INTERNAL ARCHITECTURE OF MIOCENE

PSEUDOTABERINA AND ITS RELATION TO

CARIBBEAN ARCHAIASINS

by WILLEM RENEMANationaal Natuurhistorisch Museum Naturalis, PO Box 9517, 2300 RA Leiden, the Netherlands; e-mail: Renema@naturalis.nl, and Centre for Marine Studies,

University of Queensland, Brisbane, Qld 4072, Australia

Typescript received 31 January 2006: accepted in revised form 8 January 2007

Abstract: Well-preserved, isolated specimens of Pseudotabe-

rina were found at three localities in Indonesia, thus enabling

the study of the relationships between this genus and Carib-

bean Archaiasinae. The internal architecture of the chambers

of Pseudotaberina consists of four layers of stolon planes,

each of which is divided by superposed septula into chamb-

erlets. The chamberlets are connected to the next chambers

by foramina opposing the septula. The stolons are Y-shaped.

All other Archaiasinae have radial stolon axes combined with

an initial involute growth stage, whereas the Soritinae have

cross-wise oblique stolon axes, but with evolute growth in

the early stages. Pseudotaberina shows a combination of the

characters of these two subfamilies, confirming the need to

study more Tethyan material to elucidate phylogenetic rela-

tionships within this group of large benthic foraminifera.

Key words: Archaiasinae, Soritinae, large benthic foramini-

fera, Indonesia, Burdigalian.

Phylogenetic analyses of the Soritidae have revealed

ambiguous results with regard to the monophyly of the

subfamily Archaiasinae. Analyses based on morphological

characters show a monophyletic grouping of all Archaiasi-

nae as a sister group to the Soritinae (Gudmundsson

1994; Richarson 2001). The analysis carried out by

Gudmundsson (1994) only involved extant taxa, but

Richarson (2001) included fossil taxa as well. In the latter

analysis both Nemophora floridana (=Miarchaias florid-

anus) and N. malabarica (= Pseudotaberina malabarica)

were included and they grouped as sister taxa in a clade

ancestral to the Recent species of Archaias. This would

imply a palaeobiogeographical connection between the

Middle and Far East and the Caribbean for which hardly

any other evidence is available. However, Holzman et al.

(2001) found in a molecular phylogeny of the Soritacea,

obviously restricted to living taxa only, that taxa without

an endoskeleton form a paraphyletic group within the sub-

clade Archaiasinae. Taxa with an endoskeleton are derived

independently from basal taxa without endoskeletons.

To understand the phylogenetic relationships in fossil

and Recent large benthic foraminifera (LBF) an analysis

of the complex internal structure is necessary. This com-

plicates the study of fossil taxa, since only exceptionally

well preserved, isolated specimens allow for this kind of

detailed investigation. Well-preserved, isolated specimens

from shallow, in particular reef-associated, habitats are

even rarer. One of the groups in which phylogenetic rela-

tionships are unclear is the soritids, and especially the

archaiasins. At least six species of the Archaiasinae occur

in the Caribbean province, whereas only one genus occurs

in the Indo-West Pacific (Hallock 1988; Hallock and

Peebles 1993; Holzman et al. 2001). All species harbour

chlorophytes as symbionts and occur in shallow

environments.

Three genera of Archaiasins have been described from

the Upper Miocene of the Dominican Republic (Hottin-

ger 2001). The preservation of this material was sufficient

to allow a full description of the microstructure of these

species, as well as of some co-occuring soritins. Hottinger

concluded that these species are unlikely to be candidates

for a direct ancestry of the species living today in the

Caribbean province, and he described several new species

and genera from this material. The taxa reflect differences

in the apertural face and the respective arrangement of

radial partitions (Hottinger 2001). One of them, Miarcha-

ias meander Hottinger, 2001, develops cyclical agamonts

with a diameter up to 1.5 cm and with meandering struc-

tures covering the lateral surface of the disc, while smaller

species do not develop cyclical growth. The Tethyan

genus Pseudotaberina Eames, in Davies 1971 (emended by

Banner and Highton 1989) has the same basic architec-

ture. These descriptions are based on material in thin sec-

tions. In the present study, isolated specimens of

Pseudotaberina, allowing study of the internal architec-

ture, are described from the Miocene of Java. These turn

out to have a different architecture, and thus the genera

are not synonymous.

[Palaeontology, Vol. 51, Part 1, 2008, pp. 71–79]

ª The Palaeontological Association doi: 10.1111/j.1475-4983.2007.00731.x 71

AGE AND ORIGIN OF THE MATERIAL

In the collection of the Nationaal Natuurhistorisch

Museum, Leiden, samples collected by Martin in

1910 ⁄ 1911 contain isolated, sediment-free specimens of

Pseudotaberina malabarica. They were labelled and stored

under the name Orbiculina cf. adunca Fichtel and Moll,

1798, which is also the name under which they were pub-

lished by Rutten (1917).

The samples were collected at Kembang Sokkoh and

Gunung Spolong (Yogyakarta, Indonesia; Text-fig. 1), two

localities separated by less than 4 km. The lower part of

the Jonggrangang Formation comprises a thick succession

of Oligocene–earliest Miocene andesitic volcaniclastics,

overlying the Middle–Late Eocene Nanggulan marls. The

upper part of this formation consists of an alternation of

marls and limestones, and grades into a bedded lime-

stone. A detailed description of the sample sites and their

relative position was published by Martin (1911). In total,

at least 103 species of molluscs (Martin 1917), 15–17 spe-

cies of corals (Gerth 1921), and nine species of benthic

foraminifera (Rutten in Martin 1917; Tan Sin Hok in van

Bemmelen 1949) have been described from the Jonggran-

gang Formation. The setting was interpreted as reefal,

with Kembang Sokkoh situated near a (small) river-

mouth (van Bemmelen 1949). Accompanying LBF were,

among others, Lepidosemicyclina thecidaeformis, Austrotril-

lina howchini, Flosculinella globulosa and Planogypsina.

This is a typical LBF zone Tf1 fauna, approximately

equivalent to early Burdigalian.

Though rare, Pseudotaberina malabarica has also been

found in some other localities. Van der Vlerk (1924) fig-

ured a specimen from the Njalindung beds at Ci Talahab

(West Java, Indonesia; Text-fig. 1). The accompanying

fauna at this locality indicates a slightly younger age: Tf2

or late Burdigalian. An assemblage very similar to that on

Java was described by de Neve (1947) from the Sangkuli-

ran area in East Kalimantan (Text-fig. 1). The age of this

locality is Tf1. Just as in the Jonggrangan Formation, the

specimens of Flosculinella globulosa were very small and

spherical. De Neve (1947) described his specimens as a

new species, Archaias vandervlerki.

In East Kalimantan, P. malabarica was also found at a

locality near Bontang (Renema, unpublished data) where

it is present in dark marls containing numerous Flosculi-

nella bontangensis (Rutten) and Lepidosemicyclina bifida

(Rutten), as well as a limited number of Nephrolepidina

angulosa (Rutten). Flosculinella bontangensis is a fusiform

alveolinid species (length approximately twice height).

The specimens found at Bontang (the type locality) show

very limited variation in elongation. In this friable marl,

very rare specimens of Pseudotaberina malabarica were

found in one sample. The age of these samples is Tf2, lat-

est Burdigalian.

Wonders and Adams (1991) reported another occur-

rence of Pseudotaberina from the Darai Limestone (Papua

New Guinea: PNG). They determined the stratigraphical

position of their samples as basal Tf1, earliest Burdigalian.

Other localities outside Indonesia include Saipan (Mari-

ana Islands) in the West Pacific (Cole, 1957) where

Pseudotaberina was found in cores in association with

Austrotrillina howchini, Flosculinella globulosa and

Nephrolepidina spp., suggesting an age very similar to that

of the Javanese specimens. The illustrations are not of

sufficient quality to allow identification to species level

but they suggest Pseudotaberina vandervlerki.

In Flosculinella, a general trend from spherical to fusi-

form is observed, with intermediate variable populations

120° E

0 200 400

km

10° S

10° N

0°

140° E

1

2

34

N

TEXT-F IG . 1 . Map showing the

localities mentioned in the text. 1,

Menoreh Mountains, Yogyakarta; 2,

Bontang, East Kalimantan; 3, Ci Talahab,

West Java; 4, Darai Limestone, Papua

New Guinea.

72 P A L A E O N T O L O G Y , V O L U M E 5 1

(Reichel 1936–37; Wonders and Adams 1991). The associ-

ation with both basal spherical and derived fusiform pop-

ulations shows that Pseudotaberina occurred over a longer

period. The faunal associations fit well within Tf1–Tf2 of

Burdigalian age. Thus Pseudotaberina has a very wide geo-

graphical distribution but is restricted to a short strati-

graphic range through most of the Burdigalian.

TERMINOLOGY

A complete description of the architectural elements in

archaiasin tests has been provided by Hottinger (2001,

2005). Here the elements used in the description of Pseu-

dotaberina are repeated. A schematic overview of the

structural elements is given in Text-figure 2. Archaiasins

are porcellaneous, imperforate foraminifera. In this group,

subdivisions of the chambers by structural elements are

produced by localized thickening of the chamber wall.

Apertures, openings in the chamber wall through which

the foraminiferan communicates with the outside envi-

ronment, develop on the apertural face, the area of the

chamber wall where single or multiple apertures are posi-

tioned. During the formation of the next chamber, the

apertural face is covered and the apertures transform into

intercameral foramina through which there is communi-

cation between successive chambers. The axes of aligned

apertures in successive chambers are called apertural axes.

These can be parallel to the radius of the test: in this case

the apertural axis does not cross the previous or the next

apertural axes. This pattern is called radial. In a cross-

wise-oblique pattern the apertural axes are oblique to the

a

a

pi

su

ap

f

f

ap su

s

ma

lw

A

B

af

so

TEXT -F IG . 2 . The structure of archaiasinid frontal and lateral

chamber margins. Over part of the chamber, the frontal and

lateral walls are cut off to show the septal face and partitions

below the lateral chamber wall. Stereographs, schematic, not to

scale. A, Pseudotaberina sp.; B, Miarchaias sp. (after Hottinger,

2001). Abbreviations: a, aperture; af, apertural face; ap, annular

passage; f, foramen; lw, lateral chamberwall; ma, marginal

aperture; pi, pillar; s, septum; so, socculus; su, septula.

so

bts

su

su

A

B

f

f

s

s

s s

lw

TEXT -F IG . 3 . A, scanning electron microscope (SEM)

micrograph of the ultimate chambers of Pseudotaberna

vandervlerki, Kembang Sokkoh, Menoreh Mountains,

Yogyakarta, RGM 508454; white hatched lines represent the

direction of the passages through the foramina; solid white lines

represent the test radius. B, schematic interpretation of the

structural elements in Text-figure 2. Abbreviations: as for Text-

figure 2 and bts, basal thickening of the septula. Scale bar

represents 0.1 mm.

R E N E M A : I N T E R N A L A R C H I T E C T U R E O F M I O C E N E P S E U D O T A B E R I N A 73

radius of the test, and cross each other regularly (Text-

fig. 3). In archaiasins the chambers can be subdivided by

septula (continuous walls) or free-standing pillars. Septula

and pillars tend to thicken at their proximal contact with

the basal layer. These structures have been called socculi

(Seiglie et al. 1976; Hottinger 2001). The annular passage

through the septula connects chamberlets of the same

chamber.

During growth, planispiral, involute archaiasinids tend

to extend their apertural face by peripheral extension of

their spiral chambers during ontogeny: a first spiral,

peneropline growth stage ends with the first chamber over-

growing the spiral periphery of the previous chamber. This

growth stage ends where a chamber produces a complete

annulus for the first time (Hottinger 2005). To increase

further the extension of the apertural face, two structures

can be formed: (1) forward growing: meandrine chambers

growing from the apertures of the last involute chambers

form lateral supplementary chambers overgrowing the

annular growth stage (Text-fig 4A); or (2) backward

growing: the marginally placed apertures produce lateral

supplementary chambers overgrowing the central part of

the test at the point where the distal part of the apertural

face touches the more proximal part (Text-figs 4B, 8).

SYSTEMATIC PALAEONTOLOGY

Superfamily SORITACEA Ehrenberg, 1839

Family SORITIDAE Ehrenberg, 1839

Genus PSEUDOTABERINA Eames, in Davies 1971, emend.

Banner and Highton, 1989

Type species. Orbitolites malabarica Carter, 1853a.

Pseudotaberina malabarica (Carter, 1853a)

* 1853a Orbitolites malabarica Carter, pp. 142–144, pl. 2A.

1853b Orbitolites malabarica Carter; Carter, pp. 425–427,

pl. 16B, figs 1–4.

1989 Pseudotaberina malabarica (Carter); Banner and

Highton, pp. 113–129, pl. 1, figs 6–8.

? 1923 Orbiculina malabarica (Carter); Wayland and

Davies, pp. 591–592, pl. 28, figs 1–4.

Description. Banner and Highton (1989) formally erected the

genus Pseudotaberina, which up to then had been a nomen nu-

dum, and emended the type species P. malabarica based on the

specimens used by Carter (1853a). The size of the topotype

specimens (Banner and Highton 1989, pl. 1, figs 6–8) varies

from 2.9 to 6 mm. The figured apertural face of the 6 mm speci-

men shows multiple, 5–7, rows of apertures. The alternating,

rimmed apertures are positioned in depressions. At a diameter

of 2.9 mm the alar prolongations are 0.6 mm. The chamber

height averages 0.04 mm.

In the macrospheric generation, specimens reach a diameter

of up to 6.5 mm and a thickness of 0.7 mm. Microspheric speci-

mens can reach a diameter of up to 15 mm, and attain annular

chambers at a diameter of c. 8 mm. Larger specimens develop

forward-growing meandrine chambers from the apertures of the

ultimate involute chamber.

The internal architecture, as shown in the oblique sections, is

very similar to that of the Indonesian specimens. Both have alter-

nating septula that reach approximately half-way to the chamber

just below the lateral wall, and increase in length and width, cut-

ting deeper into the test. Laterally flattened socculi subdividing

the chamberlets are also present in the Indian specimens.

Remarks. Wayland and Davies (1923) recorded a larger

maximum diameter (up to 21 mm) than that for the

specimens described by Banner and Highton (1989). The

specimens attain cyclical growth when they reach a diam-

eter of 3–5 mm, which is slightly smaller than in the

specimens figured in Banner and Highton (1989). Way-

land and Davies (1923) were the first to discuss the differ-

ence between the Indian and Indonesian specimens. They

also observed that at the same size, the Indonesian speci-

mens do not attain cyclical growth. Interestingly, they

mentioned (but did not figure) a single specimen of

P. vandervlerki from India. By contrast, Tobler (1927)

mentioned a Meandropsina from Kalimantan, a genus that

externally resembles Pseudotaberina in attaining cyclical

growth, but the internal structure is different. So far, it is

known only from the Cretaceous (L. Hottinger, pers.

comm. 2005). The dimensions of the Kalimantan speci-

mens equal those of P. malabarica from India, although

the largest specimens are larger than those mentioned by

Wayland and Davies (1923). In the samples from Kali-

mantan, the small, probable macrospheric, specimens

measure 5–7 mm, the largest, probable microspheric,

specimens 25 mm (Tobler 1927).

A B

TEXT -F IG . 4 . Schematic drawing of the outside of the test. A,

forward-growing Miarchaias, after reaching annular growth;

lateral extensions of the apertural face of the alar prolongation

of the ultimate chamber before reaching annular growth. B,

backward-growing Pseudotaberina vandervlerki, extensions of the

apertural face grow over the previous chamber starting at the

point where the distal part of the apertural face touches the

more distal part at the onset of evolute growth (see also Text-

fig. 8); thin lines indicate sutures, thick black line indicates

apertural face where growth occurs. Not to scale.

74 P A L A E O N T O L O G Y , V O L U M E 5 1

Pseudotaberina vandervlerki (de Neve, 1947)

Text-figures 2A, 3–9.

1917 Orbiculina cf. adunca Fichtel and Moll, 1798;

Rutten, pp. 276–285, pl. 5, fig. 142.

1924 Orbiculina? sp.; van der Vlerk 1924, pp. 27–28,

fig. 25.

* 1947 Archaias vandervlerki de Neve 1947, pp. 14–15,

text-figs 1–4.

? 1957 Archaias vandervlerki De Neve; Cole, p. 335,

pl. 103, figs 5–9.

Material. More than 100 isolated specimens, of which only two

are microspheric, from Kembang Sokkoh and Gunung Spolong,

Yogyakarta, Indonesia (RGM 3267, 3481, 47264, 47265, 508454–

508456, 508479, 508481, 508483–508484). Type specimens

(unnumbered) of Archaias vandervlerki de Neve, 1947, deposited

in the Geological Museum in Bandung, Indonesia. Unnumbered

single specimen labelled Orbiculina sp. from the Njalindung

beds, figured by van der Vlerk (1924), also deposited in the Geo-

logical Museum in Bandung.

Description. Lenticular test, planispiral involute throughout

ontogeny. In the macrospheric generation, specimens can reach

a diameter of 5 mm and a thickness of 0.7 mm. Microspheric

specimens can reach a diameter of 12 mm. Annular stages have

not been observed. The chambers proximally embrace the previ-

ous whorls but distally they are evolute. They are low but long,

with a strong posterior curvature so that they cover at least half,

but usually more, of the previously formed test, and cover more

as growth continues. Chamber height increases gradually from

58–71 lm in the first whorl to 65–86 lm in the last whorl. The

last chambers in the microspheric generations become 90–

120 lm in height. The alar prolongations extend 4 mm in a

specimen of 12 mm (microsphere), and 1.2–1.4 mm in macro-

spheric specimens of 3.1–3.3 mm diameter. The embracing com-

ponent (length of alar prolongation ⁄ radius) in specimens with a

diameter of 3–4 mm is 0.76–0.84. Apertures are round and

rimmed. They are placed in four alternating superposed rows;

the shape of those in the marginal rows is the same as those in

the equatorial layers (Text-fig. 5A). The lateral chamber wall is

thin and slightly pitted.

The length of the chambers increases with growth. In the later

chambers test growth occurs along the entire circumference, but

without becoming annular as in Miarchaias. At the place where

the distal part of the apertural face meets the more proximal

part, the marginal apertures of the proximal part form back-

ward-growing meandrine lateral chambers (Text-figs 4, 8, 9E).

The occurrence of this structure at exactly the transition from

involute to evolute growth, and never at other positions along

the apertural face, indicates that this backward-growing is not

the result of shell repair. No forward-growing lateral extensions

of the alar prolongations have been observed.

The megalospheric embryo consists of an approximately

spherical megalosphere 0.25–0.30 mm in diameter followed by a

wide flexostyle of c. 0.1 mm and about one-quarter volution in

length. The flexostyle has two apertures at the proximal part: the

first spiral chamber does not show a subdivision in all thin sec-

tions, but the second and later chambers always have subdivi-

sions. There are 3–3.5 volutions in macrospheric specimens. The

lateral test wall is very thin, and the septula are visible from the

outside.

The chambers consist of four stolon planes and are radially

divided into chamberlets by septula. An annular passage, of

which the diameter is about one-half to one-third of the cham-

ber height, is positioned distally in the septula directly below the

test wall (Text-fig. 2A). The septula are somewhat irregularly

placed and are thicker proximally than distally. The laterally flat-

tened bases, which can possibly be referred to as socculi, physi-

cally separate the stolon planes. The chamberlets are laterally

divided into four stolon planes by the flattened bases of the sep-

tula (soculi; Text-fig. 2A).

Between the septula are round, rimmed foramina in the sep-

tum, with a diameter of c. 0.025–0.031 mm. The foramina are

placed opposite the septula in the previous chamber, resulting in

Y-shaped stolon passages (Text-fig. 3). Owing to this pattern,

every chamberlet is connected to two chamberlets in the more

distal chamber. This results in the absence of stolon axes, pro-

viding a direct line of transport from the central part of the test

to the apertural face.

Remarks. The specimens described here differ from those

described by Banner and Highton (1989) in that the

embracing component of the alar prolongations is much

larger. Other differences relate to the apertural face, which

does not show the rimmed apertures positioned in depres-

sions, and the smaller diameter of full-grown tests. Also,

annular chambers and forward-growing lateral extensions

of the alar prolongation as described for P. malabarica

(Banner and Highton 1989) have not been observed in

Indonesian specimens.

Stratigraphic distribution. Burdigalian of Java, Borneo, PNG and

India.

DISCUSSION AND CONCLUSIONS

Henson (1950) found Taberina [Archaias, Orbiculina, Or-

bitolites] malabarica Carter, 1853a in Iraq. However, no

isolated specimens are available and his material could

not be assigned with certainty to either of the two species

described here.

The type material of Archaias vandervlerki de Neve, 1947

is deposited at the Geological Museum in Bandung and

was available for comparison. The specimens concerned

are identical to those from Java; thus the name P. vander-

vlerki (de Neve, 1947) is available for this species.

Based on external morphology, Pseudotaberina resem-

bles Miarchaias, described by Hottinger (2001) from the

Miocene of the Dominican Republic. However, the stolon

axes in Miarchaias are radial instead of Y-shaped as in

Pseudotaberina (Text-fig. 3). The stolon axes in each sto-

R E N E M A : I N T E R N A L A R C H I T E C T U R E O F M I O C E N E P S E U D O T A B E R I N A 75

lw

af

a

a f

f

s

su

su

bts

bts

so

C

B

1

2 3 4

A

D

TEXT-F IG . 5 . SEM micrographs of details of the test of

Pseudotaberina vandervlerki, RGM 508484. A, oblique view

of the apertural face showing four rows of irregularly

superposed apertures. B, oblique view of chamberlets in the

ultimate and penultimate whorl; white arrows indicate two

superposed foramina, the top one with a stolon passage to

the left, the lower one to the right; black arrow indicates the

annular passage. C, schematic interpretation of B;

abbreviations as for Text-figures 2 and 3. D, axial view of

broken specimens showing four stolon planes. Scale bars

represent 50 lm in A–B, 0.1 mm in D.

76 P A L A E O N T O L O G Y , V O L U M E 5 1

lon plane in all of the genera described by Hottinger

(2001) are parallel instead of Y-shaped. Another morpho-

logical difference is the laterally partitioned chambers.

The partitioning possibly originates from the fusion and

lateral flattening of the socculi (or swollen feet) of the

septula. Lateral partitioning of the chambers was not

observed in any of the American Archaiasinae.

Hottinger (2001) used the presence of cross-wise-obli-

que apertural alignment as a character to separate the

subfamilies Soritinae and Archaiasinae. Soritines tend to

reduce the spiral juvenile stage to form evolute, early con-

centric shells (Hottinger 2001). The annular chambers are

subdivided by radial partitions. Archaiasins, by contrast,

develop extensive endoskeletons consisting of pillars and

septula, remain involute, and enlarge the apertural face by

successively more extended apertural faces (Hottinger,

2001); only in a late stage of growth do some taxa attain

annular growth. Archaiasins can be separated into taxa

lacking an endoskeleton, e.g. the extant Laevipeneroplis

(Text-fig. 10), and those with an endoskeleton, e.g. the

extant Archaias and Miocene Miarchaias.

Pseudotaberina shows a unique combination of charac-

ters, because Y-shaped stolon configurations have not

been observed in any other soritid. The endoskeletal

structure of Pseudotaberina confirms the conclusions of

Hottinger (2001) that a revision of all Tethyan species

with an archaiasin growth form is necessary to develop a

suprageneric classification. Only comparison of more

material will elucidate whether the Soritidae can best be

separated on growth form or endoskeletal characters, but

the generic rank differentiation of Pseudotaberina is sub-

stantiated.

The undoubtedly high number of morphological differ-

ences between extant Archaias and both the Caribbean

and Indo-Pacific Miocene forms shows that during the

TEXT -F IG . 7 . Photograph of thin section of microspheric

specimen of Pseudotaberina vandervlerki, Gunung Spolong,

Menoreh Mountains, Yogyakarta, RGM 508533. Scale bar

represents 1 mm.

f

apB

A

TEXT -F IG . 8 . SEM micrographs of Pseudotaberina

vandervlerki, Kembang Sokkoh, Menoreh Mountains,

Yogyakarta, RGM 508483. A, habitus. B, detail of the extensions

of the apertural face overgrowing the previous chambers. Scale

bars represent 0.5 and 0.1 mm respectively. Abbreviations: f,

foramen; ap, annular passage. Arrows indicate the direction of

growth at the apertural face.

TEXT -F IG . 6 . Detail of Pseudotaberina vandervlerki showing

the very thin, transparent, lateral test wall, below which the

annular passages (indicated by arrows) and alternating septula

are clearly visible; Kembang Sokkoh, Menoreh Mountains,

Yogyakarta, RGM 508532. Scale bar represents 0.1 mm.

R E N E M A : I N T E R N A L A R C H I T E C T U R E O F M I O C E N E P S E U D O T A B E R I N A 77

Miocene this group was highly diverse. At present, no

extant Archaiasinae with an endoskeleton are known from

the Indo-West Pacific. Pseudotaberina and the Middle

Eastern taxa described by Henson (1950) became extinct

at the end of the Early Miocene. Two separate radiations

can be hypothesized, one during the Oligocene–Miocene

in the Tethyan realm and another in the Middle–Late

Miocene in the Caribbean. The stratigraphical use of the

occurrence of Pseudotaberina has long been recognised in

the Indo-West Pacific. The Indonesian specimens

reported here are restricted to the Burdigalian. The

assemblages in which Pseudotaberina was found are typi-

cal of very shallow, epiphytic depositional environments.

Acknowledgements. Frank Wesselingh and Steve Donovan read

earlier drafts of this manuscript and their constructive remarks

improved the paper. Comments by Lukas Hottinger and an

anonymous reviewer further improved its quality.

REFERENCES

B A N N E R , F. T. and H I G H T ON , J. 1989. On Pseudotaberina

malabarica (Carter) (Foraminiferida). Journal of Micropalaeon-

tology, 8, 113–129.

B E M M E L E N , R. W. van 1949. Geology of Indonesia. Govern-

ment Printing Office, The Hague, 732 pp.

C A R T E R , H. J. 1853a. A description of Orbitolites malabarica,

illustrative of the spiral and not concentric arrangement of

A B

C D E

TEXT -F IG . 9 . Habitus of Pseudotaberina vandervlerki. A, small specimen. B–E, large specimens; arrows in C indicate length of alar

prolongation. A–C, Kembang Sokkoh, West Progo Mountains, Yogyakarta. A, RGM 508487. B, RGM 508484. C, RGM 508530. D–E,

Gunung Spolong, Menoreh Mountains, Yogyakarta. D, RGM 508479. E, RGM 508531. Scale bar represents 0.5 mm in A–C, 1 mm in

D–E.

B A

TEXT -F IG . 10 . SEM micrographs of Laevipeneroplis

malayensis Hofker. Both specimens are from Maratua, off

Tanjung Redeb, East Kalimantan, Indonesia. A, small specimen,

RGM 508462. B, large specimen, RGM 508463. Scale bars

represent 0.1 and 0.2 mm respectively.

78 P A L A E O N T O L O G Y , V O L U M E 5 1

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