene and between Lyell’s
Eocene and Miocene.)
It is worth noting that at about the time Lyell defined
these ages, W.B. Carpenter had just published his micro-
scope studies of Nummulites and other genera (1850) and
clarified the structure of these fossils, although they were
then placed in the Cephalopoda and thereby considered
a form of mollusc (but not used in Lyell’s percentage
method).
By the end of the 19th Century the first synthesis of
European stratigraphy, at Epoch or Series level, was vir-
tually complete and work was then focused on applying
these Epochs in the relative dating and correlation of the
lithostratigraphic stages defined on various stratotypes
across Europe.
One of the many over-lapping terms proposed for
the older Tertiary was the Nummulitique of Emile Haug,
proposed in 1902 for the Palaeogene (Palaeogene itself
introduced in 1853 but only becoming a practical term
in 1874 when it could cover a combined Paleocene and
Eocene). The term Nummulitique survived for some fifty
years and is directly comparable to the more diversely
fossiliferous Indonesian Tertiary a through d. This wider
use of the term Nummulitic was one of the reasons that
the name Nummulites was adopted in preference to the
original (senior) term Camerina.
SUBDIVISIONS OF THE TERTIARY IN SE ASIA
In Indonesia, where the Tertiary is a more significant
part of the sedimentary geological record than it is in Eu-
rope, the 19th and early 20th Century workers attempted to
utilise the new stratigraphic schemes. Professor K. Martin
(1852-1944), studying the mollusc faunas of Java and sur-
rounding areas was particularly outstanding in this field.
His work was able to follow that of Lyell in Europe in uti-
lising the ratios of extant to extinct faunas to indicate rela-
Biostratigraphy of Indo-Pacific larger forams: page 2
tive ages. Two major contributions to Tertiary stratigraphy
came from the work of Martin. Firstly he argued that the
mollusc faunas of the East Indies developed separately
from those of Europe and secondly, as a consequence of
this, the stage names of Europe could not be correlated
with confidence to the Indonesian region.
The birth of the Letter Stages Eames, 1975) arrived at the conclusion that “even in the
Martin's view that the Indonesian area could not be Oligocene [separated from the top Eocene after the time
correlated with the European stratigraphy gradually won of Lyell’s first work], Recent species never exceed 1 per
acceptance between the first publication of Martin on the cent” of fossil forms.
famous Eocene section at Nanggulan in 1914, and the
comprehensive review of Indonesian geology by Rutten While mollusc studies had been important in under-
in 1927. This break with European stratigraphy was the standing the stratigraphy of the Indonesian area, and
reason the Survey of the Dutch Indies began to set up specialists like Martin could reliably arrange faunas in
their own stratigraphic system, and there is fair reason to stratigraphic order based on ratios of extant species, it was
acknowledge the Nanggulan section as the place of origin the larger foraminifera that were to be the foundation of
of the Letter Stages. There is, however, a twist to this tale the new stratigraphic system. By the late 1920s they had
that is not widely known. become the preferred fossil group for biostratigraphy, pri-
The original Nanggulan samples were collected by a marily as they were more abundant than molluscs, but also
coal prospector, van Dijk in about 1872, but these were a they did not require extensive knowledge of large numbers
mixed bag from the Eocene inlier and a Miocene forma- of living and fossil species. A scheme was developed that
tion (Jonggrangan Fm.) that unconformably overlies the utilised assemblage zones rather than percentages of extant
older beds. Consequently the first work by Martin, used forms. The larger foraminifera assemblage zones could be
by Verbeek and Fennema (1896) in their review of Java identified by the presence of a few key taxa, often with a
and Madura, arrived at a value of 28% extant species. hand-lens in the field.
This was considered a typical Miocene ratio, but with the In 1927 Van der Vlerk and Umbgrove published the
old Tertiary foraminifera Discocyclina and Nummulites Letter Classification of the Indonesian Tertiary, based on
present, a compromise age of Oligocene age was assigned larger foraminifera. This scheme subdivided the Tertiary
to the Nanggulan beds. In 1920 Martin did his own field into seven parts. Six parts were labeled “a” through “f”
work and began a number of detailed descriptions of the (e.g. Tertiary “a”, or “T.a” for short), and a seventh section
exceptionally well preserved fauna of the site (106 gastro- of non-orbitoidal Tertiary between Tf and the Quaternary
pods, 23 bivalves, 3 scaphopods and 4 foraminifera, - the was noted. This scheme was immediately successful and
most diverse old Tertiary mollusc fauna in the east Asian was rapidly adopted as a workable biostratigraphic scheme
region). However no species at Nanggulan were consid- for the the Far East, even if the correlation with the Eu-
ered extant, in contrast to the 5% of the Eocene molluscs ropean epochs and stages was not adequately known.
in Europe, plus it contained some genera not known from The authors pointed out that the taxonomy and detailed
Europe before the Oligocene. Martin
asked the expert on the Eocene of the

Quaternary
Distribution of larger foraminifera TERTIARY
Paris Basin, Cossman, to compare
in the "East Indian" Tertiary
the faunas (cf. Rutten 1927, chapter Van der Vlerk & Umbgrove, 1927 a b c d e f
VI and Oppenoorth & Gerth 1929).
Cossman confirmed Martin’s obser- Assilina
vations that the Nanggulan fauna Pellatispira
consists of forms closely related but Nummulites (Camerina)
not identical with those of the Paris Discocyclina (Orthophragmina)
Eocene. This observation “led Martin Alveolina
to the conclusion that the fauna of Flosculina*
Nanggulan belongs to the younger Heterostegina
Eocene, but that a common con- Lepidocyclina
nection between the European part
Lep. (megalospheric generation >7mm)
Biostratigraphy of Indo-Pacific larger forams: page 3

of the Tethys geosyncline and the

Spiroclypeus**
region now occupied by the Indian
Archipelago no longer existed.” (p.5 Miogypina***
Oppenoorth & Gerth). Trillina howchini
Alveolinella
It is ironic that this break from
A. bontangensis****
European stratigraphy may not
Cycloclypeus
?

have been necessary. Reviews of

the Eocene faunal ratios first de-
scribed by Lyell, summarised by
Morley Davies (1935, revised by
* This genus covers the heavily flosculinised species of the modern genus Alveolina
** In 1928 this would have covered both Spirocylpeus and Tansinhokella
*** would have covered both Miogypsina and Miogypsinoides.
**** Now Flosculinella bontangensis. Note also that Alveolina would include the
modern Borelis lineage and that "megalospheric Lepidocyclina" would be Eulepidina.
Figure 1. The original scheme of the Letter Stages.
Van der Vlerk and Umbgrove 1927.
biostratigraphy of the genera used was not fully under-
stood, and suggested that future work would increase the
number of recognisable subdivisions.
The original proposal of the Letter Stages was:
Tertiary a, the oldest larger foraminifera assemblage
known in the region, characterised by Nummu-
lites, Discocyclina, Pellatispira, “Alveolina” (i.e.
the forms that now fit under Alveolina s.s. [or Fas-
ciolites of some authors]) “Flosculina” (i.e. what
would now be regarded as heavily flosculinised
species of Alveolina), and Assilina.
Tertiary b faunas also contain Nummulites, Discocyclina,
Pellatispira and “Alveolina” (which in this case
would be the plexus that later became known as
Clausulus, then Neoalveolina, but now under the
senior name, Borelis). This stage differs from the
Tertiary a by the absence of Assilina and “Floscu-
lina”.
Tertiary c contains Nummulites but Discocyclina, and
Pellatispira are absent.
Tertiary d still contains Nummulites, overlapping with the
first appearance of the lepidocyclinid forms. Van der
Vlerk and Umbgrove divided their lepidocyclinid
forms into two groups. “Lepidocyclina” and “Lepi-
docyclina megalospheric generation, tests >7mm)”.
These taxonomic concepts equate to Lepidocyclina
(isolepidine, nephrolepine and related forms) and
Eulepidina respectively. These “megalospheric gen-
eration >7 mm” forms are noted to have a limited
stratigraphic range that is identical to Spiroclypeus.
It is surprising that the term Eulepidina was not
used as it had already been proposed in 1911 by
Douvillé working in the Philippines.
Tertiary e is identified by the “Lepidocyclina megalospher-
ic generation” (=Eulepidina) and Spiroclypeus men-
tioned above, as well as by Miogypsina, a term that
in 1927 included both the modern Miogypsina and
Miogypsinoides, and by “Trillina howchini” (= the
whole genus Austrotrillina at that time).
Tertiary f is characterised by Alveolinella - which then
included “A.” bontangensis (= the modern Floscu-
linella bontangensis). The forms Eulepidina and
Spiroclypeus were absent from this stage as was
“Alveolina” (Borelis forms).
Shortly after this, in 1929, the 4th Pacific Science Con-
gress was held in Bandung at the newly opened building
of the Mijnbouw or Mining Bureau. The director of the
Survey, de Jongh, in the foreword to the first mapping
report on Java (Koolhoven 1933), noted that this important
conference was an incentive to beginning the systematic
mapping of Java, not least so as to have well documented
field trips for the Congress participants. It was already
known that Java was richer in microfossils than Sumatra
where mapping efforts had been focused, but Java also
appeared to have more facies variation, and close co-ordi-
nation between field geologists and palaeontologists was
considered crucial. Beginning in 1928, this work, together
with similar efforts in Borneo, found many of the sections
used to develop the Letter Stages. It was also this mapping
work that correlated, then replaced, the field geologists "e
1 to 3" and "n 1 to 5" mapping stratigraphy with Letter
Stages. The "e" and "n" annotation however continued as
map symbols until the 1940's, and modern readers of these
maps appear to have forgotten the annotation is shorthand
for larger foram and mollusc biostratigraphy.
The correlation is;
e1 = contains Assilina (=T.a)
e2 = No Assilina, but Pellatispira pressent (=T.b)
e3 = lower part contains Nummulites fichteli-intemedi-
us, and an upper part has the Lepidocyclines, the
top being based on the presence of Spiroclypeus
s.l. (inc. Tansinhokella); T.c to basal T.e. Effec-
tively most of the Oligocene, and it was called
this by the field geologists.
n1 = Most of T.e, on Spiroclypeus and Miogypsi-
noides.
n2 = T.f, with Alveolinella, C. annulatus, Lepidosemi-
cyclina, mid Early to mid Middle Miocene, both
Lower and Upper T.f in the original scheme.
n3 = Lepidocyclina of the trybliolepidine type and
molluscs 20-45% extant, later Middle and Late
Miocene (Upper Tf of modern usage)
n4 = considered lower Pliocene with 45-60% extant
molluscs, the trochid gastropod Solariella am-
blygoniata possibly being diagnostic.
n5 = later Pliocene and younger based on beds with
either mammal fossils, or the molluscs Natica
sulcifera, Turris gendingganensis, Ringicula
arctatoides and Pecten tjaringinensis.
This field geologist's scheme was devised by workers
who were both geologists and capable palaeontologists
such as Tobler and Rutten. The "e" and "n" notation com-
ing from the terms “Eogenic” and “Neogenic”, where
the former is equivalent to the Nummulitique of Haug
in Europe
Biostratigraphy of Indo-Pacific larger forams: page 4
In 1931 Leupold and van der Vlerk published a detailed
revision of the Letter Classification. This new version had
sixteen pre-Quaternary divisions or subdivisions to replace
the previous seven. The primary divisions were a through
to h and for the first time were were named Stages although
they are much closer in principle to the European Epochs
or Series. Subdivisions, such as T.a1, T.a2 etc., were called
zones. It is significant to note that the new T.g and T.h
units were not defined on larger foraminifera and are not
detailed in the text of this paper. The accompanying figure
however noted the percentages of extant molluscs for the
T.g, T.h1 and T.h2 divisions.
 Figure 2
The revised Letter-classification proposed by Leupold & Van der Vlerk, 1931
TERTIARY STAGES a b c d e f g h

Quaternary
Zones 1 2 123 45 1 2 3 1 2

20
30
35
45
50
60
18
Percentage of Mollusc species still living

8
NUMMULITES (Camerina)
N. nuttalli
N. javana var B
N. djokdjokartae
N. pengaronensis (=nanggoelani)
N. bagelensis
N. fichteli-intermedius
ASSILINA
PELLATISPIRA
HETEROSTEGINA
H. borneensis
SPIROCLYPEUS
S. pleurocentralis
S. tidoenganensis
S. leupoldi, margaritatus
CYCLOCLYPEUS
DISCOCYCLINA
LEPIDOCYCLINA
Isolepidina
Eulepidina
E. papuaensis
E. dilatata
Pliolepidina
Nephrolepidina
N. isolepidinoides
N. borneensis
N. angulosa
N. inflata / ferreroi
N. verbeeki
N. sumatrensis
Trybliolepidina
T. rutteni
Lepidocyclina -microspheric specimens
L. flexuosa
L. acuta
MIOGYPSINA
Miogypsina without lateral chambers
Biostratigraphy of Indo-Pacific larger forams: page 5

M. dehartii
Miogypsina with lateral chambers
M. thecideaeformis
M. polymorpha
FASCIOLITES [= Alveolina]
F. wichmanni [= Lacazinella wichmani]
Clausulus pygmaeus [= Borelis pygmaeus]
ALVEOLINA [= Alveolinella & Flosculinella]
. A. bontangensis [= F. bontangensis]
. A. boscii [= A. quoyi]
TRILLINA HOWCHINI
 This 1931 paper marked the peak of the Letter Clas-
sification. Over the next twenty years the scheme was
periodically modified, with a tendency to lump the 1931
“zones” rather than split them. By 1955 Van der Vlerk (cf.
Rutten in Van Bemmelen, 1949; and van der Vlerk 1955)
had reduced the number of pre-Quaternary divisions to
eight. Tertiary a and b, as well as T.e1 to T.e4 had been
reduced to single units after criticism from Tan (1939) and
others, as were T.f2 and T.f3 and T.g and T.h.
This apparent failure to develop the zones to a high
Figure 3
The revised Letter-classification at the time of Van Bemmelen (1949),
based on Rutten (1947)
TERTIARY STAGES a/b
resolution zonation is due to the inability to consistently
recogise individual species' evolution and extinction da-
tums in a shallow marine, hiatus-rich and facies controlled
sedimentary setting.
Nethertheless for more than a decade after Leupold &
van der Vlerk's 1931 paper the larger foraminifera were
the premier biostratigraphic tool in the Dutch East Indies,
and were also being used more elsewhere. However in mid
1942 the Japanese Imperial forces took over the Mijnbouw
and field-work virtually ceased. This was the end of the
efforts of the Opsporingsdient or exploration department
of the Minng Bureau, but not by the main researchers,
c d e 1-4 e5 f1f2,3 g / h

Quaternary
Approximate % of living mollusc species � � �� � � ��� �� � ���
Approximate correlation with Europe Eocene Oligocene Miocene Pliocene
NUMMULITES (Camerina)
N. javanus = N. perforatus
N.djokdjokartae, N. vredenburgi
N. fichteli-intermedius
ASSILINA
PELLATISPIRA
BIPLANISPIRA
HETEROSTEGINA
H. borneensis
SPIROCLYPEUS
S. vermicularis
other species of Spiroclypeus
CYCLOCLYPEUS
C. koolhoveni
Approximate ranges C. oppenorthi
according to Tan, 1932 C. eidae
C. indopacificus
CYCLOCLYPEUS (Katacycloclypeus)
ALVEOLINA
Flosculina [= flosculinised Alveolina spp]
Neoalveolina pygmaea [= Borelis pygmaeus]
FLOSCULINELLA
ALVEOLINELLA
Biostratigraphy of Indo-Pacific larger forams: page 6

AUSTROTRILLINA
LEPIDOCYCLINA
L. (Polylepidina)
L. (Lepdiocyclina)
L. (Nephrolepidina)
L. (Trybliolepidina)
L. (Eulepidina)
L. (Multilepidina)
MIOGYPSINA
M. (Miogypsinoides)
M. (Miogypsina)
DISCOCYCLINA
 Figure 4. The BPM zonation,
From Brouwer 1957 (published 1966)
The Letter classification of the Tertiary, after van

Tertiary g
to Recent
der Vlerk & Umbgrove, 1927, and Leupold & van
der Vlerk, 1931, modified by Brouwer. At the time
e 2/3

f 1

f 2

f 3
e 5
e 4
e 1

of his report the following taxonomic conventions

were adopted:
Austrotrillina howchini 1. Austrotrillina was considered a monospecific
Cycloclypeus posteidae genus so the A. howchini range given here is
equivalent to all modern species of this genus.
Cycloclypeus indopacificus
2. Miogypsina s.l. included Miogypsinoides,
Cycloclypeus cf. guembelianus
Miogypsina s.s. is added from the definitions in
Flosculinella bontangensis Muhar 1957.
Flosculinella borneensis 3. Miogypsinoides “praeubaghsi” is now referred
Heterostegina borneensis to as Neorotalia mecatepecensis and the inter-
Lepidocyclina (Eulepidina)
mediate forms from this species that grade into
Miogypsinoides.
Lepidocyclina s.l.
4. M. ubaghsi is the primitive Miogypsinoides
Miogypsina s.l. (i.e. all Miogypsinids) with a short fan of gerontic (later adult stage Mio-
Miogypsina s.s. (added from Muhar 1957) gypsinoides) chamberlets
Miogypsinoides "praeubaghsi" 5. M. complanata is a simple Miogypsinoides
Miogypsinoides ubaghsi form with a fuller fan and less pronounced juve-
nile whorl, thin lateral walls and therefore in large
Miogypsinoides complanata
part overlapping with the concept of Miogypsinel-
Miolepidocyclina excentrica la of some authors. Miogypsinoides dehaarti
Spiroclypeus & Tansinhokella was used by the BPM for the thicker, lammelar
walled form, which is this scheme is included in
“Miogypsina s.l.”
Note: Flosculinella borneensis is generally regarded as synonymous with Alveolinella praequoyi or A. fennemai, the
early form of A. quoyi with incomplete development of multiple attic tiers of chamberlets.

who returned to academic posts in the Nederlands to write
up their work. A great loss were the premature deaths of
Tan Sin Hok in 1945 and Rutten in 1946. After this time
geological work in Indonesia passed into the hands of the
new Indonesian geological survey and the BPM [Bataaf-
sche Petroleum Maatschappij] and related oil exploration
companies, who did not always publish their findings.
In Irian Jaya in the 1950's and 60's, the scheme used
by the NNGPM [Nederlandsche Nieuw Guinea Petroleum
Maatschappij] was based on an internal report by Mohler
which did not differ much from Rutten (1947) and van der
Vlerk (1948), except to extend the range of Austrotrillina
higher as the particular facies preferred by this genus is
found much more often in younger rocks in east Indonesia.
The quality of material in this area allowed NNGPM to
he carefully observed the mid Oligocene to Early Miocene
of Java and the evolution of the Neorotalia to Miogypsina
lineage, and the development of Spiroclypeus s.l. from
Heterostegina borneensis. This allowed subdivision of
the T.e Stage by selecting the best evolutionary events
in the complex range chart of Leupold and van der Vlerk
(1931).
In 1962 Eames, Banner, Blow, and Clark reviewed
the basic principles of mid-Tertiary stratigraphy in a
landmark contribution, making considerable advances in
sorting out the complex stage and biozonal systems from
Eocene through to the Miocene, and correlating most
of this with a planktonic stratigraphic framework (the
planktonic review only covered sediments up to the mid
Early Miocene). This work included a world-wide review
Biostratigraphy of Indo-Pacific larger forams: page 7

re-separate T.a from T.b. The NNGPM team also started
integrating early planktonic foram data with larger foram
ranges, but this was hampered by the rather indurated
limestone facies that dominated Irian Jaya through much
of the Tertiary, plus having only the late 1950’s rudiments
of the developing science of planktonic biostratigraphy.
In western Indonesia most oil was being found in the
Miocene and it is this section that was most studied by the
BPM. In 1957 Muhar completed an internal report that
was never published, but nethertheless was an important
contribution on the Oligo-Miocene section. In this work
of mid Tertiary larger foraminifera and a consideration
to the Far East Letter Stage system. Two unconformities
were suggested for the Letter Stages; first between T.a1
and T.a2, with no Early Eocene represented in the Letter
Stage system. Secondly an unconformity was noted in
mid-Oligocene times.
In 1970 the Letter Classification was reviewed by C.G.
Adams who extended the breadth of coverage of larger
foram genera and added notes on some planktonic zonal
datums. In 1984 this same author detailed the Neogene
with additional planktonic zonal control and clarification
 Figure 5
The Far East Letter-classification
correlated to European Stages by Eames et al. 1962
Time scale biased to current dating of stratigraphic units

Pliocene and younger not considered by Eames et al.

1&2
Ta1

Ta2

Tf3
Tb

Te

Tg
Td
Tc
TERTIARY STAGES

Tf
Eocene Oligoc. Miocene
SERIES Middle Upper Lower Mid. Upper

Burdigalian
Paleocene

Auversian

Sarmatian
Vindobian
Rupelian
Bartonian

Lattorfian
No

Lutetian
Ypresian
& European Stages reported
Aquitanian

Distichoplax (algae)
Miscellanea
Ranikothalia
Discocyclina s.l.
Nummulites
N. obtusus-perforatus
N. acutus - djokdjokartae
Alveolina oblonga
Alveolina
Assilina
N. pengaronensis-nanggoelani
N. fabianii
Biplanispira & Pellatispira
N. intermedius
Borelis pygmaea
Eulepidina
Spiroclypeus vermicularis
Spiroclypeus [Miocene spp.]
Lepidocyclina (Nephrolepidina)
Miogypsinella
Miogypsinoides
Miogypsina s.s.
Pliolepidina (= Multilepidina)
Austrotrillina howchini
Flosculinella reicheli
Flosculinella globulosa
Pseudotaberina malabarica
Flosculinella bontangensis
Alveolinella
Cycloclypeus annulatus
Marginopora
"Miogypsinella" of these authors are the forms of Miogypsinoides with distinctly thin lateral walls

of taxonomic concepts. on Lepidocyclinids and Miogypsinids up until 1937.

Biostratigraphy of Indo-Pacific larger forams: page 8

The application of biometric analysis in systematics
and biostratigraphy
In was probably the work by Tan Sin Hok in 1932,
on the genus Cycloclypeus, which highlighted the de-
tailed and gradual evolutionary changes in the complex
foraminifera over geological time. The changing charac-
ters could be counted or measured, and the values derived
used to define species with discrete ranges. In effect, this
allowed a measured specimen (almost always an average
of many specimens) to indicate a stage of evolution and
thereby an age. Tan Sin Hok continued with similar work
After the War period it was the Dutch school of work-
ers, primarily van der Vlerk, Drooger and MacGillavry,
who continued this theme of study. Van der Vlerk concen-
trated on Nephrolepdina - Lepidocyclina, such as a com-
parison of grades of evolution and planktonic zonations
in van der Vlerk and Postuma (1967). Drooger published
work on many genera but is best known for his work on
the Miogypsinids.
With the advent of modern ideas on punctuated equilib-
rium, after Gould and Eldridge (1977), many metaozoan
examples of “gradual” evolution were shown to be series
of punctuated steps with evidence for any gradual nature wide range of genera, in a single morphological trend over
being the product of a poor (macro)fossil record. Larger very long periods of time can only lead to the conclusion
foraminifera were one of the few remaining examples of that this is a fundamental aspect of larger foraminferal
gradual evolution, however the Dutch school was largely evolution and not a change in environmental morphocline.
ignored by American and English macro-palaeontologists. Drooger's comments on environment playing a dominant
The work of Ozawa (1975), however, on simple biomet- role in morphology, can be therefore be modified to en-
rics (the size of the proloculus) in Permian fusulinids vironment playing a minor role in the longer geological
was cited by Gould and Eldrige as a possible example of term.
gradual evolution. Rather than crystallising around this
example, and arguing that some life evolves gradually, One of the major gaps in larger foraminifera research
foraminiferologists left the debate, and it was left to Peter is to measure biometric character in association with
Sheldon, working on trilobites, to demonstrate the real- planktonic zonations and modern strontium dating. Van
ity of gradualism (ref). Drooger in his 1993 “farewell to der Vlerk & Postuma did this in 1967, and Raju and van
arms” summarises the position of the remaining Dutch Vessem contributed a little such work in their detailed
school at that time. He cites the results of a study on Re- studies on Miogypsinids and Lepidocyclinids respectively
cent Operculina ammonoides in the Gulf of Aqaba (Fer- (1974 and 1978).
mont et al. 1977), where there is a change in proloculus
size with depth comparable to the change observed by
Ozawa in Permian Lepidolina. He therefore believes that
this undermines any gradual morphometric changes as
merely indicating gradual changes in environment, even
over geological time. “Such a type of explanation would
be consistent with some (but not all) observed correla-
tions between morphoclines and depth gradients in radial
foraminifera” and "Whatever their value as mathematical
excercises on palaeontological collections, the results are
not really elucidating evolution; actually environmental
changes seem to play an important if not dominating role"
(p.21, Drooger, 1993). Drooger, his co-workers and stu-
dents publish very little on the stratigraphic distribution of
their material, and it is by applying a detailed stratigraphic
test that this negative view of gradual evolution can be
challenged. Observation shows that biometric characters
are highly reliable in determining stratigraphic position,
both in shallow carbonates and in allochthonous faunas in
calc-turbidites, where faunas are mixed from many depths
of origin (that is original depth ranges in the upper tens
of metres where larger forams lived). Also, the thirty mil-
lion years of evolution of Cycloclypeus noted by Tan Sin
Hok (1932) is hard to imagine as a massively prolonged,
one-way, gradual migration of the generic niche through
an environmental morphocline.
There are natural limits to the biometric technique
in biostratigraphy (cf. Drooger and Raju 1978) possibly
Biostratigraphy of Indo-Pacific larger forams: page 9

brought on by the practical limits in meaningful measure-

ments of such simple features, and maybe local effects
such as environmental morphoclines and reworking.
However, at a resolution of about a million years or more
observations of biometric characters can, repeatedly and
over a wide area, correctly predict the order of a series
of samples and correlate them to other biostratigraphic
schemes.
The features observed to evolve in a gradual manner
are invariably those linked to the heterochonous mode
of evolution (number of chambers in the juvenile stage
etc.). Such gradual heterochronous changes are seen in a
Figure 6. The Tertiary Letter Stages, as reviewed here
Lr.Te U.Te Lr. Tf U. Tf

SBZ10

SBZ12

SBZ13
SBZ14
SBZ15
SBZ16

SBZ17
SBZ18
Tb SBZ19
SBZ20
SBZ zones from Serra-Kiel et al 1998, from west Tethys

SBZ11
SBZ 9
SBZ 1

Quaternary
not yet calibrated to the Indo-Pacific

to

"Th"
Te2-3
Indo-Pacific

Tf3
Te5
Tf1
Td
Tc

Tf2
Te4
Te1
Province Only Ta1 Ta2 Ta3
Lt.
SERIES Pal Eocene Oligoc. Miocene Pl.
Early MIDDLE LATE EARLY LATE EARLY Mid. LATE
Amphistegina
Wilfordia sarawakensis
Sphaerogypsina
Alanlordia types
Discogypsina Discogypsina saipanensis
Linderina
Halkyardia
Dictyoconus
Others

"Dictyoconus" melinauensis Planorbulinella solida

Planorbulinella
Sporadotrema
Fabiania
Calcarina ? ?
Schlumbergerella
Baculogypsina
Quasirotalia
Neorotalia mecatepacensis ranges much older in C. American realm
Miogypsinoides
3-layered orbitoids

Miogypsina
Lepidosemicyclina
Discocyclina
Discocyclina omphalus
Asterocyclina
ranges much older in C. American realm
Lepidocyclina (Nephrolepidina) (as the ancestral Lepidocyclina)
Lepidocyclina (Trybliolepidina)
Lepidocyclina (Multilepidina)
L. (Nephrolepidina) ferreroi
Eulepidina ranges slightly older in C. American realm
Nummulites ...
N.: septa meandrine, very large size
often granulate ?*
N.: septa reticulate, generally small
N.: septa meandrine to sub-reticulate
Nummulites & related forms

Late Tertiary involute forms sometimes referred

Ranikothalia usually smooth to a super-genus concept of Nummulites, but
N.: septa radial or sigmoid better placed in Palaeonummulites.
Palaeonummulites Palaeonummulites / Operculina plexus
Operculina [& Planoperculina]
Spiroclypeus s.s.
Tansinhokella
Heterostegina (Vlerkina) ?
?
Heterostegina s.s. [& Heterocyclina]
Planostegina ? ?
may be polyphyletic
Cycloclypeus Large proloculus C. eidae
forms
C. with partings
C. annulatus in sidewalls
Grzybowskia

Planocamerinoides (ex. Assilina)

?
Pellatispira
Vacuolispira
Biplanispira
Orbitolites
Lacazinella
Alveolina
Borelis
Biostratigraphy of Indo-Pacific larger forams: page 10

F. globulosa F bontangensis
Flosculinella
"A. praequoyi"
Alveolinella
Miliolids

or "fennemai"

Fabularia ovata unpub. record

Austrotrillina T. Allan

Amphisorus
Sorites
Pseudotaberina malabarica
Marginopora
Peneroplis

in Papua New Guinea, in

Distichoplax biserialis [alga] same terrains as Pellatispira
Halimeda [alga]

?* = uncertainty stems from one report of reticulate = extinction or evolution diagnostic of zonal boundary
Nummulites with Tansinhokella.
Time scale based on Berggren et al 1995. Note that stage definitions are biased to planktonic biostratigraphic observations. Thus top Middle Eocene is based on top
P14 (c 38.4 Ma, Morozovella / Acarinina extinction, rather than the top Bartonian stage, c 1 ma younger), base Middle Miocene is based on the Orbulina datum
(15.1 Ma) not base Langhian at c 16.4 ma, top Middle Miocene is top of N14 at 11.4 Ma, not top Serravallian and top Miocene, on top of N17 /NN11 not top
Messinian.
 Martini Zones

Letter Stages
Figure 7 Summary of the Letter Stages

et al. Zones

Bukry Zones
Blow Zones
Magnetic pol.
calibrated to the GPTS of Cande and

Berggren
SERIES
Kent (1992/1995) and planktonic
MYBP

biostratigraphy of Berggren et al. (1995)

0 20/21
C1n
PLIOCENE PLEIST.

N 22

13-15
1 - N 23 PT 1

CN
C1r NN 19

PL 6
C2
Early (Z) Late (P/G)

originally
CN 12
N 19 - N 21

PL 5 NN
C2An PL 3-4
18-16

"T.h"
C2Ar PL 2 NN 14-15
CN 11
NN 13
5 C3n PL 1
C3r N 18 NN 12 CN 10 Top of Upper T.f: Extinction of last of the Lepidocyclines (Trybli-
Mess.

M 14
M. MIOC. LATE MIOCENE

(b)

C3An olepidina). A minor extinction during a period of gradual radiation

(b)
NN 11

C3Ar
N 17

CN 9

of larger foraminiferal types in shallow marine carbonate faunas

Upper T.f
C3B
(b)
(a)

C4n dominated by coral, green algae and molluscs.

(a)
Tortonian

C4r
M 13

NN 10 CN 8
C4A
N 16

(a)

10 NN 9 CN 7
C5n NN 8
Lower T.f to Upper T.f: (Originally the top of T.f of the early
N 15 M 12 CN 6
C5r N 14 M 11 NN 7 workers). Extinction of Miogypsina, but also Cycloclypeus annula-
Langh. Serravallian

N 13
C5An M 8-10 tus. Generally a reduction in the abundance of larger foraminifera
CN 5
NN 6

N 11-12
C5Ar
C5AB in shallow marine carbonates and an increase in green algae and
N 10

M7

C5AC corals. In Java this was the top of the old "OK" [Orbitoiden Kalk]
NN 5

CN 4

C5AD
unit with common larger forams in limestones. Appearance of
Lower T.f

15 C5Bn N9 M6

C5Br faunal turnover exaggerated in SE Sundaland due to major se-

N7 N8

M 4 M5

C5Cn quence boundary and widespread facies change at this time.

NN 4

CN 3

C5Cr
Burdigalian
EARLY MIOCENE

C5D Steady radiation in L. (Nephrolepidina) to L. (Trybiolepidina),

M3
N6

C5E NN 3
CN 2 Miogypsina, Cycloclypeus and Borelis to Flosculinella lineages
in lower Tf. Records of individual datums appear slightly dia-
C6
M2
N5

20
chronous due to facies control.
C6An
Aquitanian

T.e to T.f: Extinction datum of Eulepidina and Spiroclypeus /

NN 2

Upper T.e

C6Ar
Tansinhokella.
CN 1

C6AA
M1
N4

C6B
C6Cn NN 1 Radiation of Tansinhokella / Spiroclypeus, Miogypsinoides and L.
C6Cr T.e 4 (Nephrolepidina) lineages define sub-zones of T.e
Lt. OLIG.

C7
Chattian

T.e 2-3
P 22

P 22

NP 25

C7A
C8n
C8r
CP 19

C9n
Te1 T.d to T.e: extinction datum of Nummulites s.s. may be dia-
(a) (b)
P 21

P 21

C9r
NP 24

C10n
chronous and overlap much, if not all of T.e 1.
Lt. EOCENE E. OLIGOCENE

C10r
T.d

30 C11n P 20 P 20 Radiation from T.c through T.d in Planostegina - Heterostegina to

Rupelian

C11r
Cycloclypeus lineage. Slight biometric change in Nephrolepidina.
CP 18

C12n
NP 23
P 19

P 19

C12r CP 17 T.c to T.d: rapid and widespread influx of Central American forms
T.c

NP 22 Eulepidina, Nephrolepidina and Neorotalia mecatepacensis.

P 18

P 18

CP 16
NP 21

C13n
C13r T.b to T.c: very close to extinction of several groups of plank-
Priabonian

P 17 P 17
Biostratigraphy of Indo-Pacific larger forams: page 11

C15
P 16 P 16
19-20
tonic foraminifera at top P16/17. Some species of Nummulites,
CP 15

Discocyclina s.l. and Pellatispira / Biplanispira all become extinct.

T.b

C16
NP 18 Tansinhokella appears to become "temporarily extinct"
P 15

P 15

C17
Bartonian

T.a to T.b: very close to extinction of several groups of plank-

NP 17

(b)

tonic foraminifera at top P14. Highly dimorphic Nummulites with

P 14

P 14

C18n
40 complex sutural traces, most granulate species of Nummulites,
CP 14
MIDDLE EOCENE

P 13 P 13
C18r Alveolina species and Planocamerinoides (=Assilina of previous
T.a (T.a 3)
NP 16

C19 workers) all become extinct.

(a)
P 12

P 12

C20n
Extinction event.
Lutetian

Period of steady radiation in

P 11

P 11

C20r
NP 15

CP 13

larger foram morphologies

C21n
P 10

P 10

NP 14

CP 12

C21r
C22n
9

9
3. A MODERN SYNTHESIS
OF THE LETTER STAGES
After more than seventy years of use and adaptation, a
basic eight-fold division of the Tertiary of the Indo-Pacific
area remains valid. These units are Tertiary a to e, upper &
lower f, and post-f "stages". These biostratigraphic units
are comparable in principle and scale to the Paleocene,
Eocene, Oligocene etc., of Europe.
Within the Letter Stages there are often secondary
markers for upper, middle and lower parts offering age
determination to between one and two million years for
much of the Oligo-Miocene. This chapter summarises cur-
rent knowledge on the main units, the changes between
then and outstanding problems.
PRE-TERTIARY
In the pre-Tertiary, Permian larger foraminifera are
known from Thailand and neighboring areas (many refer-
ences), and rarely from East Indonesia (Kepala Burung
area).
In early to mid-Cretaceous times, the marine mar-
gin of Sundaland may be traced on the distribution of
bioclastic limestones containing the arenaceous larger
foram Orbitolina, which is reported from Kalimantan
and areas north and west (Hashimoto et al., 1975, and
encountered in oil-well samples), plus a few locations
in Central Java, namely float samples east of Yogyakarta
(Bothé 1929), and a few small carbonate lenses in the
Lukulo Melange (Harloff, 1935).
In Late Cretaceous times only three areas are known
with shallow marine, larger foram bearing facies; - in
Papua New Guinea not far from Port Moresby there is an
outcrop of limestones and sandstones with Campanian -
Maastrictian larger forams (Glaessner, 1960; see Image 1
re-sampled by T. Allan in 2001), in Misool and the western
Bird’s Head part of Irian Jaya there is a similar regres-
sive sedimentary suite of this age (Visser and Hermes,
1962, and Rusman, 1989) and in Luzon in the north of
the Philippines is a similar facies (Hashimoto et al., 1978,
see notes below on Palaeocene). These three occurrences
contain Lepidorbitoides and Pseudorbitoides faunas with
molluscs and bryozoa, sometimes overlying or adjacent to
Globotruncana facies, indicating regression or uplift.
THE PALEOCENE [T.A1] & EY. EOCENE [T.A2]
Paleocene and lower (Early to mid Middle) Eocene
faunas are very rare in Southeast Asia due to the regional
geological setting, with widespread erosion or non-ma-
rine deposition (poorly dated red beds are known in many
places on Sundaland).
Eastern Indonesia had several sedimentary basins that
existed throughout the Paleocene and early Eocene. These
were mostly deep marine, but locally shallow conditions
are known (e.g. around Misool Island). These faunas have
to be compared with the better known sections in the Mid-
dle East and Pakistan.
There are a few reports of Paleocene to Early Eocene
sediments in western Indonesia, on the flanks of the Sunda
craton and in the Philippines, some of which are disscused
below.
THE RANGE OF THE ALGA DISTICHOPLAX BISERIALIS
- PALEOCENE, OR YOUNGER?
An algal species, Distichoplax biserialis, has been
used to indicate Paleocene both in Indonesia and Papua
New Guinea (e.g. APC 1961, Eames et al., 1962, and
Carman, 1990). This species was probably first used as
an age marker in the late 1950's when British Petroleum
and their affiliates were drilling the Omati-1 well in P.N.G.
The distinctive algae was identified in thin sections from
the basal part of the Mendi Limestone in a “medium grade
detrital algal and bryozoan limestone with small Milioli-
dae, Rotalidae and Textularia.” Based on their experience
in the Middle East, these workers used it to indicate a
Paleocene age.
At about the same time, workers in the Philippines
thought a broader age assignment likely. Villavicencio
and Andal (1964) reviewed the occurrence of D. biseri-
alis in the Philippines. They found it in association with
Discocyclina and the Paleocene marker Miscellanea in
Palawan, but other locations were not as clearly dated. In
1969 Ishijima and Hashimoto reviewed this older work,
noting some possible Palaeocene planktonic forams at the
Palawan location, and described some new sediments from
Marinduque, a small island just south of Luzon, in a paper
entitled "Discovery of Distichoplax biserialis (Dietrich)
in Upper Eocene Limestone lenses". However while D.
biserialis from this site is well illustrated, the text and il-
lustrations note that it is within fragmented blocks within
the limestone. Although the authors said it also occurred
in the matrix, this is not obvious in the plates said to be
of matrix specimens. Ishijima and Hashimoto summed up
the work of Villavicencio and Andal by saying "In Cebu
Biostratigraphy of Indo-Pacific larger forams: page 12
and Bicol ... Distichoplax biserialis is found reworked
into Eocene rocks containing Flosculina, Miscellanea,
Nummulites, Discocyclina .... They [V & A] mentioned
that the specimens found in the Philippines always occur
with some Eocene foraminifera, although definate T-b
rocks were found barren of Distichoplax."
This confusion over reworking, and of considering
Miscellacea as being an Eocene fossil clouds the issue.
Consequently for some time after, workers in the region
considered it likely that D. biserialis did range into the
Eocene. This notion was supported by the abundance of D.
biserialis in the supposedly Late Eocene limestone at Jati-
 Image 1. The Late Cretaceous Barune Sandstone of eastern Papua.
This sample is from the same location studied by Glaessner (1952, 1960) and other workers, from a small outcrop near Port Moresby
in Papua New Guinea. Glaessner (1960) gave a Campanian age to the Barune Sandstone based on the presence of Pseudorbitoides
israelskii and Orbitoides tissoti, with thin sections also showing double keeled Globotruncanid and biserial heterohelicid types.

Biostratigraphy of Indo-Pacific larger forams: page 13

Image 2. The Jatibungkus Limestone

Picture of the rarer, Miscellanea-bearing facies in the Jatibungkus limestone, C. Java. Also present in this view are fragments of
Distichoplax biserialis.
The axial section through this form on the right is 1.8 mm across
 Biostratigraphy of Indo-Pacific larger forams: page 14

Image 3. The Jatibungkus Limestone.

Two pictures of the typical Nummulitid - algal association found in the limestone. Note the swollen marginal cord which, with
involute coiling, identifies the genus Ranikothalia. In both images there are common fragments of the alga Distichoplax biserialis
with distinct herring-bone appearance in some cross sections. The lower image also has the equally distinct solenoporacean alga
Parachaetetes with nodular growth and subquadrate texture of the cellular tissue. All three of these genera do not range younger
than Paleocene. Microspheric specimens of this Ranikothalia species are relatively rare and up to 7 mm in diameter. A fragment of
such a form is in the top right of the first image.
 Image 4. a & b. "Nummulites nuttalli" from the type
location in the Mangkalihat Peninsular (collection
GRDC, Bandung) = N. borneensis (Caudri)

bungkus in
Central Java.
This limestone
is dominated by
Discocyclina and a nummulitid with an inflated marginal
cord similar to the Late Paleocene Ranikothalia. However,
also present in this limestone, in small numbers, are good
specimens of Miscellanea and the distinctive alga Para-
chatetes. Both these last two genera became extinct close
to the end of the Paleocene (Adams 1970, Wray 1980).
A study of many dozens of Eocene samples from Java
has not found any sign of Distichoplax biserialis. How-
ever samples considered Eocene from Papua New Guinea
locally have frequent D. biserialis. Some samples have
Middle Eocene T.a forms such as Nummulites bagelensis
and T.a to T.c Discogypsina saipanensis, so although this
work is not yet completed it appears that D. biserialis
does appear to range into the "mid" Eocene, at least in
the higher latitude faunas of this time (cf. the Paleocene
and Eocene records of Lacazinella, which show a similar
distribution and are as yet unproven from low latitude
Eocene around Sundaland, Lunt 2003).

T.A1 AND THE OCCURRENCE OF RANIKOTHALIA

Apart from the Jatibungkus olistolith there are two
other place in SE Asia with reports of the distinctive num- Image 4. c & d. Nummulites
mulitid genus Ranikothalia (named after the Ranikot area thalicus from the type location
in Pakistan where Late Paleocene outcrops). in the Mangkalihat
Peninsula
(collection GRDC,
The Taballar beds of east Kalimantan
Bandung) = N.
taballarensis (Caudri)
In the upper Taballar River section on the Mangkalihat
Peninsula Leupold and van der Vlerk (1931) described
“Taballar marls with a small layer at the base with
[Nummulites] bagelensis and Discocyclina dispansa”
which overlies a mixed sandstone and marl unit containing
N. thalica, N. nuttalli, N. kelatensis and N. variolarius. This she considered identical with van der Vlerk's Nummulites
unit in turn overlies a red sandstone and basal conglom- nuttalli, and could distinguish these from true N. nuttalli,
Biostratigraphy of Indo-Pacific larger forams: page 15

erate (pebbles of radiolarian chert) lying unconformably
on the pre-Tertiary Danau formation (radiolarian cherts).
These authors compare their N. nuttalli and N. thalicus
with the publications on the Ranikot beds (Davies, 1927)
where the distinctive species N. nuttalli, with its inflated
marginal cord would eventually be awarded its own ge-
neric name of Ranikothalia Caudri 1944. Leupold and van
der Vlerk concluded that the sandstone-marl unit is “the
very oldest part of the Lower Tertiary”. They consider the
base of the overlying Taballar marls to be Tertiary b, and
that the highest part of the section could range into T.c.
In 1943 Caudri examined specimens from Sumba that
– an opinion shared by Adams (1970), who examined
syntype material from Borneo and Pakistan. Caudri's
specimens were found with other species of Nummulites
as well as Planocamerinoides orientalis. She re-named
van der Vlerks' form to Camerina borneensis, and the A
form from Camerina thalica to C. taballarensis.
The work of Caudri and Adams broke the link between
the Ranikothalia forms of the Middle East and the reports
from Indonesia, but the assignment of the Indonesian
forms was not clarified, nor the age of the rocks in which
they occur.
Pinugay Hill, the Philippines
In 1978 Hashimoto et al. published an account of some
Ranikothalia specimens from Pinugay Hill, about 35 kms
east of Manila, Philippines. These authors had been guided
to a location previously known for fossiliferous Creta-
ceous sediments, including Lepidorbitoides and Pseu-
dorbitoides, as well as a younger Distichoplax biserialis
- Operculina-Miscellanea assemblage. At this location
they found a form with the swollen marginal cord diag-
nostic of the genus Ranikothalia, which they named R.
bermudezi (Palmer). In the same sample were “a fragment
of Assilina, rare Asterocyclina (?) and Discocyclina (?),
abundant Distichoplax biserialis”. “Lepidorbitoides … is
thought to be reworked from the underlying Maastrich-
tian.” On this basis the sample was dated as Paleocene,
probably Middle Paleocene. Their D. biserialis is well
illustrated, but the figure referred to as Lepidorbitoides is
annotated as Pseudorbitoides on the plate, and the illus-
tration could as well be of Discocyclina. This rare fauna
remains an anomaly but could well be equivalent in age
to the Jatibungkus limestone on Java.
THE EARLY EOCENE, T.A2
Here I follow the definition of Adams (1970) who
considered the Ta.2 to be roughly the Early Eocene in his
3-fold division of Tertiary a, rather than the two-fold divi-
sion of Eames et al. (1962). Sediments of Early Eocene
age are very rare in the Indo-Pacific area as this period is
before the mid Middle Eocene, basin-forming tectonic
events.
Some good examples of T.a2 age carbonates occur
in Central Java in the olistostrome deposit at Kali Sana
(a few hundred metres from the very large olistolith of
Jatibungkus). Not only are there limestones boulders
with apparently T.a2 fossils, but there are also boulders
of Early Eocene mudstones dated from nannofossils and
planktonic foraminifera. There are no known occurrences
of Early Eocene sediments with both plankton and larger
foraminifera occuring as regular beds in contact, but at
least at this locality we can be certain that sedimentation
of Early Eocene age did occur in the vicinity. Nannofossils
in the mudstone include Ellipsolithus macellus and Tri-
brachiatus orthostylus indicting an NP11 to NP12 age,
and planktonic foraminifera include Morozovella arago-
nensis in the absence of Hantkeninids and Turborotalia
cerroazulensis s.l. suggesting P7 or P8 (unpublished work
by Baky & Lunt). The limestone sample of suspected T.a2

Biostratigraphy of Indo-Pacific larger forams: page 16

Image 5. Tertiary a2? Limestone from Kali Sana.

This limestone is an olistolith clast in Middle Eocene mudstones not far from the Paleocene olistolith of the Jatibunkus limestone.
Mudstone slumps / clasts at this location have ages as old as Early Eocene. Therefore the age of this boulder can only be said to
not be younger then Middle Eocene. The assemblage in the boulder is quite distinct with these exceptionally large, flat yet involute
Nummulitids with enlarged marginal cords. These compare very closely with Nummulites crasseornatus HENRICI from Timor, which
were assigned an Early Eocene (Ypresian) age based on a comparision of this species and N. nuttalli kohaticus DAVIES from the
uppermost Ranikot Beds in Thal, NW. India.
 Image 6. Tertiary a3 Limestone from Halmahera.
This limestone and adjacent samples contain both Alveolina as well as Planocamerinoides (=Assilina of old workers) . The strongly
dimorphic Nummulites species were often assigned to N. javanus by early workers, but the A forms do not have the same large
proloculus seen in N. bagelensis, and the coiling of the B-forms also differs from Javanese specimens of this age.

age contains common Nummulites crasseornatus (Hen-

rici), a species first defined in Timor in beds considered
Early Eocene (Ypresian) in age. This age assignment is
based on the similarity, noted by Henrici, between N.
crasseornatus and N. nuttalli kohaticus Davies, the latt-
ter being found in the uppermost Ranikot Beds, in Thal
NW India.
Other locations are assigned an Early Eocene age but
the evidence for the age is often tenuous. For instance
THE REGIONAL TRANSGRESSION IN TERTIARY A3
In Sundaland and surrounding areas mid- to later
Eocene sediments usually directly overly basement or
meta-sediments. There are a very large number of records
of Middle Eocene or Ta3 faunas in such transgressive
sediments, far more than the very rare observations of
Ta1 and Ta2 faunas. In some areas the transgression took
slightly longer and it may be Late Eocene (T.b) before the
Biostratigraphy of Indo-Pacific larger forams: page 17

Hashimoto et al, (1979) in a paper subtitled “Lower
Eocene foraminifera of the Philippines” describe the
Masungit Limestone as “Lower Eocene” based on the
occurrence of Nummulites burdigalensis, which is stated
to range over the Cuisan and Ypresian in Europe, corre-
lating these stages to Tertiary a2. However the other spe-
cies noted by these authors as T.a2 markers were Assilina
spira, Orbitoides cf. biplanatus and Alveolina javana, and
these forms are also common in T.a3, or Middle Eocene.
These authors also noted Nummulites perforatus [although
mentioning that a limited number of specimens made iden-
tification difficult]. Again, this is a T.a3 fossil (cf Adams
1970, Eames in Davies, 1971).
first marine sediments are observed.
A typical example of this succession is the detailed
work of Adams (1965) on the Melinau Limestone in
Sarawak, which has eroded slatey shales and quartzitic
sandstones of the Mulu formation being overlain by thick
limestone. The lowest thin limestone has T.a3 faunas, iden-
tified by the giant Nummulites javanus, and an absence of
T.b forms. Overlying this are thick T.b (1,000-2,000 feet),
T.c & d (combined up to 2,000 feet), and a similar 1,000
to 2,000 feet of T.e limestone.
 Image 7. Tertiary a3 Limestone with Planocamerinoides (=Assilina ).
This sandy limestone is from the Bagelen Beds near the Worowari River north Central Java. An Early Oligocene pebbly mudstone
olistolith with common boulders of Middle Eocene and rarely older lithologies left on the surface after the soft host clay has been
weathered away. This image shows a large microspheric Planocamerinoides and smaller macrospheric specimens, all with the
diagnostic evolute coiling. Nummulites bagelensis is at the top of the image

Biostratigraphy of Indo-Pacific larger forams: page 18

Image 8. Tertiary a3 Limestone with Alveolina.

This specimen from the N. Lukulo area is rich in Alveolina, clearly visible here with the diagostic pre- and post-septal passages. Note
the involute Operculina -type in the lower left, possibly showing affinities towards a transition to Tansinhokella with small lateral
chambers, as old as Middle Eocene.
 Image 9. Tertiary a3 Limestone from Karangsambung, C. Java.
This limestone is a typical Middle Eocene carbonate with the giant Nummulites javanus, partly visible at the bottom of the picture,
This is one of the largest foraminifera found in the region, frequently reaching 2 cms in size for the microspheric generation.
Equatorial and axial sections through individual specimens of this species are
left and below. The corresponding A or macrospheric generation is
seen in the above image as the form from 1 to 2 mm in size
with a proloculus over 500 µm in diameter. This form has its
own species name of Nummulites bagelensis. These highly
dimorphic and complex (in the B generation) forms are
typical of T.a3. After the end T.a extinction the species of
Nummulites lack these "advanced" evolutionary
chartacters

Biostratigraphy of Indo-Pacific larger forams: page 19

 Pellatispira

Image 10. Tertiary a3 Limestone with early Pellatispira.

A limestone from the Jiwo Hills, C. Java, interbedded with coarser grained lithologies rich in the Nummulites javanus shown in
the previous figure (note the N. bagelensis A form on the left with a proloculus nearly a millimeter in diameter). This fine grained
facies also has fragments of early Pellatispira forms which are planispiral, thick-walled and coarsely perforate and showing early
development of the marginal crest (P. provalei). This section also contains many planktonic foraminifera, includuing forms with a
keel which would be members of the genus Morozovella that became extinct at the top of Zone P14. Also under higher magnification
the coarsely rugose wall structure of the genus Acarinina can be seen, another form that became extinct to the top of P14. While
Berggren et al. (1995) use the intra Zone P15 top Bartonian Stage as their top for the Middle Eocene, workers in SE Asia have used
the mass extinction at top P14 to define top Middle Eocene, which still still seems both practical and logical as the Eocene itself is a
biostratigraphic unit.

ON THE TA3 / TB LETTER STAGE BOUNDARY
The transition from Tertiary a to Tertiary b saw the
extinction of the genera Assilina / Planocamerinoides,
Alveolina, and all the large complex Nummulites. The
Pellatispira occurs with Assilina / Planocamerinoides
and T.a Nummulites such as strongly granulate species
and N. bagelensis. He also notes Alveolina in a few of
these samples but it is likely that at that time his generic
concept also included the younger ranging Borelis. The
Biostratigraphy of Indo-Pacific larger forams: page 20

complex Nummulitids were a rock-forming bioclasts and
the rapid extinction of so many species was an important
biological event.
Following the influential paper of Adams (1970) many
workers have identified T.b on the presence (total range
zone) of Pellatispira and the related Biplanispira. In ad-
dition to Adams, Eames (in Davies 1971) noted no known
co-occurrences of Assilina with its possible descendants
Pellatispira and Biplanispira. However there are several
old references from Indonesia with Pellatispira occuring
before the end T.a extinctions. Umbgrove (1928) is quite
specific in this matter, noting several locations where
figure included here (Image 10) from the T.a beds of
Watu Perahu in Jiwo, C. Java, has common N. javanus
in the same facies. The image shown here has several
specimens of Pellatispira aff. provalei, with N. bagelensis
and other granulate forms the secondary T.a marker Lin-
derina, as well as the planktonic foraminifera Acarinina
and Morozovella, demonstrating that Pellatispira not only
evolved with T.a3 but that this was within the Middle
Eocene (Zone P14 and older).
Elsewhere other T.a limestones with age markers such
as Orbitolites are reported to contain Pellatispira, for ex-
ample in south Sulawesi (Dollfus, 1915).
 Image 11. Early part of Tertiary b Limestone from West Java.
This limestone is from a mineralised area in West Java (Cimuncang site of Koolhoven 1933) hence the limestone is dark grey in
hand specimen and lacks contrast in thin section. Local stratigraphy places this site as close to the T.a to T.b boundary. This was
the one site in West Java that Koolhoven recorded "Assilina" but so far has only been found to contain a flat, mostly evolute and
small Operculina species. Planktonic forams from mudstones abve and below this limestone bed (debris flow) indicate Zone P15. The
limestone samples are special in that they contain granulate Nummulites (locally called N. hoogenraadi) with advanced species of
Pellatispira [P. ?fulgeria] (left).

Until we have a detailed taxonomy of the Nummulites
species from the Eocene showing the multiple extinctions,
the end T.a extinction event will always be under-repre-
sented on range charts. The easiest way to emphasise the
severity of the T.a3 to T.b transition is the fact that it takes
only a few moments to distinguish a thin section of T.b
from T.a3 age as the whole fauna has shifted in character,
even though we currently lack the species level taxonomy
to adequately describe this transition.
There is one location where part of the transitional
T.a3 to T.b fauna is known. This is from the Cimuncang
contain fairly large, moderately dimorphic and slightly
granulate Nummulites species, including N. hoogenraadi
(Doornink), the type location for which is nearby in a
section lacking any plantonic fossils. In both the Cimun-
cang and the N. hoogenraadi type location the limestones
contain advanced species of Pellatispira (see Image 11),
which have a much larger late adult or gerontic stage com-
pared to the upper T.a3 forms illustrated from Watu Perahu
at Jiwo Hills (Image 10). Both the early T.b locations also
have common Palaeonummulites gerthi (Doornink), a spe-
cies possibly restricted to the T.b Stage. This survival of
a few granulate species into T.b is not without precedent
Biostratigraphy of Indo-Pacific larger forams: page 21

site of Koolhoven (1933) in West Java. Originally this

location was the single site in West Java dated as T.a on
the presence of Assilina. Recent unpublished work has
relocated the single limestone bed in a mudstone sequence.
In the 1930's Koolhoven and the Mijnbou were not able
to carry out planktonic biostratigaphy and even now it is
hard because of the slightly mineralised and baked nature
of the lithologies. However the few samples with good
planktonic faunas lack Morozovella and Acarinina, and are
dated as Zone P15. Elsewhere in the region (eg. the Cipatu-
lah well to the south) this mudstone-dominated sequence
has a similar age. The limestone at Cimuncang has a flat,
evolute Operculina species but no sign of Assilina. It does
as Eames (in Morley Davies 1971) notes that in western
Tethys a single granulate species (N. yawensis) survives
after the extinction of most granulate species.
TERTIARY B
Following the end T.b there is a radiation of new larger
foraminifera types, particularly in the Pellatispirine and
Heterostegine lineages. Fairly large, flat but involute
Operculina species are noted in T.a 3 (Image 10) but in
the T.b subdivision of chambers evolved in a number of
different ways to give rise to simple Operculina eniweto-
 Image 12. Tertiary b Limestone from Nanggulan.
This limestone is below P16 mudstones in Nanggulan and similar limestone found as xenoliths in the Sangiran Dome mud volcano
have yielded specimens of Porticulasphaera semiinvoluta identifying Zone P15. There are several fragments of Pellatispira as well as
Nummulites gerthi, a species morphologically un-exceptional, typical of this age, without the strong dimorphism and distinct physical
features (complex septal traces, granules etc.) seen in T.a faunas. Above the 1/2 mm scale dot is an involute specimen with clear
secondary septa of the genus Heterostegina (Vlerkina).

kensis, with pinching-out of adult chamber growth in the
median plane, the rare Grzybowskia (Christmas Island,
Lunt 2003), true secondary septa in Heterostegina (Vlerki-
na) which developed into Tansinhokella. This radiation of
the Heterostegines parallels a similar record in the western
Tethyan area (Banner and Hodgkinson, 1991)
The evolution of the Pellatispines saw a rapid develop-
Forms that were present in Ta but become more com-
mon in T.b are Amphistegina and Asterocyclina, and pos-
sibly the abundance of corals.
On the Australian Plate, which was much further south
in Eocene times, the Pellatispirines are virtually unknown
and T.b is often characterised by the species Lacazinella
wichmani, which appears to be restricted to the Australian
Biostratigraphy of Indo-Pacific larger forams: page 22

ment of the adult or gerontic stage and the separation of the
Biplanispira lineage. The work of Hottinger et al (2001)
has greatly improved out knowledge of the morphology
of these complex forms but the new taxonomy has not yet
been applied to the sections in the Indo-Pacific. Within P15
times (from material dated with Porticulasphaera semi-
involuta from Sangiran, Gamping Barat and Nanggulan
in Central Java) forms had arisen with the spiral stage
reduced to a single whorl and the bulk of the test being
composed of minor chamberlets or cubiculae derived
from an evolutionary exaggeration of the late adult stage
of ancestral forms.
plate. This is known to occur in with Nummulites javanus
and Alveolina (T.a), as well as with Nummulites fichteli
(cf. Adams, 1970, Bursch 1947, confirming T.c) but the
majority of records of this species are considered to be
equivalent to the T.b Stage (Lunt 2003).
ON THE T.B / T.C LETTER STAGE BOUNDARY
At the moment it is widely assumed that the T.b-T.c
boundary is equivalent to the top of the Eocene as defined
by other means, most commonly planktonic foraminifera
and nannofossils. The planktonic marker datum planes are
 Image 13. Tertiary b Limestone from Pasir E. Kalimantan.
A limestone of T.b age showing better detail in the Pellatispira tests than the previous examples, especially the delicate chamberlets
of interlamellar spaces on the outer sides of the test. The marginal crest is strongly developed in a form that may be Pellatispira
fulgeria. Some Discocyclina and involute Operculina specimens are present, the latter being ancestral to Heterostegina (Vlerkina) and
Tansinhokella known in this area.

correlated to be just below the base of the normal polarity
of magnetostratigraphic chron C13 (ref.). However just
within the base of the normal polarity of chron C13 is
a marked oxygen isotope shift that has been interpreted
as the result of an abrupt cooling event (Mei, 1991). It
is possible that the shallow marine, tropical carbonate
faunas were catastrophically affected by such an abrupt
cooling .
Tan Sin Hok 1932). Secondly a gradual evolutionary suc-
cession from these slightly involute Heterostegina forms
to Tansinhokella is seen in mid-Oligocene sediments, ulti-
mately evolving into Spiroclypeus s.s. in latest Oligocene
(Lunt and Rennema in prep).
TERTIARY C
Biostratigraphy of Indo-Pacific larger forams: page 23

On any stratigraphic range chart for larger forams,
this T.b to T.c transition stands out as one of the most
significant extinctions. Most notably the dominant
Eocene larger foram, Discocyclina, and its relatives
such as Asterocyclina, abruptly became extinct, as did
the varied forms of Pellatispira and Biplanispira. The
Eocene Heterostegina (Vlerkina) to Tansinhokella line-
age seems to become extinct and is therefore unrelated to
the Oligocene forms with the same name. This example
of iterative evolution has two lines of evidence. Firstly
the Early Oligocene has no records of Tansinhokella, just
Planostegina and rare, slightly involute Hereostegina (cf.
The next larger foram association (T.c), lasted only
about 2 Ma. This was the period after the terminal Eocene
extinctions but before a migration of new forms arrived
from Central America. It is during this period that Sr iso-
topic dating becomes practical and is now being applied
to date these and younger limestones.
Carbonate faunas with foraminifera from this period
are characterised by Nummulites fichteli-intermedius
with their distinctive reticulate septal traces. Other forms
found in this Stage include large species of Planostegina
(P. bantamensis and praecursor Tan). Some genera are
found more commonly in this stage than before or after,
 Image 14. Tertiary c Limestone from SW Java.
Following the major end Eocene extinction, limestones no longer contain Discocyclina, a form than occurs in every image of the
letter stages so far. Also missing from T.c faunas are the Pellatispirines and the rarer Tansinhokella. Although species of Nummulites
are still not well defined there is a clear faunal turnover at this time. A few simple small species better placed in Palaeonummulites
occur both before and after the extinction but among the larger Nummulites it is the reticulate Nummulites fichteli-intermedius that
dominate Early Oligocene assemblages. The above image, from Cikalong, West Java shows several N. fichteli in oblique view that
show the reticulate septal traces.

Biostratigraphy of Indo-Pacific larger forams: page 24

Image 15. Tertiary d Limestone from Papua.

The left of this image has N. fichteli next to Eulepidina, the overlap of which defines T.d. The right side of the image has fragments of
peneroplid miliolid forms; one of the oldest records of this type.
 Image 16. Tertiary e1 Limestone.
A limestone from the Cicarucup area of West Java, unconformably overlying Eocene beds. These limestones have Eulepidina (top
left) without Nummulites and common Heterostegina forms as shown above. These are slightly involute with extensive adult flanges
which clearly have secondary septa in equatorial view. At this evolutionary stage fragments of these involute forms can be mistaken
for Cycloclypeus This sample is from below the stratigraphic datum (T.e1 to T.e2-3 boundary) where these forms develop into
Tansinhokella by acquistion of lateral chambers, through subdivision of the alar prolongations.

but are not index species. These include Borelis pygmaeus
and Halkyardia.
THE TRANSITION FROM LETTER STAGE TC TO TD
The original definition of van der Vlerk and Umbgrove
reports in the late 70’s and 80’s was that the T.d stage
was an artifact of reworking of Nummulites following the
large mid-Oligocene eustatic sea level fall. This followed
the influential papers on eustacy and stratigraphy by Vail
and others at that time. Note that no such cited cases had
any other genera reworked; that is, erosion from the pro-
Biostratigraphy of Indo-Pacific larger forams: page 25

(1927) defines the Tertiary d on the co-occurrence of Ne-
phrolepidina and Eulepidina, together with Nummulites.
The first appearance of the family Lepidocyclinidae in the
Indo-Pacific region, from the much older central American
stock, has been argued to be diachronous by Adams (1984)
because of assumed slow migration of the genera from
America and across Tethys. This model would suggest
the base of T.d is a poor zonal boundary.
In recent years it has become acceptable to lump Terti-
ary C and D together, hence the informal name “CD lime-
stones” for Early Oligocene carbonates in eastern Java.
A more extreme opinion suggested in some oil industry
posed 150 metres sea level fall only seemed to affect Early
Oligocene and never seemed to reach Eocene strata.
New work in the Cimanggu section of West Java (Lunt,
Allan and Baky, in prep.) dates the T.c to T.d transition and
also notes that a third taxon, Neorotalia mecatepecensis,
may all have migrated from Central America at this time.
The arrival of these forms was at about 32 MYBP, within
planktonic zone P19, which appears to be well before the
mid-Oligocene sea-level fall. This major sea-level fall was
proposed to have been eustatic by Vail et al. (1977), with
revisions by Haq et al (1987) who dated it at 30 MYBP,
hence its popular title of "the 30 million year sea-level
 Neorotalia
mecatepecensis

Image 17. Tertiary e2-3 Limestone.

This is a poorly preserved limestone as it contains clay contamination and is interbedded with mudstones in a fore-reef calc-
turbidite or debris flow setting in NE Java. Weathering has allowed etching of the bioclasts through humic acids and darkened the
rock. However this location, both in mudstones and limestones has common Neorotalia mecatepecensis, which is seen in samples as
old as T.d but becomes more common in T.e2-3. The lower right corner has two poor specimen of Tansinhokella in which the alar
prolongations have developed into layers of lateral chamberlets. Tansinhokella ranges much younger but the presnce here of so many
specimens of N. mecatepecensis without any sign of the descendant Miogypsinoides identifies T.e 2-3.

fall". Later work on the Tertiary time scale culminating
in Berggren et al (1995; the same time scale used here)
re-dated the level given in Haq et al. at Chron C10n, intra-
P21 and NP24 event to about 28.5 MYBP.
There is little published data precisely dating the Vail
/ Haq mid-Oligocene event. Some early efforts such as
used strontium dating and backstripping techniques to
study Oligocene sea levels. Using the same time scale as
Beggren et al (1995) these workers did not recognise a
single outstanding eustatic event in mid-Oligocene times.
Instead their data (cf. their figures 2 and 8) show a mod-
erate sea level fall of about 45 metres between 27.9 and
28.3 MYBP (both ± 0.7 MA), and only smaller, fluctuating
Biostratigraphy of Indo-Pacific larger forams: page 26

Olssen et al (1980), on well cuttings from the Atlantic mar-
gin of the USA, suggested a slightly older Early Oligocene
age although a revision of this work compared to the
Irish Atlantic margin by Miller et al (1985) suggested the
unconformity studied was closer to Vail's mid-Oligocene
level (c. 31 MYBP). The problem with such studies was
their reliance on cuttings-samples, as only deep boreholes
penetrate the para-conformable expression of the event,
and also the weakness of planktonic biostratigraphic zones
in the early to mid-Oligocene.
More recently Kominz and Pekar (2001) studying the
profile of the eastern seaboard of the US Atlantic margin
rising and falling, sea-level changes in the period 31.5 to
33 MYBP.
This data suggests that the outstanding major (c. 150
metre) sea-level fall shown on oil-industry sequence
stratigraphic charts in mid-Oligocene times, if it ever
existed, was not likely to have been the cause of the
mass-migration of central American larger foraminifera
into the Mediterranean and Indo-Pacific areas. Also the
migration reached the Far-east at an older date than pre-
viously estimated, so Adam's (1984) hypothesis of slow,
diachronous migration over millions of years is probably
not correct.

THE TOP OF T.D
Unlike the sudden appearance of the Lepidocyclinids
at the base of T.d, Nummulites seems to disappear more
gradually. There are records of this species as high as the
extinction of Chilguembelina cubensis in field samples
from the center of the Kujung Anticline in NE Java (28.5
MYBP), and as young as 28.3 MYBP at the Pelang limestone
C-E Java (Pelang Lst 87Sr/86Sr = 0.70804, NBS987 =
0.710235, Age of 28.3 mybp, combined precision & corre-
lation error 27.59 to 28.89 Ma. Center of Kujung Anticline
T.d / top P21(a) sample: 2K1/9/21, 87Sr/86Sr = 0.70800,
NBS987 = 0.710235, Age of 29.3 mybp, combined preci-
sion & correlation error 28.58 to 29.95 Ma.). However in
the Kujung anticline we have found single specimens of
Nummulites slightly higher into the basal Late Oligocene,
but not yet after the evolutionary appearance of Tansin-
hokella (base Te2-3). However Hashimoto et al. (1977)
working on the Bugton Limestone of east Mindoro in the
Philippines have clearly illustrated Nummulites next to
Tansinhokella. This unique sample was a loose boulder
of limestone, so while their photograph is clear and shows
no difference in preservational type, the location or strata
cannot be resampled.
Therefore it is likely that the disappearance of this very
important genus was gradual and took place over one or
two million years. If Nummulites did become extinct after
the evolution of Tansinhokella then Letter Stage Te1 is
not strictly viable, although the most significant reduction
in Nummulites abundance at about 28.5 MYBP remains a
useful biostratigraphic datum.
SUBDIVISION OF T.E
The previously unpublished work of Muhar (1957) of
the BPM, shown in the previous chapter, remains the best
method to subdivide the T.e Letter Stage. This scheme
simply traces the evolutionary development of Neorotalia
mecatepecensis to Miogypsinoides and to Miogypsina, and
Heterostegina (Vlerkina) to Tansinhokella. These events
are approximately fixed to biostratigraphy and some Sr
isotopic ages. Only the final transition from Tansinhokella
to Spiroclypeus remains uncalibrated.
ON THE UPPER T.E / T.F LETTER STAGE BOUNDARY
The position of the top of Letter Stage T.e, with respect
to planktonic zones or absolute ages, is still not adequately
fixed. Most publications (Chaproniere 1975 & 81, Adams
1984) suggest the top occurrences of Spiroclypeus and
Eulepidina are at about the N5 to N6 zonal boundary,
which current estimates place at about 19 MYBP. This is
difficult to prove in Indonesia as the N5 to N6 boundary
is hard to recognise. The defining evolutionary event, of
Globigerinatella insueta, is a poor datum because this
species is rare in the region. In addition a major regional,
tectono-stratigraphic event at about this time dominates
the stratigraphic record, with strong facies changes.
This regional geological event is characterised by a
change from widespread carbonate development (a pro-
longed flooding period) to the appearance of a new and
highly active clastic sedimentary source. In Sundaland this
severe facies change includes at least a hiatus, or deposi-
tion of a substantial clastic section before the occurrence
of the next carbonate related facies. Isolated reefs seen in
oil exploration may have T.e carbonate separated by only a
small hiatus from basal T.f carbonate (e.g. the Kujung fol-
lowed by Rancak limestones in offshore NE Java) however
these are coral dominated reefs, often pinacle reefs with
limited foraminiferal grainstones, and they have only been
encountered in oil wells with cuttings and limited core
samples. They are not known in outcrop onshore. Such a
break or change in the geological record can be expected
to emphasise the contrast in evolving assemblages.
In Irian Jaya and Papua New Guinea there is less tec-
tonic or facies contrast during this mid-Early Miocene
period but the same faunal turnover can be seen. This is
known as the change from Kereruan to Kikorian faunas
in Papua New Guinea. Not only do two locally important
components of the larger foram assemblages disappear
but there is subsequently a radiation of new forms. Note
also that in eastern Indonesian and Papuan areas the rela-
tively rare genus Austrotrillina, previously thought to be
a T.e marker, is seen to continue into younger sediments.
In addition some of the forms that radiated and became
important in the T.f have occasional records alongside
upper T.e markers (e.g. Flosculinella, and Marginopora).
The distinction between T.e and T.f Letter Stages appears
to becomes slightly less abrupt in areas away from Sunda-
land where there is maximum expression of the mid-Early
Miocene tectono-stratigraphic event.
The T.e to T.f boundary is therefore not comparable to
the end T.a or T.b mass extinction events. The surviving
T.f fauna is much more a continuation of the end-T.e fauna
rather than the major and widespread T.a to T.b, or T.b to
T.c faunal turnovers.
TERTIARY F
The Tertiary F Letter Stage has, by far, the most confu-
sion in its useage between generations of workers. The top
of the Stage has shifted considerably, and the two or three
subdivisions also have inconsistent definitions.Therefore
Tf.3 of Adams (1984) is T.g of workers in the 1930's and
40's; and the old Dutch Tf, 1,2,3 are very different to
Adams' divisons of the same name.
Tertiary f was first defined as the interval between the
end T.e extinctions [Spiroclypeus and Eulepidina] and the
 primitive
Miogypsinidoides

primitive Images 18 & 19. Tertiary e4 Limestone.

Miogypsinidoides These limestone both contain specimens with Neorotalia juveniles
but a flange of Miogypsinoides stage chambers, identifying evolution
into the latter genus. Miogypsina is absent from the samples and
Biostratigraphy of Indo-Pacific larger forams: page 28

both are from locations with good Sr isotopic dating confirming an

age of latest Oligocene.
The picture on the left is important as it contains both Tansinhokella
(the high contrast specimen) and true Spiroclypeus (the largest
specimen). Spiroclypeus was defined on a microspheric specimen,
and until recently there was doubt about whether this was a
valid genus or just a modified microspheric generation of late
Tansinhokella. In this regard the flattish specimen just right and
above the image center is important as it shows true spiroclypid lateral chambers in a macrospheric tests. Spiroclypeus is therefore
a valid, morphologically defined genus. Both images same scale: left from the lower Prupuh limestone in NE Java; right from the
Rajamandala Limestone west of Bandung.
 Image 20. Tertiary e5 (Upper T.e) Limestone.
The type Prupuh Limestone in N.E. Java. This sample is from a bed of calcarenite debris which occurs between beds of chalky
limestone. In this image is a large, ?microspheric specimen of Miogypsinoides (top right) and elsewhere are smaller macrospheric
members of the same genus. This dimorphism in the Miogypsinoides is only seen in the Early Miocene, as Late Oligocene members
of the genus are of roughly equal size. These contrast with the specimens on Image 18 that are from near the base of the Prupuh
and close to the evolutionary development of the genus, dominated by the ancestral Neorotalia stage with only a slight flange of
Miogypsinoides stage adult chambers. Several specimens of Miogypsina are in this picture, as is the embryont of Eulepidina (near
bottom, left of center)

mass extinction of Lepidocyclina, all Miogypsinids, Flo-
sculinella bontangensis and Austrotrillina (van der Vlerk
& Umbgrove 1927, see Figure 1). By 1931 Leupold and
van der Vlerk had noted that Austrotrillina became ex-
tinct before the others [i.e. within Tf]. In 1962 Eames et
al. noted that Flosculinella also did not range as high as
the more cosmopolitan Miogypsina and Lepidocyclina.
deviated from the original Dutch concept of the Letter
Stage, which was based on this mass extinction.
We can be sure that this was the intention of the
early workers as their foraminiferal stratigraphy was
integrated with molluscan stratigraphy, often sampling
the same formations. The Top T.f of the early workers
Biostratigraphy of Indo-Pacific larger forams: page 29

However it was Adams (1970) who recognised that the
range of Lepidocyclina extended well after the extinction
of Miogypsina. By 1979 Adams et al. had found Lepidocy-
clina as young as base Pliocene. Adams therefore extended
T.f to the base of the Pliocene and first established Lower
T.f [up to the extinction of Miogypsina and a few rarer
species] and Upper Tf to the extinction of Lepdiocyclina.
As the Letter Stages are assemblage zones, not defined
on individual evolution / extinction datums this was not
an incorrect approach. However this demoted a mass
extinction and faunal turnover event from occuring at an
assemblage stage boundary to within such a stage. It also
was the division between molluscan faunas with less than
35% extant species, and those with more. Examples of
the older Molluscan Stage, called the Rembangian and
Preangerian by Oostingh (1938, also in van Bemmelen
1949) contain Miogypsina, Cycloclypeus annulatus etc.
and the type locations for both these stages are now
dated with modern planktonic biostratigraphy as latest
Early and base Middle Miocene respectively (pers. obs.,
unpublished). The molluscan stage with 35-50% extant
species (Tjiodeng or Ciodeng Stage) has 42% extant
species at type locality in W Java, which was considered
Late Miocene by Martin 1919). The Genteng Beds of SW
 Image 21. Lower Tertiary f Limestone - T.f1.
This picture shows two well preserved Austrotrillina howchini tests, with bifurcating alveolae in the walls. Right of center is a
specimen of Miogypsina with the rounded chmbers of the median layer and numerous "lateral chamberlets" now termed cubiculae.
Several L. (Nephrolepidina) specimens are present, the one in the lower right showing the four-cornered distribution of pillars
diagnostic of L. (N.) ferreroi, a Lower T.f marker
From a thin, un-named limestone above the Prupuh Limestone, NE Java.

Java are also in Oostingh’s Ciodeng Stage, and these beds
overly the Bojongmanik Fm. which contains limestone of
T.f3 age (trace Lepidocyclina with Alveolinella) which has
recently been dated as between 10-11 MYBP by strontium
isotopic methods. Another T.g location are the Lawak
Beds in Central Java, such as the lower NN11 sample at
Margamukti (Image 24).
The algae often preserved as recognisable plate fragments,
or as an increase in micrite and fine bioclastic products
from the early breakdown of its aragonitic needles.
Larger foraminiferal grainstones are generally rarer after
this event, most often as as deeper photic Cycloclypeus
facies, or minor Operculina / Amphistegina or Alveolinella
calcarentites.
Biostratigraphy of Indo-Pacific larger forams: page 30

As the works of Adams (1970 and 1984) ae so widely

quoted it would be confusing to revert now to the origi-
nal concept of Tf. However it should be stressed that the
original Tf to Tg boundary was first defined as a mass
extinction in mid-Middle Miocene times.
This important biostratigraphic event is also an impor-
tant faunal turnover for other carbonate facies organisms.
The result was that shallow marine biohermal limestones
changed from mixed coral and coralline algal boundstones
with larger foram grainstones to dominant coral reefs with
a marked increase in Halimeda green algae and molluscs.
FAUNAL DEVELOPMENT IN LOWER T.F
Within the Lower T.f (particularly its lower part, T.f1),
there are consistent occurrences of Miogypsinoides, which
then becomes extinct at or just before the Early to Middle
Miocene boundary. This contrasts with the Mediterranean
area where Miogypsinoides is hardly seen at all above the
basal Miocene evolution of Miogypsina. In basal T.f1 are
several records of “Conomiogypsinoides” or Miogypsi-
noides dehaartii var. cupulaeformis. In some samples (e.g.
the Lutut Beds of Central Java) these variants appear to be
associated with a Miogypsinoides from with incipient lat-
eral chambers, well after the evolutionary transition from
Miogypsina to Miogypsinoides (see secion on Miogypsin-
odes). Many of these examples may be "Miogypsinodella",
which is thought to be the microscopic generation of later
Miogypsina forms. Also in the lower Tf the Miogypsina
variant Lepidosemicyclina is found, suggesting a radiation
in diversity of the Miogypsinidae at this time.
Austrotrillina is rare in Sundaland after T.e but
frequent in Irian Jaya and Papua New Guinea, where it
survives until just after the Orbulina datum that is usually
used to define the base Middle Miocene (Adams, 1970 &
84). This extinction is used by Adams (84) to mark the top
of T.f1. The fusiform miliolid Flosculinella bontangensis
grades into Alveolinella forms about the same time as the
disappearance of Austrotrillina, occurring alongside the
intermediate form Alveolinella fennemai or “praequoyi”
within the upper part of Lower T.f (=T.f2). Also in T.f1
are forms such as L. (Nephrolepidina) ferreroi, which
has a distinct profile in thin section, and the rare (espe-
cially in western Indonesia / Sundaland) Pseudotaberina
malabarica.
The sub-zone T.f2 is relatively short lived (less than
3 Ma compared to just over 5 Ma for T.f1) but is well
represented in Southeast Asia, especially western areas,
as regional transgressive conditions developed into late
transgression (fewer clastic sources, more drowned land
areas) and prograding highstand, with common carbonate
development. Apart from the absence of Miogypsinoides,
which faded from faunas through the later Early Miocene,
and Austrotrillina, which was never a major faunal com-
ponent except in very specific facies, there are few fea-
tures to distinguish T.f2 from T.f1. The distinctive species
Cycloclypeus annulatus (so distinctive it has been often
placed in its own genus or sub-genus of Katacycloclypeus)
is a common component of T.f2 assemblages. However
is does overlap in range with both Miogypsinoides as well
as Austrotrillina, although it almost never occurs with the
latter, as the two genera lived in opposite extremes of the
carbonate platform.
In well preserved early Middle Miocene assemblages
it is not rare to find representatives of the genus Planos-
tegina. Such forms are found in the upper Ngrayong Fm.
sediments of NE Java and Madura. Chaproniere (1980)
reported it from N10-N12 sediments of New Zealand and
Hashimoto (1977) observed it in Letter Stage T.f1 in the
Philippines.
THE TOP OF T.F2 (FAUNAL TURNOVER); THE EXTINCTION
OF MIOGYPSINA.
Since the modification of the concept of top Tf / top Tf2
by Adams the most important marker at this mid-Miocene
boundary is the extinction of the previously ubiquitous
genus Miogypsina. Secondary to this in deeper photic
facies is the widespread marker Cycloclypeus annulatus,
while the common for L. (Nephrolepidina) ferreroi also
becomes extinct at about this time.
In other areas, Cycloclypeus annulatus has been de-
scribed from sediments in Fiji, dated by Adams and Frame
(1979) as N14. From their published data it is possible to
interpret a slightly older, N13, age for these Fiji samples
(the absence of G. subquadratus /ruber does not clearly
identify N14).
LETTER STAGES AFTER TF2
The Letter Stage T.f3 (or Upper T.f of Adams, 1970)
is the period following a mass-extinction of larger
foraminifera. Limestones of T.f3 age are dominated by
green coralline algae, coral and molluscs, all of which are
composed of aragonite and prone to break-down during
early diagenesis. Larger foraminifera are less often found
as the calcarenite grainstones so frequently seen in the
older Tertiary. The most comon genera are Cycloclypeus,
Palaeonummulites / Operculina, Amphistegina, localised
Lepidocyclina, and, in shallower settings, Alveolinella
quoyi, and Marginopora. The end of the T.f Letter Stage
was defined by Adams on the extinction of Lepidocyclina,
which appears to be just above the modern definition of
the Miocene-Pliocene boundary.
The Letter Stages T.g and Th are not practical units
for foraminiferal stratigraphy as they were defined on the
molluscan percentage method and, as explained above, T.g
is equivalent to the modern Upper T.f or T.f3.
The Tertiary h was first defined in 1931 with two parts;
the lower having 50% extant molluscs and the upper 60%.
These equate to the Cheribonian and Sondien Molluscan
Stages of Oostingh respectively, which are roughly equiva-
lent to the Lower and Upper Pliocene. The Quaternary or
Pleistocene is represented in Oostingh’s mollusc scheme
by the Bantamien Stage with 70 % or more extant species.
The Quaternary therefore has no equivalent in the Letter
Stages (cf. Leupold & Van der Vlerk 1931).
Biostratigraphy of Indo-Pacific larger forams: page 31
T HE L ATEST C ENOZOIC RADIATION OF LARGER
FORAMINIFERA
The general scarcity and eventual extinction of Lepido-
cyclina though T.f3 is a secondary biostratigraphic event
for foraminiferologists. Starting before the last record of
this genus there is an often overlooked raditation of new
larger foraminifera. Carbonates were still dominated by
aragonitic bioclasts, and foraminifera were not as common
as in older Tertiary times. However in the Late Miocene
[within T.f3] a number of new forms began to appear.
This radiation is slow compared to earlier diversifications,
such as in early T.b., however, the evolution of several
 Image 22. Lower Tertiary f Limestone - T.f2.
A sandy limestone from Gunung Geger on Madura, known to be above the N9, Orbulina datum, but well below the N12 extinction of the Globortalia fohsi lineage. In this view are several
specimens of Miogypsina, and fragments of hexagonal median chambers thought to be from Lepidosemicyclina (specimens with the enlarged embryont also diagnostic of this lineage are known
from this formation), one of the youngest records of this genus. A large specimen of Cycloclypeus annulatus with its distinct ring-like thickening is present. L. (Nephrolepidina) ferreroi is also
present in this sample, but not in this view.

Biostratigraphy of Indo-Pacific larger forams: page 32

 Image 23. Lower Tertiary f Limestone - T.f2.
A sandy limestone from the Annulatus Beds at Jatiluhur, West Java, dated in surrounding mudstones as lower Middle Miocene (N11-N12). This is close [107.3909˚E, 6.95156˚S; within about 2
kms] to the type location of Cycloclypeus annulatus MARTIN 1880 (Die Kluft des Citarum bei Cikao nordlich vom Gunung Parang” West Java.
Note the very large lepidocyclinids. These are microspheric forms (probably Lepidocyclina dilatata) and not Eulepidina. Also in the sample are Multilepidina forms. Concentrations of very
large microspheric lepidocyclines to several centimeters are most common around the latest Early to basal Middle Miocene (upper Tf1 / Tf2). This may be a parallel trend to the increasing
divergence in A and B generations seen in of other lineages (eg Middle Eocene Nummulites) but this is hard to demonstrate as a genuine evolutionary trend on the limited data available.

Biostratigraphy of Indo-Pacific larger forams: page 33

new forms mean that it is now possible to recognise lime-
stones as being from either latest Miocene / Pliocene or
Pleistocene sediments.
First among this radiation was the appearance of the
ornate, but relatively small, trochospiral genus Calcarina,
in the latter part of T.f3. This strongly ornamented, carbon-
ate facies preferring form is thought to be the ancestor to
the Baculogypsina, Baculogypsinoides, Schlumbergerella
plexus, with a tendency to increased size and radial mor-
phology. These last three taxa are only known from the
latest Pliocene to Recent. Adams (1984) has described
Calcarina with Baculogypsina from the Middle Miocene
Futuna limestone of Fiji but he has cautioned that these
could be recent contamination. I have seen Calcarina
spengleri in thin sections from Halmahera occurring with
the distinctive sections of Globorotalia tumida and Glo-
boquadrina altispira, and therefore of Early to basal Late
Pliocene age. Calcarina also occurs in the Karren Lime-
stone of Java and Madura, a unit that has been dated by
strontium isotopic methods as very latest Miocene (6 to 71⁄2
MYBP: Karren Lst with Calcarina (1): 87Sr/86Sr=0.708922
(NBS987 - 0.710235), age 7.5 MYBP, combined precision
and correlation error 6.61 to 9.16 MYBP. Karren Lst with
calcarina (2): 87Sr/86Sr=0.708912 (NBS987 - 0.710140),
age 5.5 MYBP, combined precision and correlation error
5.12 to 5.74 MYBP).

Borelis and Heterostegina are other genera that re-ap-

pear in later Pliocene times, either by iterative evolution,
or possibly, for Borelis, by migration from outside the
Indo-Pacific area. (Borelis disappeared from the Indo-Pa-
cific near the end of Tertiary e but continued to live in the
newly isolated Mediterranean realm.) Planostegina is hard
to recognise unless a particular oblique section is enounc-
tered (it would easily be mistaken for either Operculina or
Cycloclypeus is the plane of section is too close to axial
or equatorial respectively). Good Planostegina has been
found in the latest Miocene Kapung limestone in Central
Java (dated by correlation to oil wells and planktonic zo-
nations). The genus Alanlordia is recorded from a single
location in the T.f2 (Banner and Samuel, 1995) but is
far more frequently seen in latest Miocene through Late
Pliocene carbonates in Sumatra and Java. Similarly the
Biostratigraphy of Indo-Pacific larger forams: page 34

genus Quasirotalia is seen in latest Miocene and Pliocene

carbonate of the Indo-Pacific area.
 Image 24. Upper Tertiary f Limestone - T.f3.
Limestone from the river Cicabe, Margamukti N. C. Java. This site was first descibed by Hetzel (1935) in the mapping quadrangle reports for Sheet 54, Majengang. The sediments here were
mapped as the “Lawak Beds”, consisting of "flaggy" (beds up to 20 cm thick) grey-yellow foraminiferal limestone, interbedded with brittle mudstone and minor volcaniclastic sandstone. Tan Sin
Hok determined the following foraminifera from the limestone: Cycloclypeus indopacificus Tan (with very variable sculpture), Radiocycloclypeus sp., Trybliolepidina radiata Martin, Trybliolepidina
stellata (Scheffen). He dated the sample as Middle Miocene as Cycloclypeus most commonly has 6 nepionic chambers (range 4 to 8). Recent work on nannofossils re-dates this unit as the lower to
mid Late Miocene (lower part of Zone NN11, on the presence of Discoaster quinqueramus, D. berggrenii and Helicosphaera orientalis. Note the large specimen of Sporadotrema, and specimens of
Palaeonummulites near the center.
 Image 25. The radiation of new larger foraminifera in latest Miocene to Recent.
This is the Karren Limestone of NE Java and Madura, where it occurs over an erosional angular unconformity in a generally reefal
facies (hence the meteoric phreatic diagenesis in the above image, with clear equant spar locally becoming drusy mosaic cement). The
limestone has been dated as very latest Miocene by strontium isotopic analysis (c. 6 MYBP). This assemblages contains much mollusc
and bryozoan debris, as well as involute but relatively flat Operculina, Alanlordia and Calcarina. Also in this area Alveolinella and
rare Heterostegina is known.

Biostratigraphy of Indo-Pacific larger forams: page 36

 Image 26. The radiation of new larger foraminifera in latest Miocene to Recent.
This is an un-named limestone found SW of Semarang, N. C. Java, mapped as "N4x" following the field mapping designation for
the early Pliocene, as explained in the section on history. It has been dated from strontium isotopic analysis at about 6 MYBP. This
facies contains abraded bioclasts, but with no evidence for stratigraphic reworking. Amphistegina is common, but thee are also many
specimens of Palaeonummulites, Quasirotalia and a few Alanlordia. At the bottom of the image is Alveolinella.
Biostratigraphy of Indo-Pacific larger forams: page 37
4. CAUSES OF LARGER FORAM growth of ice sheets, and the related production of cold,
deep water masses and temperature stratification of the
ASSEMBLAGE TURNOVER oceans.
As one of the main themes of this report is that the On the figure below the earlest of these (top T.a) is not
Letter Stages are periods equivalent to European Epochs, as obvious as the later events, partly as only emphem-
mostly separated by faunal turnovers, it is a fair question eral ice sheets were forming at the poles (Zachos et al.,
to ask what caused such important events. 2001). However this time was the first development of
significantly different deep and shallow water records
The most abrupt extinctions affecting multiple gen- (Douglas & Savin 1975, Shackleton & Kennett, 1975).
era of larger foraminifera are the top T.a, top T.b and top Also Morley (2000) has pointed out that cooling effects
of the original T.f, now called top Lower T.f for reasons were first seen at this time in the floras of S.E. Asia. One
explained above. After each of these events there was a of the sections where this is seen is the Nanggulan site
radiation of many new forms of foraminifera into un-oc- in Central Java where Lunt and Sugiatno (in press) have
cupied niches, although in the case of the top Lower T.f dated the influx of the southern gymnosperm Podocarpus
non-foraminiferal organisms may also have adapted to (polystachyus type) reported by Morley at the T.a to T.b,
empty niches. top P14 boundaries.
Palaeotemperature / climate The climatic shifts at the Eocene - Oligocene bound-
By far the most convincing correlation to these faunal ary and mid Middle Miocene are well documented (eg.
turnovers is climatic change, notably palaeotemperature. Prothero & Berggren eds., 1992; Zachos et al, 2001).
The three main extinction events correlate to the three Berggren and Prothero (1992) describe how Middle
largest changes in the oxygen isotopic record, and in par- Eocene climate deteriorated in steps through to the Early
ticular the separation of deep-water isotope records from Oligocene, the first major step being close to the Middle
surface (plankton) records. The increase of 18O in deep to Late Eocene boundary, a second step virtually on the
marine fossil tests reflects both loss of lighter isotopes to Eocene-Oligocene boundary and a pronounced isotope

“Lr. Tf” “U. Tf”

Lr. Te

“Th“
Quaternary
Ta1

Ta3
Ta2

Td
LETTER STAGES
Tb

Ur.
Te
Tc

Tf1

Tf3
Tf2
Lt. EOCENE OLIGOC. MIOCENE
SERIES Pal EARLY MIDDLE LATE Ey. LATE EARLY M. LATE
Pl.
87 86
Sr / Sr 0.7090
60 Ma

40 Ma

10 Ma
Beginning of inc-

continental crust
from erosion of
Global oceanographic conditions

Blue line: Strontium isotope 0.7085

rease in Sr87

curve reflecting weathering

of Himalayas (increase in 0.7080
sialic derived Sr87) oceans N. Hemisphere
4 Ratio of Sr87 to Sr 86 in
0.7076
Ice-sheets
Red line: Antarctic Ice-sheets
Oxygen isotope curve, benthic 3
ocean waters, reflecting global W. Antarctic
cooling and development of ice � 18 O (‰) Ice-sheets
Warmer

caps. Solid line from Zachos et al.

2
*Mass extinctions of the *
(2001), dotted line Douglas &
Savin (1975) and Shackleton & 1 * larger foraminifera
Kennett (1975) modified to new
global migration of Lepidocyclinids
*
GPTS. = abrupt deviation of
deep sea from planktonic record
0
L ong term se
a le v el
Biostratigraphy of Indo-Pacific larger forams: page 38

Rising
Black line:
Sea level curves from Vail et al. � 18O Falling
‰ 3rd order curve. 100m

(1977), Haq et al (1987). Scale Short term sea level

at right is modern day s.l. at 0 Peak carbonates 0
across S.E. Asia

Tectono-stratigraphic Lt. Cret. - m. Eoc. m. Eoc. - E. Olig. mid- Olig to Miocene Lt. Mio
base. Mioc. sediment. to Rec.
sequences in sedimentary sedimentary
sediment. megasequ. sediment.
megasequence megasequence
west Indonesian area megaseq. megasequ.
Notes
The calibration of the Letter Stages, Series, isotopic data & eustatic curve to numerical ages is corrected as far as possible to the
GPTS of Cande & Kent (1995) / Berggren et al. (1995). The sharp cooling that is possibly the cause of mass extinction among the
larger forams (=top T.b) is known from DSDP data Wei, 1991) to be basal-most Oligocene, close to base Chron C13n..
The blue shading in Oligo-Miocene times represents the period of widespread carbonate deposition over Sundaland (Batu Raja,
Kujung / Prupuh limestones etc.) and other parts of S.E Asia.
shift close to the base of the normal polarity of magneto-
stratigraphic chron C13 (Mei, 1991). This is now dated as
33.55 MYBP (Berggren, et al. 1995), while the extinction
datums for Hantkenina and Turborotalia cerroazulensis
are 33.7 and 33.8 MYBP according to the same source.
These authors and Aubry in the same volume note how
some organisms such as nannofossils had a greater extinc-
tion event within the earliest Oligocene rather than at the
Eocene-Oligocene boundary. Similarly Keller et al. in this
same volume note how the supposed mass-extinction of
planktonic foraminifera at the Eocene-Oligocene bound-
ary has been exaggerated, and in fact the extinctions follow
a more stepped pattern as climate cools and changes.
The mid-Miocene cooling also appears to be spread
over a few million years. Work by Vincent et al. (1985)
suggests that in latest Early Miocene times deep marine
benthic conditions were at their warmest for the Neogene
followed by a gradual drop until late Middle Miocene cool
times. The fall in palaeotemperatures begins at about 14.6
MYBP and flatten out at about 12.5 MYBP. This slightly pre-
cedes current data on the faunal turnover between Lower
T.f to Upper T.f assemblages. However, as discussed in
the text and the entry for Miogypsina there is still work
to be done precisely fixing the extinction datums of the
foraminifera at this time, as in much of Indonesia the
record is overprinted by a regional tectono-stratigraphic
event and an inadequate fossil record due to facies
changes. This apparent slight mis-alignment of the data
does not weaken the case for the link between palaeo-
temperature and faunal turnover, as the three outstanding
events in both data sets have a one to one relationship for
40 million years.
Strontium isotopes
Strontium isotope ratios are shown on the chart on the
previous page not because they have any relationship to
the causes of faunal change in larger foraminifera, but
because they are a tool that can help clarify this problem.
Ever since the base Oligocene the 87Sr/86Sr ratio in sea-
water and marine calcite tests, has a single gradient, so
that measuring this ratio in fossils can directly date unal-
tered calcite or aragonite fossils. Such work has already
helped calibrate the Letter Stages to the GPTS and future
work will help in more precise correlation around times
of climate change. Notes elsewhere in this report suggest
that top Lower Tf may be slightly older than currently
thought based on several dates of the terminal T.f event
in Java represented by the Platten Limestone closer to 14
MYBP than 12 MYBP.
Eustasy
Also shown in the figure overleaf are the long and
short term sea level changes of Haq et al. (1987). At one
time is was thought possible that the outstanding sea-level
fall in mid-Oligocene times might have initiated oceano-
graphic changes that allowed Eulepidina, Lepidocyclina
and Neorotalia mecatepecensis to migrate out of Central
America to eastern Tethys. New biostratigraphy with
strontium isotope dating at the Ciapus and Cimanggu
sections in West Java (Lunt et al. in prep) now dates this
migration event as old as 32 mybp. This is significantly
before the "30 MYBP" event that has been corrected to the
Cande and Kent (1995) / Berggren et al. (1995) GPTS at
about 28.5 MYBP.
The sea level curve does, however, correlate with a
S.E.Asia wide transgression and abundance of carbonates,
between Late Oligocene and early Middle Miocene (cf.
Netherwood 2000). During this ten million year period the
larger foraminifera evolve at a steady rate. Only the ex-
tinction of Spiroclypeus and Eulepidina (top T.e) requires
explanation. This must wait until samples un-affected by a
regional tectono-stratigraphic event and unconformity can
first demonstrate if the two extinctions are simultaneous,
and then accurately date them for comparison with the
timing of other events.
SUMMARY
The original Letter Stages have survived some sev-
enty years of usage with minimal change, and appear to
represent fundamental Tertiary biostratigraphic periods
reflecting faunal turnover, migration (for base T.d) and
evolutionary development in the larger foraminifera.
This is an interesting contrast to mollusc based
stratigraphy where there are no recognised faunal turno-
vers, and the percent extinct to extant method still appears
valid where large enough faunas are found. If this contrast
is found to be true then it is curious that complex meta-
zoans such as molluscs, which are more likely to evolve
through punctuated equilibrium, change as a fauna in a
gradual manner, yet protozoans such as foraminifera,
with good examples of gradual evolution in individual
lineages, evolve as a fauna though stages of punctuation.
Possibly the life history strategies of the extreme K-se-
lected larger foraminifera, in highly stable environments
that periodically collapse, may be the answer. In that case
it emphasises the importance of life history strategies as
Biostratigraphy of Indo-Pacific larger forams: page 39
fundamental characters of species that can be inferred
from stratigraphic evidence.
Increased integration of planktonic zonations and
strontium dating will better improve our biostratigraphy
of carbonates, and such work will also have important
applications in palaeoclimatic studies, and models of how
different organisms respond to climate changes. The larger
foraminifera remain a unique, and still understudied group
of organisms where a detailed evolutionary history is sen-
sitive to both environmental and stratigraphic changes.
5. NOTES ON THE MAJOR TAXA, BY GENUS
Following the method of Adams (1970, 1984), the following is a list of genera with notes on their
stratigraphic ranges and other comments, modified with new data and focused more on forms found in the
Indo-Pacific area. Note that the definitions of epoch boundaries do not always follow Berggren et al (1995)
who adjust these to European based stage or age definitions. Instead the SE Asian oil industry accepted
defintions based on microfossil datums are used, namely; top Middle Eocene is the multiple extinction at
the top of Zone P14, top Early Miocene is the Orbulina datum (base N9), top Middle Miocene is the top of
N14, extinction Neogloboquadrina mayeri, top Miocene at the top of NN11 or CN9 on nannofossils, which
is very close to base N18 on foraminifera.
Data from strontium isotopic analyses are from a number of laboratories but all ratios of 87Sr / 86Sr are
corrected to a a value of 0.710235 for the Standard NBS 987 (after 87Sr / 86Sr normalised to 86Sr / 88Sr =
0.1194). Instrument error is used to determine high and low ranges of precision and the median and both
error values are converted to numerical ages via a calibration curve based on McArthur et al. (2001) to give
a median age, plus and minus a combined analysis and calibration error range. This gives values that should
be compatable with the time scale of Berggren et al. (1995) used for planktonic biostratigraphy.

Acervulina Schultze, 1854

Type species Acervulina inhaerens Schultze, 1854
The type species has been recorded in sediments as old as Early Miocene, and is extant. Becomes more
common after end Tf2 carbonate faunal turnover.
Test attached, early chambers coiled, later irregularly encrusting. No aperture but coarse perforations
in chamber walls. Similar to Gypsina except the chambers are vermiform - irregular and somewhat
contorted.

Alanlordia Banner and Samuel, 1995

Type species: Alanlordia niasensis Banner and
Samuel, 1995
The type location for this genus is in the
later Pliocene of Nias Island near Sumatra
and it is also found in latest Miocene and
Early Pliocene limestone on Java. Banner
100µm
and Samuel also illustrate Alanlordia from
the Middle Miocene Tf2 on Banyak Island.
An apparent homeomorph of this test type
occurs in the Middle Eocene T.a of Central Java. Above and left. Specimens of
Alanlordia from the latest Miocene
Karren limestone in NE Java (Kali
Suwuk, 2K1/09/02) illustrating
the asymmetrical formation of
cubiculae, with well formed spines
and coarse pores in the later
Biostratigraphy of Indo-Pacific larger forams: page 40

cubicular walls.
Date of this locality; 7.56 MYBP ±
combined precision & correlation
error 6.61 to 9.16 MYBP. Repeat
analysis from sample on strike =
5.46 with error ranging 5.123 to 5.74
MYBP.
Alveolina d'Orbigny, 1826
= Fasciolites Parkinson 1811. Alveolina nom. conserv. ICZN 2356 (petition pending). Some books e.g. Van
Bemmelen (1949) used the name Borelis (Fasciolites).
Type species: Oryzaria boscii Defance in Bronn, 1825
First appears in the later Paleocene (Serra-Kiel, 1998), last occurrence at the top of Ta3, close to the
Middle to Late Eocene boundary, as defined by the extinction at the top of P14
An often thick-walled (flosculinised) fusiform miliolid that has a senior name of Fasciolites, but this
name has been proposed to be suppressed. In the scarcity of older sediments in the Indo-Pacific region
the presence of this genus is often used to indicate Ta3. The strongly flosculinised forms are mostly
of this age, but the genus does occur in older rocks. An important diagnostic feature is the presence of
both pre and post-septal passages. More examples of Alveolina are on Image 8.

An Alveolina limestone from East Indonesia (Lengguru fold belt, Irian Jaya). Note the well developed thickening of the chamber
floor (flosculinization), and the dominance of this one genus, with a few smaller or juvenile miliolids.
The field of view is 14 mm across. These dark (porcellanous) bioclasts are bright white in reflected light.

Off-center equatorial sections of

Biostratigraphy of Indo-Pacific larger forams: page 41

Alveolina. In several instances the

diagnostic postseptal passage can
be distinguished. Enlarged from
image above. The specimen is
about 2 mm in diameter.
Alveolinella H. Douville, 1907
Type species: Alveolina quoyi dʼOrbigny, 1826 (as quoii, nom. imperf.)
First appears as intermediate forms between Flosculinella and Alveolinella in latest Letter Stage Tf1,
before the extinction of Austrotrillina (well illustrated in Eames et al. 1962, Plate VI, Darai imestone
Papua New Guinea) close to top Early Miocene, as Alveolinella fennemai or A praequoyi (see figure,
next page). The CSIRO Sr laboratories have noted samples with Alveolinella older than the Orbulina
datum, at least as old as 87Sr/86Sr = 0.708700 or >16.6 MYBP. The more distinct A. quoyi form, which
is for most of its range the only recognised species of Alveolinella, first occurs near the base of upper
Tf (=Tf3).

Alveolinella (right of center and bottom left) from the Tf3, Late Miocene of the Lengguru Fold belt Papua. On the left-centre is a
specimen of Marginopora. Field of view about 7 mm across

Amphistegina d’Orbigny, 1826

Type species: Amphistegina quoyii dʼOrbigny, 1826 (= A. lessonii; see Loeblich and Tappan, 1987)
Probably evolved in early Eocene times but a minor component of faunas until the Late Eocene /Tb,
becoming a cosmopolitan carbonate facies form from Tc or Td times. Extant, with many species defined,
but hard to assign consistently reliable species names in random thin sections of such a simple test.

Biostratigraphy of Indo-Pacific larger forams: page 42

Axial section through a specimen of Amphistegina, about 1.5 mm across, showing the development of a secondary septum or
toothplate (right, in final chamber) that extends from the previous apertual face to about half way down the ventral side dividing the
chamber lumen. This, plus the resulting slight asymmetry, which may include a more pronouced umbonal pillar on the ventral side,
and a more acute margin, distinguishes this form from any species of Nummulites or Palaeonummulites.
 ALVEOLINELLA &
FLOSCULINELLA
Four images from same sample,
Lengguru fold belt of Papua. Left are
two enlargements from the central image,
which has a 7 mm field of view.

upper or outer tier

additional tier
appearing
lower or principle
tier the largest
chamberlets
Biostratigraphy of Indo-Pacific larger forams: page 43

The Alveolinella (bottom) has acquired more than the two layers of
chamberlets found in the ancestral Flosculinella (top). Note how the
outermost tier of small chamberlets in each chamber is more regularly
aligned than those below it. This is a feature retained from the
Flosculinella ancestor.
Specimen to right: oblique section showing multiple tiers of chamberlets
but still a dominant inner set and small regular outer set; intermediate
between the two generic morphotypes. This form is Alveolinella
praequoyi.
Archaias deMontfort, 1808
Type species Archaias spirans de Montfort, 1808 = Nautilus angulatus Fichtel & Moll, 1798
Range: Adams (1970) suggests from sparse data that the oldest Archais worldwide is mid-Eocene, but
in Southeast Asia it is recorded very rarely, the oldest being Early Oligocene, Letter stage Td. The ge-
nus is extant. It is more commonly encountered in Tf3 sediments, but still scarce, even in the extensive
carbonates of this age in the Papuan region.

Nephrolepidina

Pillars

Proloculus

Archais from the a Miocene limestone in Papua (Irian Jaya). Note the involute coiling and the pillars between septa. This specimen is
slightly over 2 mm in diameter and has a rather large proloculus. The relatively small size of this specimen and the thin nature of the
most mature chambers (upper right) suggests this specimen in Archais and not Pseudotaberina. This generic difference is best seen in
equatorial section.

Assilina: see Planocamerinoides

Asterocyclina Güembel 1870

Type species: Calcarina? stellata dʼArchiac 1846, syn: Asterodiscus pentagonalis Schafhautl 1863
First occurrence: In the Mediterranean as old as Early Eocene (Fermont, 1982) The summary of
Drooger (1993) suggests it may occur in the Paleocene.
Last occurrence: Top of Letter Stage T.b, very near the top of the
Eocene. An abrupt extinction considering how abundant this
genus was in carbonate facies up to this event.
Biostratigraphy of Indo-Pacific larger forams: page 44

Serra-Kiel et al (1998) show a fine zonation based on species

of Asterocyclina but such a scheme has yet to be proven in
SE Asia, especially in rocks examined in rhin section. There
is visible variation in morphological characters, both in the
size of the embryont and the form of the test as a whole but
none of these seem yet to have stratigraphic significance and
could be entirely ecophenotypic.

Asterocyclina, sketch of stellate body plan and the multi-layered thickening of the median layer
that produces it
Austrotrillina Parr, 1942
Type species: Trillina howchini Schlumberger, 1893
First occurrence: In both the Middle East and Indo-Pacific areas the genus is first recorded in mid-
Oligocene Letter Stage T.d (Adams, 1965, 68). It is rare in these beds and is more usually seen In Letter
Stage Te and overlying strata. It is seen in Te1 beds in outcrop in NE Java. Tony Allan has recently found
several Austrotrillina specimens in rich Ta, Middle Eocene faunas in parts of Papua New Guinea.
Last occurrence: At top of Letter Stage Tf1. Continues after the appearance of Orbulina at base Middle
Miocene (Adams 1968). Note that modern work such as that by Adams (1968, 1984) gives discrete
ranges to species of Austrotrillina, based on increasing complexity in the alveolar structures. The transi-
tion from simple to branching alveolae (A. striata to A. howchini) occurs close to the top of Te.
Work prior to 1968 works (such as Dutch mapping of Indonesia, and van Bemmelen 1949, Marks
1957) uses the taxon A. howchini for what was then the sole species in the genus, not the more narrowly
ranging A. howchini sensu Adams and others. See Image 21 for an example of the modern concept of
A. howchini.

Austrotrillina striata, with simple, undivided alveolae, next to a specimen of Tansinhokella,

from a core in the Te of Dermawu-1 well, Java

Baculogypsina Sacco, 1893

Type species: Orbitolina concava Lamark var. sphaerulata Parker and Jones, 1860
A Pleistocene to Recent form, found in high energy environments (Cole 1957 -Saipan, Cole 1963). It
is mostly found in the Pacific Islands (eg. Saipan, Guam) where is can form calcarenite sands. Differs
from Baculogypsinoides as it has 5-7 spines roughly in a single plane, whereas the later has a larger
test, and four spines arranged
in a tetrahedron.

Right, Baculogypsina with the larger

Biostratigraphy of Indo-Pacific larger forams: page 45

Calcarina and a frament of a soritid

miliolid (top) as a stereo-paired view of a
Pacific beach-sand.
Baculogypsinoides Yabe & Hanzawa, 1930
Type species: Baculogypsinoides spinosus Yabe & Hanzawa, 1930
Ranges Pleistocene to Recent. There are some records of Baculogypsinoides from later Eocene (T.b,
e.g. Samata 1978 in west India / Pakistan) but these are the species B. tetraedra which are now placed
in Silvestriella.
Hanzawa (1952) has stated that the juvenile of Baculogypsinoides cannot be distinguished from small
Calcarina specimens. Baculogypsinoides has a similar west tropical Pacific distribution to Baculogypsina
and is most common in high energy Pacific island locations.
Below. Baculogypsinoides from Loeblich & Tappan 1987. On the left is a juvenile resembling Calcarina, the other two figures are of
the adult tetrahedral form.

1mm

1mm

1mm

Biplanispira Umbgrove 1937

Type species: Heterospira mirabilis Umbgrove 1936 (“Heterospira” already used for a Triassic gastropod
so Umbgrove selected the new name Biplanispira in 1937).
Ranges within Letter Stage Tb. Specimens with later adult stages of biserial chambers either side of a
continuous marginal crest are found in what is thought to be the lowest Tb outcrop on Java at Cimuncang
(Koolhoven,1933 restudied by the writer), occuring here with P15 planktonic foraminifera, granulate
Nummulites species. The nearby Cicarucup section, mapped as slightly higher in the Late Eocene, contains
better developed Biplanispira specimens. Unlike the ancestral Pellatisipra, there are no known examples
of this genus in latest Ta sediments. Biplanispira mirabilis is only known from the Indo-Pacific region,
whereas B. absurda is known from across Tethys, as far west as Spain (Hottinger et al 2001).

Biplanispira absurda,
from a xenolith ejected
from a mud volcano in the
Sangiran Dome, Central
Java. The upper, smaller
specimen is a Pellatispira-
like juvenile stage with a
Biostratigraphy of Indo-Pacific larger forams: page 46

well developed marginal

crest.The lower specimen
is larger, relatively flat, and
has an obviously bi-serial
set of chambers either side
of the marginal crest.
Borelis de Montfort, 1808 = Nautilus melo Fichtel and Moll, 1798
Neoalveolina A. Silvestri, 1928 is a junior synonym.
Type species: Borelis melonoides de Montfort, 1808 = Nautilus melo Fichtel and Moll, 1798
First appears in Tb, Late Eocene but rare. Many good specimens found by the authors with Discocyclina
and Tb Nummulites in Papua New Guniea [IO Asta 29/2]. Note that many apparent records in old Dutch
reports, especially Van Bemmelen, are clouded by the usage of Borelis meaning Borelis (Fasciolites) [=
Alveolina] occuring with Flosculina [= Alveolina with flosculinisation] in what in the late 1940ʼs was
a lumped T.ab, mid to late Eocene Letter Stage (sensu Rutten, 1947). Cole (1957) reports rare Borelis
in T.b assemblages in the Pacific island of Saipan, although not in his 1953 report on the same area.
Adams (1970) regards this as the only authentic T.b record, but notes the genus occurs in the Eocene
in Europe.
Borelis is common in Early Oligocene (T.c) with Nummulites fichteli-intermedius, before the appearance
of Eulepidina / Lepidocyclina (pers obs. in several oil wells NE Java). Borelis inflata has been found
in outcrop near the Kujung-1 well in NE Java, in T.d sediments at the base of the Late Oligocene (just
above extinction Chiloguembelina cubensis, near samples with some of the youngest known Nummulites).
The type location of Borelis pygmaeus Hanzawa is the Rajamandala limestones just west of Bandung,
with a Te1 fauna (Hanzawa 1930).
Last occurrence: Borelis schlumbergeri and B. pulchra are living in the Indo-Pacific today. However
there is a possibility these have migrated or evolved from a different source. After Borelis pygmaeus in
Upper T.e, Borelis is not recorded from sediments in the Indo-Pacific area until the very end of Letter
Stage T.f. The sub-spherical species Borelis melo is found in lower Tf (Early Miocene) sediments the
western Indian Oocean but not reliably in the Indo-Pacific. Eames et al found B. melo in Kenya and the
Middle East at this time, along with Flosculinella. Adams (1984) notes that it is likely that closure of the
Persian Gulf, separating the western Tethyan
/ Mediterranean area from the Indo-Pacific in
mid Early Miocene times (near basal Tf) may
account for the fact that after this time Borelis
is found in the Mediterranean but not Indo-Pa-
cific, and the descendant Flosculinella is found
in the Indo-Pacific but not Mediterranean.

Borelis from two locations in Tc of Java. With the single

row of chamberlets in each chamber / whorl there is a
strong resemblance to Alveolina, however the upper left
image shows clearly that there are passages (in & out of
the plane of the section) behind the septa but no post-
septal passage
Biostratigraphy of Indo-Pacific larger forams: page 47
Borodinia Hanzawa, 1940
Type species Borodinia septentrionalis Hanzawa, 1940
An encrusting foraminifera similar to Acervulina, range uncertain but defined in the Early Miocene of
the Philippines. Hanzawa and Hasimoto (1970) illustrate an example where Acervulina and Borodinia
are growing together so the Borodinia can be seen to have more inflated chambers with more pinching
and swelling. Also thought to have stolons connecting adjacent chambers, unlike Acervulina which has
coarse pores only. However these stolons are not always obvious in Borodinia, and gaps in the cement-
ing of Acervulinid chambers can look like stolons.

Calcarina d’Orbigny, 1826

Type species: Nautilus spengleri Gmelin,1788
Has been found with Globorotalia tumida and Globoquadrina altispira (=Early Pliocene) in east Indo-
nesia (Halmahera). Adams 1984 regards with suspicion his own observation of several Plio-Pleistocene
genera mixed with Middle Miocene carbonates in Fiji, regarding the location as possibly young beach
sands filling old karst cavities. The oldest locations known to me are the Karren and Kapung limestone
in northeast Java, which, as noted under the entry for Alanlordia, are dated by strontium isotopes as
latest Miocene, about 5.5 to 7.5 MYBP, an age that matches off-reef chalky sediments above the initiating
unconformity dated as mid N17, mid NN11.

Calcarina from the Latest Miocene

Karren limestone above Kali Suwuk,
NE Java. Note the trochospiral test
and robust spine. Specimen is 2.5 mm
across.

Cellanthus Montfort, 1808

Type species: Nautilus craticulatus Fichtel & Moll, 1798
Both Rögl & Hansen (1984), redescribing type material, and Loeblich and Tappan (1987) consider
Cellanthus as a junior synonym of Elphidium. Haynes (1981) kept the genera separate on the basis of
different degrees of development of the septal flap. Both Elphidium craticulatum and Cellathus crat-
iculatus are valid names depending on if one ranks the reported slight variability in septal flap and its
outstandingly large size compared to the rest of the Elphidiidae as either of specific or generic rank.
After some time studying larger foraminifera I am biased to rank size as an important property of an
Biostratigraphy of Indo-Pacific larger forams: page 48

organism, especially the magitude of difference seen in the robust test of Cellanthus. I would therefore
interpret the more restricted geographic range of Cellanthus to fully marine, low latitude environments
as evolutionary specialisation rather than ecophenotypic gigantism. Note that this genus is more often
found in non-carbonate sediments than most larger foraminifera. Ranges from Eocene to Recent.

Chapmanina
Adams (1970) noted that the majority of records of this Tb genus were from East Africa and the Middle
East and Europe. He checked and discounted some old reports from Borneo. His conclusion that this
genus may never have reached the Indo-Pacific area still seems a valid conclusion.
Cycloclypeus Carpenter 1856
Type species Cycloclypeus carpenteri Brady, 1881
Ranges Tc to Recent. While species of this genus are better understood than most, they are still difficult
to identify in rock thin-section. The exception is Cycloclypeus annulatus (Katacycloclypeus annulatus
of some) with its distinctive annular thickenings, which is restricted to upper Tf1 and Tf2. The single
record suggested by Adams (1984) for Katacycloclypeus in Te5 was based on Hashimoto et al. 1977b
from the Angat Limestone in Luzon. While this limestone is placed in Te on the correlation figure, all
the samples with records of Katacycloclypeus noted in the text of this paper are without Te markers and
contain Tf forms such as Austrotrillina howchini and Miogypsina polymorpha.
Evolution is from the series Operculina - Planostegina - Cycloclypeus, with remnants of the ancestors
preserved in the juvenile of the descendant. Throughout its evolution the embryonic pair of cham-
bers increases in size, the Operculinid growth phase rapidly disappears, and also the Heterosteginid
(=Planosteginid) phase becomes much reduced. Dimorphism between macro- and microspheric forms
is significant only in the Miocene (Tf letter stage) to Recent forms. In Oligo-Miocene populations the
size overlap between micro- and macrospheric forms has been suggested (MacGillavry, 1962) to be the
cause of confusion in biometric analysis of older assemblages.
The lower Tf (later Early Miocene through to mid Middle Miocene) was a period of particular abundance
of Cycloclypeus in Indonesia with the locally abundant Cycloclypeus annulatus. Initial observations on
the annulatus species suggest it has the same morphometric values as the non-annulaus tests (see below)
and hence I consider, for now, annulatus as a minor species variant rather than a separate genus.
In Late Miocene times C. carpenteri, with its large embryont, is the domiant form, along with variations
such as stellate types (Radiocycloclypeus of Tan), and forms with partings or pseudo-lateral chambers
in sidewall laminae (Kali Lawak, N.C. Java, of mid Late Miocene age, lower part of Zone N17 before
evolution of Pulleniatina unpub. obs. on same beds as original work of Tan 1932). Cycloclypeus is

Biostratigraphy of Indo-Pacific larger forams: page 49

Cycloclypeus koolhoveni or oppenoorthi Tan (the former has a central boss not visible in equatorial section) from the para-
type location at Ciapus, West Java, associated with Tertiary d foraminifera and with Early Oligocene (P20-P21a) planktonic
foraminifera.
This specimen has about 27 nepionic chambers [1 operculine, and about 26 heterostegine chambers], before adult annular
growth is attained. Note the proloculus is about 150 microns in diameter, which distinguishes C koolhoveni / oppenoorthi from
the Late Oligocene C. eidae forms which have small proloculi about 80-90 microns in diameter. Image 3.0 mm across.
 Well preserved mid Middle Miocene bed within the Ngrayong sandstone East Java (Sample GW-6, planktonic Zone N10-11). Left:
external view showing fine pustules in typical cycloclypid arrangement, as well as boss above embryont. Right: broken inner surface of
Cycloclypeus eidae, with 12 nepionic chambers, a number comparable with Tan Sin Hok's Kebon-Matingan location which is nearby.
The Ngampel location of Tan is a limestone strata overlying this location, without planktonic species but thought on other data to be
less than a million years younger, where the modal number of nepionic chambers is only 6. This specimen with a small embryont,
about 80 microns, is thought to be a late member of the small embront C. eidae lineage. Subsequent specimens with protoconchs 200
µm or more in diameter are assigned to C. carpenteri.

locally abundant in Pliocene and younger carbonate facies limestones (e.g. some facies of the Kapung
Limestone, Central Java).
From the Early Oligocene to Recent there is a well documented and apparently gradual reduction in
the number of nepionic chambers in macrospheric forms (work pioneered by Tan Sin Hok, 1932). In
Early Oligocene there are typically 30 to 24 nepionic chambers in a very obviously heterostegine /
planostegine juvenile. In Late Oligocene times this growth stage had reduced to 24 to 18 chambers, in
Early Miocene Upper Te, to between 17 and 15. In lower Tf (later E. and ey. Middle Miocene) 14 to 6
nepionic chambers are typical, and the small juvenile test no longer appears distinctly heterostegine.
Later Middle Miocene to Recent forms usually have only 3 to 5 nepionic chambers before full, adult
annular growth is achieved.
While this progression sounds gradual, it probably is not. Tan proposed a model of minor saltations with
an overall smooth transition, although his minor saltations were not found by later workers (Drooger
1955, or MacGillavry 1962). However these latter workers were active before progress in planktonic
or nannofossil biostratigraphy allowed accurate age dating of samples not taken in sequence, and they
could only estimate age for the measured biometric values. On-going work on samples dated with both
planktonic biostratigraphy and SIS dating strongly suggests the evolution of Cycloclypeus has significant
periods of stasis then rapid change.
Biostratigraphy of Indo-Pacific larger forams: page 50

Regardless of the probability of saltations or punctuations, an important point that Tan Sin Hok and
following workers appear to have demonstrated is an orthogenetic trend, comparable to that seen in
other larger foraminifera. That is, a trend that is apparently independant of external factors, directing
evolution over nearly thirty five million years. This is the sort of macroveolutionary trend that Schinde-
wolf (1950) described in Ammonites and other metazoans, and was derided as anti-Darwinian by the
prevalent Anglo-American, strict NeoDarwinist schools of evolutionary thought.
As a consequence of the evolutionary relationships described above, it is often very difficult to distin-
guish Planostegina / Heterostegina from juvenile stages of older specimens of Cycloclypeus, especially
in random thin sections. On the other hand it is easier to identify the adult Cycloclypeus stage even in
small fragments because of a change in the canal system. Both Planostegina and juvenile Cycloclypeus
have an intraseptal canal system based on a marginal cord derived from the Nummulitid ancestor. Upon
Figure (Plate III, re-coloured) from Tan Sin Hok 1932 showing variation in the embryonic and nepionic chambers of Cycloclypeus
specimens at the same magnification (except number 2 which is 2.8 times as large as the others).
Fig 1. Center of a Cycloclypeus indopacificus, protoconch coloured and the first part of the subsequent whorls that bears a marginal
cord also coloured (a = apertures). The 7 nepionic stages are highlighted red and showing some of the stolons or passages (p)
between chamberlets.The last whorls shown are the first of the annular neanic stages. M = end of marginal cord. F and L =
first and last chamberlets of neanic stages.
Fig. 2. Detail showing canal system in the embryont developing into the marginal cord (M .... M) in the nepionic chamberlets
Fig.3. Center of a very advanced "Katacycloclypeus" (C. annulatus) with an unusually large embryont (P1 & P2), and only 2 rows of
nepionic chamberlets. Note the additional stolons (o) from the deuteroconch (P2) communicating with later chamberlets.An
unusual feature of this form is the aperture from the first nepionic stage (a1) is coiling in the reverse direction to the aperture
(a) from the deuteroconch (P2)
Fig. 4. Center of a Cycloclypeus indopacificus with 5 nepionic stages.The fifth nepionic stage (R) is interupted.
Fig. 5. Center of an advanced microspheric Cycloclypeus with a protoconch (coloured) and then 10 small operculine chambers
(including an indistinct secondary chamber that is therefore not called a deuteroconch by Tan Sin Hok), after which there
are 11 heterostegine chambers, making a total of 21 nepionic stages.
Fig. 6. Center of a very advanced megalospheric Cycyloclypeus (C. cf. guembelianus) with 3 nepionic chambers. Note the incomplete
Biostratigraphy of Indo-Pacific larger forams: page 51

or irregulat neanic cycles (R).

acquisition of annular chamber growth the marginal cord is replaced by what Tan Sin Hok called a
septal cord as it is located in the primary chamber septum. This canal system, and a related system
within the secondary septa, are of paired or twinned canals, often clearly visible in thin section.
Cycloclypeus is mostly an Indo-Pacific and Australian genus which appears to have always favored a
deep photic environment. There are frequent Mediterranean fossils but these are mostly in the Oligocene
and possibly earliest Miocene, after which it seems to disappear from this western area (NB the clo-
sure of the sea link between the Mediterranean Sea and Indian Ocean was at about this time). Middle
eastern records are rare. Murray (1987), summarising living larger foraminifera, shows Cycloclypeus
to be recorded as far west as the Maldive Islands and as far east as mid Pacific atolls.
 High magnification photographs of the canals system in neanic chambers of
Cycloclypeus. Left, shows the paired canal system within the septal wall, and also
some faint pores in the thin lateral walls. Below, an oblique section shows the paired
canals in both primary and secondary septa, as well as a stolon or intercameral
foramina between sucessive chamberlets.

STOLON

Cycloclypeus, from the Middle Miocene "Platten" limestone at Ngampel -

Lodan area. The fauna is dominated by Cyclcoclypeus annulatus with the
image above also having L. (Nephrolepdina) ferreroi. The close up of the
embryont shows the large embryonic chamber pair and 5 or 6 juvenile
(ana-nepionic) chambers before the annular growth stage.
Black dot is 100µm.
Biostratigraphy of Indo-Pacific larger forams: page 52

Cycloclypeus appears to be an environmentally specialised larger foraminifera. The genus was first
discovered in “water of a considerable depth off the coast of Borneo” by Carpenter in 1856. During a period
of prolific documentation all forms of living creatures, Cycloclypeus was the last major genus of liv-
ing larger foraminifera to be found [cf. Peneroplis and Sorites (1775), Calcarina (1791), Operculina
and Amphistegina (1798), Heterostegina (1826), Marginopora (1830), and alveolinids (1839)]. All the
 other genera can occur in near reefal or sea grass communities, at relatively shallow depths of less than
about twenty meters, and hence are easily found. The second recorded occurrence of Cycloclypeus
was the now invalid species Cycloclypeus guembelianus (synonym of C. carpenteri) which was more
accurately documented by Brady, in 1881, as coming from a dredge in two hundred and ten fathoms of
water (388 meters) off the coast of Fiji.
Cushman (1921) found abundant Cycloclypeus as shallow as 24 fathoms in the Buton Strait, Philip-
pines, and continuing as deep as 565 fathoms. The Siboga Expedition dredged them from one instance
at 31-36 metres but also as deep as 1595 m (See Tan Sin Hok, 1932. p.80). Chapman (cited in Tan Sin
Hok 1932) found them in comparative abundance from 50 to 200 fathoms around the Funafuti Atoll,
Tuvalu. Cushman et al. (1954) found Cycloclypeus only in dredges around the Marshall Islands, in water
depths of 580 to 800 feet (175 to 250 meters), where it could be very abundant, and these specimens
tested positive for live protoplasm with rose bengal stain. In contrast the lagoonal, beach, and shallower
marine areas (less than a hundred metres deep) around the Marshalls contained no Cycloclypeus.
In a study of the distribution of larger foraminifera in the northwest South China Seas (Li & Wang, 1985)
the presence of Cycloclypeus in the vicinity of the study islands was noted, however the detailed data in
the main part of their study, from 67 samples in water less than 20 meters deep, records no Cycloclypeus.
These samples cover a wide range of facies from reefal and near reefal, to carbonate beach sands.
The shallowest published occurrence of Cycloclypeus in recent studies is probably Leutenegger (1984),
in a sample from the Maldive Islands taken from seventy meters.

Dictyoconus Blanckenhorn, 1900

Ranges from Mesozoic into Tc
Dictyoconus is a high conical arenaceous form, up to 5 mm in size, with both vertical and horizontal
plates in the marginal (cortical) zone, and a central area with a moderately regular spongiform pattern
of intra-septal pillars. The coiled juvenile chambers contain a notable proloculus. Wide circular open-
ings connect successive layers and form the apertures..
Outside Southeast Asia Dictyoconus occurs in beds as old as Early Cretacous, but in the Indo-Pacific
area it is rarely recorded. Adams' (1965) observations of the form D. melinauensis in Boreo is the only
known example from the Early Oligocene (T.c). In the Eocene the species D. chimbuensis Binnekamp
(1973) has been recorded from Papua New Guinea.

Discocyclina Güembel, 1870

Type species: Orbitolites prattii Michelin 1846
Ranges from Late Paleocene to the top of Tb, effectively top Eocene. During this period it is a common
to abundant and cosmopolitan member of the tropical carbonate foraminiferal assemblage.
Work by Brönnimann (1940,1945) found that there were two major lineages within the group, based
Biostratigraphy of Indo-Pacific larger forams: page 53

on the juvenile stage of the microspheric form, as he could find little meaningful variation in the
macrospheric generation. These were his families Discocyclinidae and Orbitoclypeidae. Since then
Less (1987, 1997) has established a diverse taxonomy and biostratigraphy on the two groups, for the
European region. This work has yet been tested in the Indo-Pacific area where forms are usually just
lumped as "Discocyclina" and very few species have regular distribution, with the possible exception
of Discocyclina omphalus (var "selliformis" of some) which has a slight central thickening (centrum)
with a marked depression in the center (omphalus = navel). This morphotype is known from many Tb
locations on Kalimantan (van Bemmlen, 1949, p. 138, 139) but not Ta sediments there, or on Java.
Fragments or random thin sections through specimens can sometimes be confused with other orbitoids
with thin median layers, especially Lepidocyclina (Nephrolepidina). The diagnostic feature is the invari-
ably rectangular shape of the median chambers which is visible in equatorial or oblique sections.
 Discocyclina, a specimen in near-equatorial section, showing the annular septa with a smoothly sub-circular outline and the thinner
radial septula forming rectangular equatorial chamberlets diagnostic of this genus. The fine, scaly texture of the lateral chambers
seen in the upper right are also typical of the genus and differ from the younger homeomorph L. (Nephrolepidina).

Discogypsina A. Silvestri, 1937

Type species: Discogypsina vesicularis A. Silvestri, 1937 = Tinoporous vesicularis (Parker & Jones) of
Goës, 1882
Ta to Recent. A simple discoidal form with a weak 3-layered symmetry in which the middle layer is
usually not very well distinguished from the lateral cubiculae, except in some Middle Eocene (Ta) to
Early Oligocene (Tc) examples. These older forms are Discogypsina saipanensis (Hanzawa) in which
a middle layer is dominant, sometimes increasing
in thickness towards the periphery, with relatively
small numbers of lateral cubiculae. Wall fairly
coarsely perforate, no stolon system. The type
location for D. saipanensis is the Matansa Fm.
on Saipan Island, in association with T.b fossils in-
cluding Biplanispira absurda and B. mirabilis.

Biostratigraphy of Indo-Pacific larger forams: page 54

Two specimens of Discogypsina saipanensis

(Hanzawa) from limestone thought to be
later Ta age, Papua New Guinea. Note the
fragments of Distichoplax biserialis.
Specimen top right 1.4 µm, above 1.67 µm.
Right: Discogypsina vesicularis from
Northwest Kangean Island. In Late Oligocene
deposits characterised by very large, platy
Eulepidina (right) and Cycloclypeus (left). Note
there is less distinction of the median layer
than in D. saipanensis.
Eulepidina H. Douville, 1911
Type species: Orbitoides dilatata Michelotti, 1861
First appears in the Cimanggu section of West Java in the earlier part of zone P19, shortly above the
highest Tc bed dated by SIS as 31.9 MYBP (86Sr/87Sr = 0.707897, analytical and calibration error ranges
age from 31.41 to 32.31 MYBP, work in progress). From its earliest appearance it is common to abundant
in larger foraminiferal assemblages and its extinction, with Spiroclypeus, defines the top of Te. This
extinction is not yet well defined but according to Allan et al (2000) and subsequent work the young-
est 86Sr/87Sr values for whole rock limestonewith Eulepidina and Spiroclypeus are around 0.708450
with is close to 20 MYBP, assuming the youngest such samples are not affected by diagenesis, which is
a significant risk for defining the limits of composited sample sets not collected in order. Unpublished
work with Allan from the NE Java Kali Suwuk section has a high number of samples with Eulepidina
up to at least 0.708360 (c. 21.4 to 21.5 MYBP, plus or minus less than 0.5 MA as multiple samples repeat
closely). The planktonic foraminiferal zone at this point is N4 with Globorotalia kugleri. Slightly up
section, but across a major sequence boundary samples are Tf Letter Stage, Zone N5, and the single
sample analysed for Sr has a value of 0.708416 (20.45 MYBP with analytical and calibration error giving
an age range from 20.1 to 20.83 MYBP). Until higher resolution work is available the working range for
Eulepidina, and T.e, is therefore considered to range from about 31.7 MYBP to between 20.5 and 21.5
MYBP, (probably closer to the younger of these last two values).

.
A large Eulepidina (top) and a smaller Nephrolepidina specimen from the type Prupuh Beds in NE Java, field of view 5 millimeters
across. The layer of median chambers in Eulepidina is ovr 300 µm thick and shows the numerous diagonal stolons that give this
genus its distinctive serrate or crenulate dark line within the median chamber walls. The median layer of the Nephrolepidina is about
100µm thick

Usually Eulepidina is easy to recognise in random thin section, however there are a few potential pitfalls.
Some fragments of the thickened median layer of Asterocyclina can look like fragments of Eulepidina,
although usually with Asterocyclina there are other Eocene fossils present which can reassure the in-
experienced. More difficult to separate are the genera Lepidocyclina (Multilepidina) [lower Tf] and the
Biostratigraphy of Indo-Pacific larger forams: page 55

trybliolepidine forms of Lepidocyclina (Nephrolepidina). In good equatoral sections the multilepidine

and trybliolepidine embryonts can be seen to be different, but in oblique sections they can be mistaken
for the embracing eulepidine sort. Also, while Eulepidina has a median layer typically close to 400 µm
or more thick, both the other genera have thinner median layers close to the embryont (c. 200-250µm)
although this can flare to 600µm in later adult stages.
Fabiania A. Silvestri, 1924
Type species: Patella (cymbiola) cassis Oppenheim, 1896
A variable or irregular cone-shaped test. Relatively regular outer chambers, umbilicus with little or-
ganised structure and often deeply excavated. Bilocular embryont may be well developed. In horizontal
section. The outer cortex has usually well developed radial partitions
Ranges from the Middle Eocene (T.a3) of Assam (Wilson & Metre, 1953) and the same age in Papua
New Guinea (Bain & Binnelamp, 1973), up to Letter Stage T.b in the Melinau Limestone Sarawak
(Adams, 1965) and in the T.b of Papua New Guinea (Bain & Binnelamp, 1973).
In Southeast Asia most records are assigned to Fabiania saipanensis but Binnekamp (1973) argues that
this is a junior sysnonym of F. cubensis. Widely distributed, from Papua New Guinea to Saipan, but
rarely a common component of larger foram assemblages.

Fabiania cubensis from Lengguru Fold belt, West Papua

Fabularia Defrance, 1820

Type species: Fabularia discolites Defrance, in Bronn, 1825 = Nummulites ovata de Roissy, 1805
A distinctive form that is essentially biloculine, but with a quinqueloculine juvenile stage in microspheric
forms. Microspheric specimens are possibly the more common type. Macrospheric forms have just two
Biostratigraphy of Indo-Pacific larger forams: page 56

adult biloculine whorls. Adult chambers are composed of very thick walls with thick partions subdiving
the chambers. These elongate divided chambers are parallel to the axis of coiling linked at their ends
by tubes or canals.
Ranges from within the Middle Eocene to the top of Tb, effectively top Eocene.
Probably evolves from a quinqueloculine miliolid ancestor. A rare form in the Indo-Pacific region. The
figures shown here are both from the Lengguru fold belt of Papua (Irian Jaya) where they only occurred
in a single sample.
 Fabularia ovata from West Papua, Lengguru
Fold Belt

Flosculinella Schubert, 1910

Type species: Alveolinella bontangensis L.Rutten, 1912
The almost spherical Flosculinella reicheli and slightly larger, oval F. globulosa have been recorded
from the Upper Te, with Spiroclypeus (Hanzawa 1957, Cole 1954, 1957a&b). However the genus,
especially the elongate oval to cigar shaped F. bontangensis is mostly a Lower T.f (T.f 1-2) form. This
latter species evolves into and is gradually replaced by Alveolinella in latest T.f1 through Tf2 (end Early
to mid Middle Miocene). It is found above the Orbulina datum in NE Java (pers. obs) and in western
Biostratigraphy of Indo-Pacific larger forams: page 57

Australia (Chaproniere 1975, 1981).

It is presumed that the genus evolved from Borelis, which is known to have some septula (secondary
septa) in a Y-profile, with secondary chambers above. However the transition to Fosculinella is a mor-
phologically sudden and distinct shift, and also included a change from septula that were aligned across
the chamber septum (subdivisions of each chamber correlated straight across from one chamber to the
next) to an alternating pattern in Flosculinella. However use of septula alignment in identification in
randon thin section is very difficult and not always practical.
 Flosculinella bontangensis, from shortly below the Orbulina datum in the Lodan Anticline NE Java. The defining two levels of
chamberlets are clearly seen in this near axial section. Also in this specimen are Miogypsina (left) and part of Cycloclypeus annulatus
(at base of image).

Gypsina H.J. Carter, 1877

Type species: Polytrema planum H.J. Carter, 1876
Early Oligocene or possibly Eocene to Recent.
Test usually attached on one, flattened, side. Inflated chambers are closely packed and arranged some-
what irregularly to give a polygonal (rather than hexagonal) form in section. No obvious structured
juvenarium. Coarsely perforate with no apertures.

Biostratigraphy of Indo-Pacific larger forams: page 58

Halkyardia Heron-Allen and Earland, 1918
Type species: Cymbalopora radiata von Hagemow var. minima Leibus, 1911
A small conical form with umbilical boss, usually around 1mm in size. Trochospirally coiling, radially
arranged, tubular-conical chambers (about 15 "spokes" in the adult whorls), around a wide, finely per-
forate plug. Wall thickened on the spiral side by addition of lamellae, but densely perforate. No apertures
known, communication via pores. Macrospheric form has hemispherical protoconch and deuteroconch
with two auxiliary chambers making embryont.
According to Adams (1970) this genus has been recorded from Ta3 (Middle Eocene) faunas in India
(Kutch, by Tewari, 1956). Data is sparse, at least any with illustrations. In the Sundaland area there are
no records of this genus older than T.c. Halkyardia ranges as young as the basal part of the Rajamandala
Limestone, with a Te1 assemblage dated by SIS (87Sr/86Sr=0.708027 (NBS987 of 0.710235) as 28.6
MYBP within a range or precision and correlation error of 28 to 29.1 MYBP.

Adams (1970) mentions he knows records from "several lower Te limestones in Borneo", but in 1984
Adams only expands on this claim by quoting the work of Hashimoto et. al (1978) on beds of "Tc to
Te1-4 age" in the Philippines. In this latter study there are samples with Tc faunas, that include Halk-
yardia, but the Lower Te faunas, mentioned do not. In PNG Belford (1984) finds this genus mostly in
samples in the Tc of the OK-Tedi area, but in one instance (p.20 op. cit.) he records Halkardia with
Eulepidina and Borelis in what is a Td, or possibly lower Te age sample.

Halkyardia, from Central Java, Early Oligocene, specimen upper left is 0.7 mm maximum width, upper right fractionally under
1mm. Lower pair of photos from same set of samples, same magnification, - oblique sections roughly on line marked with red line on
upper-left photo. These lower pair might be confused with reticulate Nummulites, except Nummulites would not have a distinct plug,
and wall thickness would be similar all around the specimen.

Halkyardia, sectioned
roughly along plane of
red-line above. Note
the pores in thickened
spiral side wall. Also
the radially arranged
Biostratigraphy of Indo-Pacific larger forams: page 59

chambers are sectioned

close to parallel on spiral
side but very obliquely on
distal side
Heterostegina d’Orbigny, 1826 and closely related genera
Type species Heterostegina depressa dʼOrbigny, 1826 emend. BANNER & HODGKINSON 1991
Nearly identical to Operculina except the chambers are subdivided by multiple secondary septa.
There are a number of heterostegine forms which have been given generic status, but which have been
lumped together in many oil industry and published reports. Most recently the work of Banner & Hodg-
kinson (1991) established the following:
Heterostegina (Heterostegina) D'ORBIGNY s.s. - Initially involute becoming evolute in adult ("maturo-
evolute")
Planostegina BANNER & HODGKINSON - Wholly evolute, flat. Type species: Planostegina costata (from
the Middle Miocene of Austria)
Heterostegina (Vlerkina) EAMES ET AL. Wholly involute, thereby usually lenticular and not as com-
pressed as H. (H.). Type species H. (V.) borneenisis VAN DER VLERK, from the Late Oligocene of
NE Kalimantan. this form evolves into Tansinhokella BANNER & HODGKINSON by extension of the
lateral chambers extending over umbilical area - producing more than one layer of lateral cham-
bers per whorl.
The later Eocene, Heterostegina-like form Grzybowskia BIEDA has recently been recorded from Christ-
mas Island (Lunt 2003).
Planostegina appears to evolve from flat Operculina by subdivision of the primary chambers several
times, and is the ancestor of Cycloclypeus, which is considered possibly polyphyletic. Heterostegina
(Vlerkina) also probably evolves from involute Operculina several times. One two occasions it is seen
to evolve into Tansinhokella; in Late Eocene and Mid Oligocene times. The stratigraphic distribution
of the various types is shown on the main range chart.

Biostratigraphy of Indo-Pacific larger forams: page 60

HETEROSTEGINA
Distinguishing Heterostegina s.l. from Tansinhokella in
oblique section. Image top-left (?Oligocene of Lengguru,
Irian Jaya) is of Heterostegina, with secondary septa and
chamberlets clear, no lateral chamberlets and spiral not annular
T
cycloclypid growth even in this apparently mature specimen. ANSINHOKELLA
Image to right is a similar section but note the pattern created
by the "lateral chamberlets" (but not cubicula). This is a specimen of Tansinhokella
Note that on the range chart (Figure 6) the range of Heterostegina (Vlerkina) is shown extending to
about the top of the Oligocene. This follows my own observations and Muhar (1957, also in Brouwer
1957/1966), but runs contrary to a clear statement in BouDagher-Fadel, Noad and Lord (2000) which
claims to have observed many Heterostegina (Vlerkina) in earliest Miocene Te, that is, co-occuring the
Miogypsina and descendants. However none of the samples listed in their paper with Heterostegina
(Vlerkina) are noted to contain Miogypsina, only Miolepidocyclina, a related genus but one not known
from SE Asia (cf. Loeblich and Tappan 1988). Furthermore the related paper by BouDagher-Fadel, Lord
and Banner (2000), based on the same NE Borneo material, reviewing the whole family Miogypsinidae,
does not mention Miolepidocyclina at all (and also contain errors such as not ranging Miogypsinoides
into the Late Oligocene). Data in this second paper also does not list Miogypsina specimens in any
sites that contain H. (Vlerkina). In the 40 km wide area they study, sites in the west contain Miogypsina
and sites in the east contain H. (Vlerkina) and Miogypsinoides. Therefore, in spite of the claim of Bou-
Dagher-Fadel, Noad and Lord (2000) that H. (Vlerkina) ranges up into the basal Miocene, it remains
unproven on doccumented evidence.

Biostratigraphy of Indo-Pacific larger forams: page 61

 Heterostegina
(Heterostegina)

Figure left is reverse side of stereo-

pair above

Alar prolongations, not containing secondary septa or chamberlets

Stereopairs of Heterostegina (Heterostegina) aff. depressa d'Orbigny, Recent, from a dredge in about 40 meters of water off Sakala
island, eastern Java Sea. The specimens above have strongly involute coiling, with alar prolongations extending over previous whorls.
Within these prolongations there are no secondary septa. In the middle and upper specimen the whorls can be seen be coming less
involute (maturo-involute form) but later stages are lost by abrasion.
Background dots are 100µm in diameter; a row of 8 dots = 1mm.
Biostratigraphy of Indo-Pacific larger forams: page 62

Heterostegina (Vlerkina) borneensis external

view of test, showing alar prolongations over
central umbonal thickening, with much
weaker development of secondary septa in
these spaces.
Lower T.e (Late Oligocene) Citarate
Limestone, West Java
Lacazinella Crespin, 1962
Type species: Lacazina wichmanni Schlumberger, 1894
At first glance in hand-specimen, Lacazinella appears similar to alveolinid miliolids. However it is
soon noted that they are more spherical than the fusiform alveolinids and, with a hand lens, one can
see that growth is not in a spiral coil but by “onion skin” addition of sphere-within-sphere layers. More
detailed examinations shows that these layers alternate so that the aperture is at opposite poles on each
successive whorl. Longitudinal ribs separate the chambers. The maximum size is about 3 millimeters.
Lacazinella is recorded from the Paleocene in North Africa the Middle East and Turkey (Eames, 1971;
White, 1994). In S.E. Asia it is reported from T.a3, Middle Eocene (Crespin, 1962, Rutten 1936 and
possibly by Bain & Binnekamp, 1973) where it is infrequently found with Nummulites javanus (B form)
/ bagelensis (A) and Alveolina. The genus is mostly a later Eocene index form, however Bursch (1947)
described Lacazinella occuring with reticulate Nummulites in Kai Besar (Larger Kai Island, near Aru
Islands of Irian Jaya). Adams (1970) reports confirmation of Nummulites fichteli with Lacazinella in
Burschʼs material, which extend the range of the genus into Letter Stage T.c, Early Oligocene.
Little is known on the phylogeny of this form. The proloculus is a spherical chamber with no traces of
ancestry in embryonic or nepionic stages. Two species are known: L. wichmanni and L reicheli, differ-
ing on shape of the test.
Lacazinella is almost unknown from outside the (Papua) Irian Jaya / Papua New Guinea and related areas.
Its northwestern-most record appears to be from the eastern arm of Sulawesi (Koolhoven, 1930). When
it does occur it can be very common and a rock building bioclast. Adams (1970), in a review from the
Middle East to the Pacific Islands, notes the genus is known only from the Moluccas and New Guinea
areas. So far all reliable data indicates that wherever Eocene carbonates with Lacazinella occur, Assilina,
Pellatispira and Biplanispira are absent, and vice-versa. Lunt (2003) has mapped out this distribution
and used these faunal differences to identify microplates of low and high latitude Eocene origins.

Biostratigraphy of Indo-Pacific larger forams: page 63

Lepidocyclina (Nephrolepidina), Lepidocyclina (Multilepidina) & Lepidocyclina (Trybliolepidina)
The revision of the Lepidocyclininae by BouDagher-Fadel and Banner (1997) is followed here with
two exceptions. This 1997 review was important as it maintained the use of Lepidocyclina as the main
genus because microspheric forms could not be assigned to any sub-genus, yet were obviously of the
same lineage group. Therefore, following van der Vlerk (1928), there is Lepidocyclina which covers
the earliest forms found in SE Asia (as well as Central American ancestors) including both microspheric
forms and the macrospheric (A generation) forms with isolepidine embryonts [Isolepidina not avail-
able as a sub-genus name as it is a simple synonym of Lepidocyclina], Lepidocyclina (Nephrolepidina)
for forms with reniform deuteroconchs, and Lepidocyclina (Trybliolepidina) for forms with enclosing
deuteroconch. In addition I use Lepidocyclina (Multilepidina) for the lineage placed by others in its
own genus.This follows van der Vlerk (1928) except he uses the term Pliolepidina, which is an older,
Central American form. The second point of difference with the work and BouDagher-Fadel and Ban-
ner is retaining L. (Trybliolepidina) which they said should be abandoned. It is true that after van der
Vlerk's clear original description, but lacking a type specimen, Berry (1929) proposed a type specimens
for Trybliolepidina but which was from Eulepidina, pointed out by later workers to clearly not meet the
original intentions of van der Vlerk's Trybliolepidina. This alone does not invalidate the sub-generic
name. BouDagher-Fadel and Banner (1997) claimed their observations of Trybliolepidina suggested it
was polyphyletic with differing varieties and hence the term should be abandoned. On this last point I
disagree, for reasons too detailed to specify here other than the morphotype has a clear range from near
the base of the Middle Miocene to basal Pliocene with variation being part of the normal increase of
deuteroconch embracing the protoconch through time.

Schematic block diagram of Lepidocyclinid from Eames et al 1962(b). Chambers underneath the median layer (equatorial
chambers) not shown.
Biostratigraphy of Indo-Pacific larger forams: page 64

a: transition between arcuate, ogival ("engine-turned"), and spatulate equatorial chambers

b: increased elongation of the spatualte chambers
c: "hexagonal" equatorial chambers
d: exterior surface of the equatorial chambers showing their paried interiomarginal apertures
e: pillar forming no exterior tubercle
f: pseudopillar
g: pillar forming an external pillar
h: ray formed in both equatorial and lateral chambers, periphery stellate
i: ray formed solely in equatorial chambers, periphery stellate
j: ray formed solely in equatorial chambers, periphery polygonal
k: ray formed in both equatorial and lateral chambers, periphery polygonal
l: embryont or nucleoconch (l1 protoconch, l2 deuteroconch)
From Eames et al 1962(b)
 The S.E Asian Lepidocyclininae are a side-branch of a Central American stock, having migrated east at
the base of T.d, along with the related Eulepidina and unrelated Neorotalia. Note that some workers (e.g.
Dooger and Rohling, 1988) have proposed more than one eastward migration of the genus across the
Atlantic, based on fluctuations in biometric measurements of populations in the Eur-african area. This
is still unproven, and requires a wider study of the migration event(s), particularly accuate age dating
and correlation with potential causes. Drooger (1993) sums up the work at that time by saying (p.127)
"Only the last wave at about the middle of the Oligocene would have been really successful, because
the descendants of these immigrants managed to spread over the entire length of the Tethys belt as far
as its eastern, west Pacific end." The data presented here (Cimanggu & Ciapus sections) shows that
the main T.d migration actually reached S.E. Asian by 32 MYBP, Early Oligocene.
The L. (Nephrolepidina) lineage evolved over time and gave rise to several descendants. In Early Miocene
times a form with an increasingly irregular embryonic apparatus was known first as L. (N). transiens,
and it is thought that this gave rise to the more extreme form L. (Multilepidina), which survived until
nearly the end of Lower T.f. By a heterochronic process of increasing radial symmetry and enclosure
of the protoconch by the deuteroconch the L. (Nephrolepidina) lineage graded into the L. (Trybliol-

Stereopaired images of L. (Nephrolepidina). Both recovered from a mudstone (Tuban Fm. latest Early Miocene Java). These
specimen are probably L. (N.) martini, (see text). Background white dots are 100 µm diameter.

Adauxiliary chambers (AAC II)

Spatulate to hexagonal chambers,
diagonal and annular 6 or more stolon system.
Diagonal stolons on several levels.
Annular stolons appear in proximal part
of common wall.

DII Ogival chambers,

diagonal 4 stolon system evolves
as adjacent chamber walls impinge
"Engine-turned" appearance
DI
Biostratigraphy of Indo-Pacific larger forams: page 65

(AAC I) Auxiliary Simple arcuate chambers,

a pair of stolons connect each chamber
chamber to adjacent chamber of next cycle
Details of morphology of Lepidocyclinid embryonic stage and median chamberlets (modifed from Haynes 1981).
LEFT: Embryonic chambers
RIGHT: Schematic view of median chambers
Biometric Factor A is usally defined based on the two parts of the blue line in
and stolon development, as sometimes used in
the protoconch. The value used is the proportion of the circumference of the
taxonomy.
common wall as a percentage of the whole inner circumference.
Factor B is the proportion on the embryont (DI & DII covered by auxiliary
chambers (red lines, as a percent of the whole inner circumference).
Factor C is the number of auxiliary chambers, excluding the rare AAC I
chambers, in the example above this value is 2.
 Lineages and biometric characters of nepionic chambers
in Miogypsinids and Lepidocyclinids
N.B. embryonic chambers of Miogyspinids and Lepidocyclinids not shown at same scale.
On Lepidocyclinids only the ad-auxiliary chamberlets (=factor "C") are shown

Stage &
SERIES
Martini Zones
Letter Stages

Magnetic pol.
Blow Zones

MYBP
5 N 18
Nephrolepidina species concepts
following Van Vessem (78).

Mess.
� � ��� Biometric parameters are

N 17
NN 11
averages for populations
L. (T.) rutteni
(52.5%<A; C>6.5)
NN 10

N 16
10

Lt. MIOCENE
NN 9

Tortonian
UPPER Tf (=Tf3)
L.

N 15
�� ������
NN 8

Miogypsina
N 14
NN 7
N 13
L. (Nephrolepidina)

N 11-12 L. (N.) martini

transiens lineage
(Multilepidina)

Range of

NN 6
antillea (52.5%<A; 4.75<C<6.5) �� ��� ?

(Tf2)
Lepidosemicyclina

N 10
Miogypsinoides
"Conomiogypsinoides"
�� ���

M. MIOC.
NN 5
15 N9

or M. cupulaeformis variants
L. (N.). angulosa
cushmani (52.5%<A; C<4.75)

N8

Serravallian Langh.
N7
with "ferreroi" or

NN 4
indica
cruciform ornament

globulina
Nephrolepidina sometimes

(Tf1)

N6
LOWER Tf (=Tf1 & 2)
NN 3 globulina exentrica �� ���

dehaartii droogeri

Burdigalian
20

N5
thecideaeformis L. (N.) sumatrensis
tani (40%<A<52.5%; C<3.75)

NN 2
bantamensis

EARLY MIOCENE
gunteri

N4
Upper Te

Aquitanian
NN 1

�� ���

Te4

NP 25

P 22
L. (L.) isolepidinoides
formosensis complanatus (A<40%; C<2)

Te2-3

Chattian
L. (Trybliolepidina & L. (Nephrolepidina)

?
Eulepidina

Te1 �� ���

Lt. OLIGOC.
NP 24
= Acme of giant microspheric Lepidocyclina

P 21
ancestral
Neorotalia

(b) (a)
mecatepecensis
L. (Lepidocyclina)

Td
30 P 20

Rupel.
NP 23
Nummulites

Biostratigraphy of Indo-Pacific larger forams: page 66

Two images of L. (Nephrolepidina ferreroi) from a unit
dated within nannofossil zone NN 4 (based on interbedded
mudstones), from the Lutut Beds, north Central Java.

L. (Nephrolepidina) "crucifera". This

form, seen in stereo, is associated with
Cycloclypeus annulatus and is from a
later Early Miocene bed in Central Java
(Widoro River section, about N6 to N7).
These specimens which were sectioned and
found to have a normal nephrolepidine
embryont. The degree of enclosure of the
protoconch by the deuteroconch (biometric
factor A) is just less than 50% which is
consistent with the age of this sample based on
planktonic foraminifera (see Kadar 1986; this
sample adjacent to his sample KYF 38, one
or two samples below the first Praeorbulina
specimen and above the evolution of
Globorotalia peripheroronda).

Below L. (Multilepidina). In random thin section. On left and top right, from
the Ngrayong Sandstone in east Java (early Middle Miocene). Both specimens
on the left show the typical Mulitlepidine embryont. Bottom right, from the
Lutut Beds (late Early Miocene, nannofossil zone NN 4,
Central Java). These forms are clearly distinguishable from L.
(Nephrolepidina). The width (relative to hight) of the embryont,
and the thinness of the equatorial layer distinguishes these
forms from Eulepidina.

Biostratigraphy of Indo-Pacific larger forams: page 67

 epidina) form. The name comes from "tryblios" for small cup, and the cup-like way the protoconch is
surrounded on three sides by a quadrate deuteroconch. In Upper T.f, Late Miocene times, this was the
surviving lepidocycline form.
The ranges of the Lepidocyclines (Nephrolepidina, Multilepidina and Trybliolepidina) are shown in a
later Figure, which shows sketches of the development of the embryonic and juvenile chambers through
time. Lepidocyclina (Trybliolepidina) became extinct in basal Pliocene times. Van Vessem (1978) gives
a weak description of a planktonic fauna associated with larger forams including "Heterostegina ...
Baculogypsina ... Lepidocyclina". The presence of the distinctive Pulleniatina suggests latest Miocene
or younger, but the reporting of taxonomically difficult forms such as Sphaeroidinella dehiscens, but
not the usually ubiquitous and distinct Globorotalia tumida suggests re-checking would be worthwhile.
Similarly Adams (1984) cited Adams and Frame (1979) to claim that beds of N19 age in Fiji contained
Lepidocyclina but the original reference assigns a significantly older age.
Lepidocyclina (Trybliolepidina) has been found in the Tapak Limestone of north central, Java (ter
Haar, 1935, original specimens in the collection of the GRDC, Bandung) and in the equivalent Bodas
Limestone to the east that can be correlated on seismic (van Bemmelen, 1949 and personal re-sampling
of the type location). These limestone shave recently been dated using strontium isotopic methods and
tied to the stratigraphy of an oil well (KRG-1). This is 87Sr/86Sr=0.709011 and 0.708959 respectively
(NBS987 of 0.710235), with combined error and precision for each result of 4.8 to 5.78 MYBP and 5.98
to 6.86 MYBP. These ages are virtually on the Mio-Pliocene boundary that on the same time scale is
placed at 5.2 mybp (Berggren et al, 1995).

Species names in the Lepidocylinids

The literature is rich in species names for the different lineages of lepidocyclinids. These are mostly
based on the influential revision of van der Vlerk (1928), but these species may equally reflect ecophe-
notypic variants as species with stratigraphic evolutions and extinctions. More continuous sections in
oil wells and the advent of stronitum datings now offers the chance to separate environmental effects
from evolutionary effects and establish a thorough taxonomy for the group in the near future. For the
record the system of van der Vlerk was:

van der Vlerk 1928, modified

- Initial chamber completely or nearly completely surrounded by second chamber, embryont greater than 700
microns, diameter of test greater than 6 mm. = EULEPIDINA
- Embryont in the shape of a cupule (tryblios), i.e. a quadrate tendancy with the second chamber enclosing the
initial chamber on three out of the four sides. = TRYBLIOLEPIDINA
- A reniform second chamber surrounds about a half of the initial chamber = NEPHROLEPIDINA
- The embryont is composed of two semi-circular chambers = ISOLEPIDINA [generally included in Nephro-
lepidina in S.E. Asia]
- Embryont multicellular = MULTILEPIDINA
Biostratigraphy of Indo-Pacific larger forams: page 68

Determination table for Isolepidina:

Only one species known.[L(I) boetonensis vdV 1928 is a junior synonym] L.(N.) isolepidinoides

Determination table for Neprolepidina:

Forms with peripheral flange
Prominent ridges passing from central boss into slight peripheral processes
Columns spread over whole surface, thickness of columns ± 90 µ. L.(N.) radiata Martin
No prominent ridges
Columns present
Columns spread over whole surface.
Test diameter to thickness ratio 6-7 to 1, equatorial chambers arranged in circles:
L.(N.) verbeeki Newton & Holland
 Test diameter to thickness ratio about 3 to 1, equatorial chambers arranged in polygons:
L.(N.) japonica Yabe
Test diameter to thickness ratio about 1-11⁄2 to 1, equatorial chambers arranged in polygons:
L.(N.) sumatrensis Brady
Columns absent
Diameter 2-4 mm. L.(N.) sumatrensis Brady var. inornata Rutten
Diameter 11⁄2 to 2 mm. L.(N.) sumatrensis Brady var. minor Rutten
Forms with no peripheral flange
Test stellate, 6 to 9 rays, thickened equatorial chambers in rays extending as pseudospines.
L.(N.) martini Schlumberger
Test not stellate.
Columns present, spread over whole surface
Outline polygonal.
L.(N.) douvillei Yabe
Outline round.
Diameter 1.2 to 2 mm. L.(N.) parva Oppenoorth
Outline round.
Diameter 2.5 to 4 mm, thickness of central columns ± 160 µ. L.(N.) borneënsis Provale
Diameter 2.5 to 4 mm, thickness of central columns ± 1000 µ. L.(N.) atjehensis Oppenoorth
Few columns present, only in central part
3 to 5 columns forming an umbonate central prominance, thickness of columns ± 600 µ.
L.(N.) angulosa Provale
Columns not in center, but in corners of a polygon, thickness of columns at least 150 µ.
L.(N.) ferreroi Provale

Columns over centrum, thickness of columns ± 300 µ. L.(N.) brouweri Rutten

Only one column in central part of test L.(N.) inflata Provale
Columns absent L.(N.) epigona Schlumberger

Determination table for Trybliolepidina:

Forms with peripheral flange
Columns present
Columns spread over whole surface, thickness of columns ± 90 µ. L.(T.) orientalis van der Vlerk
Columns only in the central part of the test and thickness of columns ± 150 µ.
L.(T.) talahabensis van der Vlerk
Columns absent L.(T.) ephippoides Jones & Chapman
Forms with no peripheral flange, columns spread over the whole surface L.(T.) rutteni van der Vlerk

Determination table for Multilepidina:

Forms with peripheral flange, columns absent, diameter 6-8 mm L.(M.) luxurians Tobler
Forms with no peripheral flange, columns over whole surface, diameter ± 25 mm L.(M.) stigteri van der Vlerk

Determination table for Eulepidina:

Forms with peripheral flange
Biostratigraphy of Indo-Pacific larger forams: page 69

Columns present
Columns spread over whole surface E. papuaensis Chapman
Columns only in centrum
Test diameter to thickness ratio about 7 to 1: E. mediocolumnata van der Vlerk
Test diameter to thickness ratio about 21⁄2 to 1: E. stereolata Oppenorth
Columns absent
Diameter about 7 mm, thickness of walls bordering the equatorial layer ± 15 µ:
E. andrewsiana Jones & Chapman
Diameter 15-30 mm, thickness of walls bordering the equatorial layer ± 30 µ:
Embryont ± 1000 µ E. formosa Schlumberger
Embryont > 1000 µ, as much as 4500 µ E. formosa Schlumberger var irregularis Rutten
Forms with no peripheral flange
Columns present
Columns spread over whole surface, thickness of columns ± 150-200 µ E. dilatata Michelotti
Columns absent
Test diameter to thickness ratio nearbly 3:1 E. atuberculata van der Vlerk
Test diameter to thickness ratio nearbly 10:1 E. planata Oppenoorth

Van der Vlerk’s determination table for microspheric (B) forms is complex and applies to types that are rarely en-
countered. It is basically a list of most of the permutations of present / absence of peripheral flanges, columns, size
and ratio of diameter to thickness. Species used by van der Vlerk were:
L. bonarellii (Provale) L. perornata H. Douvillé
L. verbeeki Newton & Holland L. orientalis van der Vlerk
L. soebandii van der Vlerk L. papuaensis Chapman
L. mediocolumnata van der Vlerk L.acuta Rutten
L. formosa Schlumberger L. luxurians Tobler
L. blanfordi Nuttall L. inaequilateralis Jones & Chapman
L. flexuosa Rutten L. provalei Osimo
L. cebuensis Yabe & Hanzawa L. gallieni Lemoine & R.Douvillé
L.dilatata Michelotti L.dilatata var. tidoenganensis van der Vlerk
L. gigantea Martin L. papulifera H. Douvillé
L. euglabra H. Douvillé L. subradiata H. Douvillé
L. atuberculata van der Vlerk L. insulaenatalis Jones & Chapman
L. sumatrensis Brady var minor Rutten L. dekroesi van der Vlerk
L. glabra Rutten

Biostratigraphy of Indo-Pacific larger forams: page 70

Lepidosemicyclina Rutten, 1911
Type species Orbitoides (Lepidosemicyclina) thecidaeformis Rutten, 1911
Ranges from uppermost Te5 (Adams, 1984 and Raju 1974) where it is hard to identify, as the earliest
species still have the ogival chamberlet shape of the ancestral Miogypsina in the early adult stage. The
distinctive hexagonal chamberlets and especially the enlarged embryonic chambers are most clearly
noticable in Tf1 and the basal part of Tf2. Van Vessem has reported this genus from strata of N7/8 age
in Borneo. These forms are found above the Orbulina datum in the Ngrayong Formation of NE Java
but appear to become extinct before the disappearance of Miogypsina at the top of Tf2. Chaproniere
(1981) also recorded this genus from limestones of possibly N8 to N9 age in NW Australia.
Originally defined as a lineage of Miogypsina that developed rhombic and then hexagonal chamberlets.
Raju (1974) argued that the lineage also shows a more rapid rate of evolution though nepionic accel-
eration than the main Miogypsina lineage and this justified maintaining a separate genus. The juvenile
stage of the oldest species in this lineage, L. thecidaeformis, hardly differs from Miogypsina globulina,
the distinction being the hexagonal chamberlets. The enlarged embryont in younger Lepidosemicyclina
species is especially developed in the the deuteroconch. Multiple apertures from the inflated deutero-
conch in L. excentrica leads to chamberlets growthing across the apex, and the embryonic chambers
are shifted slightly from the periphery of the adult test.

Image above and blow of Lepidosemicyclina from early Middle Miocene NE Java. Above a whole specimen about 2 mm is length with
the highly enlarged embryont characteristic of younger members of this lineage. Below, at twice the magnification are fragments of
?Lepidosemicyclina on the left, but in the center is a fragment with clearly hexagonal chamberlets from the median layer.

Biostratigraphy of Indo-Pacific larger forams: page 71

Linderina Schlumberger, 1893

Type species: Linderina brugesi, Schlumberger, 1893
A rotaliid form distinct in thin section because of its thick lamellar lateral walls that bulge over the
central embryont. Wall porous even in thickest part. These often distinct pores give it quite a different
appearance from the much younger genus Miogypsinoides, also the median layer in Linderina is rela-
tively thicker , and the chambers are more inflated in axial section. It differs from Planorbulinella as
the lateral walls in Linderina are substantially thicker and bulge in the center.
Range: This genus has been recorded in Early Eocene strata from East Africa, Pakistan and India (Sil-
 vestri, 1942; Eames 1952). Most records in Southeast Asia are from the Middle Eocene. Adams (1970)
noted this form occurs with Pellatispira in Pakistan and suggested a T.b extinction, however this report
shows that Pellatispira occurs frequently in the T.a, so a Late Eocene or T.b age is still unproven.
Little is known about the ancestry and evolution of this genus. It is a rare component of Middle Eocene
carbonate faunas in Southeast Asia.

Two examples of Linderina from the Middle Eocene of Central Java. Bagelen Beds, nr. Worarwari (top) and from West Papua (Irian
Jaya, below)
Biostratigraphy of Indo-Pacific larger forams: page 72

Marginopora Quoy & Gaimard, 1830

Type species Marginopora vertebralis Quoy & Gaimard, 1830
A discoid miliolid, large (up to a centimeter for macrospheric forms and 3 cms for microspheric forms)
and biconcave with a thickening periphery. The microspheric forms have a small peneropline juvenile
stage before growth of annular chambers. The megalospheric embryont is a circular, flattened box or
pill shape, and consists of a large deuteroconch, which encloses a small protoconch. This embryont is
surrounded by an annular ring of chamberlets. These cyclically growing chambers are, for a few whorls,
simple, but after this “juvenile” phase a three-fold differentiation appears with an upper and lower set
of relatively small lateral chamberlets separated by a wedge of more irregular equatorial chamberlets.
The upper and lower layers of lateral chamberlets remain roughly the same small size throughout the
adult phase, and it is the expanding wedge of annular chamberlets that produces the biconcave shape.
In the adult phase the equatorial chamberlets have a vertical septula that is irregular, wavy, due to the
 cross-diagonal streaming paths of protoplasm towards the multiple rows of coarse stolons or pores on
the outer peripheral wall. In external view these coarse pores are irregualry arranged.
The only forms that could be mistaken for Marginopora would be very oblique, near equatorial sections
of Sorites / Amphisorus, neither of which has the lateral chambers, thickening periphery and biconvex
shape (or such a large embryont). While in the Treatise and text books the upper and lower layers of
lateral chamberlets are well illustrated, in many fossil samples these features are obscured or poorly
developed.
Hanzawa (1957) figured a good specimen of Marginopora in association with Miogypsina from the
Tinian Island in the Marianas group, in a formation dated as Upper Te (Tagpochau Limestone). Cole
(1969) confirmed such ancient occurrences of Marginopora in an Upper Te limestone from the Midway
Atoll, occurring with Spiroclypeus and other markers. However it is mostly a later Middle Miocene,
Tf3, to Recent form.
An Indo-Pacific form, cosmopolitan, but prefering sea-grass environments. Endosymbiotic dinoflagel-
lates are known in the modern specimens.
In modern forms a final reproductive stage is known where about half a dozen undivided and presumably
fragile whorls are added to the test. These reproductive chambers are not comparable to gerontic stages
of, say, the giant Mid-Eocene Nummulites. They are more like the thin “kummerform” final chambers
in planktonic forams, which are also associated with reproduction before death. Gerontic adult stages
are usually features that have the potential to be increasing drawn into the normal life cycle through
heterochrony (hypomorphic development).

Miogypsina Sacco, 1893

Type species: Nummulites globulina Michelotti, 1841
A three-layered orbitoid-like test which can be distinguished from other orbitoid forms in equatorial
section as the juvenile and embryonic chambers are at the periphery, not in the center (i.e. growth is not
truly "orbital"). The equatorial chambers of Miogypsina, in axial or near axial section, are distinct from
other three-layered orbitoids, being distinctly inflated, that is, the septula of the equatorial chambers grow
as well-formed "arches" that extend in an arc from the upper to the lower walls of the median layer (see
images). This contrasts with nearly all other orbitoids, which have flat upper and lower median layer
walls, nearly straight septula and perpendicular contacts at upper and lower lateral walls.
Otherwise Miogypsina can be variably flat, robust, with or without ornament, and even sometimes with
greatly reduced numbers of lateral chambers. Well developed pillars and pustules can form, however
these external features appear to have no stratigraphic or taxonomic importance. Like Miogypsinoides
the embryont consists of a spherical protoconch and reniform deuteroconch, and these are larger in
later Miogypsina species. The juvenile may be a short, single rotaliid-like spiral, or, reduced through
evolution, a second juvenile spiral may be formed on the opposite side of the embryont.

The evolution datum of Miogypsina hs long been suspected to be close to the base Miocene and base
Biostratigraphy of Indo-Pacific larger forams: page 73

N4 planktonic zone but, as pointed out by Adams (1984) detailed data is lacking. This is probably be-
cause large parts of the Indo-Pacific were undergoing transgression at this time and platform carbonates
are common, but with few occurrences of planktonic foraminifera. Once again the unique geological
position of Java yields suitable interbedded mixtures of the two assemblages, in the Prupuh Fm. type
location and similar sections along strike. Here, within the lower part of the Prupuh limestone, a consist-
ent series of SIS measurements dates the lowest Miogypsina (above samples with Miogypsinoides) as
86
Sr/87Sr = 0.708257, which correlates to a a median age of 23.8 MYBP, and a combined analytical error
and age calibration range [2Sd] of between 23.34 MYBP and 24.17 MYBP. Mudstone samples below this
are P22 but there is a significant gap is good planktonic faunas until N4 above this. Note this is from the
same traverse as van der Vlerk and Postuma 1962, Krandji section, located from original BPM maps
and their samples h280 and h266, the later with a few Gt. cf. kugleri and common Globigerinoides.
 Apical line
st
ed te
p

a
-sh
elta
of d
end
ical
of ap
Z
ne
Outli

Schematic sketch of the earliest chambers in older Miogypsinoides and Miogypsina species. There are 18 spiraling chambers (biometric
factor X). The biometric factor "Y" is the number of spiral chambers up until the first chamber with two foramen (stolons) - the blue
coloured chambers in the example above, i.e. Y = 10 in this sketch. The biometric factor "Z" is the number of nepionic chambers (excluding
embryonic pair) up to and including the largest spiral chamber, - 13 in the sketch above. The angle γ is the angle of rotation between the
apical line of symmetry and the central line between the protoconch and deuteroconch. For long-spired juveniles (>1 juvenile whorl) this
value is given as a negative number, so the example above would be -330°. The number is considered positive when there is less than one
juvenile whorl, and when there is a second nepionc spiral (see below).
With a value for factor X of 18 and 13⁄4 whorls in the juvenile spiral it is likely that a specimen such as the one above would be from the Late
Oligocene and would probably be referred to the species M. complanatus.

Apical line

est
dt
pe
ha
a-s
elt
of d

�
nd
al e

�
apicf
ine o
Outl

Biostratigraphy of Indo-Pacific larger forams: page 74

Schematic sketch of the earliest chambers in stratigraphically young Miogypsina species. The two embryonic chambers are larger than in
the older (ancestral) example above. The diameters of the embryonic chambers are measured perpendicular to the imaginary axis between
them, and are measured from mid-chamber wall so that any overgrowth on these walls does not have an effect. The diameter of the
protoconch is called DI and the deuteroconch DII, given in microns (µ).
After this embryonic stage there are two spirals of juvenile chambers, one on each side (starting with each "A"= auxillary chambers).
As above the value of factor X is the total number of spirally coiled nepionic chambers. More useful, once a second spiral appears, is the
biometric factor "V" or 200 α/β, which is the degree of symmetry between the two spirals. This is low in older species, and high in the the
nearly symmetrical example above, which would be a young form. This symmetry is measured as the ratio of alpha (α) to beta (β) defined
as 200α over β. The factor gamma (γ) is the positive angle of deviation of the line of symmetry of the embryont from the apical line (line of
symmetry of the adult test). The example above has M 200α/β of 75 to 80 suggesting a form such as M. cushmani, from close to the Early
to Middle Miocene boundary.
The base of G. kugleri is used by Berggren et al 1995 to define the Oligo-Miocene boundary (p.173
op.cit) and is dated as 23.8 MYBP. These workers record common Globigerinoides from 24.3 MYBP. Thus,
considering the range of error, this lowest record of Miogypsina correlates very closely with the base
Miocene and the base N4 planktonic zone. On the range chart (Figure 7) the base N4 is drawn at the
original defining datum of "evolution" or lowest common occurrence of Globigerinoides, which is still
the most frequent interpretation among biostratigraphers in SE Asia. The lowest strontium ratio values
in samples with Miogypsina recorded by Allan et al (2000) from PNG is 0.708280 which converts on
the same scheme used here to a median values of 23.3 MYBP (combined error and calibration range from
22.8 to 23.8 MYBP.)

The extinction of Miogypsina is also well preserved in the interbedded shallow carbonate and planktonic
rich faunas found on the fringe of the Sundaland craton in what are now Sumatra and Java. Clarke and
Blow (1969), Adams (1970), Bauman (1975) and van Vessem (1978) all make observations at many
localities in this region of Miogypsina as high as "N11 to N12", "N12 probably N13", or "N11". It must
be remembered that the base N11, N11-N12 boundary and top N12 are all based on the Globorotalia
fohsi lineage. There have been alternative interpretations of the G. fohsi subspecies and their zonal use
and annotation (cf. Blow 1969, 1979, Stainforth et al 1975, Bolli and Saunders 1985 and Berggren et al
1995). The observations of the above workers are that Miogypsina is found above the base N11 evolu-
tion of G. fohsi s.l. and that it is not found with the later G. fohsi robusta forms. Bauman leaves open
this upper limit suggesting he did not find G. fohsi above the highest Miogypsina, which, as explained
below, is possible because of a regional sequence boundary at this time.
The most recent view of Berggren et al (1995) is widely accepted now, with the base G. fohsi s.l. at
12.7 mybp, the extinction of the whole group being at 11.9 mybp, and with the extception of G. fohsi
robusta from 12.3 to 12.1 MYBP the fohsi subspecies having little use in zonation. Therefore using GPTS
calibrated planktonic biostratigraphy the extinction of Miogypsina is probably between 12.7 and 11.9
MYBP.

An attempt has been made to clarify this important extinction event by visiting the Nyalindung forma-
tion of Adams, the Jiwo Hills and Wonosari section of Bauman, and many location in NE Java including
several of those of van Vessem. These visits confirm that the fohsi group evolves before the extinction
of the Miogypsina, however only NE Java has a relatively unbroken succession into younger beds. This
sucession include a thin, flood and sequence boundary defining limestone called the Platen Member.
Above this limestone are deep marine marls without any trace of larger foraminifera but with well
preserved N13-14 planktonic assemblages. There is rare Miogypsina in the Platen but no planktonic
zonation is possible so a range of Miogypsina into N13 cannot be ruled out. An attempt to date the very
well preserved, hyaline bioclasts in several samples with G. fohsi and Miogypsina, the Platen limestone
samples at several sites, and the basalmost N13-14 marls of the Wonocolo Fm all tended to return ages
older than acceptable to biostratigraphic control. These ages were basal Middle Miocene, and some
were later Early Miocene in samples clearly above the 15.1 MYBP Orbulina datum. Analytical precision
and repeatability on adjacent subsamples was only slightly higher than usual for this type of analysis
Biostratigraphy of Indo-Pacific larger forams: page 75

so the SIS data cannot be entirely discounted, but for the time being the disappearence of Miogypsina
and other larger foraminifera in the area is considered within N11 or N12 (?N13) which is rounded to
a GPTS age of about 121⁄2 MYBP, plus or minus at least half a million years.
The highest ratio measured in Miogypsina in PNG by Allan et al. 2000 is 86Sr/87Sr = 0.708850, which
correlates to 10.9 MYBP (precision and calibration error 10.1 to 11.7 MYBP), slightly younger than the
disappearance in Java, straddling the N14-N15, Middle to Late Miocene boundary. However the tech-
nique of Allan et al is for large numbers of whole rock SIS analysis to be plotted with generic ranges,
with obvious out of series diagenetic errors to be discarded. There is no way to identify slight diagenetic
errors (or large errors from mid-ranges plotting slightly off the true isotopic limits) and blurring at the
highest and lowest ends of the series. As a result the analysis of eight locations in West and Central
Java, from mostly un-named carbonate formations (but including three above the Nyalindung Formation
 mentioned above), are important as they have tightly grouped strontium ratios ranging from 86Sr/87Sr =
0.708835 to 0.708874 (11.4 to 10.2 MYBP median age range, analysed by T. Allan) and these carbonates
are without any sign of Miogypsina (or C. annulatus, or L. ferreroi). This supports the dating of the top
of Tf 2 within a range of 111⁄2 to 13 MYBP.
The inflated median chambers and simple arched cubicula of Miogypsina are quite distinctive, even
in small random thin sections. Note that Miogypsinodella Boudagher-Fadel, Lord & Banner (2000) is
considered a junior synonym of Miogypsina. This is because a sample rich in these forms, with unstacked
partings in the lateral wall, unlike typical cubicula, could be examined both in thin section and as as
loose individuals. This sample was from the Badui Beds of West Java of very similar age to the type
level of Miogypsinodella, being mid or later Early Miocene (nannofossil zone NN4, Tf1, and 86Sr/87Sr
= 0.708466, median age 19.7 MYBP, range within error and precision 19.3 to 20.1 MYBP). In this sample
there is a population of typical Miogypsina and a population of the "Miogyspinodella" forms but all
specimens of the latter were found to be microspheric.

Badui Beds West Java. The larger specimen has "partings" in the lateral walls rather than cubiculae [Miogypsinodella of BouDagher-
Fadel et al], but all specimens of this type are found to be microspheric (B generation). The A or macrospheric generation are
represented by typical Miogypsina specimens such as the two in the lower right of this image. The generic name Miogypsinodella is
therefore considered invalid, as representing a dimorphic variation only.

Genus Miogypsinella Hanzawa, 1940

Biostratigraphy of Indo-Pacific larger forams: page 76

Type species: Miogypsinella borodinensis Hanzawa, 1940

Range: within Te4 only. A precise definition of Miogypsinella has yet to be arrived at. Some workers
such as Boudagher-Fadel et al. (2000) simply say that Miogypsinella has a weaker trochospire and
thinner walls than Miogypsinoides. They consider it intermediate between the robust, thick walled
Miogypsinoides and the thinner-walled Miogypsina with pores in the reduced wall leading to lateral
chamberlets or cubiculae. New, unfinishised work suggests that Miogypsinella is not intermediate be-
tween Miogypsinoides and Miogypsina and these last two have a single biometric series of nepionic
acceleration, whereas Miogypsinella has a different embryonic and nepionic evolutionary series as
compered in the same samples. Miogypsinella appears to be a distinct lineage well before the evolu-
tionary appearance of cubicula.
 Miogypsinella from the Tagogapu Beds of west Java, near Bandung, photographed without sectioning or grinding. Immersed in oil
and with transmitted light it becomes transparent. The axial thickness (in the direction of the view) of this specimen is just over
a hundred microns, about a quarter of the thickness of Miogypsinoides in the same sample. The diameter of the protoconch in 88
microns, slightly smaller than the average for the thicker Miogypsinoides forms, and the angle γ is about 345 degrees, close to the
average for Miogypsinella, but which is at the high end of the range of values for γ in Miogypsinoides in the same sample

Miogypsinoides Yabe & Hanzawa, 1928

Type species: Miogypsina dehaartii van der Vlerk, 1924, Miogypsinoides Chapman, 1932 nom. transl.

Miogypsinoides with part of juvenile whorl visible at top of specimen, oblique view of median chambers and thick lateral wall at
bottom with lamellar growth

Biostratigraphy of Indo-Pacific larger forams: page 77

Three contrasting examples of Miogypsinids from the Lutut Beds of Central Java. It is uncertain if these specimens are in-situ, as
these beds are mid Early Miocene in age, with common Late Oligocene reworking. The specimen on the left clearly has started to
develop a lateral miogypsinid fan of chambers, but the trochospiral juvenile is weak and thin-walled. Such a specimen might be
referred to Miogypsinella. The central specimen has a juvenile with a thicker axial dimension, thicker lateral chamber walls and a
compound umbilical plug. This is a more typical juvenile Miogypsinoides, before advanced growth has had a chance to thicken the
lateral walls into a specimens like the one on the right, which is typical adult Miogypsinoides.
Miscellanea Pfender 1935
Type species: Nummulites miscella dʼArchiac & Haime, 1854
While the genus is placed in the Nummulitacea by Loeblich & Tappan (1988) it does not have the same
spiral marginal cord of other Nummulites but a reported ʻinternal spiraling canal of triangular section”.
Authors such as Haynes (1981) place the genus in the Rotaliacea (associated with the outwardly similar
Daviesina).
Miscellanea is a lenticular form with two to three whorls, usually with a slightly compressed margin.
Sutures are straight, and in external view there is a distinct ornament along sutural traces comprising
ridges or semi-fused radial rows of small pustules with additional rows of pustules between these el-
evated sutures. A few (typically 5 to 7) larger pustules occur near the umbo.
Restricted to Tertiary a1, the middle and Late Paleocene. Mostly known from the Middle East.
Very rare in Southeast Asia, because sediments of this age are rare. More usually reported from western
Tethys, Europe to Pakistan. Adams (1970) has confirmed the presence of this form in Sarawak (Engkilili
Fm. Lupar Valley) but is understandably skeptical on the occurrence in the East Indonesian Kai Islands
reported by Bursch (1947) as the illustrations are ambiguous and the occurrences are in beds of Late
Eocene to Early Miocene age. It has been reported but not described from Palawan (Villavicencio &
Andal, 1964), from Luzon by Hashimoto et al. 1978, and from the Jatibungkus Limestone of Central
Java, illustrated here (Image 2).

Neorotalia Bermudez, 1952

Type species: Rotalia mexicana Nuttall 1928
Neorotalia mecatepecensis (Nuttall, 1932)
1932 Rotalia mexicana NUTTALL var. mecatepecensis NUTTALL
1953 Streblus saipanensis COLE
1957 Rotalia mecatepecensis NUTTALL. - HANZAWA
1996 Pararotalia mecatepecensis (NUTTALL) MATSUMARU
The name Neorotalia mecatepecensis is rarely encountered in Southeast Asian references, although the
microfossil it is applied to has long been recognised in the region. In the 1950's the name almost be-
came established, appearing first as Miogypsinoides pre-ubaghsi, then as "Miogypsinoides pre-ubaghsi
(Rotalia mexicana)" (eg. Muhar 1957). Other workers have used names such as Pararotalia sp., Rotalia
mecatepacensis, Rotalia tuvuthalensis, Rotalia tectoria, Rotalia viennoti / Daviesina viennoti or Streblus
saipanensis. It is an assumption that all the reports of all these taxa, which are usually not illustrated, are
the same, but they all share a similar, short stratigraphic range in limestone facies of T.d to T.e (prob-
ably lower T.e) age. Streblus saipanensis Cole (1953) occurs in association with Te4 faunas on Saipan
Island and Te2-3 on Guam (Cole 1963).
Biostratigraphy of Indo-Pacific larger forams: page 78
 Stereopaired images of Neorotalia
mecatepecensis, a robust, large rotaliid from
the latest Oligocene Kranji Mudstone from
northeast Java.
In this area the form disappears just before
the first appearance of Miogypsina and near
the highest occurrence of the nannofossil
Helicosphaera recta.
These specimens show excellent detail of the test
ornamentation.
Dorsal images at top and bottom, umbilical in
middle.
Background dots are 100µm in diameter; a row
of 8 dots = 1mm.

Biostratigraphy of Indo-Pacific larger forams: page 79

Images of the robust, large rotaliid Neorotalia mecatepecensis, from the basal Late Oligocene of Central Java (Letter Stage T.d,
nannofossil Zone NP24), type locality of the Pelang formation. Parts of the coarse intraseptal canal system and umbilical canal can
be seen
Nummulites Lamark 1801
Type species Camerina laevigata Bruguiere, 1792
Nummulites should not be confused with Palaeonummulites. Nummulites is much larger, especially in
the microspheric form, and with a distinctly larger proloculus in the macrospheric form (0.2 to 1 mm).
Palaeonummulites has no significant variation in size between micro- and macrospheric generations,
which are both typically 2mm in diameter and the proloculus of the macrospheric form is less than 0.2
mm in diameter. In Nummulites the chambers, viewed in equatorial section, are about 1.5 times as high
as they are long (in the direction of coiling) or less. In Palaeonummulites they are are at least greater
than 1.5, often more than twice as high. These two genera need to be separated as they have very dif-
ferent stratigraphic ranges.
Ranges T.a1 to T.d (Paleocene to mid Oligocene)
From a simple Palaeonummulites ancestor the genus becomes more complex and ornamented, reach-
ing a peak in the Middle Eocene. Ornamentation evolves from smooth to granulate (i.e. pillars forming
granules on the surface). Nearly all granulate forms became extinct at the end of T.a. There is also a
progression from radial septal traces to sinuate, meandrine, semi- and then fully reticulate ones.
In the ontogeny of the “advanced”, large Middle Eocene forms there can be a juvenile spiral that is
regular with simple septa (such a juvenile stage is not clear in the species found in SE Asia), a prolonged
middle spiral phase of often slightly irregular whorls and complex alar prolongations, and a terminal
phase, more narrowly built and more slowly increasing in size, still with complex septa traces but
without a distinct marginal cord.
The literature on European and Middle Eastern Nummulites is vast and includes many works on phyl-
ogeny and speciation. Important examples are: Boussac (1911 a & b), Douvillé (1919), Abrard (1928),
and Schaub (1953 & 1966).
The taxonomy of species found in South East Asia is currently under review. However the following
notes summarise species concepts at the moment.

Historical notes and species of Nummulites / Palaeonummulites in Indonesia

The Nummulitids were studied by Verbeek at the end of the 19th Century, reviewed in a short work
by Doornink 1932. The study of Nummulites pre-dates the recognition of dimorphism (A and B, or
macro- and microspheric generations) so the earliest morphotypes established as species included both
generations with separate names. After the discovery of dimorphism it became a practise to hyphenate

Biostratigraphy of Indo-Pacific larger forams: page 80

The generalised test structure of Nummulites, from Racey 1992.

 names such as N. fichteli-intermedius. Such compound names are incorrect under a strict view of the
laws of nomnclature, and there has been a trend to follow the code and revert to a single name for a
dimorphic pair, using only the senior name. This of course assumes that one can reliably know which
pairs are dimorphic. For some taxa this is fairly well established, but for others there is probably more
doubt than some workers would like to admit.

Legally available names.

Names in red are commonly accepted as occuring in SE Asia, or were defined here.
In order of seniority
Nummulites gizensis Forskål, 1775 [originally placed in Nautilus?]. Reports of N. gizensis (often mis-spelt gizehensis)
in older reports from Indonesia refer to N. javanus, which has been shown to be a separate species, (cf.
summary in Renema et al. 200?)
Nummulites laevigatus (Bruguière 1792) THE TYPE FOR NUMMULITES [originally placed in Camerina]. The B
form has sub-reticulate to reticulate septal traces, with granules on the filaments, size is about 15 mm.
The B form has simpler, sigmoidal to sub-reticulate traces, granules, more n the center, and is typically
31⁄2 mm The B form has about 15 whorls and the A form about 5. This strong dimorphic variation is con-
sidered one of the diagnostic charatcters of the genus, hence its separation from the much longer ranging
Palaeonummulites. Reports of N. laevigatus in older reports from Indonesia refer to the microspheric or
B form of N. djokjokartae.
Nummulites laevigatus Lamarck 1801 -Name conserved as the generic name by ICZN 1945.
Nautilus mammilla Fichtel & Moll 1798. The original now considered Amphistegina, see Rogl & Hansen 1984
Palaeonummulites venosus (Fichtel & Moll 1798). Type from Recent from the Arabian side of the Indian Ocean.
[originally placed in Nautilus]. Probably the senior synonym to P. cumingii.

Biostratigraphy of Indo-Pacific larger forams: page 81

Above: Nummulites variolarius

Palaeonummulites variolarius (Lamarck) 1804 [originally placed in Lenticulites] (above, from Doorninck 1932, all
x10, fig 11 being a B form and almost 4mm diameter)
Not Camerina variolaria Doornink 1932
= Camerina semiglobula Doornink 1932
As Doornink pointed out in 1932, the early literature of this species is very poorly illustrated and there is the
possibility of confusion. Doornink himself suffers from this lack of illustrated examples as he applies the name
to forms with granules or pustules. He claims to follow the examples of Lister (1905), Rozlozsnik (1929), and
 van der Vlerk (1929), and justifies this by pointing out that “why did Lamarck name the species ‘pocked’ or
‘spotted’? Spots can be caused in Camerina only by column heads”. Lamarck was referring to the overall small
size and form of the species when he named the species “variolaria”. To quote Lamarck’s type description (in
translation);
“It is very small, not greater than 1 mm in diameter, and looks like the newly formed pustules /
spots from measles or syphilis. I thought the test belonged to the family of N. planulata but the
two faces are more convex and the septal filaments (cloisons) which form outside it are radial
striae, more numerous and more frequent. The aperture of the last chamber is less angular than
in N. planulata.”
(N.B. Nummulites planulatus (LAMARCK) is smooth, without granules.) In mis-identifying N. variolarius Doornink
needed to establish a new species, Camerina semiglobula, to cover the forms closest to the accepted type of N.
variolarius. They are very small with both A and B forms included in this definition as there is very little differ-
ence in size and form apart from in the protoconch. The septa are curved, radial with a more or less developed
umbilical boss. A few examples were placed by this worker in N. mamilla (Fichtel & Moll).
The range of Eames in Morley Davies, of Mid Middle through Late Eocene fits our current data in Indonesia.
The forms mis-placed by Doornink in N. variolarius probably belong in N. taballarensis (cf. Renema et al.
2003)

Nummulites orbignyi Galeotti 1837, suspected B form to Nummulites prestwichianus (Jones) 1862. In Indonesia this
species has been noted by Doornink (1932) in the lower Nanggulan river sections, but these specimens
are probably better placed in N. crasseornatus.
Nummulites irregularis Deshayes 1838, B form, (Kali Watupuru and K. Balong Nanggulan according to Doornonk)
but these specimens are probably better placed in N. crasseornatus.
Nummulites acutus (Sowerby 1840) [originally placed in Nummularia] Maybe senior dimorphic equivalent to N.
djokdjokartae. See below.
Nummulites fichteli Michellotti 1841, A form
Nummulites intermedius d’Archaic 1846, B form
Nummulites beaumonti d’Archiac & Haime, 1853, emend Davies 1940. A “B” form, about a centimeter, septal fila-
ments straight, to slightly curved in the polar area, weak pillaring in the polar region, does not cause
external pustules, and may not be seen in adult whorls. Appears to be a valid species, recorded at several
SE Asian locations, Ta restricted.
Palaeonummulites cumingi Carpenter 1860, Type from Recent in the Philippine Sea. [originally placed in Amphiste-
gina]. Considered by Rogl & Hansen (1984), after examining type material of both, to be synonymous
with P. venosus.
Nummulites kelatensis Carter 1861.Type material re-examined by Davies 1940, as some workers had identified this
form from the “Dutch East Indies, which, in certain cases, seem to be really N. beaumonti d’Archiac &
Haime.” ... “This species seems. as Carter said, to be quite distinct from N. Beaumonti, which is also a
much later form”. Considered here a junior synonym to N. beamonti. Biostratigraphy of Indo-Pacific larger forams: page 82
 Above: Nummulites pengaronensis
Nummulites pengaronensis Verbeek 1871, (above, from Doonink, 1932, specimen 1 from Jiwo Hills Gamping, and 2
& 3 A forms from Nanggulan all x10). Original illustration appears as B form but according to some, may
be A form to N. nanggoelani (type desc in German)
Numulites subbrongniati Verbeek 1871, defined as B form, but at the same time a “var. a” was also described as mac-
rosperic with same locations, Kalimantan. A reticulate form. (type desc in German)

Biostratigraphy of Indo-Pacific larger forams: page 83

Above: Nummulites djokdjokartae

Nummulites djokdjokartae Martin 1881. Considered the dimorphic A form to the B form N. acutus by Eames / Morley
Davies 1975, and this has been a widspread view in recent times. However Samanta (1982) & Renema et
al. (2003) consider the type of N. acutus to differ from N. douvillei [=N. vredenburgi] a subjective junior
synnym of N. djokdjokartae. On the other hand, outside comfort factors with various statistical views on
species concepts, the range of N. acutus appears similar to N. djokdjokartae so my view is that one can
assume dimorphism and call them both N. acutus or remain unconvinced and maintain N. acutus for the
B and N. djokdjokartae for the A form.
Nummulites javanus var α Verbeek 1891. A morphotype of N. javanus.
Nummulites javanus var α (soloensis) Verbeek 1891. A morphotype of N. javanus, with very little or no internal
pillaring.
Nummulites javanus var β Verbeek 1891. A morphotype of N. javanus, but illustrations appear varied.

Biostratigraphy of Indo-Pacific larger forams: page 84

Nummulites javanus var γ Verbeek 1891. This flat form now considered Nummulites boninensis Hanzawa.
Nummulites javanus var δ Verbeek 1891. A morphotype of N. javanus.
 Above: Nummulites bagelensis
Nummulites bagelensis I Verbeek 1891. An A form, larger than II, possibly the A form for the larger N. boninensis.
Nummulites bagelensis II Verbeek 1891: an A form. Bagelen residency, Kali Gorang, Karangsambung and Pesawahan
villages, Gunung Lodang & Lamouk, all the terrain of Lukulo and Bagelen. The village of Sampang west
of Bagelen.
Nummulites bagelensis II Verbeek 1891, var megaspherica Rutten 1915: an A form. Rante Boea, Makale, Central
Sulawesi.

Above: Nummulites nanggoelani

Nummulites nanggoelani Verbeek, 1891 B form, considered a junior synonym to N. pengaronensis.
Palaeonummulites niasi I Verbeek 1896, proposed as new name for Nummulina ramondi (defrance) var verbeekiana
Brady 1875. Probably a junior synonym of P. venosus.
Palaeonummulites niasi II (Verbeek 1896), type from Recent sediments, may be couple to N. niasi I [originally placed
in Nummulites]. Probably a junior synonym of P. venosus.
Biostratigraphy of Indo-Pacific larger forams: page 85

Palaeonummulites doengroeboesi Verbeek 1908, Both A and B recognised. Pliocene type from the river Ngetos, kam-
pung Doungbroubrous, near the boundary of the residecies of Madiun and Rembang East Java. [originally
placed in Nummulites]
Nummulites vredenburgi Prever 1908
Nummulites nuttalli Davies 1927 apparently B-form to N. thalicus. The type for Ranikothalia, a Paleocene genus.
Reports of this form in Indonesia have been amended to N. borneensis.
Nummulites thalicus Davies 1927 apparently A-form to N. nuttalli. Reports of this form in Indonesia have been
amended to N. taballarensis
Nummulites semiglobula Doornink 1932. A junior synonym for N. variolarius. Doornink notes this form from Nanggulan
in the Kali Watupuru, Songo and Balong, as well as the Gamping area in the Jiwo Hills. A and B forms.
Both location in the upper part of the Middle Eocene.
Nummulites densa Doornink 1932, (above, all x10) Kali Dowo in the Jiwo Hills area, B forms only, probably a junior
Biostratigraphy of Indo-Pacific larger forams: page 86

synonym of Nummulites beaumonti.

Nummulites gerthi Doornink 1932, (above, all x10) Found in Gunung Gamping and Cikarang, Bantam ["fig 21"
above], and the Pellatispira limestone at the top of the Nanggulan section C. Java, all Tertiary B locations.
A “B” form which is probably a junior synonym of N. pengaronensis.

Biostratigraphy of Indo-Pacific larger forams: page 87

Nummulites hoogenraadi Doornink 1932, (above, all x10) A form, TjiTjukarang. As described also by Koolhoven
1933, this form is found in a stream in the Cicarucup beds north of Bayah (near Langkop), in early Tertiary
B stage, and at the Cimuncang sections of similar age to the west. In both locations it is recorded as fairly
nummerous, as A forms only. Preservation is not good in this area due to a high degree of mineralisation
and thermal decomposition of sediments. An examination of topotypic material at the GRDC notes a
simple lenticular form, 3-4mm, curved radial septa, weak trabeculae and traces of pillaring forming 9-10
small pustules in the center of the test. It may be a new species or a poorly preserved N. gerthi (A form
of N. pengaronensis) or possibly N taballarensis, which, if the latter would be the youngest record yet of
this form.
Nummulites divina Doornink 1932, A form, Cijengkol (probably the more fossiliferous Ciapus-Cimanggu area). Similar
to N. fichteli, an A form, but thought to be dimorphic with N. subbrogniarti rather than intermedius. The N.
subbrongniarti-divina pair having a coarser reticulate mesh and larger size than N fichteli-intermedius.
Nummulites absurda Doornink 1932, A form Tji sukarama. Probably a junior synonym of N. fichteli (A).
Nummulites crasseornatus Henrici, 1934, Timor. Recently found in olistoliths in the Middle Eocene slump deposits
at Kali Sana, near Karangsambung, C. Java (Figure X)
Nummulites borneensis (Caudri) 1934. The name that replaced N. nuttalli, misapplied to SE Asian types. This is the
rare B form. This species was described by Caudri immediately before the much more comon A form
taballarensis and therefore has priority should the two names be considered dimorphs.
Nummulites taballarensis (Caudri) 1934. The name that replaced N. thalicus, misapplied to SE Asian types. This is
the much more common A form.
Nummulites discoidea (Caudri) 1934. A rare form which could be either oblique sections through, or a variant of N.
crasseornatus.
Nummulites kemmerlingi (Caudri) 1934. Only a few specimens found by Caudri on Sumba, in Ta letter stage, and only
single axial and equatorial sections illustrated. Uncertain affinity. possibly N. crasseornatus
Nummulites boninensis Hanzawa 1947, A form like a flattened N. javauns.both A and B forms included in definition.
Type from the Eocene on the island of Haha-Jima of the Ogasawara [Bonin] group, Japan.

In addition, the collection of the GRDC in Bandung has a large collection of specimens labelled for a work that was
never published, possibly never completed, and is now apparently lost. These names are therefore not
valid but occur on many samples and some notes at the GRDC.
Nummulites balongensis Tan (unpub) - An "A" form, probably N. crasseornatus. Usually 21⁄2 to 3mm. From Kali
Balong Middle Eocene of Nanggulan.
Nummulites canaliferoides Tan (unpub) - A "B" form8 mm in size, with more pillaring than N. indistncta, probably
N. crasseornatus. From sites at Nanggulan.
Nummulites crijptospira Tan (unpub) - "A" & "B: forms like N. crasseornata. From sites at Nanggulan.
Nummulites doorninki Tan (unpub)
Nummulites indistincta Tan (unpub) - A "B" form 6-7 mm in size, probably N. crasseornatus
Nummulites spirifera Tan (unpub). A small form not unlike P. variolarius. From sites at Nanggulan.
Nummulites subpelius Tan (unpub) Kalimantan. Reticulate septal filaments, "A" generation and close to N. divina in
form.
Nummulites zwierzyckii Tan (unpub)

Biostratigraphy of Indo-Pacific larger forams: page 88

DETERMINATION TABLE FOR MAIN SPECIES OF S. E. ASIAN NUMMULITES
For each proposed dimorphic pair the senior same is underlined. The junior name is valid if applied to the correct
morhotype, but its use implies that there is doubt about the association of the two generations.
"B" Forms with meandrine septal traces
• With fine, but distinct granules
Maximum diameter typically 20 mm but some larger forms reported
Nummulites djokdjokartae (Martin)
[=N. acutus Sowerby of some authors]
Highly likely A form; with sub-reticulate septal traces and strong, protruding,
granules over most of test [not final whorl] over septal filaments, proloculus 0.5 to 0.8 mm
Nummulites djokdjokartae (Martin)
• Granules visible in axial section but not reaching the surface
(Granules may be visible in hand-specimen when wet, but form no relief on the surface)
Maximum diameter typically 40 mm, flat (<5mm max. axial dimension)
Nummulites boninensis Hanzawa
[=N. javanus var γ Verbeek of some authors]
A form; found in association, lenticular small and with radiating, curved
septa and some granules over septal filaments, proloculus 0.5 to 1.0 mm
Nummulites boninensis Hanzawa
Maximum diameter typically 25 to 30 mms, lenticular with axial dimension
c. 1/3 rd of equatorial dimension. Nummulites javanus Verbeek
[=N. javanus vars α, possibly part of β and δ of Verbeek]
A form; found in association, lenticular small and with radiating, curved septa,
very similar to A for of N. boninensis. Proloculus 0.5 to 1.0 mm and maybe more than this,
but affinities of these megalospheric A forms unclear Nummulites bagelensis Verbeek
"B" Forms with reticulate septal traces
• Test typically 10 to 15 mm maximum diameter.
Flatter form than A form, with axial dimension 1/4 of equatorial.
Nummulites intermedius d’Archiac
Highly likely A form; with reticulate septal traces, lenticular up to 4 mm,
Proloculus 0.25 to 0.35 mm. Consistently associated with intermedius, within both
stratigraphic and geographic limits. The senior name of the dimorphic pair.
Nummulites fichteli Michellotti
• Test typically 15 to 25 mm maximum diameter. Coarser spaced reticulation than N. intermedius
Similar to N. intermedius and subjective junior synonym, except for coarser mesh of reticulate
septal traces, slightly tighter coiling and larger size
Nummulites subbrongniarti Verbeek
Highly likely A form; with relatively coarse reticulate septal traces, lenticular up to 5 mm.
Proloculus 0.25 to 0.35 mm. In axial section each instar of chamber growth is, in the adult
phase, 3 to 5 times as long in the spiral direction as height between whorls. For the very similar
fichteli this ratio is rarely more than 3. Fairly consistently associated with subbrongniarti,
within both stratigraphic and geographic limits. The junior name of the dimorphic pair.
Nummulites divina (Doornink)
Biostratigraphy of Indo-Pacific larger forams: page 89

Some workers do not differentiate these pairs of species, in which case the senior names are
N. fichteli (and N. intermedius if dimorphic forms are being kept separate)

"B" Forms with radial, more or less curved septal traces (included in Palaeonummulites by some)
• Septal traces straight, but terminate at polar area possibly becoming curved. Max 9 mm.
Similar to N. pengaronensis, except for a much higher numbers of chambers per whorl, which
leads to straighter forming (more packed) septal traces in equatorial section. In the fifth whorl of
the lectotpe there are 39 chambers. Also fine pillars in axial section but only in inner part of test,
not on surface.
Nummulites beaumonti (d’Archiac & Haime)
[=N. kelatensis of several authors in Indonesia]
Equivalent A form not established.
 Septal traces slightly curved. Max 9 mm.
Similar to N. beaumonti, except for wider spaced, slightly curved septal traces which, in equa-
torial section appear curved back between the previous whorl and the periphery. (for example,
in the type illustration of Verbeek the fifth whorl contains 21 chambers.)
Nummulites pengaronensis Verbeek
[=N. nanggoelani Verbeek]
A form; found in association, lenticular small (31⁄2 mm) and with radiating, curved septa. Proloculus
0.2 to 0.25 mm. Naming of A form has unclear history. Is probably in part the form called
Nummulites gerthi (Doornink)
Septal traces curved to S-shape. Distinctly thin, with inflated marginal cord. Max 7-8 mm.
Coarse granules are present in the central, slightly thickened, part of the test, aligned in a
rough spiral trend. Distinctly thin form, with sometimes irregular periphery (not completely flat edge)
In equatorial section chamber lumen are much higher in radial dimension, than in direction
of spiral growth (2 or 3 x). A rare form in SE Asia.
Nummulites crasseornatus (Henrici)
A form; found in association, lenticular small (3 mm), with pronounced marginal cord,
coarse granules in central (polar) area, Proloculus 0.25 to 0.3 mm. Also called N. crasseornatus.

Septal traces gently curved to S-shape. Lenticular,becoming flatter in last whorl. Max 5 mm.
Still taxonomically confused. The B form is rare, hence the preference of some authors to use the name of
the much more common A form, although technically the junior name.
Nummulites borneensis (Caudri)
A form; found much more commonly. Lenticular, small (21⁄2 -3 mm) and with radiating,
only slightly curved septa. Coarse (for such a small test) granules as pillars in rough spiral
alignment over central part of test. Proloculus 0.35 to 0.5 mm.
Nummulites taballarensis (Caudri)

"A" & "B" Forms almost indistinguishable, with radial, slightly curved septal traces
(included in Palaeonummulites by most modern authors)
No granules, but an umbilical pillar present. Max 3 (rarely 4) mm.
A simple small, slightly inflated lenticular form, with very little dimorphic variation.
Proloculus (check Eames et al.) 0.15 mm in A generation.
Nummulites OR Palaeonummulites variolarius (Lamark)
[=N. bagelensis, in part, of Verbeek, and N. semiglobulus of Doornink]

Operculina d’Orbigny 1826

Type species Lenticulites complanatus Defrance, 1822 Biostratigraphy of Indo-Pacific larger forams: page 90

Assilina, involute (but without alar prolong- Operculina,evolute, faster

gations), slow rate of whorl growth. rate of chamber growth
(spiral opening rate about 1.4) (spiral opening rate about 2.1)

Specimens of Operculina that are like the type species O. complanta are easily identified, but there is
a range, or plexus of forms that contains more and less involute types, some may be ornamented but
 most are smooth. Fully involute forms (esp. in the Paleocene) overlap with the definition of the closely
related genus Palaeonummulites. In Neogene to Recent times fully involute forms may have thinner
walls than older Palaeonummulites and a slightly faster rate of whorl growth, but otherwise the generic
definitions grade into each other.
Range: Paleocene, Ta1, forms are reported by Adams (1970), to Recent (extant)

Orbitolina d'Orbigny 1850

Type species Orbulites concava Lamark 1816
A large conical arenacous foraminifera with a fine-gained dark walled appearance in thin section. Is is
a rare Early Cretaceous form in SE Asia, requiring specialist knowledge to study in detail. Oil wells
drilled to basement in parts of Kalimantan sometimes recover fragments of this fossil in the cuttings,
as shown below.

Above and right; images of Cretaceous

Orbitolina in a thin section made of ditch
cuttings from TD of an exploration well NW
Biostratigraphy of Indo-Pacific larger forams: page 91

of the Meratus Mountains of S.E. Kalimantan.

Identification of this formation defined
economic basement for this well.
The multiple subdivisions of the marginal zone
are clearly seen, but the zig-zag to sinusoidal
radial pattern of intraseptal beams is not
obvious, -certainly not as obvious as pencil
drawings suggest. Some wavy fabric can be
seen elsewhere in this thin section.
Orbitolites Lamark, 1801

Type species: Orbitolites complanatus Lamark, 1801 (nom. imperf as O. complanata)

A large Alveolina (?A. javana) with Orbitolites to the left, shown enlared at the top. From E. Kalimantan (GRDC coll.) maximum
diamter of Orbitoides 3.2 mm

Large discoidal test, flat to slightly biconcave, large embryonic chambers that differ from Marginopora
by being more “nephrolepidine” in shape while that latter has “eulepidine” shape. Unlike any of the later
discoid miliolids there is no sign of a peneropline juvenile phase. Ogival to arched chamberlets surround
the embryont and grow in a similar fashion to Sorites and Amphisorus and in equatorial section in is
difficult to distinguish these forms (especially from macropsheric, non- peneropline specimens of the
last two named genera). In oblique sections Amphisorus has its distinct two layered structure and Sorites
a relatively planar single layer. Orbitolites has a more strongly biconcave profile and internal partitions
complicate the chamberlets they do not form completely separate layers of chambers. Orbitolites does
not have the three fold apparent layering of Marginopora.
Ranges from Ta 2 to Ta3, possibly into Tb as records from South Sulawesi indicate it overlaps with
Pellatisipra (Dollfus, 1915) but authors such as Eames (in Davies 1975) have flagged this as requiring
Biostratigraphy of Indo-Pacific larger forams: page 92

clarification and it is pointed out here that Pellatisipra occurs in latest Ta3 sediments. Orbitolites is
found in Somalia (Azzaroli, 1952) with Tb Palaenummulites forms specifically P. fabianii.
Not a very common genus in Indonesia, largely as Middle Eocene marine strata are relatively rare. Known
from South Sulawesi (Tonasa limestone [Dollfus, 1915] and later authors) and from the Masungit Lime-
stone on Luzon Island in the Phillipines (Hashimoto et al, 1979). It is interesting to note in Hashimotoʼs
data that samples with Orbitolites were without any Nummulites or Assilina and vice versa, suggesting
strong facies control. Records of this genus on Java (van Bemmelen 1937) have been checked but van
Bemmelen's own slides contain a different and as yet un-named form not a miliolid.
Palaeonummulites Schubert, 1908
Type species Nummulina pristina Brady, 1874
Like Nummulites except smaller, with negligible variation between microspheric and macrospheric
generations and without the ornament and complex septal arrangement of Nummulites. Some authors
include small, simply ornamented forms such as N. variolarius as being within Palaeonummulites.
In cases where involute Palaeonummulites and Nummulites are in doubt, Nummulites is larger, espe-
cially in the microspheric form, and with a distinctly larger proloculus in the macrospheric form (0.2 to
1 mm). Both micro- and macrospheric generations of Palaeonummulites are typically no more than 2
or 3 mm in diameter, the proloculus of the macrospheric form is less than 0.2mm in diameter, and there
are no more than about 5 whorls in the adult A form. The spiral opening rate in Palaeonummulites is
higher than in nearly all Nummulites, with whorls being about 1.5 times greater diameter than preceeding
whorls, consequently chambers can be up to twice as high as long, as viewed in equatorial section.
Palaeonummulites ranges from middle Paleocene to Recent and is a cosmopolitan form. Published
summaries might suggest it to be a European and Indo-Pacific organism, but this is probably only a
result of the traditional taxonomic approach of Caribbean - American workers to place such forms in
Nummulites s.l.
Hottinger (1977) has argued that Amphistegina cumingii, the type species of Operculinella, has an
identical structure to that of Nummulites. Eames et al. (1962) regard both Operculinella and Oper-
culinoides to be synonyms of Palaeonummulites so all three genera are regarded in the Loeblich and
Tappan treatise as being Nummulites. Palaeonummulites however remains a practical and legal genus
for biostratigraphers. Palaeonummulites is locally common in later Miocene and younger limestones
in Indonesia, sometimes occuring with Trybliolepidina and sometimes as the dominant member of an
assemblage.

Palaeonummulites

Biostratigraphy of Indo-Pacific larger forams: page 93

L. (Trybliolepidina) Cycloclypeus Palaeonummulites

Palaeonummulites from an un-named late Miocene limestone, north central Java, occurring with Nephrolepidina and Cycloclypeus.
They can be distinguished from Cycloclypeus as the latter has such a distinctly thin layer of chamber lumena, as well as the overall
coiling form and symmetry seen in more complete specimens.
Pellatispira Boussac 1906
Type species: Pellatispira douvillei Boussac 1906 = Nummulites madaraszi Hantken 1876
Found in later Ta3 in many locations (cf. Umbgrove 1928, and Jiwo Hills, Java herein). Is the ances-
tor to the complex Biplanispira and Vacuolispira forms which, with the exception on one record of
Vacuolispira in Ta3 illustrated here (Halmahera, under Vacuolispira), are both Tb restricted. In later
Tb these complex forms are more common than Pellatispira. Lunt (2003) has noted that Pellatispira,
Biplanispira and Vacuolispira (as well as the possiblly ancestral Planocamerinoides or Assilina) are
almost never found on the Australian Plate, being predominantly low latitude genera.
Examples of Pellatispira are shown on Images 10-13. An important review of the morphology and
taxonomy of the genus and whole subfamily was recently published by Hottinger, Romero and Caus
(2001) . These workers recognise three species of Pellatispira , the widespread, large (3 - 4+ mm) P.
madaraszi with its A generation proloculus larger than 200 µm, the smaller (1.5 to 2 mm), often more
lenticular and robustly pillared P. provalei Yabe 1921, with its A form proloculus invariably less than
100µm, also is the Indo-Pacific restricted P. fulgeria Whipple 1932 with its considerable extension of
the marginal crest into a complex supplimentary skeleton around a rapidly opening spiral of chambers.
this often large form (7mm +) is also very flat, although it may develop spiral rims and similar orna-
ment. It has not been found as old as Ta.

Pellatispira fulgeria with a distinctive highly developed supplimentary skeleton, in this example forming a bi-concave, dumb-bell
form, from a xenolith in the Sangiran Dome, Central Java. Width of specimen is 4 mm
Biostratigraphy of Indo-Pacific larger forams: page 94

Genus Penerolpis deMontfort, 1808

Type species Nautilus planatus Fichtel & Moll, 1798
Peneroplis is almost unknown from the fossil record of the Indo-Pacific area, but it is recorded in
small numbers from studies of Recent faunas, e.g. Cushman, Todd and Post's (1954) extensive study
of the Marshall Islands. Peneroplis is a simple planispiral, generally flat, evolute miliolid form. There
are multiple coarse pores on the apertural face. Externally Peneroplis usually has fine striations in the
direction of growth, but it does not have any internal divisions or pillars within the chambers. It was
probably the ancestor to Sorites, Amphisorus, Archais and Marginopora.
In the Caribbean, the Middle East and East Africa it has been recorded from sediments as old as Eocene
(e.g. Henson,1950). It is considered a back-reef facies form.
Planocamerinoides (=Assilina of older workers)
Planocamerinoides Cole 1958
Type species: Nummulites exponens Sowerby 1840
Recent publications by Haynes (1988) and Haynes et al (in press) have reclassified these genera with
the effect that possibly all forms previously considered Assilina in the literature on SE Asia are now
placed in Planocamerinoides (cf. Renema et al 2003). The difference is that Assilina sensu stricto is
evolute with no lamellar or spiral sheets extending over the previous whorls, with the effect that with
chamber growth the test is flat to biumbilicate. Planocamerinoides has multilamellar thickenings over
the umbonal area, except in P. umbilicata which has thickening over the previous whorl only, and is
consequently strongly biumbilicate. Serra-Kiel et al. (1998) do not distinguish Assilina from Planocam-
erinoides and also include a number of species which range higher than top Ta3 (top of their SBZ17)
in Assilina. These are all species which workers in Indonesia would have assigned to Operculina, with
much faster rates of whorl growth. Examples of Planocamerinoides are on Image 7.
Planocamerinoides ranges from Late Paleocene to the top of Ta, near top Middle Eocene.

Genus Planorbulinella Cushman, 1927 (inc. Planolinderina Freudenthal 1969)

Type species Planorbulina vulgaris dʼOrbigny, 1865
Planorbulinella consists of a single layer of chamberlets growing as a relatively irregular flat disk.
Chamber walls are finely porous. The embryont has three chambers, distinguished from post embryonic
chambers by having thicker walls. There may be a short juvenile spiral and there is an inverse corre-
lation between the size of the embryont and the numbers of chambers in the juvenile spire.
In the juvenile stages the chambers are connected by single stolon but, as part of the move to radial
growth, chambers with two stolons soon appear; - as is the case for most larger foraminifera. However
a characteristic of this genus is that sometimes there may be a few subsequent quasi-nepionic cham-
bers that revert to having one stolon, slightly interupting the radial symmetry. These are called relapse
chambers.
There is no evidence that Planorbulinella, which is extant, contains any photosythesising symbionts.
This might explains its relatively small size and frequent distribution in non-carbonate sediments.
A closely related species, Planolinderina was defined in 1969 by Freudenthal. This differs from Planor-
bulinella as apertures are said to be a series of basal openings, instead of a single slit, and there are no
records of relapse chambers. Such a difference is difficult to apply in examination of fossil material.
Ranges from within the Oligocene to Recent. One species, Planorbulinella solida has distinctly thick-
ened lateral walls and is found from Middle Miocene times onwards.
Biostratigraphy of Indo-Pacific larger forams: page 95

Planorbulinella from the Late

Oligocene Citarete Limestone
of West Java (T.e1 or T.e2-3).
Image at left has an upper part
with axial view and lower part
with near equatorial section with
a fragment of Eulepidina.
 Specimens of Planorbulina solida from
the Bodas limestone, Central Java (Late
Miocene).

Genus Planostegina Banner & Hodgkinson 1991

Type species Heterostegina costata d'Orbigny1839
Biostratigraphy of Indo-Pacific larger forams: page 96

See also entry under Heterostegina.

This genus was erected to cover the forms of Heterostegina that descend from flat, wholly evolute
"Operculina" (=Planoperculina Hottinger 1977 emend Banner & Hodgkinson 1991) by the develop-
ment of secondary septa. The type species is from the Middle Miocene of Austria and equivalent Mid-
dle Miocene forms are found in Indonesia. There are also large and flat Planostegina species frequent
in the Tertiary c, in particpular P. bantamensis and P. praecursor (the latter having a central, umbonal
boss). These forms ere ancestral to Cycloclypeus and the juvenile stage of early Cycloclypeus contains
a Planostegina phase.
Planostegina costata from the Middle Miocene (planktonic
foram N10-11) of East Java (sandy mudstone within the
Ngrayong Formation).

The stereopair and the image of the left are the same
specimen. Background dots are 100µm in diameter; a row
of 8 dots = 1mm.

Below: reflected and transmitted light views of same

specimen. Of the material available this has the most
completely developed secondary septae.

They differ from the latest Tertiary to Recent Operculina

heterosteginoides as the latter has incipient but not truely
developed secondary septa. They can be distinguised from
the much larger Planostegina operculinoides illustrated
later, by their small size, lack of orament and often
irregular initial secondary septa.

Biostratigraphy of Indo-Pacific larger forams: page 97

 Planostegina bantamensis and P. praecursor (the latter with central umbonal boss) from the type location
at Cimanggu (Tjimanggoe) West Java, associated with Tertiary c foraminifera and with Early Oligocene
ratios of stable isotopes of strontium.Near centre axial section on left, and stereopair of specimens above
clearly showing the flat, evolute Planostegine nature, similar to its flat descendent Cycloclypeus.

Biostratigraphy of Indo-Pacific larger forams: page 98

Right: Planostegina bantamensis or praecursor

TAN from close to the type location, from the
Ciapus section West Java, either Tertiary c or
possibly Td age. The maximum dimension of this
individual is almost exactly 4.5 mm. An equatorial
section.
This specimen is from the same beds Tan Sin Hok studied in
his 1932 monograph on Cycloclypeus. The large, flat Cycloclypeus
specimens in the Tertiary c and d have prolonged heterostegine juvenile phases (or nepionic stage), which can be mistaken for
Planostegina. In Tan's Ciapus 14 sample the Cycloclypeus specimens had between 16 and 30 (most commonly 21) neopionic or
heterostegine chambers, before developing annular Cycloclypeus chambers. The specimens above has at least 50 heterostegine
chambers and therefore is clearly not a juvenile Cycloclypeus. The diameter of the initial chamber (protoconch) is almost 130
microns.
 Biostratigraphy of Indo-Pacific larger forams: page 99

Stereopairs of Planostegina operculinoides HOFKER 1927, Recent, from a dredge in about 40 meters of water
off Sakala island, eastern Java Sea. (Type specimen and paratypes found by Hofker in the same general area,
in 59m to 90 m of water.
Specimen above is photographed from both sides.
The specimens above have completely evolute coiling, typical of this genus.
Background dots are 100µm in diameter; a row of 8 dots = 1mm.
Genus Praerhapydionina Van Wessem, 1943
Type species Praerhapydionina cubana Van Wessem, 1943
This elongate, circular miliolid form is mostly known from the Middle East, and although Adams (1970)
gives a distribution as far east as the East Indies, no positive examples of this genus are known to the
authors in the Indo-Pacific area. Its range according to Adams (1970) is Tc to lowermost Te.

Genus Pseudotaberina Eames, 1971

Type species Pseudotaberina malabarica (Carter) 1853
An infrequent Early to Middle Miocene form most commonly encountered in Southeast Asia in Papua
(Irian Jaya) and Papua New Guinea. The monospecific genus was poorly documented until Banner and
Highton (1989). It is an extreme variant of Archais, characterised by greater size (up to 2 cms for B-
forms), greater differences between A and B forms, with B forms often achieving many cyclical whorls
in the adult phase (A-forms up to 1 cm & may achieve limited cyclical growth). Most importantly,
the interseptal pillars fuse to form evenly-spaced subdivisons of the chamber. These divisions are not
always completely formed but are distinct enough in equatorial section. Apparently a descendant of
Archais by acceleration and increase in size, becoming highly planispiral and ultimately cyclical, with
better subdivision of the chambers. As with other larger foraminifera the descendant shows greater
differentiation between A and B generation.
Occurs with Miogypsina, Flosculinella bontangensis and Austrotrillina howchini. The summary in
Adams (1984) and Banner and Highton (1989) gives justification for considering its range to be from
within T.f1 to probably basal T.f2.

Genus Quasirotalia Hanzawa, 1967

Type species: Quasirotalia guamensis Hanzawa, 1967
Test plano-convex with a nearly flat spiral side, sometimes tending towards lenticular. Up to about 2
mm in diameter. Three or four whorls in adult with 15 or more chambers in the final whorl. The adult
has a few layers of chamberlets / vacuoles added to the periphery and the umbilical side. Pillars often
present on umbilical side. Coarse pores usually visible in the thick wall, best seen or best developed
on spiral side.
A rare genus originally found in the Pliocene of Guam. Specimens illustrated here from a latest Miocene
limestone (dated by SIS at about 6 MYBP) S.W. of Semarang, Central Java.

Biostratigraphy of Indo-Pacific larger forams: page 100

Quasirotalia ?guamensis from the same latest Miocene N4x limestone illustrated in Image 26
 Quasirotalia ?guamensis from the same latest Miocene N4x limestone illustrated in Image 26

Genus Schlumbergerella Hanzawa, 1952

Type species: Baculogypsina floresiana Schlumberger, 1896
A globular form typically 2 to 3 mm in diameter with slightly projecting spines or tubercles. In the
microspheric generation there is a planispiral coil of about 2 whorls without spines. Megalospheric
forms with an irregular, sub-spherical embryont composed of three chambers. No significant difference
in size between the two generations. Most of the test is composed of dome-like lateral chamberlets or
cubiculae. These fuse at sutures to form pillars which usually do not reach the juvenile chambers. Many
dozens of these small pillars may be formed, to varying degrees. Stolons or pores connect the lateral
chamberlets by some longer stolons or short canals may form along the edge of the pillars. These canals
only occur on the outer edge of the spines, which are otherwise solid.
It is possible to confuse Schulmbergerella with Baculogypsinoides (="Baculogypsina tetraedra") as some
specimens of the former have a tetrahedral shape. In Schlumbergerella the corners of the tetrahedra do
not contain prominent spines, and none of the pillars in this genus have an internal canal system.
Similar to the rare Eocene Wilfordia
Ranges Pleistocene to Recent (extant)
Ancestry uncertain. There is no reason to consider a descent from Calcarina (as in Baculogypsina and
Baculogypsinoides) in view of the absence of a canal system in the spines, and the detachment of the
pillars or spines from the juvenile.
Found in abundance on beaches in Bali (image below), Flores and other islands in Indonesia.
Biostratigraphy of Indo-Pacific larger forams: page 101
Genus Sphaerogypsina Galloway, 1933
Type species: Ceriopora globula Reuss, 1848
Paleocene to Recent
Test can large, up to 2 mm and is roughly spherical. Juvenarium small and not well ordered. Cham-
bers added in poorly defined spherical layers. Chamber roofs are perforate but walls imperforate and
slightly thicker, although not forming pseudopillars. Should not be confused with Schlumbergerella
or Baculogypsina as these are larger, with real or pseudo-pillars and larger and more complex juveile
stages. Wilfordia, which is not yet known outside the type locality in Sarawak also has pseudo-pillars
and complexembryont & juvenile but is of similar size to Sphaerogypsina. May be monospecific (S.
globula)
A cosmopolitan carbonate facies form being found in small numbers in most facies, with the exception
of restricted, miliolid rich, back-reef faunas.

Eulepidina and Sphaerogypsina globula. Northwest Kangean Island. In Late Oligocene deposits characterised by very large, platy
Eulepidina and Cycloclypeus.

Genus Spiroclypeus H. Douville, 1905

Type species Spiroclypeus orbitoideus H. Douville, 1905
and
Genus Tansinhokella Banner & Hodgkinson, 1991
Type species Spiroclypeus yabei van der Vlerk, 1925, from the Late Oligocene of NE Kalimantan.
Tansinhokella and Spiroclypeus are heterosteginid forms that are practically indistinguishable from
Heterostegina in perfect equatorial section. They differs by the presence of "lateral chamberlets" or
cubiculae which can be seen in axial or oblique sections. Both are completely involute, and consequently
Biostratigraphy of Indo-Pacific larger forams: page 102

thicker and more robust than even the broadest Heterostegina (Vlerkina) forms.
Banner and Hodgkinson (1991) defined the genus Tansinhokella, named after the eminent Indonesian
palaeontologist, as the descendant of Heterostegina (Vlerkina) and the ancestror of Spiroclypeus sensu
stricto. The morphological series was interpreted to occur as follows:
Heterostegina (Vlerkina) had long alar prolongations which are roughly radial from the umbonal areas,
and the alar prolongations are mostly un-divided by secondary septa (some secondary septa may cross
over into the outer most part). Thus in axial section a single whorl of the spiral sheet corresponds to
a single alar prolongation chamber division over the central part
Tansinhokella has obvious subdivisions deep into the alar prolongations and the numbers of layers of
divided alar prolongations does not readily match the recognisable primary chambers in the median
 layer. The alar prologations are more or less strongly curved and weave in and out of the plane of
section to help produce such complex lateral chamberlets.
Spiroclypeus does not have lateral chamberlets like those above but true cubicula which may stack
up in rows vertically out from the median plane and have thin walls, and are not formed as simple
subdivisions of a single "instar".
Spiroclypeus sensu stricto resembles a three-layered orbitoid such as Lepidocyclina, and care needs to
be taken in identification of less well preserved material.
The type species Spiroclypeus is S. orbitoideus, which is defined on a microspheric type specimen.
For some time after Banner & Hodgkinson's paper it was not clear if such cubicula development was
associated only with microspheric generations, which would invalidate this taxonomy. However sev-
eral examples have now been found that show cubicula on macrospheric or A generation species (See
Image 18).
A species of Tansinhokella, T. vermicularis, is recorded from many locations in sediments of the T.b
Letter Stage (Late Eocene). No records of this genus are known from the Early Oligocene and it re-
appears within lower T.e (the evolution defining the base of T.e 2-3), shortly after the first occurrence
of Eulepidina (Leupold & van der Vlerk, 1931). While Eulepidina overlaps with Nummulites in Letter
Stage T.d, there is are only two records of Tansinhokella occuring with T.c or T.d Nummulites faunas.
One of these is in the Lutut Beds of Central Java, where both these genera are reworked. The second
occurrence is from the Bugton Limestone in the Philipines (Hashimoto et al 1977) which needs re-ex-
amination, but unfortunately is of float material, not bedded outcrop.
The Late Oligocene lineage of Tansinhokella appears to evolve gradually from Heterostegina (Vlerkina)
borneensis, which has been called the “Spiroclypoid Heterostegina of Rutten” (Leupold & van der Vlerk,
1931). Heterostegina (V.) borneensis has thin, but deep alar prolongations as a result of involute coiling.
The work of Muhar (1957), re-sampled by myself, plus mid-Oligocene sediments from the Bayah area
of West Java, apparently shows this gradual transition from simple Heterosteginids with continuation of
septae and septulae into the alar prolongations to acceleration in the acquisition of these lateral chambers,
recticulation of the septa traces and rapid development in complexity in lateral chamber form.
The development of Spiroclypeus sensu Banner and Hodgkinson is not yet as firmly fixed. Specimens
of this type certainly exist in latest Oligocene times, based on Sr dating and the evolution datum of
Miogyspina. It has not yet been determined if Spiroclypeus s.s. evolved before or after Miogypsinoides
from N. mecatepecensis which defines the T.e2-3 and T.e 4 boundary.
The extinction of both these this genus appears to be coincident with that of Eulepidina, and this event
defines the top of Letter Stage Te. Many authors have noted that facies control on the distribution of
the both genera means that continous records are scarce and the relative order or synchrony of their
extinctions is not yet accurately established.
Since the work of Krijnen in 1931 there have been five main species recognised in the Indo-Pacific area.
More recent work suggsts these species are not all valid, but Krijnen's work was the basis for taxonomy
for many years, and his species concepts are therefore incorporated into many existing reports.
Biostratigraphy of Indo-Pacific larger forams: page 103

Krijnen's work, on samples from Java, Sumatra, Christmas Island, Borneo and the Philippines, and the
range charts of Leupold and van der Vlerk (1931), and Rutten (1947), suggests that, with the excep-
tion of S. orbitoideus which is defined by him as a microspheric B form, all these taxa occur through
nearly all of the stratigraphic span of the genus. All except S orbitoideus would now be considered
Tansinhokella.
By the 1970's the concept of these five species had changed, and it is uncertain as to when this happened.
A table by Baumann in 1971 uses very different criteria to Krijnen and also hints at a gradual series from
architecturally "simple" S. pleurocentralis to "complex" S. orbitoideus. The associated stratigraphic
succession such a development theory implies has not yet been observed. A direct conseqence of this
change in concepts is that the later observation to species level are not consistent, as it is not known on
 what basis each workers defined his or her species. Consequentely environmental interpretations based
on species is not yet possible.

Species pleurocentralis tidoenganensis margaritatus leupoldi orbitoideus

Pustulate
External texture Pustulate Pustulate reticulate Reticulate Reticulate

Walls between Much thicker Much thicker than About the same Thinner than Thinner than
lateral chambers than lateral lateral chambers thickness as the lateral chambers lateral chambers
chambers lateral chambers
Shape of lateral
chambers Irregular long Irregular long Becoming regular Regular short Regular short

Approx numbers
1 to 30 30 to 70 70 to 110 110 to 180 more than 180
of lateral chambers

Determination table for species of Oligo-Miocene Spiroclypids in axial section, based on the five species established by Krijnen
(1931). The above definitions were used by oil industry biostratigraphers in Indonesia in the 70's and 80's. From Baumann, 1971.
This table is for reference on reading older reports. It is not a receommended taxonomy for the Spiroclypids

Genus Sporadotrema Hickson, 1911

Type species Polytrema cylindricum H.J.Carter, 1880
A homotremid form, attached, with cyclindrical branches. Chambers initially planisprial but then loosely
spiraling aournd a core of irregular passages that open distally. Outer wall thick and coarsely perforate
but these perforations change to muliple fine pores before penetrating the inner wall. Hickson's photo-
graphs shows how some chambers can be inflated near the distal ends (where they may not have thick
secondary lamellae obscuring the sutures). Overall the chambers are notable less inflated that Victo-
riellids. There are no pillars or pustules. Early upright spiralling chambers have a terminal aperture
that remains as a foramen to subsequent chambers before the development of the complex network of
stolons of passages in the central core.
Ranges from within the Eocene to Recent (extant). A good example is shown in Image 24.

Genus Vacuolospira Tan Sin Hok 1936

Type species: Pellatispira inflata Umbgrove 1928.
Latest Ta and Tb of the Indo-Pacific region
Generally globose, very thick walled forms. While small interlamellar spaces are seen in the gerontic
stage of Pellatispira forms, in Vacuolospira the primary chambers become diminished soon after the
first or second whorl and the adult or gerontic test is composed of interlammellar spaces, usually in-
flated into lateral chambers. Unlike Biplanispira these are often well outside the plane of coiling. As
Biostratigraphy of Indo-Pacific larger forams: page 104

with the transition from gerontic Pellatispira to Biplanispira, there are intermediate forms with sparse
interlamellar spaces, depending on the degree of development of the gerontic stage. Hottinger et al.
suggested three species based on size of proloculus, development of radial canals, which in V. glabra
can be especially wide, and degree of flattening of the test. These authors note that this last aspect may
not be a valid taxonomic character, and observations of examples from Java (Gamping Barat) suggest
that there are unclear graditions between all three species.
 Axial section of Vacuolispira glabra from the northeast arm of Halmahera, in the same thin section as several specimens of
Planocamerinoides and Alveolina. So far this is the only record known of Vacuolispira with Ta markers. Note the A form Nummulites
at the top of the image. Field of view is 3 mm.

Specimens of Vacuolispira from the Gamping Barat

location just outside of Yogjakarta. In a Tb, Late Eocene
assemblage. Like V. glabra these have coarse radial
canals and highly developed secondary skeleton, with
little left of the originla spiral whorls

Biostratigraphy of Indo-Pacific larger forams: page 105

Genus Wilfordia Adams, 1965

Type species: Wilfordia sarawakensis Adams, 1965
Only known from the Melinau limestone in Sarawak (Adams 1965), where it occurred in 2 samples at the
top of Tb. Alanlordia banyakensis Benner & Samuel from the Tf2 of Sumatra may be a homeomorph.
REFERENCES
Adams, C.G., 1965, The foraminifera and stratigraphy of the Melinau Limestone, Sarawak, and its importance
in Tertiary correlation: Quarterly Journal of the Geological Society of London, v. 121, p. 283-338.
—, 1970, A reconsideration of the East Indian letter classification of the Tertiary: Bulletin of the British
Museum of Natural History (Geology), v. 19, p. 1-137.
—, 1984, Neogene larger foraminifera, evolutionary and geological events in the context of datum planes,
in Ikebe, N., and Tsuchi, R., eds., Pacific Neogene Datum Planes: Tokyo, University of Tokyi Press,
p. 47-68.
Adams, C.G., and Belford, D.J., 1974, Foraminiferal biostratigraphy of the Oligo-Miocene limestones of
Christmas Island (Indian Ocean): Palaeontology, v. 17, p. 475-506.
Bain, J.H.C., and Binnekamp, J.G., 1973, The foraminifera and Stratigraphy of the Chimbu Limestone, New
Guinea: Bureau of Mineral Resources and Mines Geological Papers Bulletin, v. 139, p. 1-12.
Bakx, L.A.J., 1932, De genera Fasciolites en Neoalveolina in het Indo-Pacifische gebied: Verhandelingen
van het Koninklijk Nederlands Geologisch
Mijnbouwkundig Genootschap, v. 9, p. 205-266.
Banner, F.T., and Hodgkinson, R.L., 1991, A revision of the foraminiferal subfamily Heterostegininae.:
Revista Española de Micropaleontologia, v. XXIII, p. 101-140.
Banner, F.T., and Samuel, M.A., 1995, Alanlordia, a new genus of Acervuline foraminifera from the Neogene
of Indonesia: Journal of Micropalaeontology, v. 14, p. 107-117.
Berggren, W.A., Kent, D.V., Swisher, C.C., III, and Aubry, M.-P., 1995, A revised Cenozoic geochronol-
ogy and chronostratigraphy, in Berggren, W.A., Kent, D.V., Aubry, M.-P., and Hardenbol, J., eds.,
Geochronology Time Scales and Global Correlation, Volume 54, SEPM Special Publication, p.
129-212.
Berggren, W.A., and Prothero, D.R., 1992, Eocene-Oligocene climatic and biotic evolution: an overview, in
Prothero, D.R., and Berggren, W.A., eds., Eocene-Oligocene climatic and biotic evolution: Princeton,
Princeton University Press, p. 1-28.
Berry, E.W., 1929, Larger foraminifera of the Verdun Formation of northwestern Peru: Johns Hopkin Uni-
versity Studies in Geology, v. 9, p. 9-166.
Blow, W.H., 1969, Late Middle Eocene to Recent planktonic foraminiferal biostratigraphy, Proceeding First
International Conference on Planktonic Microfossils, Volume 1: Geneva, 1967, p. 199-422.
Bolli, H.M., and Saunders, J.B., 1985, Oligocene to Holocene low latitude planktic foraminifera, in Bolli,
H.M., Saunders, J.B., and Perch-Nielsen, K., eds., Plankton Stratigraphy: Cambridge, Cambridge
University Press, p. 155-262.
Bothé, A.C.D., 1929, Dwijo Hills and Southern Range, Excursion Guide Book for the 4th Pacific Science
Congress: Bandung, p. 1-15.
BouDagher-Fadel, M.K., and Banner, F.T., 1997, The revision of some genus-group names in Tethyan Lepi-
docyclininae: Palaeopelagos, v. 7, p. 3-16.
BouDagher-Fadel, M.K., Lord, A.R., and Banner, F.T., 2000a, Some Miogypsinidae (foraminifera) in the
Miocene of Borneo and nearby countries: Revue de Paléobiologie. Genève., v. 19, p. 137-156.
BouDagher-Fadel, M.K., Noad, J.J., and Lord, A.R., 2000b, Larger foraminifera from the Late Oligocene-
Biostratigraphy of Indo-Pacific larger forams: page 106

Earliest Miocene reefal limestones of North East Borneo: Revista Española de Micropaleontologia,
v. 32, p. 341-361.
Brouwer, J., 1966, Stratigraphy of the younger Tertiary in north-east Java and Madura (report 1957, issued
1966): The Hague, Bataafse Internationale Petroleum Maatschappij N.V.
Cande, S.C., and Kent, D.V., 1995, Revised calibration of the geomagnetic polarity time scale for the Late
Cretaceous and Cenozoic: Journal of Geophysical Research, v. 100, p. 6093-6095.
Caudri, C.M.B., 1934, Tertiary deposits of Sumba: Amsterdam, H.J. Paris, 223 p.
Cole, W.S., 1950, Larger foraminifera from the Palau Islands: U.S. Geological Survey Professional Paper,
v. 221-B, p. 21-31.
—, 1954, Larger foraminifera and smaller diagnostic foraminifera from Bikini drill holes: U.S. Geological
Survey Professional Paper, v. 260-O, p. 569-608.
—, 1956, Larger foraminifera [of Saipan, Mariana Islands]: U.S. Geological Survey Professional Paper, v.
 280-I, p. 321-357.
—, 1957, Larger foraminifera from Eniwetok drill holes: U.S. Geological Survey Professional Paper, v.
260-V, p. 743-784.
—, 1960, Upper Eocene and Oligocene larger foraminifera from Viti Levu, Fiji: U.S. Geological Survey
Professional Paper, v. 374-A, p. 1-6.
—, 1963, Tertiary larger foraminifera from Guam: U.S. Geological Survey Professional Paper, v. 403-E, p.
1-28.
—, 1969, Larger foraminifera from deep holes on Midway Atoll: U.S. Geological Survey Professional Paper,
v. 680-C, p. 1-13.
—, 1970, Larger foraminifera of Late Eocene age from Eua, Tonga: U.S. Geological Survey Professional
Paper, v. 640-B, p. 1-15.
Cole, W.S., and Bridge, J., 1953, Geology and larger foraminifera from Saipan Island: U.S. Geological
Survey Professional Paper, v. 253, p. 1-45.
Dollfus, G.F., 1917, Paléontologie du voyage à l’île de Celebes de M.E. Abendanon, in Abendanon, M.E.,
ed., Geologische en geografische doorkruisingen van Midden Celebes III. Koninklijk Nederlands
Aardrijkskundig Genootschap: Den Haag, p. 955-958.
Doornink, H.W., 1932, Tertiary Nummulitidae from Java.: Verhandelingen van het Geologisch en Mijnbou-
wkundig Genootschap voor Nederland en Koloniën, v. 9, p. 267-315.
Drooger, C.W., 1963, Evolutionary trends in Miogypsinidae., Evolutionary trends in Foraminifera: Amster-
dam, Elsevier, p. 315-349.
—, 1984, Evolutionary patterns in lineages of orbitoidal foraminifera: Proceedings of the Koninklijke Ned-
erlandse Academie van Wetenschappen, v. Ser B., 87, p. 103-130.
—, 1993, Radial foraminifera: morphometrics and evolution: Amsterdam, Afd. Natuurkunde, Eerste Reeks,
deel 41.
Drooger, C.W., and Rohling, E.J., 1988, Lepidocyclina migration across the Atlantic: Proceedings of the
Koninklijke Nederlandse Academie van Wetenschappen, v. Ser B, 91, p. 39-52.
Eames, F.E., Banner, F.T., Blow, W.H., and Clarke, W.J., 1962a, Fundamentals of mid-Tertiary correlation:
Cambridge, Cambridge University Press, 163 p.
Eames, F.E., Banner, F.T., Blow, W.H., Clarke, W.J., and Smout, A.W.H., 1962b, Morphology, taxonomy
and stratigraphy of the Lepidocyclininae: Micropaleontology, v. 8, p. 289-322.
Eames, F.E., Clarke, W.J., Banner, F.T., Smout, A.W.H., and Blow, W.H., 1968, Some larger foraminifera
from the Tertiary of Central America: Palaeontology, v. 11, p. 283-305.
Fortuin, A.R., 1970, The early ontogenetic stages in Eulepidina dilatata: Proceedings of the Koninklijke
Nederlandse Academie van Wetenschappen, v. Ser. B, 73, p. 196-208.
Glaessner, M.F., 1960, Upper Cretaceous larger foraminifera from New Guinea.: Science Reports, Tohoku
University, 2nd Series (Geology), v. Special Volume 4, p. 37-44.
Gould, S.J., and Eldridge, N., 1977, Punctuated equilibria: the tempo and mode of evolution reconsidered:
Paleobiology, v. 3, p. 115-151.
Hanzawa, S., 1930, Notes on Foraminifera found in the Lepidocyclina-limestone from Pabeasan, Java: To-
hoku Imperial University Science Reports, Series 2 (Geology), v. 14, p. 85-96.
—, 1957, Cenozoic foramnifera from Micronesia: Geological Society of America Memoir, v. 66, p. 1-163.
Biostratigraphy of Indo-Pacific larger forams: page 107

Haq, B.U., Hardenbol, P.R., and Vail, P.R., 1987, Chronology of fluctuating sea levels since the Triassic:
Science, v. 235, p. 1156-1167.
Hashimoto, W., and Matsumaru, K., 1978, Larger Foraminifera from the Philippines. 6. Larger Foraminifera
found from the Pinugay Hill Limestone, Tanay, Rizal, central Luzon: Geology and Palaeontology of
Southeast Asia, v. 19, p. 65-72.
Haynes, J.R., 1988, Massively chordate nummulitids (foraminifera) at the Paleocene/Eocene boundary in
Tethys.: Indian Journal of Geology, v. 60, p. 215-247.
Hottinger, L., 1960, Recherches sur les alvéolines paleocènes et eocènes: Mémoire de la Société Paléon-
tologique de la Suisse, v. 75/76, p. 1-242.
Hottinger, L., Romero, J., and Caus, E., 2001, Architecture and revision of the Pellatispirines, planispiral
canaliferous foraminifera from the Late Eocene Tethys: Micropaleontology, v. 47, p. 35-77.
Kominz, M.A., and Pekar, S.F., 2001, Oligocene eustasy from two-dimensional sequence stratigraphic
backstripping: Geological Society of America Bulletin, v. 113, p. 291-304.
Koolhoven, W.C.B., 1933a, Toelichting bij Blad 10 (Malingping), Geologische kaart van Java 1:100,000
(unpublished, E33-73 in files of Geological Survey): Bandung, Dienst van den Mijnbouw in Ned-
erlandsch-Indië.
—, 1933b, Toelichting bij Blad 14 (Bajah), Geologische kaart van Java 1:100,000: Bandung, Dienst van den
Mijnbouw in Nederlandsch-Indië.
Krijnen, W.F., 1931, Het Genus Spiroclypeus in her Indo-Pacifische gebied: Verhandelingen van het Geolo-
gisch en Mijnbouwkundig Genootschap voor Nederland en Koloniën, v. XI, p. 77-112.
Leupold, W., and van der Vlerk, I.M., 1931, The Tertiary: Leidsche Geologische Mededelingen, v. 5, p.
611-648.
Loeblich, A.R., Jr., and Tappan, H., 1987, Foraminiferal Genera and their Classification: New York, Van
Nostrabd Reinhold Company, 1182 p.
Lunt, P., 2003, Biogeography of some Eocene larger foraminifera, and their application in in distinguishing
geological plates: Palaeontologica Electronica, v. 6, p. 1-22.
Lunt, P., and Sugiatno, H., in press-a, The Bagelen Beds, Central Java: Bulletin of the GRDC, Bandung.
—, in press-b, A report on field work in the Rajamandala - Citarum area, West Java: Bulletin of the GRDC,
Bandung.
—, in press-c, A review of the Eocene and Oligocene in the Nanggulan area, south Central Java: Bulletin
of the GRDC, Bandung.
Lunt, P., Sugiatno, H., and Allan, T., in prep., A review of the Lutut Member in the type area, north Central
Java.: Bulletin of the GRDC, Bandung.
MacGillavry, H.J., 1962, Lineages in the genus Cycloclypeus Carpenter (Foraminifera): Proceedings of the
Koninklijke Nederlandse Academie van Wetenschappen, v. Ser. B, 65, p. 429-458.
Miller, K.G., Mountain, G.S., and Tucholke, B.E., 1985, Oligocene glacioeustasy and erosion on the margins
of the North Atlantic: Geology, v. 13, p. 10-13.
Morley Davies, A., Eames, F.E., and Savage, R.J.G., 1971, Tertiary Faunas: London, George Allen & Unwin
Ltd., 1018 p.
Muhar, A., 1956, Micropalaeontologishe onderzoek van monsters afkomstig van het geologishe onderzoek
Tuban., B.P.M. unpublished report, p. 22.
Ozawa, T., 1975, Evolution of Lepidolina multiseptata (Permian foraminifer) in East Asia: Memoirs of the
Faculty of Science. Kyushu University, Ser D. Geology, v. 23, p. 117-164.
Raju, D.S.N., 1974, Study of Indian Miogypsinidae: Utrecht Micropaleontology Bullentins, v. 9, p. 148.
Renema, W., Racey, A., and Lunt, P., 2003, Palaeogene nummulitids (Foraminiferida) from the Indonesian
Archipelago: a review: Cenozoic Research, v. 2.
Rutten, L.M.R., 1927, Voordrachten over de Geologie van Nederlandsch Oost Indië: Groningen, Den Haag,
J.B. Wolters, 839 p.
Schindewolf, O.H., 1936, Paläontologie, Entwicklungslehre und Genetik: Berlin, Bornträger.
Serra-Kiel, J., Hottinger, L., Caus, E., Drobne, K., Fernandez, C., Jauhra, A.K., Less, G., Pavlovec, R., Pign-
atti, J., Samso, J.M., Schaub, H., Sirel, E., Strougi, A., Tambareau, Y., Tosquella, Y., and Zakrevskaya,
E., 1998, Larger foraminiferal biostratigraphy of the Tethyan Palaeocene and Eocene: Bulletin de la
Biostratigraphy of Indo-Pacific larger forams: page 108

Société Géologique de France, v. 169, p. 281-299.

Stainforth, R.M., Lamb, J.L., Luterbacher, H., Beard, J.H., and Jeffords, R.M., 1975, Cenozoic planktonic
foraminifera zonation and characteristics of index forms: Lawrence, University of Kansas Paleon-
tological Contributions, 425 p.
Tan Sin Hok, 1932, On the genus Cycloclypeus Carpenter Part 1. And an appendix on the Heterostegines
of Tjimanggoe, S. Bantam, Java.: Wetenschappelijke Mededelingen Dienst van den Mijnbouw in
Nederlandsch-Indië, v. 19, p. 1-194.
—, 1937, On the genus Spiroclypeus H. Douvillé with a description of the Eocene Spiroclypeus vermicularis
nov.sp. from Koetai in East Borneo.: De Ingenieur in Nederlandsh-Indië (Mijnbouw en Geologie,
Bandung), v. No.4 pt.4, p. 177-193.
Umbgrove, J.H.F., 1928, Het genus Pellatispira in het indo-pacifische gebied.: Weteenschappelijke Med-
edeelingen, v. 10, p. 103 pp.
—, 1936, Heterospira, a new foraminiferal genus from the Tertiary of Borneo.: Leidsche Geologische Med-
edelingen, v. 8, p. 155-157.
—, 1937, A new name for the foraminiferal genus Heterospira: Leidsche Geologische Mededelingen, v. 8,
p. 309.
Vail, P.R., Mitchum, R.M., Todd, R.G., Widmier, J.M., Thompson, S., Songree, J.B., Bubb, J.N., and Hatlelid,
W.G., 1977, Seismic stratigraphy and global changes of sea-level: American Association of Petroleum
Geologists Memoir, v. 26, p. 49-212.
van Bemmelen, R.W., 1941, Toelichting bij Blad 73/74 (Semarang / Oengaran), Geologische kaart van Java
1:100,000: Baundung, Un-published, in proof at outbreak of war.
—, 1949, The Geology of Indonesia: The Hague, Netherlands, Government Printing Office, 997 p.
van der Vlerk, I.M., 1928, The genus Lepidocyclina in the Far East: Eclogae Geoloicae Helvetiae, v. 21, p.
182-211.
—, 1929, Groote foraminiferen van N.O. Borneo. Indië: Wetenschappelijke Mededelingen Dienst van den
Mijnbouw in Nederlandsch-Indië, v. 9, p. 1-46.
van der Vlerk, I.M., and Postuma, J.A., 1967, Oligo-Miocene Lepidocyclinas and planktonic foraminifera
from East Java and Madura: Proceedings of the Koninklijke Nederlandse Academie van Wetenschap-
pen, v. 70, p. 392-399.
van der Vlerk, I.M., and Umbgrove, J.H.F., 1927, Tertiaire gidsforaminiferen uit Nederlandsch Oost-Indië:
Wetenschappelijke Mededelingen Dienst van den Mijnbouw in Nederlandsch-Indië, v. 6, p. 1-31.
van Vessem, E.J., 1978, Study of Lepidocylinidae from South East Asia, particularly from Java and Borneo:
Utrecht Micropaleontology Bullentins, v. 19, p. 1-163.
Verbeek, R.D.M., 1891, Voorlopig bericht over den Nummulieten, Orbitoiden en Alveolinen van Java, en
over den ouderdom der gesteenten waarin zij optreden: Natuurkundig Tijdschrift voor Nederlansch-
Indië, v. 51, p. 101-139.
Verbeek, R.D.M., and Fennema, R., 1896, Geologie van Java en Madoera: Amsterdam, Uitgegeven op last
van zijne Excellentie den Gouveneur-Generaal van Nederlansch-Indië, 2, & Atlas p.
Yabe, H., 1921, Notes on some Eocene foraminifera: Science Reports, Tohoku University, 2nd Series (Geol-
ogy), v. V, p. 97-108.
Zuffardi-Comerci, R., 1928, Di alcuni foraminiferi terziari dell’ isola di Borneo: Bolletino della Societa
Geologica Italiana, v. 47, p. 127-148.
.

Biostratigraphy of Indo-Pacific larger forams: page 109