Geodinamica Acta

2018, VOL. 30, NO. 1, 137–162

https://doi.org/10.1080/09853111.2018.1443654

OPEN ACCESS

Bartonian orthophragminids from the Fulra Limestone (Kutch, W India) and

coeval units in Sulaiman Range, Pakistan: a synthesis of shallow benthic zone
(SBZ) 17 for the Indian Subcontinent
Ercan Özcana, Pratul Kumar Saraswatib , Ali Osman Yücela, Nowrad Alic,d and Muhammad Hanifd
a
Faculty of Mines, Department of Geological Engineering, İstanbul Technical University (İTU), İstanbul, Turkey; bDepartment of Earth Sciences,
Indian Institute of Technology–Bombay, Mumbai, India; cDepartment of Geology, University of Peshawar, Peshawar, Pakistan; dNational
Centre of Excellence in Geology, University of Peshawar, Peshawar, Pakistan

ABSTRACT ARTICLE HISTORY

Orthophragminids from the Bartonian Fulra Limestone in Kutch, India and the coeval units in Received 8 December 2017
Sulaiman Range in Pakistan suggest the establishment of a significant number of endemic species Accepted 19 February 2018
in the Indian subcontinent (Eastern Tethys). Among a total of fifteen species of Discocyclina, KEYWORDS
Orbitoclypeus and Asterocyclina, six of them appear to be confined to Indian subcontinent while Orthophragminids;
seven species are common both to the peri-Mediterranean/Europe region (Western Tethys) and Indian subcontinent;
Indian subcontinent. Two species, Asterocyclina sireli, a four-ribbed species of possibly Indo- Fulra Limestone;
Pacific origin, and Orbitoclypeus haynesi that form large populations in Fulra Limestone, appear paleobiogeography; Eocene
to have spread into North Africa and Turkey but not into European platforms as a response to
Middle Eocene Climatic Optimum (MECO). The lack of Lutetian and Priabonian fauna in the
studied sections, either due to a hiatus or unsuitable depositional environments, hampers the
establishment of the actual stratigraphic ranges of the identified taxa. Our record provides us to
characterize the orthophragminids in shallow benthic zone (SBZ) 17 for Eastern Tethys in detail
by comparing the data from the above localities with those from the North Africa, Europe and
Turkey, showing the change in diversity.

1. Introduction of orthophragminids from the Fulra Limestone and

Paleocene and Eocene shallow-marine deposits in Indian
subcontinent were traditionally regarded to have formed
under three major transgressions, Ranikot (Paleocene),
Laki (early Eocene) and Kirthar (middle Eocene), the latter
being the most significant one owing to deposition of
highly fossiliferous beds in India and Pakistan (Eames,
1952a, 1952b; Nagappa, 1959; Nuttall, 1926). The Fulra
Limestone in Kutch Basin in western India and the cor-
relative units in Pakistan, the Pirkoh and Drazinda forma-
tions in Sulaiman Range correspond to Bartonian Kirthar
transgression coeval with the Middle Eocene Climatic
Optimum (MECO) (Khanolkar, Saraswati, & Rogers, 2017).
The Bartonian shallow-marine deposits are exposed in
Kutch, Cambay and Jaisalmer basins in western India,
in the Sulaiman and Kirthar ranges in Pakistan, and in
the Assam-Meghalaya regions in eastern India. Generally
referred to as ‘Nummulitic limestone’, these sequences
contain abundant orthophragminids that are locally
more common than nummulitids (Ali et al., 2018; Özcan,
Saraswati, Hanif, & Ali, 2016b; Samanta & Lahiri, 1985).
Our studies on the taxonomy and paleobiogeography
Sulaiman Range in Pakistan revealed a significant
number of new species that appear to be confined to
Indian subcontinent (Ali et al., 2018; Özcan et al., 2016b).
This suggests a major faunal differentiation between
peri-Mediterranean/Europe region (Western Tethys)
and Indian subcontinent (Eastern Tethys) in the middle
Eocene. As opposed to Western Tethyan orthophrag-
minid assemblages, Discocyclina, along with subordi-
nate Asterocyclina, is the most common group in Fulra
Limestone, whereas Orbitoclypeus is represented by a
single species, O. haynesi. Pirkoh and Drazinda forma-
tions from Pakistan yielded predominantly Discocyclina
consisting of some new species including Discocyclina
pseudodispansa Özcan, Ali & Yücel, 2018, D. sulaimanen-
sis Özcan, Ali & Hanif, 2016, D. rakhinalaensis Özcan, Ali
& Yücel, 2018, D. zindapirensis Özcan, Ali & Yücel, 2018,
and D. kutchensis Özcan & Saraswati, 2016, the latter one
occurring also in the Fulra Limestone (Ali et al., 2018;
Özcan et al., 2016b). A new morphological structure,
bulges, observed at the test surface as semi-rounded to
rounded thickening of the lateral layers, in D. kutchensis
was introduced (Özcan et al., 2016b).

CONTACT Ercan Özcan ercanozcan034@yahoo.com, ozcanerc@itu.edu.tr

© 2018 The Author(s). Published by Informa UK Limited, trading as Taylor & Francis Group.
This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted
use, distribution, and reproduction in any medium, provided the original work is properly cited.
138  E. ÖZCAN ET AL.

We aim to describe here in detail the orthophrag-
minids from the Fulra Limestone to complement our
findings from the Pirkoh and Drazinda formations in
Pakistan (Ali et al., 2018). We make a general synthesis
of Bartonian orthophragminids from the Indian subcon-
tinent by incorporating and comparing the data from
Western Tethys (Europe, north Africa and Turkey). A syn-
thesis for shallow benthic zone (SBZ) 17 for Indian sub-
continent is made in the light of new orthophragminid
data from this region.
deepening. The Fulra Limestone is overlain unconform-
ably by the Maniyara Fort Formation.
The age of the Fulra Limestone is firmly constrained
as Bartonian by the occurrence of key planktonic spe-
cies identified at its lower part and in the upper part of
the underlying Harudi Formation (see Saraswati et al.,
2017 for details). Samanta (1970) described planktic

2. Geological setting and stratigraphy

2.1. Kutch Basin
In Kutch the Eocene succession represents the first
marine transgression following the Deccan volcan-
ism and consists of Naredi and Harudi formations and
Fulra Limestone (Biswas, 1992; Saraswati, Khanolkar, &
Banerjee, 2017) (Figure 1(A), (B)). The Fulra Limestone
conformably overlying the Harudi Formation is inter-
preted to have been deposited in a warm, oligotropic sea
during the globally recognized warming period called
Middle Eocene Climatic Optimum (MECO) (Khanolkar
et al., 2017). It comprises exclusively of shallow marine
carbonates consisting of various fossil groups including
planktonic foraminifera at its lower part (Khanolkar et
al., 2017; Samanta, 1970, 1993; Samanta, Bandopadhyay,
& Lahiri, 1990; Samanta & Lahiri, 1985; Saraswati, Patra,
& Banerjee, 2000; Saraswati et al., 2017; Sen Gupta,
1963a, 1963b). The unit is characterized by six facies, as
orthophragminid mudstone (sample FUL1, Figure 2(A)),
orthophragminid wackestone-mudstone alternation,
orthophragminid wackestone-packstone alternation,
nummulitic grainstone, nummulitic wackestone-pack-
stone alternation, and Alveolina wackestone-packstone
alternation (samples FUL2–15) corresponding to vari-
ous depositional settings on a carbonate ramp (Figures
1(C), 2(C)–(E)) (Banerjee, Khanolkar, & Saraswati, 2017).
These facies indicate a depositional spectrum ranging
from bar-lagoon to mid-ramp depositional setting.
Orthophragminid mudstone in the lower part of the
Fulra succession contains planktonic foraminifera includ-
ing Orbulinoides beckmanni and records the deepest
bathymetry, up to 60 m, corresponding to mid-ramp
setting. The cross-stratified Nummulitic grainstone
beds, with slightly convex-up geometry and common
palaeokarst surfaces, represent a high-energy deposit
in inner-ramp setting (Banerjee et al., 2017; Chattoraj,
Sarkar, Chakraborty, Banerjee, & Saraswati, 2012). These
beds record the shallowest deposits in the ramp indi-
cating low-relief bars in shoal environment. Nummulitic
wackestone-packstone alternation represents a shoal
flank environment. Orthophragminid wackestone-pack- Figure 1. Location of Kutch Basin and Sulaiman Range in W India
and Pakistan (A), simplified geological map of the Kutch and
stone alternation and orthophragminid wackestone-
position of the studied sections (B), and composite stratigraphic
mustone alternations form more distally. The repetetion section of the Fulra Limestone (C).
of above facies record repeated phases of shallowing and Note: Geological map is after Biswas (1992).
 GEODINAMICA ACTA  139

Figure 2. Field aspects of the Fulra Limestone: Transition from Harudi Formation to the Fulra Limestone (A, B), the lower beds of
the Fulra Limestone (sample FUL1) with planktonic foraminifera and LBF (B), close-up view of sample FUL6 with predominantly
Discocyclina discus and D. dispansa (C), a shallowing upward cycle with transition from wackestone-packstone alternation (FUL6–10)
to grainstone facies (FUL11–12) in the middle part of the unit (D), the upper carbonates of the Fulra Limestone of predominantly
grainstone facies (E).

foraminifera from the Fulra Limestone and assigned Dutta, and Banerjee (2014) identified Orbulinoides beck-
it to the Orbulinoides beckmanni and Truncorotaloides manni, Acarinina rohni, A. topilensis, Streptochilus martini,
(Acarinina) rohri zones. Saraswati, Khanolkar, Raju, Pseudohastigerina micra, and Jenkinsina columbiana in
140  E. ÖZCAN ET AL.
shale beds immediately below the Harudi Formation-
Fulra Limestone boundary and assigned P13 Zone to this
level. According to the scheme of Berggren and Pearson
(2005) the unit corresponds to the Zones E12 and E13.
The Fulra Limestone was assigned to shallow benthic
zone (SBZ) 17 based on orthophragminids (Ben Ismail-
Lattrache et al., 2014; Özcan et al., 2016b) and benthic
foraminiferal assemblages supplemented by the plank-
tonic foraminifera (Khanolkar et al., 2017).
2.2. Sulaiman range
The Middle to Upper Eocene sedimentary sequence
in Sulaiman Range in Pakistan consists of Habib Rahi,
Domanda, Pirkoh and Drazinda formations that were col-
lectively named as ‘Kirthar’ by Blanford (1879) (Figure 3).
The term ‘Drazinda Shale Member’ of the ‘Kirthar’
Formation was introduced by Hemphill and Kidwai
(1973) to replace the ‘Upper Chocolate Clays’ of Eames
(1952a, 1952b). The Drazinda Formation, more than 380
meters in thickness consists of dark-brown to greenish
gray shale and subordinate marl and limestone beds
containing abundant LBF, bivalves, bryozoans and echi-
noids in its lower and middle, and pale yellowish green
Pellatispira-bearing marls in upper part. The previous
studies suggested a middle and/or late Middle Eocene
age for the Drazinda Formation based on calcareous
nannofossils (Köthe, Khan, & Ashraf, 1988), planktonic
foraminifera (Afzal, Asrar, & Naseer, 1997; Samanta, 1973;
Warraich & Nishi, 2003) and orthophragminids (Ali et al.,
2018). According to Köthe et al. (1988), the age of this
unit in Rakhi Nala section is middle Eocene based on the
presence of nannoplankton zones NP 16 and 17. Afzal
et al. (1997) identified P14 and dated the lower part of
Drazinda Formation (below the ‘Pellatispira beds’) as late
middle Eocene (Bartonian) in age. The Pellatispira-beds
containing Heterostegina, reticulate Nummulites and
Pellatispira is considered Priabonian (Özcan et al., 2016a).
The Pirkoh Formation was deposited in the inner to
outer shelf, and the bulk of the Drazinda Formation has
been deposited in the shelf lagoon and shoal environ-
ments. The ‘Pellatispira beds’ of the Drazinda Formation
have been deposited in the outer shelf (Abbas, 1999).
The Drazinda Formmation is unconformably overlain
by coastal deltaic to fluviatile deposits of Oligocene
Chitarwatta Formation (Shah, 2009).
3. Material and methods
3.1. Material
The samples of the Fulra Limestone were collected
from three sections, near Kharai and Harudi villages,
100 km northwest of Bhuj, Kutch (23°30′55.97″N,
68°40′5.98″E; 23°28′45.32″N, 68°40′49.23″E) (Figures
1, 2). We have sampled 15 levels representing the
unit. The Pirkoh and Drazinda formations have been
sampled in three different outcrop areas, Rakhi Nala,
Zinda Pir and Domanda Bridge, in the Sulaiman Range
to provide the widest coverage of the orthophrag-
minids (Figures 3, 4). The Rakhi Nala A 29°57′12.80″N,
70°06′56.80″E; 29°57′13.51″N, 70°7’1.74″E), and Rakhi
Nala B (29°57′25.92″N, 70°7′0.50″E; 68°40′49.23″E,
29°57′16.10″N,70°7′11.20″E) sections are from the same
area west of Dera Ghazi Khan in the Punjab province.
The Zinda Pir section (30°20′2.38″N, 70°29′32.54″E;
30°19′56.65″N, 70°29′39.48″E), located in the Zinda Pir
anticline, is ca. 55 km north-east of the Rakhi Nala sec-
tion. The Domanda Section (31°35′0.19″N, 70°11′40.31″E;
31°35′14.43″N; 70°11′28.78″E) is located in the north-
west of both Rakhi Nala and Zinda Pir sections, and is
about 73 km west of Dera Ismail Khan in the Federally
Administered Tribal Areas (FATA) (Ali et al., 2018).
3.2. Sample preparation
Specimens extracted from the shale, marl and limestone
beds were studied for their external features, features
in the equatorial layer and axial sections. The oriented
sections of 507 megalospheric and 70 microspheric
specimens (A and B-forms respectively) have been pre-
pared through their equatorial layer. The morphometric
measurements and counts were carried out on equato-
rial sections of the megalospheric specimens. Axial sec-
tions of 83 megalospheric and microspheric forms have
been prepared for the comparisons in axial sections and
also in order to facilitate specific recognition in rock thin
sections.
4. Test features of the Tethyan
orthophragminids
Orthophragminids are bilamellar, perforate foraminif-
era characterized by a discoidal, lenticular test with an
equatorial layer consisting of cyclically arranged equa-
torial chambers and lateral layers composed of lateral
chambers and pillars on either side of the equatorial layer
(Figure 5(A)). Externally, the test surface is either smooth,
occasionally with an inflated or depressed central part,
or it is characterized by radially developed ribs and
bulges (Özcan et al., 2016b). Based on the relationship
of protoconch and deuteroconch in equatorial sections,
about ten configurations are used in the description of
the embryon (Figure 5(B)). The equatorial chambers are
divided into chamberlets of different shapes as observed
in equatorial sections. Tethyan orthophragminids consist
of five genera; Discocyclina Gümbel, 1870, Nemkovella
Less, 1987 and poorly known Asterophragmina Rao,
1942 placed in Discocyclinidae Galloway, 1928 and two
genera, Orbitoclypeus Silvestri, 1907 and Asterocyclina
Gümbel, 1870 placed in Orbitoclypeidae Brönnimann,
1946 (Ferràndez-Cañadell, 1998; Less, 1987) (Figure 6).
The obsolete genus Actinocyclina Gümbel 1870, charac-
terized by the presence of numerous ribs, is assigned to
 GEODINAMICA ACTA  141

Figure 3. Location of Sulaiman Range in Pakistan (A), simplified geological map of the Sulaiman Range (B), and stratigraphic sections
and sampling points from the Pirkoh and Drazinda formations (C).
Notes: Geological map is simplified from Kazmi and Rana (1982). PSB: Pellatispira beds (from Ali et al., 2018).
142  E. ÖZCAN ET AL.
Figure 4. Field aspects of Pirkoh Limestone and Drazinda Formation from Sulaiman Range in Pakistan and sampling points (from Ali
et al., 2018).
genus Discocyclina after their discocyclinid-type micro-
spheric juvenarium (Ferràndez-Cañadell, 1997; Less,
1987) (Figure 6.1). The microspheric forms of some of
the Tethyan genera are illustrated in Figure 6.
5. The orthophragminids from the Fulra
Limestone and Pirkoh and Drazinda
formations
The LBF in Fulra Limestone are characterized by com-
mon occurrence of orthophragminids, nummulitids
and alveolinids accompanied by rare rotaliids, and
some stratigraphically important taxa as Dictyoconoides,
Linderina, Calcarina, and smaller benthic foraminif-
era (Khanolkar et al., 2017; Samanta, 1993; Samanta &
Lahiri, 1985; Samanta et al., 1990; Saraswati et al., 2000;
Sen Gupta, 1963a, 1963b). Orthophragminids belong
mainly to Discocyclina Gümbel, 1870, a single species
of Orbitoclypeus Silvestri, 1907 and three species of
Asterocyclina Gümbel, 1870 were also recognized. The
following species are identified; Discocylina dispansa
(Sowerby, 1840), Discocyclina pseudodispansa Özcan, Ali
& Yücel, 2018, Discocyclina kutchensis Özcan & Saraswati,
2016, Discocyclina praeomphalus Samanta & Lahiri, 1985;
Discocyclina augustae van der Weijden, 1940; Discocyclina
discus (Rütimeyer, 1850), Discocyclina pratti (Michelin,
1846), Orbitoclypeus haynesi (Samanta & Lahiri, 1985),
Asterocyclina sireli Özcan & Less, 2006, Asterocyclina alti-
costata (Nuttall, 1926), Asterocyclina stellata (d’Archiac,
1846) (Figure 7(A)). The nummulitids, such as Nummulites
beaumonti (Figure 8.1–2), Nummulites sp. 1 (Figure 8.3–4),
Nummulites maculatus (Figure 8.5–6), Nummulites gr. bul-
latus (Figure 8.7–8), Operculina gr. gomezi (Figure 8.9–
10), Linderina sp. (Figure 8.11–12), Calcarina sp. (Figure
8.13–14), and Dictyoconoides sp. (Figure 8.15). Operculina
gr. gomezi, a diagnostic Bartonian-Priabonian species in
Western Tethys (Less & Özcan, 2012), is identified for the
first time from the Fulra Limestone.
Orthophragminids in the Pirkoh and Drazinda for-
mations are represented by Discocyclina and scanty
Asterocyclina while Orbitoclypeus and Nemkovella are
absent (Figure 7(B)). The Pirkoh Formation yielded only
discocyclinids, represented by D. dispansa (Sowerby,
1840), D. discus (Rütimeyer, 1850), D. praeomphalus
Samanta & Lahiri, 1985 accompanied by Dictyoconoides,
Nummulites, Assilina, Operculina, Linderina and
 GEODINAMICA ACTA  143

Figure 5. General test features in Tethyan orthophragminid genera (A) (after Less, 1987; Ferràndez-Cañadell, 1997; Özcan et al.,
2016b), qualitative parameters (B): a- types of embryon configurations, b- types of the adauxiliary chamberlets, c- different growth
patterns of the equatorial annuli, d- types of granules and lateral chamberlets on the test surface, and parameters used in the
morphometric description of orthophragminids as illustrated in D. pseudodispansa from Sulaiman Range (C). pac: principal auxiliary
chamberlets.

alveolinids, found only in one level (sample ZP.2) in the
Zinda Pir section. Orthophragminids in the Drazinda
Formation are more diverse and are characterized by D.
dispansa (Sowerby, 1840), D. discus (Rütimeyer, 1850),
D. praeomphalus Samanta & Lahiri, 1985; D. augustae
van der Weijden, 1940; ‘D.’ sulaimanensis Özcan, Ali &
Hanif, 2016, D. kutchensis Özcan & Saraswati, 2016, D.
nandori Less, 1987; D. pseudodispansa Özcan, Ali &
Yücel, 2018, D. rakhinalaensis Özcan, Ali & Yücel, 2018,
D. zindapirensis Özcan, Ali & Yücel, 2018, Asterocyclina
sireli Özcan & Less, 2006, and A. stellata (d’Archiac, 1846).
The associated LBF are represented by Nummulites sp.,
Assilina sp., Operculina sp., Calcarina sp., and Linderina sp.
The Pellatispira-beds in the upper part of the Drazinda
Formation contain scarce unidentified Discocyclina and
other LBF, such as Pellatispira, reticulate Nummulites,
Heterostegina, Silvestriella and Operculina (Ali et al.,
2018).
6. Systematic paleontology
Detailed description of some of the orthophragminids
from the Fulra Limestone was given by Sen Gupta
(1963a) and Samanta and Lahiri (1985). We here focus
144  E. ÖZCAN ET AL.

Figure 6. Microspheric juvenarium representative of some Tethyan orthophragminid genera (1–4: family Discocyclinidae Galloway,
5–7: family Orbitoclypeidae Brönnimann). 1: Nemkovella evae Less, Horsarrieu, SW France, early Cuisian. 2: Discocyclina nandori Less,
(a species with ribs), Cambay Basin (W India), late middle Eocene. 3–4: Discocyclina archiaci (Schlumberger), Patala Formation (Thal,
Pakistan), early Eocene. 5: Orbitoclypeus haynesi (Samanta & Lahiri, 1985), Fulra Limestone (Kutch, India), late middle Eocene. 6–7:
Asterocyclina stellata (d’Archiac), Soğucak Formation (Thrace Basin, Turkey), middle-late Eocene transition.

on the general features important for the genus and
morphometry of these taxa.
Order Foraminiferida Eichwald, 1830
Family Discocyclinidae Galloway, 1928
Genus Discocyclina Gümbel, 1870
Type-species: Orbitolites pratti Michelin, 1846
Discocyclina dispansa (Sowerby, 1840)
(Figures 9(B), 10)
1840 Lycophris dispansus n. sp., Sowerby, p. 327, pl.
24, Figs. 16, 16a–b.
1926 Discocyclina undulata sp. nov., Nuttall, p. 160–
151, pl. 7, Figs. 8–9, pl. 8, Fig. 5.
2018 Discocyclina dispansa (Sowerby, 1840), Ali et al.,
Figs. 9, 12, 15, 17.
Diagnosis. Discocyclina dispansa is a small to large
sized, ‘flat to saddle’ shaped, unribbed form. The small to
medium-sized megalospheric embryon is semi-nephro-
lepidine in the earliest representatives of its lineage (e.g.
D. d. broennimanni and D. d. taurica) and is trybliolepidine
in the phylogenetically advanced members. The
 GEODINAMICA ACTA  145

Figure 7. Distribution of orthophragminids and associated LBF from the Fulra Limestone (A) and Pirkoh and Drazinda formations (B).
Notes: The synthetic stratigraphic columnar section representing Pirkoh and Drazinda formations in Sulaiman Range includes data from Rakhi Nala A and B,
Zinda Pir, and Domanda Bridge sections.
146  E. ÖZCAN ET AL.

Figure 8. Common LBF associated with orthophragminids in the Fulra Limestone. 1–2: Nummulites beaumonti, 1: external view,
FUL3–27, 2: equatorial section, FUL3–29. 3–4: Nummulites sp. 1, external view and equatorial section, FUL3–28. 5–6: Nummulites
maculatus, external view and equatorial section, FUL2–33. 7–8: Nummulites gr. bullatus, external view and equatorial section,
FUL9–2. 9–10: Operculina gr. gomezi, 9: external view, FUL9–5, 10: equatorial section, FUL12–16. 11–12: Linderina sp., external view
and equatorial section, FUL2–32. 13–14: Calcarina sp., external view and equatorial section, FUL9–3. 15: Dictyoconoides sp., vertical
section, FUL2–26. 16–26: Alveolina spp., 16–21: FUL15, 22: FUL4, 23–26: FUL12–62.

adauxiliary chamberlets are moderately wide and high, Less, 1987 (Dmean < 160 μm); D. d. taurica Less, 1987
and of the ‘archiaci’ type. The equatorial chamberlets are (Dmean = 160–230 μm); D. d. hungarica Kecskeméti,
also moderately wide and high. This species includes 1959 (Dmean = 230–290 μm); D. d. sella (d’Archiac, 1850)
six subspecies in Western Tethys: D. d. broennimanni (Dmean = 290–400 μm); D. d. dispansa (Sowerby, 1840)
 GEODINAMICA ACTA  147

Figure 9. External test features of orthophragminids from the Fulra Limestone (B, F, I-N) and Drazinda Formation (A, C-E, G-H). A:
Discocyclina discus, B: D. dispansa, C: D. pseudodispansa, D: D. sulaimanensis, E: D. zindapirensis, F: D. augustae, G–H: D. nandori, I: D.
praeomphalus, J–K: D. kutchensis, L: Asterocyclina alticostata, M: A. sireli, N: Orbitoclypeus haynesi.

(Dmean = 400–520 μm); and D. d. umbilicata (Deprat,
1905) (Dmean > 520 μm) (Less, 1998).
Remarks. Discocyclina dispansa, one of the most com-
mon orthophragminid species in the Fulra Limestone,
first described from this unit by Sowerby (1840). In Fulra
Limestone and Drazinda Formation, the species is rep-
resented by a flat test with a thick umbo, surrounded
by a thin flange. In equatorial layer, a characteristic
feature is the presence of the high early chambers fol-
lowed by low chambers (Figure 10.1–2). It appears that
in the Western Tethys, saddle-shaped and flat forms
exhibiting the ‘similar’ embryonic configuration and
development of equatorial chambers appear to have
been lumped under this species as evidenced by nota-
ble overlaps in the ranges of some subspecies such
as D. dispansa hungarica and D. dispansa sella in the
Lutetian and Bartonian (Zakrevskaya, Beniamovsky,
Less, & Báldi-Beke, 2011), and D. dispansa sella and D.
dispansa dispansa in the Bartonian (Less, Özcan, & Okay,
2011). We think that a revision of this group is necessary
focusing on the distinction between the flat and saddle
forms. Based on biometry (Table 1), the species is rep-
resented by D. d. dispansa (Sowerby, 1840) in the Fulra
Limestone. This species may externally be confused with
D. pseudodispansa.
Discocyclina pseudodispansa Özcan, Ali & Yücel,
2018
(Figures 9(C), 11)
2018 Discocyclina pseudodispansa n. sp. Özcan, Ali, &
Yücel, Ali et al., Figs. 9, 12, 15, 16.
148  E. ÖZCAN ET AL.
Figure 10. Equatorial and axial sections of D. dispansa dispansa from the Fulra Limestone. 1: sample FUL12–72, 2: FUL8–23, 3: FUL7–
5, 4: FUL13–208.
Diagnosis. Flat test, small-to medium-sized, strongly
inflated with a thick umbo. The small embryon consists
of a rounded to nearly-rounded protoconch and larger
deuteroconch displaying semi-nephrolepidine to try-
bliolepidine type configuration in equatorial sections.
The annuli are typically narrow, presenting ‘archiaci’ type
growth pattern. The piles are coarser in the umbonal
part, compared to flange. Moderately high and wide
lateral chamberlets are arranged in irregular rows.
Description.
External features. The test is typically flat, small to
medium sized (1.35–6.8 mm in diameter) with a thick
umbo, surrounded by a narrow, thin flange (Figure
12). The thickness of the test varies between 600 and
2100 μm. The piles are relatively larger along the rims of
the umbo (80–160 μm) compared to those in the flanges
(50 μm), and are evenly distributed over the test surface.
A detailed description of the species is given in Ali et al.
(2018).
Remarks. Discocyclina pseudodispansa, a common
species in the Sulaiman Range, occurs only in the lower-
most sample from Fulra Limestone. It could be easily con-
fused with D. dispansa on the basis of external characters.
The axial sections of both species are also similar to each
other in terms of development of lateral chamberlets and
their size (compare Figures 10.4, 11.4–5) (Ali et al., 2018).
This species, however, differs from D. dispansa in having a
much smaller embryon and tighter early chambers (n0.5
parameters in Table 1). In equatorial sections, some of
the specimens exhibiting semi-nephrolepidine type of
embryon in the Sulaiman Range may be confused with D.
augustae. The external features and axial sections of both
species are totally different as the lateral chamberlets in
D. augustae are very low (slit-like) and small in size.
Discocyclina augustae van der Weijden, 1940
(Figures 9(F), 12)
1940 Discocyclina augustae n. sp. van der Weijden, p.
23−26, pl. 1, Figs. 4, 5, 7, 8; pl. 2, Figs. 1, 2, 11.
2018 Discocyclina augustae van der Weijden, 1940, Ali
et al., Figs. 10–11, 12, 21.
Diagnosis. Discocyclina augustae is an unribbed
form having a very small to small, semi-iso- to neph-
rolepidine embryon, narrow and low, ‘archiaci’ type
adauxiliary chamberlets and also narrow and rela-
tively low equatorial chamberlets mostly with ‘strophi-
olata’ type growth pattern. This species includes four
subspecies in Western Tethys: D. a. sourbetensis Less,
1987 (Dmean < 145 μm); D. a. atlantica Less, 1987
(Dmean = 145–180 μm); D. a. olianae Almela & Rios,
1942 (Dmean = 180–225 μm); D. a. augustae van der
Weijden, 1940 (Dmean > 225 μm).
Remarks. Discocyclina augustae is easily differentiated
externally by its flat test with small central umbo and
uniformly distributed piles (Figure 9(F)). It can be, how-
ever, easily confused with D. pseudodispansa in equato-
rial sections, whereas their axial sections are completely
different. This species has a much larger embryon than
that of D. rakhinalaensis (Ali et al., 2018). In the studied
material, D. augustae is represented by transitional stages
between D. a. atlantica and D. a. olianae, according to
the evolutionary scheme of the genus in the Western
Tethys (Table 1).
Discocyclina praeomphalus Samanta & Lahiri, 1985
(Figures 9(I), 13)
1985 Discocyclina praeomphalus n. sp. Samanta &
Lahiri, p. 272–275. pl. 5, Figs. 1–6, text Figures 5–7, 12.
2018 Discocyclina praeomphalus, Ali et al., Figs. 10, 14,
23–24.
Table 1. Statistical data of orthophragminids from the Fulra Limestone.
Outer cross diameter of the embryon Adauxiliary chamberlets Equatorial chamberlets
Deuteroconch Protoconch Number Height Width Annuli/0.5 mm Height Width
D P A H w n0.5 h w
Sample N Range Mean ± S.E Range Mean Range Range Range Range Range Range Species/subspecies
FUL1 16 270–385 329.38 ± 10.1 120–170 143.57 23–31 30–70 30–65 9–12 50–90 30–60 Discocyclina pseudodispansa
FUL2 15 180–300 223.67 ± 7.6 85–135 103.67 23–25 25–55 20–70 11–14 40–65 20–35 Discocyclina praeomphalus
FUL3 12 185–255 223.3 ± 7.2 80–105 91.11 25–28 30–50 20–35 12–16 50–85 20–35
FUL2 3 1430–1560 1480 ± 33.0 550–710 630.0 88 85–170 55–105 4–5 125–165 35–45 Discocyclina discus sowerbyi
FUL3 2 1170–1755 1462.5 615 - 65–155 65–90 3–5 85–115 35–50
FUL5 2 1140–1220 1180.0 440.0 - 150–205 55–85 3 140–150 40–60
FUL6 10 1030–1910 1396.5 ± 73.3 460–1405 761.6 - 75–200 35–180 3.5–6 130–155 30–50
FUL7 3 1180–1875 1481.6 ± 168.0 765 - 80–210 55–90 3–4 70–80 35–45
FUL8 3 1360–1420 1383.3 ± 15.5 550–620.0 585.0 - 100–165 55–90 5 100–150 25–35
FUL9 5 800–1670 1259.0 ± 125.3 390–515 447.0 - 55–200 40–100 4–5 75–120 25–65
FUL13 3 1280–1550 1400.0 ± 64.8 370–550 473.0 - 85–155 35–85 4 90–135 35–75
FUL2 2 470–550 510.0 ± 28.2 210–220 215.0 - 40–80 30–50 8–9 40–95 25–40 Discocyclina dispansa dispansa
FUL3 4 430–655 546.2 ± 43.6 170–220 200.0 41–42 40–90 20–50 7–9 85–140 25–45
FUL5 4 360–490 441.2 ± 24.3 160–220 195.0 42–46 35–90 25–50 9–11 55–95 25–35
FUL6 10 360–655 515.5 ± 23.3 135–215 187.0 35–46 70–100 35–45 7–9 50–105 20–45
FUL7 4 475–665 546.2 ± 39.3 185–260 218.7 48–49 50–100 25–70 7–8 40–75 25–50
FUL9 1 475 - 45–75 25–50 6 75–80 35–55
FUL8 5 400–690 583.0 ± 45.5 115–300 227.0 50 35–100 25–65 6–7 35–75 25–50
FUL11 1 460 155 36 50–85 25–55 9 70–85 30–60
FUL12 7 400–745 529.3 ± 43.4 180–260 210.0 39–43 30–85 25–70 7–8 60–150 35–50
FUL13 21 380–700 559.5 ± 19.0 150–300 210.8 49–55 40–100 25–60 7–9 65–90 20–50
FUL14 5 450–520 481.0 ± 10.6 150–210 185.0 - 30–55 25–45 8 75–85 25–40
FUL5 2 170–180 175.0 75–80 77.5 - - - 14 30–50 25–30 Discocyclina augustae ex. interc. atlanti-
ca-olianae
FUL6 11 155–190 174.1 ± 2.47 65–110 75.0 20–25 20–40 20–40 15–16 30–80 20–35
FUL8 4 165–175 171.2 ± 2.07 70–75 71.7 22 30–45 20–40 12–17 50–65 25–50
FUL11 1 150 70 19 25–30 25–45 18 50–65 30–45
FUL12 5 150–200 180.0 ± 8.12 70–85 80.0 21 20–30 20–35 16–18 50–55 25–50 Discocyclina augustae ex. interc. ol-
ianae-atlantica
FUL13 24 150–215 184.4 ± 3.61 60–110 86.1 22–26 25–45 20–40 15–17 25–60 25–40
FUL6 34 275–610 411.1 ± 14.6 105–250 169.4 27–47 70–110 25–50 5–6 75–100 25–30 Discocyclina kutchensis
FUL13 6 310–460 423.3 ± 21.0 150–200 177.5 28–36 80–120 25–45 5–6 80–120 25–30
FUL13 6 605–865 730.8 ± 35.8 250–395 335.0 70–74 60–120 35–65 4–7 75–110 25–35 Discocyclina pratti
FUL3 1 150 75 - 30–40 20–25 14–15 70–85 25–30 Orbitoclypeus haynesi
FUL7 2 225–260 242.5 ± 12.3 110–140 125.0 25 25–50 20–60 12–13 65 30–35
FUL8 16 215–310 251.8 ± 6.9 90–175 130.0 23–32 30–50 25–45 14–15 45–70 25–35
FUL9 1 230 115 30–45 20–45 15 35–55 25–35
FUL11 8 205–250 225.0 ± 5.5 80–115 105.0 25–30 25–55 25–40 12–14 40–90 25–40
FUL12 51 165–300 231.18 ± 3.8 85–160 115.7 26–28 35–60 20–50 13–17 40–70 25–40
FUL13 24 190–300 243.33 ± 5.6 100–165 122.5 27–30 35–65 25–45 12–15 35–95 25–40
GEODINAMICA ACTA 

FUL14 4 200–245 218.75 ± 8.3 90–135 108.7 26–29 35–50 20–50 13–15 50–100 30–40
FUL6 20 110–140 124.7 ± 2.1 60–90 74.74 25–35 25–65 23 40–85 20–25 Asterocyclina sireli

(Continued)
149
 150  E. ÖZCAN ET AL.

Diagnosis. Discocyclina praeomphalus is a medium

Asterocyclina alticostata ex. interc. alticos-

sized (2.4–4 mm), saddle shaped, unribbed form with a

Asterocyclina alticostata ex. interc. cuvil-

slight depression at the center of the test. The megalo-
spheric embryon (average 222.0–223.7 μm in diameter)

Species/subspecies
exhibits semi-nephrolepidine to trybliolepidine type
configuration. The adauxiliary chamberlets are mod-

Asterocyclina stellata
erately wide and high, and of the ‘archiaci’ type. The

lieri-alticostata
equatorial chamberlets are low and narrow in the early
tata-cuvillieri stage and are rectangular in shape in the later stages.
The lateral chambers are typically low, numerous and
not aligned in regular rows.
Remarks. This saddle-shaped, omphaloid species, first
described from the Fulra Limestone (Samanta & Lahiri,
Range
Width

20–35
20–25
30–35

35–60
35–40
25–35
30–40

25–35
25–30
25–30
w

1985), occurs abundantly in the Drazinda Formation

in the Sulaiman Range (Ali et al., 2018). It differs exter-
Equatorial chamberlets

nally from other orthophragminids from the Drazinda

Formation by its characteristic saddle shape and the
150–165
70–115

80–160

55–160

70–120
Height

Range
35–45
35–40

55–90

40–55
75–85

central depression of the test. Discocyclina sulaiman-

h

ensis, another omphaloid species from the Drazinda

Formation, has a flat and smaller test than that of D.
Annuli/0.5 mm

praeomphalus. The axial sections of D. praeomphalus

Range
21–22
23–25
11–12

10–13

10–14
n0.5

9–11

are easily differentiated from that of the saddle-shaped

8–9

25
20
9

D. discus in having very low (slit-like) chamberlets. This

is a common discocyclinid in the middle Eocene of the
Indian Subcontinent.
150–280
105–285

110–280

Discocyclina kutchensis Özcan & Saraswati, 2016

80–260

70–265

45–125
Range
Width

Notes: For the illustration and explanation of the parameters see Figure 5C. N denotes the number of specimens studied in the sample.
25–65
25–30

30–90
w

(Figures 9(J), (K), 14)

2016b Discocyclina kutchensis sp. nov. Özcan &
Adauxiliary chamberlets

Saraswati, p. 267–274, Figs. 5L, 6A–I, 7A, 8A–D, 9, 10A–D.

2018 Discocyclina kutchensis Özcan, Ali, & Yücel, Ali et
40–125

50–130
Height

Range
25–35
25–30
40–90

45–90

30–85

20–75
35–50
15–45
H

al., Figs. 9, 11, 14, 22–23.

Diagnosis. Medium sized (2 to 6 mm), flat forms with
numerous bulges. The bulges, uniformly distributed over
the test surface, are semi-rounded and rounded in shape
Number

Range

and range in size between 250 and 350 μm. The embryon
A

is large, with an average diameter of the deuteroconch

ranging between 410 and 478 μm, trybliolepidine to
umbilicolepidine in configuration. The adauxiliary cham-
Mean
78.24

262.0

284.0

235.0
245.6

254.4
135.0
100.0
84.1

375

berlets (37–53 in number) are high and archiaci-type.

Protoconch

The equatorial chambers are typically high, narrow and

P
Outer cross diameter of the embryon

rectangular in shape.
215–325

220–350

160–370
110–295

200–300
30–105
Range
70–95

Remarks. This species, originally described from the

Fulra Limestone and Drazinda Formation, differs from
most orthophragminids in Tethys by having the bulges
355.0 ± 14.5

380.0 ± 22.9

375.0 ± 35.6

uniformly distributed on the test surface. The bulges

Mean ± S.E
127.2 ± 1.7
135.5 ± 1.6

348.4 ± 9.6

346.9 ± 6.8
205.0
170.0
440

are semi-rounded to rounded, localized thickenings,

Deuteroconch

homogenously distributed over the test surface (Özcan

et al., 2016b). These structures are about 250–350 μm in
D

diameter, 100–150 μm high and 100–300 μm apart from

115–140
105–160
300–430

300–470

270–450
280–450

270–450

each other, and form an uneven test surface. Internally,

Range

the bulges consist of lateral chamberlets and coarse piles

Table 1. (Continued).

of up to 100 μm in diameter at its center, surrounded by

smaller piles (25–50 μm in diameter), forming a circu-
18
52
10

6
1
4
16

36
1
1
N

lar pattern. These piles are semi-circular to polygonal in

shape in transversal sections. The lateral surfaces of the
Sample

bulges are rather sharp. The axial sections show no vari-

FUL.13
FUL13

FUL12

FUL13
FUL8

FUL6

FUL8
FUL7
FUL9

FUL8

ation in the thickness of the equatorial layer adjacent to


Figure 11. Equatorial and axial sections of D. pseudodispansa from the Fulra Limestone. 1: FUL1–3, 2: FUL1–9, 3: FUL1–7, 4: FUL1–17,
5: FUL1–19.
the bulges and inter-bulge areas. This implies that bulges
are formed solely by the thickenings in the lateral layers
(Figure 14). The Bartonian ribbed orthophragminids in
Indo-Pakistan region are represented by four species;
Discocyclina nandori Less, 1987; Asterocyclina sireli Özcan
& Less, 2006, A. alticostata (Nuttall, 1926) and rare A. stel-
lata (d’Archiac, 1846). While they have different surface
features, internally, D. nandori is differentiated from D.
kutchensis by having a small, semi-nephrolepidine- to
trybliolepidine type embryon and by low equatorial
chamberlets (Özcan et al., 2016b). D. kutchensis is sim-
ilar to D. radians, a Western Tethyan species, in having
same type of embryon configuration, development of
equatorial chambers and their shape. Both species are
phylogenetically linked and D. kutchensis replaces D. radi-
ans in Eastern Tethys.
Discocyclina pratti
(Figure 15)
1846 Orbitolites pratti n. sp. Michelin, p. 278, pl. 63,
Fig. 14.
GEODINAMICA ACTA  151
Diagnosis. Discocyclina pratti is an unribbed spe-
cies having a medium-sized to large, tryblio- to excen-
trilepidine-type embryon, numerous moderately wide
and high, ‘pratti’ type adauxiliary chamberlets and nar-
row but high equatorial chamberlets with ‘pulcra’ type
growth pattern. This species includes three subspecies:
D. p. montfortensis Less, 1987 (Dmean < 510 μm), D. p.
pratti (Michelin, 1846) (Dmean = 510–700 μm), D. p. minor
Meffert, 1931 (Dmean > 700 μm) after the analysis of
rather rich assemblages from Turkish sections (Özcan,
Less, & Kertész, 2007).
Remarks. Some specimens with thin, fragile and
undulated test revealed embryons ranging from 600 to
865 μm. The embryons display various embryonic config-
urations ranging from umbilicolepidine to centrilepidine
where the wall of deutroconch may merge to the pro-
toconchal wall at some points. The equatorial chambers
are relatively high compared to those of D. discus. The
embryon appears to be smaller than of D. discus. This
species is rare in the studied material.
152  E. ÖZCAN ET AL.
Figure 12. Equatorial and axial sections of D. augustae ex. interc. olianae- atlantica from the Fulra Limestone. 4, microspheric; others,
megalospheric. FUL13–170, 2: FUL13–171, 3: FUL13–41, 4: FUL7–3, 5: FUL13–102, 6: FUL13–209, 7: FUL13–213, 8: FUL.13–210.
Discocyclina discus (Rütimeyer, 1850)
(Figure 9(A), 16)
1850 Orbitolites discus n. sp. Rütimeyer, p. 116, pl. 5,
Figs. 70−71, 78, 80−81.
1926 Discocyclina sowerbyi nom. nov., Nuttall, p.
149−150, pl. 3, Figs. 1–3.
1985 Discocyclina adamsi n. sp., Samanta & Lahiri, p.
229−242, pl. 1, Figs. 6–10, pl. 5, Fig. 9, text –Figs. 5, 8.
2010 Lepidocyclina sp. (L. pustulosa), Matsumaru &
Sarma, p. 550, pl. 3, Fig. 11.
2018 Discocyclina discus (Rutimeyer), Ali et al., Figs.
11, 14, 24–25.
Diagnosis. Discocyclina discus is a large, saddle-shaped
and unribbed form with a giant, mostly umbilicolepidine
(rarely also tryblio- or excentrilepidine) embryon, numer-
ous, wide and high, ‘archiaci’ or transitional ‘archia-
ci-pratti’ type adauxiliary chamberlets and wide and
high equatorial chamberlets with ‘archiaci’ or transi-
tional ‘archiaci-pulcra’ type growth pattern. This species
includes two subspecies in Tethys: D. d. discus (Rütimeyer,
1850) (Dmean < 1350 μm) and D. d. sowerbyi Nuttall, 1926
(Dmean > 1350 μm) after our present data from Fulra
Limestone.
Remarks. Discocyclina discus, widely reported from the
Fulra Limestone and the Drazinda Formation as D. sow-
erbyi, is distinguished from other orthophragminids by
its thick, saddle-shaped test and its large embryon with
irregular wall. D. praeomphalus, another saddle-shaped
species, is much smaller and thinner than D. discus (com-
pare the axial sections in Figures 12 and 16). In addition,
the equatorial chambers of D. discus are much higher
than those of D. praeomphalus. Our data show that the
thick, saddle-like forms described as Discocyclina sow-
erbyi (Nuttall, 1926) by Nuttall (1926) and Sen Gupta
(1963a) from the type locality of Lycophris ephippium of
Sowerby (1840) in the Kutch Basin, present the same mor-
phological features as D. discus (Rütimeyer, 1850). Nuttall
(1926) proposed the specific name ‘sowerbyi’ to replace
‘ephippium’, which was preoccupied by another species.
We maintain that D. sowerbyi, widely recorded from the
middle Eocene of the Indian Subcontinent, is a junior
synonym of D. discus. Discocyclina adamsi, established

Figure 13. Equatorial and axial sections of D. praeomphalus from the Fulra Limestone. 8–9, microspheric; others, megalospheric. 1:
FUL2–6, 2: FUL3–41, 3: FUL2–3, 4: FUL2–20, 5: FUL2–19, 6: FUL3–7, 7: FUL2–46, 8–9: FUL.3–8.
by Samanta and Lahiri (1985) from the Fulra Limestone,
is however, a junior synonym of D. sowerbyi, which was
established from the same unit by Nuttall (1926). We
emend here the subdivision of D. discus lineage by dif-
ferentiating two subspecies, D. discus discus and D. dis-
cus sowerbyi. This requires the replacement of ‘adamsi’ by
‘sowerbyi’. In Fulra Limestone the species belongs to D.
discus sowerbyi. Matsumaru and Sarma (2010) illustrated
an axial section of a saddle-shaped specimen with a large
and thick embryon from the late Eocene of Meghalaya
(NE India) and identified it as a Caribbean Lepidocyclina
sp. (L. pustulosa) without any illustration and description
of the equatorial layer (Pl. 3, Fig. 11). This was interpreted
to be the first record of the genus in Meghalaya and sig-
nificant for the occurrence of Caribbean lepidocyclinids
in the Eocene of Indian Subcontinent. In our opinion,
the illustrated axial section has the same morphological
features of D. discus in having a large embryon, high and
wide lateral chamberlets and saddle-shaped test.
Family Orbitoclypeidae Brönnimann 1946
Genus Orbitoclypeus Silvestri, 1907
Orbitoclypeus haynesi (Samanta & Lahiri, 1985)
(Figure 9(N), 17)
1985 Discocyclina haynesi n. sp. Samanta & Lahiri, p.
262−272, pl. 4, Figs. 1−6; pl. 12, Figs. 9−14; text-Figures
5−7.
GEODINAMICA ACTA  153
2010 Orbitoclypeus haynesi (Samanta & Lahiri), Özcan
et al., 2010, p. 59−61, Fig. 29i−n.
2013 Orbitoclypeus haynesi (Samanta & Lahiri), Ben
Ismail-Lattrache et al., 2013, p. 12, pl. 2, Fig. 9−15, fig. 8.
2017 Discocyclina dispansa Sowerby, BouDagher-
Fadel and Price (2017), Fig. 5−9.
Diagnosis. Orbitoclypeus haynesi is an unribbed spe-
cies with ‘marthae’-type rosette, small eulepidine (rarely
excentrilepidine) embryon, adauxiliary chamberlets of
‘varians’-type of average size and shape, moderately
wide and high equatorial chamberlets arranged into
strongly undulated annuli with ‘varians’-type growth
pattern.
Remarks. Orbitoclypeus haynesi is the only orbito-
clypeid in the Bartonian of Indian subcontinent. This
species, referred to Discocyclina by Samanta and Lahiri
(1985), was later assigned to Orbitoclypeus based on
the embryonic configuration, presence of undulated
annuli and shape of the equatorial chamberlets (Özcan
et al., 2010). We here show a microspheric form show-
ing orbitoclypeid type juvenarium justifying the generic
assignment (Figure 17.5–6). Although similar to O.
varians in external and internal test features, O. hayn-
esi possesses a much smaller embryon, tighter annuli
(parameter n0.5) and also commonly eulepidine-type
embryonic configuration. Some specimens may exhibit
154  E. ÖZCAN ET AL.

Figure 14. Equatorial and axial sections of D. kutchensis from the Fulra Limestone. 5, microspheric; others, megalospheric. 1: FUL6–
51, 2: FUL6–131, 3: FUL6–132, 4: FUL13–145, 5: FUL6–59, 6: FUL6–123, 7: FUL6–127, 8: FUL6–102.

Figure 15. Equatorial sections of Discocyclina pratti from the Fulra Limestone. 1: FUL9–4, 2: FUL13–174, 3: FUL13–188, 4: FUL13–172,
5: FUL13–192.
 GEODINAMICA ACTA  155

Figure 16. Equatorial and axial sections of D. discus sowerbyi from the Fulra Limestone. 1: FUL3–26, 2: FUL3–32, 3: FUL6–12, 4: FUL6–
14, 5: FUL8–57, 6: FUL6–44, 7: FUL13–44, 8: FUL13–42, 9: FUL3–31.

excentrilepidine-type configuration. O. haynesi, also Genus Asterocyclina Gümbel, 1870

recorded from Turkey (Özcan et al., 2010) and northern Type species Asterocyclina stellata (d’Archiac,
margin of Africa (Ben Ismail-Lattrache et al., 2014) but 1846)
not known from Europe, might be immigrant from the Asterocyclina alticostata (Nuttall, 1926)
Indian subcontinent because of the expansion of the (Figures 9(L), 18)
Eastern Tethyan fauna during Middle Eocene Climatic 1926 Actinocyclina alticostata sp. nov., Nuttall, p. 151,
Optimum (MECO). pl. 8, Figs. 6−8.
156  E. ÖZCAN ET AL.

Figure 17. Equatorial and axial sections of Orbitoclypeus haynesi from the Fulra Limestone. 5–6, microspheric; others, megalospheric.1:
FUL8–55, 2: FUL13–30, 3: FUL12–44, 4: FUL8–17, 5–6: FUL13–57, 7: FUL9–30.

Figure 18. Equatorial and axial sections of A. alticostata ex. interc. alticostata-cuvillieri (2–5) and A. alticostata ex. interc. cuvillieri-
alticostata (1) from the Fulra Limestone. 1: FUL13–9, 2: FUL12–11, 3: FUL12–5, 4: FUL6–24, 5: FUL6–30.

Diagnosis. Asterocyclina alticostata is a star-shaped isolepidine-type embryon, very few, very wide and mod-
species usually with five to ten rays and ‘chudeaui’ erately low, ‘alticostata’ type adauxiliary chamberlets and
type rosette. It has a medium-sized to relatively large also wide and moderately high equatorial chamberlets

arranged into asteroidal annuli with ‘strophiolata’ or
‘varians’ type growth pattern. This species includes four
subspecies as; A. a. gallica Less, 1987 (Dmean < 275 μm);
A. a. cuvillieri (Neumann, 1958) (Dmean = 275–350 μm);
A. a. alticostata (Nuttall, 1926) (Dmean = 350–450 μm);
A. a. danubica Less, 1987 (Dmean > 450 μm).
Remarks. Nuttall (1926) erected this species from Kutch
and noted the presence of about eight to twelve fairly
wide prominent ribs in twelve specimens. He assigned
these ribbed specimens to Actinocyclina (Gümbel) based
on numerous ribs. We have collected a large number of
specimens from the Fulra Limestone while this species
is extremely rare in the Drazinda Formation. The num-
ber of ribs counted in fifty-four specimens from the
Fulra Limestone is given in Table 2. This suggests that A.
alticostata commonly develops more than five ribs, the
maximum number being ten ribs in our material. Some
of these ribs develop only in the adult stage of the test
and they look like as if they are ‘secondary ribs’ on the
test surface. The consistent occurrence of more than
five ribs in this species in Indian Subcontinent as well as
lower Bartonian Reineche Limestone in North Africa and
coeval deposits in Turkey and Europe (Neumann, 1958;
Köhler, 1967; Less, 1987; Less et al., 2011; Zakrevskaya et
al., 2011; Ben Ismail-Lattrache et al., 2014) also points that
this is not an ecological feature, but a specific character.
The same feature is also observed in A. schweighauseri,
a rare species recorded in the Cuisian and lower-middle
Lutetian in Europe and Israel (Less, 1987, 1998). According
to Less (1987), always five rays start from the embryon
of A. alticostata with further development of other rays
in the adult stage (up to 10 rays), and many ribs start
directly from the embryon in A. schweighauseri. We think
that this is not a valid criterion for the distinction of these
species since in some specimens of A. alticostata more
than five rays may start to develop from the embryon
(Figure 18.1). In Fulra Limestone, this species is repre-
sented by the transitional developmental stages of A.
alticostata cuvillieri and A. alticostata alticostata (Table 1).
Asterocyclina sireli Özcan & Less, 2006
(Figures 9(M), 19)
2006 Asterocyclina sireli n. sp., Özcan et al., p. 506−507,
pl. 3, Fig. 32; pl. 4, Figs. 1−3; pl. 5, Figs. 1−5; text-Figure 12.
Diagnosis. Medium to large, flat forms with mostly
four radial ribs and ‘marthae’ type rosette. The embryon
Table 2. The variation in the number of ribs in A. alticostata
(Nuttall) and A. sireli Özcan & Less.
Number of ribs and corresponding
number of specimens
Number
Species Sample of spec. 4 5 6 7 8 9
A. alticostata FUL.13 34 – 9 9 6 7 2
FUL.6 20 – 3 6 7 3 –
A. sireli FUL.13 51 50 1 subsequently improved in Western Tethys as to include
FUL.6 29 29 – twenty-three species in SBZ 17 (Özcan et al., 2006,
RN.B.8 18 18 –
ALM.6 25 22 3 2010; Less et al., 2011). Because of the recalibration of
Note: The data from the type level of A. sireli (sample ALM.6) from Turkey
are also incorporated.
GEODINAMICA ACTA  157
is small, iso- to nephrolepidine. The deuteroconchal wall
corresponding to the position of the successive stage of
the developing rib is mostly depressed. The adauxiliary
chamberlets are few (2–4) in number, low and moder-
ately wide. The equatorial annuli are arranged usually
in four rays.
Remarks. This species was first introduced from the
Upper Lutetian of the Sivas Basin (Turkey) for aster-
ocyclinid specimens displaying mostly four ribs and
an embryon different from contemporaneous aster-
ocyclinids in Western Tethys (Özcan, Less, Báldi-Beke,
Kollányi, & Kertesz, 2006). It was later recorded from the
Bartonian Reineche Limestone in Tunisia and the Fulra
Limestone in India (Ben İsmail-Lattrache et al., 2014). In
its type material (sample ALM.6) from Sivas, twenty-two
specimens out of twenty-five have four ribs (Table 2).
In lower Bartonian Reineche Limestone in Tunisia, only
three specimens out of twenty-three specimens have five
ribs. In the Fulra Limestone, only one out of seventy-nine
specimens has five ribs and in the Drazinda Formation
all specimens are with four ribs. The consistent occur-
rence of four ribs and the wide geographic range of this
feature support that A. sireli is essentially a four-ribbed
species. In contrast to its common occurrence in the Fulra
Limestone, A. sireli is very rare in the Drazinda Formation.
Asterocyclina stellata (d’Archiac, 1846)
(Figure 19)
Diagnosis. Asterocyclina stellata is a star-shaped form
usually with five rays and ‘marthae’ type rosette. It has a
small semi-iso- to nephrolepidine-type embryon, few,
wide and low, ‘stellata’ type adauxiliary chamberlets and
also narrow and low equatorial chamberlets arranged
into asteroidal annuli with ‘strophiolata’ type growth
pattern. This species includes four subspecies in Western
Tethys: A. s. adourensis Less, 1987 (Dmean < 150 μm); A. s.
stellata (d’Archiac, 1846) (Dmean = 150–190 μm); A. s. stel-
laris (Brünner, 1848 in Rütimeyer, 1850) (Dmean = 190–
240 μm); A. s. buekkensis Less, 1987 (Dmean > 240 μm).
Remarks. A. stellata, recorded only from the Priabonian
of Assam as A. matanzensis Cole by Samanta (1965) from
the Indian subcontinent, occurs sporadically in the Fulra
Limestone. A subspecies designation is not possible
because of the rare specimens in the studied material.
7. A synthesis of orthophragminids
characterizing SBZ 17 in Tethys
The orthophragminids occurring in SBZ 17 in the first
scheme of shallow benthic zonation (SBZ) in Tethys
included thirteen species and Discocyclina pulcra bacon-
10 ica was marked as the key orthophragminid species
1 for this zone (Serra-Kiel et al., 1998). This scheme was
1
orthophragminid zones across the Lutetian-Bartonian
boundary with respect to the boundaries of SBZ 17,
158  E. ÖZCAN ET AL.
Figure 19. Equatorial and axial sections of A. sireli (1–11) and A. stellata (12–14) from the Fulra Limestone. 1: FUL13–71, 2: FUL13–152,
3: FUL13–92, 4: FUL6–66, 5: FUL6–107, 6: FUL6–135, 7: FUL8–2, 8: FUL6–138, 9: FUL6–136, 10: FUL8–1, 11: FUL8–3, 12: FUL8–20,
13–14: FUL13–153.
distribution of these taxa has also changed with respect
to SBZ zonation (Özcan et al., 2006). In updated scheme,
stratigraphic range of no any subspecies corresponds
to the full range of this zone. With the integration of six
species confined to Indian subcontinent (species shown
by yellow colour in Figure 20) and two species common
to Indian subcontinent, Turkey and North Africa (A. sireli
and O. haynesi), a total of twenty-nine species are now
recorded in SBZ 17 in Tethys. In Indian subcontinent, this
zone is characterized by the occurrence of fifteen spe-
cies; Discocylina dispansa, Discocyclina pseudodispansa,
Discocyclina kutchensis, Discocyclina pratti, Discocyclina
praeomphalus, Discocyclina augustae, Discocyclina dis-
cus, Discocyclina sulaimanensis, Discocyclina rakhinal-
aensis, Discocyclina zindapirensis, Discocyclina nandori,
Orbitoclypeus haynesi, Asterocyclina sireli, Asterocyclina
stellata, and Asterocyclina alticostata. Among these only
nine species (species shown by green colour in Figure 20)
are common in Western Tethys and Indo-Pakistan region.
As opposed to Western Tethyan orthophragminids,
the orthophragminid fauna in Eastern Tethys is domi-
nantly characterized by genus Discocyclina (Figure 21).
The genus Orbitoclypeus, represented by a single species,
O. haynesi, was only recorded from the Fulra Limestone
and it is not present in the Sulaiman Range. The true
Nemkovella does not occur in both regions. Nemkovella
daguini, which displays a different peri-embryonic cham-
ber arrangement than the true nemkovellids, occurs
only in the Bartonian deposits of Meghalaya, India
(unpublished data of E. Özcan). At present it is the only
nemkovellid species in Eastern Tethys, where generic
affiliation is also not certain. Asterocyclina sireli presents
a wide geographic distribution in Tethys, from Turkey
and Tunisia to the Indian subcontinent and probably
 GEODINAMICA ACTA  159

Figure 20. Updated scheme of orthophragminids showing the occurrence of twenty-nine species (lineages) in SBZ 17 (Bartonian)
in Tethys. Yellow color shows those taxa confined to Indian subcontinent, green color shows taxa common to Turkey and Indian
subcontinent and blue color shows taxa common to Turkey and Europe. Orbitoclypeus sp. 1 is a new orbitoclypeid from Baskil, Turkey,
not yet formally described.

Figure 21. Orthophragminid species diversity in some Western to Eastern Tethys deposits corresponding to SBZ 17. K: Kutch, SR:
Sulaiman Range, Pakistan, TUN: Tunisia, TUR: Turkey, HU: Hungary. The section in Hungary is from Dudar (sample DUDAR in Less,
1998). The combined sections from Turkey correspond to Gizliliman (Gizliliman section in Özcan et al., 2011), Çevirme (Çevirme
section in Gül et al., 2012) and Baskil (Keçili section in Özcan et al., 2006). The section in Tunisia is from Damouss Quarry in Cap Bon
Peninsula (Ben İsmail-Lattrache et al., 2014).

its range extends at least to the western Pacific region.
According to Less (1987), the asterocyclinids in Western
Pacific domain are characterized by the dominance of
four-ribbed specimens. Less (1987) considers that these
four-ribbed asterocyclinids are of ‘Caribbean- Pacific
affinity. Due to lack of detailed stratigraphic and taxo-
nomic works from this area, a reliable correlation of A.
sireli with Pacific Asterocyclina is presently not possible. It
is highly probable that this species has migrated to Indian
subcontinent and Western Tethys during the Middle
Eocene Climatic Optimum that resulted in the expan-
sion of Pacific fauna to higher latitudes in Bartonian. A
similar migration scenario is also proposed here for O.
haynesi, which occurs abundantly in Fulra Limestone, in
North Africa and Turkey but not recorded in European
platforms.
8. Conclusions
(1) The Bartonian orthophragminids characteriz-
ing the SBZ 17 from the Fulra Limestone belong
to Tethyan genera Discocyclina, Asterocyclina
and Orbitoclypeus. Among a total of eleven
species, seven of them belong to Discocyclina,
160  E. ÖZCAN ET AL.
one to Orbitoclypeus and three to Asterocyclina.
Nemkovella does not occur in Indian subcon-
tinent. Taking into account the twelve spe-
cies from the Drazinda Formation, some are
common to the Fulra Limestone, a total of fif-
teen species [Discocylina dispansa (Sowerby),
Discocyclina pseudodispansa Özcan, Ali & Yücel,
Discocyclina kutchensis Özcan & Saraswati,
Discocyclina pratti (Michelin), Discocyclina
praeomphalus Samanta & Lahiri, Discocyclina
augustae van der Weijden, Discocyclina discus
(Rütimeyer), Discocyclina sulaimanensis Özcan,
Ali & Hanif, Discocyclina rakhinalaensis Özcan,
Ali & Yücel, Discocyclina zindapirensis Özcan, Ali
& Yücel, Discocyclina nandori Less, Orbitoclypeus
haynesi (Samanta & Lahiri), Asterocyclina sireli
Özcan & Less, Asterocyclina stellata (d’Archiac),
and Asterocyclina alticostata (Nuttall)] are rec-
ognized from the Indian subcontinent. Six of
them are confined only to Indo-Pakistan region
(Figure 20). Two species, A. sireli and O. hayn-
esi, common in the Fulra Limestone and also
occurring in North Africa (Tunisia) and Turkey
but not in Europe, appear to have migrated to
Western Tethys during Middle Eocene Climatic
Optimum (MECO).
(2) In addition to ribs, bulges, rounded surface
structures corresponding to thickening of the
lateral layers, are recognized in D. kutchensis,
occurring abundantly in the Fulra Limestone as
well as in the Drazinda Formation in Pakistan.
This species is associated with D. nandori in the
Drazinda Formation, while the latter species has
not been recorded from the Fulra Limestone.
(3) We recorded two common asterocyclinid spe-
cies, A sireli and A. alticostata, and scanty A.
stellata from the Fulra Limestone. A sireli and
A. alticostata display the consistent character-
istic features by developing four and many ribs
respectively in the Tethys. This is a feature not
observed in most of the asterocyclinid species,
predominantly developing five ribs. Except for
the above taxa and A. schweighauseri, a Cuisian
to middle Lutetian species, other common
asterocyclinid species, such as A. stellata and A.
stella consistently develop mainly five ribs. Our
specific examples from A. sireli and A. alticostata
from geographically distant locations of Tethys
suggest that certain asterocyclinid taxa consist-
ently develop a specific number of ribs. Thus,
the traditional concept of Asterocyclina with
‘five-ribs’ is to be abandoned.
(4) We observed that the shape of the test (e.g.
saddle-shaped vs flat tests) is a consistent char-
acter for each species. The saddle-shape test,
for instance, is a specific feature of D. discus
and D. praeomphalus. D. dispansa has always
a flat test as D. augustae. This further requires
re-evaluation of some D. dispansa populations
in Western Tethys, which include both saddle
and flat tests, interpreted to be ecological vari-
ations of the same species.
Acknowledgements
We thank Gyorgy Less (Miskolc) and an anonymous reviewer
for helpful suggestions in the review of the manuscript.
Fieldwork in India was possible due to funding by TUBA
(Turkish Academy of Sciences) and INSA (Indian National
Science Academy) Exchange of Scientists to E. Özcan and
funding for laboratory work to P. K. Saraswati.
Disclosure statement
No potential conflict of interest was reported by the authors.
Funding
This work was supported by TUBA (Turkish Academy of
Sciences); and INSA (Indian National Science Academy).
ORCID
Pratul Kumar Saraswati http://orcid.org/0000-0001-9115-
8951
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