Afzal Et Al. 2011, Lethaia, Vol. 44

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EvolutionofPaleocenetoEarlyEocenelargerbenthicforaminiferassemblagesoftheIndusBasin,PakistanARTICLEinLETHAIANOVEMBER2010ImpactFactor:2.19DOI:10.1111/j.1502-3931.2010.00247.x

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MarkWilliamsUniversityofLeicester183PUBLICATIONS2,160CITATIONS

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MelanieJ.LengBritishGeologicalSurvey375PUBLICATIONS5,857CITATIONS

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Availablefrom:MarkWilliamsRetrievedon:27August2015

Evolution of Paleocene to Early Eocene larger benthicforaminifer assemblages of the Indus Basin, Pakistan

JAWAD AFZAL, MARK WILLIAMS, MELANIE J. LENG, RICHARD J. ALDRIDGE AND

MICHAEL H. STEPHENSON

Afzal, J., Williams, M., Leng, M.J., Aldridge, R.J. & Stephenson, M.H. 2011: Evolution ofPaleocene to Early Eocene larger benthic foraminifer assemblages of the Indus Basin,Pakistan. Lethaia, Vol. 44, pp. 299320.

The PaleoceneEarly Eocene carbonate successions of the Indus Basin in Pakistanformed on the northwestern continental shelf margin of the Indian Plate in the easternTethys Ocean. Based on larger benthic foraminifera (LBF), eight Tethyan foraminiferalbiozones (SBZ1SBZ8) spanning the Paleocene to Early Eocene interval are identified.The base of the Eocene is identified by the first appearance of Alveolina sp. Other strati-graphically important LBFs that are characteristic of the earliest Eocene are Ranikothalianuttalli, Discocyclina dispansa and Assilina dandotica. Stable isotope analysis throughthe PaleoceneEocene (PE) boundary interval identifies more positive d13C values forthe Late Paleocene (+3.4& to +3.0&) and lower values (+2.7& to +1.6&) for the ear-liest Eocene. However, there is insufficient sampling resolution to identify the maxi-mum negative d13C excursion of the PaleoceneEocene Thermal Maximum. DuringLate Paleocene times LBF assemblages in the Indus Basin were taxonomically close tothose of west Tethys, facilitating biostratigraphic correlation. However, this faunal con-tinuity is lost at the PE boundary and the earliest Eocene succession lacks typical westTethys Nummulites, while Alveolina are rare; LBFs such as Miscellanea and Ranikothaliacontinue to dominate in the Indus Basin. The absence of Nummulites from the earliestEocene of Pakistan and rarity of Alveolina, elsewhere used as the prime marker for thebase of the Eocene, may imply biogeographical barriers between east and west Tethys,perhaps caused by the initial stages of India-Asia collision. Later, at the level of theEocene SBZ8 Biozone, faunal links were re-established and many foraminifera withwest Tethys affinities appeared in east Tethys, suggesting the barriers to migrationceased. h Biostratigraphy, Eocene, India-Asia collision, larger benthic foraminifera, palae-oecology, Paleocene.

Jawad Afzal [[email protected]], Department of Geology, University of Leicester, Leices-ter, LE1 7RH, UK and National Centre of Excellence in Geology, University of Peshawar,Peshawar, Khyber Pakhtunkhwa, Pakistan; Mark Williams [[email protected]], Depart-ment of Geology, University of Leicester, Leicester, LE1 7RH, UK and British Geological Sur-vey, Keyworth, Nottingham, NG12 5GG, UK; Melanie J. Leng [[email protected]], NERCIsotope Geosciences Laboratory, British Geological Survey, Keyworth, Nottingham, NG125GG, UK; Richard J. Aldridge [[email protected]], Department of Geology, University ofLeicester, Leicester, LE1 7RH, UK; Michael H. Stephenson [[email protected]], British Geo-logical Survey, Keyworth, Nottingham, NG12 5GG, UK; manuscript received on25 03 2010; manuscript accepted on 26 07 2010.

The PaleoceneEarly Eocene shallow marine carbon-ates of the Lockhart and Dungan formations in theIndus Basin of Pakistan contain stratigraphicallyimportant larger benthic foraminifera (LBF), whichhave been used for local (Indian subcontinent) andregional (especially Tethyan) biostratigraphy (e.g.Hottinger 1960, 1971; Adams 1970; Kureshy 1978;Schaub 1981; Serra-Kiel et al. 1998; Jauhri & Agarwal2001; Green et al. 2008; Afzal et al. 2009 and refer-ences therein). The most recently establishedPaleoceneEarly Eocene LBF biostratigraphic schemesfor the Indus Basin in Pakistan include the TertiaryLetter Stage biozones (Adams 1970), the TertiaryLBF biozones (e.g. Kureshy 1978; Butt 1991), theEarly Eocene alveolinid biozones of the Salt Range

(e.g. Sameeni & Butt 2004) and the LBF assemblagesof Weiss (1993). Most of these schemes are fragmen-tary and difficult to correlate with standard biostrati-graphic schemes. Other LBF studies are mostlycoarse-resolution or deal only with phylogenetic lin-eages or taxonomies (see Afzal et al. 2009 for a sum-mary of problems). The precise biostratigraphicposition of the PaleoceneEocene boundary (PE) inthe shallow marine successions of this region was,prior to the present study, ambiguous.

The Tethyan shallow marine benthic foraminiferabiozonation (SBZ) scheme of Serra-Kiel et al. (1998)provides a basis for correlation of carbonate platformsand pelagic successions of the Paleocene to Eoceneby establishing 20 LBF biozones (SBZ1SBZ20),

DOI 10.1111/j.1502-3931.2010.00247.x 2010 The Authors, Lethaia 2010 The Lethaia Foundation

correlated with the planktonic foraminifera P1P18biozones of Berggren et al. (1995) and nannofossilbiozones (e.g. the NP1NZ21 biozones of Martini1971). This SBZ scheme is based on the earlier LBFbiostratigraphic studies of Hottinger (1960), Hottin-ger et al. (1964), Schaub (1981) and Hottinger & Dro-bne (1988), and is applicable to the Tethyan shallowmarine realm as far south as Somalia and southeast tothe Indian subcontinent (Hottinger 1971; Pignatti1994; Jauhri 1998; Scheibner & Speijer 2008a). Innorthern India and Pakistan (east Tethys), the LBFs ofthe Late Paleoceneearliest Eocene show unusual fau-nal associations, where the typical latest PaleoceneMiscellanea and Ranikothalia co-occur with typicalearliest Eocene Alveolina, while Nummulites, a markerof the earliest Eocene in west Tethys, occurs only later(Hottinger 1971; Jauhri 1998; Akhtar & Butt 2000;Jauhri et al. 2006; Afzal et al. 2009; Tewari et al. 2010).Earlier studies on LBFs in Pakistan either did not rec-ognize these faunal differences between east and westTethys at the PE boundary or paid little attention tothis interval. Therefore, clarification of the formal suc-cessions through the PE boundary interval in thisregion is vital for regional and international correla-tion of important geological events (e.g. PaleoceneEocene Thermal Maximum, PETM and India-Asiacollision tectonics). It is now apparent that, despitesome faunal differences (e.g. in Hottinger 1971;Scheibner & Speijer 2008a; present study), the LBFs ofthe Indus Basin in Pakistan contain many Tethyan ele-ments (e.g. Hottinger 1960; Hottinger & Drobne1988; Serra-Kiel et al. 1998; Scheibner et al. 2005;Scheibner & Speijer 2008a, 2009). Consequently, theycan be used to establish a refined biostratigraphicbiozonation correlated to the standard Tethyan SBZscheme (e.g. Serra-Kiel et al. 1998; Scheibner et al.2005; Scheibner & Speijer 2009). Here, we aim to: (1)establish a LBF biostratigraphic scheme for the Paleo-ceneEarly Eocene carbonate platform succession inthe Indus Basin of Pakistan; (2) precisely resolve thePaleocene Eocene boundary in these successions; (3)discuss the biostratigraphic significance of LBFs thatshow variable stratigraphic ranges across thePaleoceneEocene boundary interval in this regionincluding species of Miscellanea, Ranikothalia andNummulites, and (4) discuss LBF faunal differences inthe Tethyan provinces (east and west) and the roles ofIndia-Asia collision and the PETM in generating thesedifferences.

Geological setting

The geological setting of the Greater Indus Basin ofPakistan is discussed in the study by Afzal et al.

(2009; herein see Fig. 1). The tectonic history andstratigraphical framework of the region are stronglyinfluenced by the collision of the Indo-Pakistan andAsian plates, and estimates of the timing of the initialcollision vary from 65 to 45 Ma (Beck et al. 1995;Rowley 1996; Hodges 2000; Afzal et al. 2009). A hia-tus equivalent to planktonic foraminiferal biozonesupper P5P7 in the Indus Basin has been related tothe early collision of India with Asia (see Afzal et al.2009) (Fig. 2). Carbonate platform deposition wasinterrupted by this tectonic event, for example, thetermination of platform sedimentation in the UpperIndus Basin during the latest Paleocene (see Afzalet al. 2009 for summary). However, in parts of theLower Indus Basin, platform sedimentation contin-ued through the PaleoceneEarly Eocene interval,represented, for example in the Muree Brewery,Hanna Lake and Zranda sections of the present study(Fig. 1).

Stratigraphy

The earliest marine Cenozoic sedimentation in thebasin seems to have commenced with the Palaeogenetransgression depositing continental near-shore faciesto shallow marine-deltaic facies of the Hangu

Fig. 1. Map of the Upper Indus Basin and parts of the LowerIndus Basin showing distribution of PaleoceneEocene sedimen-tary rocks and key stratigraphical sections (modified after Eames1952). The Lower and Upper Indus Basins collectively form theGreater Indus Basin.

300 Afzal et al. LETHAIA 44 (2011)

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LETHAIA 44 (2011) Paleogene larger benthic foraminifera 301

Formation in the Upper Indus Basin and mixed car-bonate siliciclastic to open-marine facies of the Dun-gan Formation in the Lower Indus Basin (Afzal et al.2009 and references therein). This marine floodingwas succeeded by widespread carbonate platformdeposition represented by the Lockhart Formation inthe Upper Indus Basin and the upper Dungan For-mation in parts of the Lower Indus Basin (Afzalet al. 2009) (Fig. 2). By contrast, the planktonic fora-minifer-bearing ( = biozones P3P6) shales of theDungan Formation in the northwestern SulaimanRange (e.g. Rakhi Nala and Zinda Pir areas) weredeposited in an open marine environment (Afzalet al. 2009).

The carbonate platform was buried by deep andshallow marine clastics of the Patala Formation( = biozones P4cP6) in most of the Upper IndusBasin and by shales of the Ghazij Formation ( = bioz-ones P7P10) in parts of the Lower Indus Basin (Afzalet al. 2009) (Fig. 2). However, carbonate sedimenta-tion in the eastern parts of the Lower Indus Basin (e.g.Balochistan Province) continued through the LatePaleocene to Early Eocene (i.e. Dungan Formation)(Afzal et al. 2009; Kassi et al. 2009). During the Ypre-sianearly Lutetian (biozones P7P10) in the UpperIndus Basin, shallow marine carbonates of the Margal-a Hill, Nammal and Shekhan formations with themudstone shale-rich Panoba Formation were depos-ited (Fig. 2). The higher parts of the successioninclude evaporites (the Bahadar Khel Salt-Jatta Gyp-sum) and, finally, the continental red bed sandstonemammal-bearing Kuldana Formation in the northernpart and carbonates and shales of the Sakesar Forma-tion in the southern part of the Upper Indus Basin(Afzal et al. 2009).

The Sakesar Formation in the Salt and SurgharRanges is overlain by Miocene-Recent terrestrial sedi-ments derived from the Himalaya, marking the clo-sure of the Tethys Ocean in the southeastern UpperIndus Basin (Afzal et al. 2009). The late Lutetian-Priabonian is represented by the upper KirtharFormation in the Lower Indus Basin and by the cor-relative uppermost Kohat Formation in the UpperIndus Basin. This followed closure of the TethysOcean in the north to northwestern parts of theUpper Indus Basin (e.g. Kohat area, Kala Chitta andHazara Ranges). The gradual retreat of the TethysOcean continued south-southwest through late Lute-tian to Bartonian time and it finally closed in the Pri-abonian (Biozone P15; Afzal et al. 2009). Oligocenemarine sedimentation was restricted to the south ofthe Lower Indus Basin, while the rest of the GreaterIndus Basin in Pakistan remained non-depositionallowland until the formation of Neogene molasse(Afzal et al. 2009).

Materials, methods and studiedsections

Materials and methods

We use the terminology of Afzal et al. (2009) for thePaleoceneEocene lithostratigraphy and that of Shah(1977) for the Palaeozoic and Mesozoic. We have col-lected and logged the Late Paleocene Lockhart Forma-tion in one outcrop section, and one petroleum wellcore in the Upper Indus Basin (Fig. 1), and the Paleo-ceneEarly Eocene Dungan Formation in four out-crop sections of the Lower Indus Basin (Fig. 1). Some199 limestone samples were collected including 179from five outcrop sections. We have prepared stan-dard limestone thin sections of 2.5 5 cm and2.5 7 cm dimensions for petrographic microscopy.The identification of LBFs in these thin sections isbased on foraminifera that have not been re-worked(based on preservation and palaeoecological context)and using well-oriented sections (of LBFs) only, com-plemented by study of limestone slabs. The LBFs werephotographed in plane-polarized light with a digitalcamera on a Nikon petrographic microscope in theDepartment of Geology, University of Leicester. Dis-crete specimens of LBFs could not be collected as thesediments are indurated. For biostratigraphic determi-nation, the SBZ scheme of Serra-Kiel et al. (1998),with amendments for the placement of the PEboundary at the boundary between biozones SBZ4and SBZ5 (see Scheibner & Speijer 2009), has beenadopted. The boundary between biozones SBZ4 andSBZ5 coincides with the base of the Ilerdian stage(Pujalte et al. 2009a). To allow precise correlation andavoid confusion, we have adopted the standard SBZscheme names (e.g. SBZ1SBZ8). The biozonal mar-ker species of both local and Tethyan realms havebeen used to identify these biozones.

For stable isotopes (d13C), bulk carbonates fromthree sections (Zranda, Muree Brewery and HannaLake) were selected. Some 58 samples were analysedcovering the Late PaleoceneEarly Eocene interval.We used only well-preserved limestone for stable iso-tope analysis that appeared not to be re-worked ordiagenetically altered (based on the type, preservation,fragmentation and ecological mode of containedbiota). The sample material was ground in agate andthe equivalent of 10 mg of carbonate was reacted withanhydrous phosphoric acid in vacuo overnight at aconstant 25C. The CO2 liberated was separated fromwater vapour under vacuum and collected for analysis.Measurements were made using VG Optima massspectrometer at the NERC Isotope Geosciences Labo-ratory, Keyworth, Nottingham, UK. Overall analytical


reproducibility for these samples is normally betterthan 0.2& for d13C. Isotope values (d13C) arereported as per mil (&) deviations of the isotopicratios (13C 12C) calculated to the VPDB scale using awithin-run laboratory standard calibrated against NBSstandards.

Studied sections in the Upper Indus Basin

Kotal Pass section. The Kotal Pass section is situatedin the northernmost part (333801N and712821E) of the Upper Indus Basin in the KohatHill Range of northern Pakistan (Fig. 1). The sectionexposes a thick succession of Jurassic to Paleocenerocks, which constitutes the hanging wall sequence ofthe Main Boundary Thrust (MBT) along which Juras-sic rocks are thrust southward over the EoceneMiocene succession of the Kohat Foreland Basin(Khan et al. 1990). In the Kotal Pass section, 78samples at 1- to 2-m interval spacing from the LatePaleocene Lockhart Formation have been collected(Fig. 3). The Lockhart Formation is 144 m thick andis composed of limestone rich in dasycladaceans andLBFs (mainly smaller rotaliids, agglutinated forms,miliolids, rare miscellanids etc.) producing wackstoneand packstone with rare grainstone and patch-reefboundstone textures (Fig. 3).

Shakardara Well-1. The Shakardara Well-1 wasdrilled in the Shakardara-Nandrakki Exploration leaseof the Oil and Gas Development Company Limited,Pakistan (OGDCL). The well is approximately 70 kmsouth of Kohat (331324N and 712939E), KhyberPakhtunkhwa Province, Pakistan (Fig. 1). The wellwas drilled to 4548 m depth, with Cretaceous to Plio-cene sediments recorded. Two cores (Core-1 andCore-2) from the Late PaleoceneEarly Eocene strati-graphic interval ( = Lockhart and Patala formations)were recovered. Core-1 from interval 4299.3 to4303.5 m, is dominantly composed of dark-grey pela-gic limestone rich in planktonic foraminifera. Thereare no PaleoceneEocene LBFs in Core-1, probablydue to the deep pelagic environments it represents(Fig. 3). However, interval 44684474 m of Core-2consists of dasycladacean- and LBF-rich packstonesand wackstones that have produced a number of age-diagnostic LBF taxa. Thirteen core samples (polishedthin sections) have been studied, which yielded anumber of marker species of the SBZ3SBZ4 biozones(Fig. 3).

Studied sections in the Lower Indus Basin

Mughal Kot section. This section is located near theMughal Kot Post (312629N and 700507E),

155 km west of Dera Ismail Khan city (Fig. 1). TheMughal Kot section represents the northeastern Sulai-man Range in the Lower Indus Basin (Fig. 1). Theexposed deposits range from marine Jurassic to conti-nental Recent. The Paleocene Dungan Formation is200 m thick, composed of a mixed carbonate-silici-clastic lower unit succeeded by a LBF- to coral-algalrich rudstone, bindstone, grainstone and packstoneupper unit (Fig. 4). A total of 21 surface rock samplesat 5 and 10 m intervals were collected (Fig. 4). Thelower unit contains rare LBFs. When present, forami-nifera of this lower unit are characterized by smallerrotaliids of the SBZ1 Biozone. However, the upperunit yields a number of age-diagnostic LBFs and cor-alline algae of the SBZ2SBZ3 biozones (Fig. 4).

Zranda section. The Zranda section is located nearZranda village (302827N and 673707E), 25 kmnortheast of Kach, Balochistan (Fig. 1). The litho-stratigraphic succession is similar to that of theKachZiarat valley, comprising the Late Cretaceousvolcanoclastic-sedimentary Bibai Formation, LatePaleoceneearliest Eocene shallow marine carbonatesof the Dungan Formation and Early Eocene openmarine clastics of the Ghazij Formation (Kassi et al.2009). In the Zranda section, the Dungan Formationis 222 m thick, consisting of LBF-rich wackstone,packstone and grainstone containing abundant age-diagnostic taxa of the SBZ3SBZ6 biozones (Fig. 5). Atotal of 39 rock samples at 23 m interval spacingwere collected (Fig. 5).

Muree Brewery section. The well exposed MureeBrewery section is situated 8 km west (301122Nand 665644E) of Quetta city in Balochistan Prov-ince (Fig. 1). The section represents the western Sulai-man Range of the Lower Indus Basin and exposes athick succession of Late Cretaceous (Parh, HannaLake Limestone and Fort Munro formations) to EarlyEocene (Dungan and Ghazij formations) marinedeposits (Kassi et al. 2009). In the Muree Brewery sec-tion, the Dungan Formation is 19.2 m thick, and ismainly composed of LBF-rich grainstone and pack-stone containing stratigraphically significant markersof the Early Eocene ( = biozones SBZ5SBZ8)(Fig. 6). A total of 20 rock samples at intervals of

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northeast of Quetta city, Balochistan (Fig. 1). TheHanna Lake section has a similar stratigraphic settingto the nearby Muree Brewery section (see Kassi et al.2009). However, the thickness of the Dungan Forma-tion here is 22 m and it is composed largely of LBF-rich packstone, wackstone and grainstone with variedamounts of coralline algae and planktonic foraminif-era (Fig. 7). A total of 21 rock samples at intervals of1 m spacing have been collected. A number of LBFmarker species of the SBZ5SBZ8 biozones are repre-sented.

Stable isotope stratigraphy

The prominent geochemical feature in the lower Pal-aeogene stratigraphic record is a negative (of )2.5 to)4&) carbon isotope (d13C) excursion (CIE) (Zachoset al. 2001). This excursion reflects a major perturba-tion of the global carbon cycle resulting from a dra-matic global warming event, the PETM (e.g. Zachoset al. 2001). The PETM CIE has been accepted as themarker criterion for the Global Stratotype Section andPoint (GSSP) for the PaleoceneEocene (PE) bound-ary in Egypt (Aubry & Ouda 2003; Aubry et al. 2007).The CIE is detectible in marine and terrestrial recordsworld wide (e.g. Bowen et al. 2001; Zachos et al. 2001;Magioncalda et al. 2004; Thiry et al. 2006). In theGSSP section of Dababiya, Egypt, a sharp decrease ind13Corg from )24& to )26.5&marks the onset of theCIE which is followed by a gradual decrease until itreaches its lowest values (i.e. )27.5&) (Dupuis et al.2003; Aubry et al. 2007). The d13Corg values thenincrease progressively until pre-excursion values arereached (Dupuis et al. 2003). Carbon isotope recordsderived from carbonates (d13Ccarb) in the Dababiyasection and in nearby sections are of similar amplitude(e.g. Aubry et al. 1999; Dupuis et al. 2003).

Here, we describe for the first time a detailedd13Ccarb stratigraphy for the early Palaeogene of theIndus Basin. In the Zranda section, Late Paleocened13Ccarb values vary between +3.0& and +3.4&(Fig. 5). The CIE peak in our sections has not beendetected, probably because of the spacing of samples(ca. 23 m apart). A d13C spike in one sample (ZRD-11) just below the PE boundary in the Zranda sec-tion appears to be associated with fresh and brackishwater facies (Fig. 5). A decrease in d13Ccarb values tobetween 2.4& and +3.1& in the Zranda section andless positive values of +1.8& to +2.6& in the MureeBrewery section and +1.6& to +2.7& in the HannaLake section indicates a marked change in the earliestEocene (post-PETM CIE maximum) (Figs 57).In the succeeding Early Eocene succession, d13Ccarbvalues fluctuate between +2.1& and +1.2& in theMuree Brewery and Hanna Lake sections (Figs 6, 7).

The Late Paleocene d13Ccarb values of the IndusBasin are comparable with those recorded in coevalplatform carbonate sections (i.e. around +3.0&) inEgypt by Scheibner et al. (2005) and Scheibner &Speijer (2009). The lower d13Ccarb values from theEocene in the Indus Basin succession may reflectthe later stages of the global decrease of d13C duringthe earliest Eocene (Dupuis et al. 2003). The d13Ccarbvalues of the earliest Eocene as recorded in Egypt arearound +1.0& (e.g. Scheibner et al. 2005; Scheibner& Speijer 2009), which are lower than Indus Basin

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Fig. 4. Lithostratigraphy and biostratigraphic ranges of largerbenthic foraminifera of the Paleocene Dungan Formation inMughal Kot section, Lower Indus Basin, Pakistan. Index as forFigure 3.


d13Ccarb values (between +1.6& and +3.1&). How-ever, d13Ccarb values of the Late PaleoceneEarlyEocene limestones of the Indus Basin are similar tovalues measured on well-preserved marine planktonicand deep-benthic foraminiferal tests and bulk carbon-ate elsewhere in Tethys, probably because these lime-stones were indurated during early diagenesis andbecame closed systems with respect to carbonexchange (Schmitz et al. 1997, 2001).

LBF biostratigraphy

We recognize the PaleoceneEarly Eocene LBF bioz-ones SBZ1SBZ8. These biozones are identified on thebasis of one or more key markers and their first andlast occurrences (Figs 8, 9). A short summary of keyLBFs that delineate these biozones in our sections isgiven below and detailed species occurrences areshown in Figures 37.

The first occurrence of Laffitteina bibensis marks theSBZ1 Biozone. The boundary between the SBZ1 andSBZ2 biozones is not straightforward, owing to a lackof marker LBFs and the variable stratigraphic range ofLf. bibensis in the Indus Basin succession (Fig. 4).However, the first occurrence of Idalina sinjaricatogether with the coralline alga Distichoplax biserialissuggests the SBZ2 Biozone (Fig. 4). The SBZ3 Biozoneis clearly identified by the biostratigraphic ranges ofCoskinon rajkae, Fallotella alavensis, Vania anatolicaand Miscellanea juliettae (Figs 35). Other key LBFsfirst appearing within Biozone SBZ3 include Raniko-thalia sindensis, R. sahni, Lockhartia haimei, L. conditi,Lf. erki, Kathina selveri, Setia tibetica and Orbitosiphonpunjabensis. The boundary between biozones SBZ3and SBZ4 is marked by the appearance of Miscellaneameandrina, M. miscella and Discocyclina ranikotensis(Figs 3, 5). The appearance of Alveolina is taken as thebase of the earliest Eocene (biozones SBZ5 6, Hottin-ger 1960; Serra-Kiel et al. 1998; Scheibner & Speijer

Fig. 5. Lithostratigraphy, stable isotopes and biostratigraphic ranges of larger benthic foraminifera of the Late PaleoceneEarly Eocene Dun-gan Formation in the Zranda section, Lower Indus Basin, Pakistan. Index as for Figure 3.


2009; Fig. 5). In addition to Alveolina, the appearanceof other marker species including Assilina dandotica,Discocyclina dispansa and Ranikothalia nuttalli marksthe earliest Eocene (Fig. 5). The last occurrences of R.sindensis and R. sahni were recorded within biozonesSBZ5 6 (Figs 57). Opertorbitolites, Orbitoclypeus sp.,D. sella and Somalina first appear within the SBZ5 6 biozones (Figs 57).

Identification of Biozone SBZ7 has proved difficultdue to the stratigraphically late appearance of Num-mulites in the Indus Basin, the low diversity of thefauna through this interval, and the generally poorlyoriented sections of alveolinids and assilinids makingtaxonomic determination complex. Nevertheless, thepresence of a continuous carbonate succession throughthe SBZ6SBZ8 biozones in the Muree Brewery andHanna Lake sections indicates that the interval of theSBZ7 Biozone is an interregnum, demarcated by diag-nostic LBFs from the underlying and succeeding strata

(Figs 6, 7). Thus, the base of Biozone SBZ7 in oursections is taken immediately above the last occur-rences of R. nuttalli and A. dandotica (Figs 6, 7). Thelast occurrence of M. miscella is recorded within Bioz-one SBZ7. The first Nummulites in our sections are N.atacicus and N. globulus which are typical of BiozoneSBZ8 elsewhere in Tethys platform carbonate succes-sions (e.g. Serra-Kiel et al. 1998) (Figs 6, 7). The top ofBiozone SBZ8 cannot be identified due to succeedingopen-marine clastic sediments (inhospitable lithofaciesfor LBFs) of the Ghazij Formation in our sections.

Comparison of LBF ranges withthose in other regions of Tethys

Larger benthic foraminifer marker species used in thisstudy are index taxa described for the SBZ1SBZ8

Fig. 6. Lithostratigraphy, stable isotopes and biostratigraphic ranges of larger benthic foraminifera of the Early Eocene Dungan Formation inthe Muree Brewery section, Lower Indus Basin, Pakistan. Index as for Figure 3.


biozones by Serra-Kiel et al. (1998). LBFs which showranges different from those described by Serra-Kielet al. (1998) are evaluated based on their faunal asso-ciates in the Indus Basin (Figs 37) and on theiroccurrences elsewhere. Key LBFs which were notdescribed by Serra-Kiel et al. (1998), but which arecritical for biostratigraphic analysis in the Indus Basin,have been compared here with data from other Teth-yan carbonate platform successions.

Paleocene LBFs

According to Serra-Kiel et al. (1998), Lf. bibensis isrestricted to the basal Paleocene SBZ1 Biozone. How-ever, in the Mughal Kot section, its association with I.sinjarica and Selandian coralline algae (Aguirre et al.2007) suggests it also occurs within the SBZ2 Biozone.Similarly, longer biostratigraphic ranges for Lf. bibensishave been reported from the Paleocene of Iran (Bignot& Neumann 1991) and the Danian to Thanetian suc-cessions of NE Turkey (Sirel 1998; Inan et al. 2005).

Serra-Kiel et al. (1998) described I. sinjarica asranging from biozones SBZ3 to SBZ6. However, its

first appearance, together with the coralline alga D.biserialis that is typical from the Selandian onward(Aguirre et al. 2007), and the lack of biozonal markersfor the SBZ3 Biozone in the Mughal Kot section, sup-ports an early appearance of I. sinjarica in the SBZ2Biozone (Fig. 4). Drobne et al. (2002) also reportedthe first appearance of I. sinjarica within the SBZ2Biozone from the Indo-Pacific, Tethyan and Carib-bean regions.

According to Hottinger (1960), Serra-Kiel et al.(1998), Pignatti et al. (2008) and Scheibner & Speijer(2009), Glomalveolina primaeva is restricted to Bioz-one SBZ3. However, in the Zranda section, the associ-ation of G. primaeva with M. juliettae, Coskinon rajkaeand F. alavensis and later with M. miscella suggests italso ranges into Biozone SBZ4 (Fig. 5). Similarly, Ja-uhri & Agarwal (2001) reported G. primaeva in associ-ation with G. levis ( = Biozone SBZ4) in NE Indiaand White (1992) recorded a SBZ3SBZ6 range for G.primaeva in Oman.

In Tethyan platform successions (including Paki-stan and India), a biostratigraphic range of biozonesSBZ3SBZ4 has been recorded for L. haimei (e.g. Butt

Fig. 7. Lithostratigraphy, stable isotopes and biostratigraphic ranges of larger benthic foraminifera of the Early Eocene Dungan Formation inthe Hanna Lake section, Lower Indus Basin, Pakistan. Index as for Figure 3.


1991; Weiss 1993; Serra-Kiel et al. 1998; Akhtar &Butt 1999; Jauhri & Agarwal 2001; Mathur et al.2009). However, in the Zranda section, L. haimeitogether with L. conditi are associated with Alveolina,A. dandotica, D. dispansa, R. sindensis and R. nuttalliwhich suggest a stratigraphically higher occurrencewithin the SBZ5 6 biozones (Fig. 5). Similarly, Butter-lin & Fourcade (1989) reported longer biostrati-graphic ranges for L. haimei and L. conditi, showingtheir highest occurrences in the EarlyMiddle Eocene.We have not recorded L. conica and L. diversa in theEocene of the Indus Basin, but they frequently occurin the Late Paleocene (in biozones SBZ3 and SBZ4)(Figs 37). Similar biostratigraphical ranges for L. co-nica and L. diversa have been shown in Qatar (Smout1954), and for L. diversa in western Tethys (Butterlin& Fourcade 1989) and in Turkey (Sirel 1998). How-ever, various studies in the wider Tethyan region(including Pakistan) indicate that L. conica is notrestricted to the Paleocene and that this taxon extendsinto the Early Eocene (e.g. Butterlin & Fourcade 1989;Weiss 1993; Akhtar & Butt 1999).

Rotalia trochidiformis is a commonly occurring LBFin Tethys and ranges from the Paleocene to the EarlyEocene (biozones SBZ3SBZ8, and possibly intoSBZ9) (e.g. Qatar, Smout 1954; Pakistan, Weiss 1993;Akhtar & Butt 1999; NE Turkey, Inan et al. 2005; NTurkey, Ozgen-Erdem et al. 2005; Northern India,Mathur et al. 2009). In the Zranda section, the bio-stratigraphic ranges of Laffitteina erki and K. selveri areshown as biozones SBZ3SBZ4 based on their faunalassociation (Fig. 5). This is in accordance with occur-rences in Slovenia (Ozgen & Akyazi 2001) and Turkey(Inan et al. 2005; Ozgen-Erdem et al. 2005), where arange of biozones SBZ3SBZ4 for Lf. erki and K. sel-veri is indicated.

According to Serra-Kiel et al. (1998), R. sindensisfirst appeared at the base of Biozone SBZ4 in Tethys.However, in the Zranda and Kotal Pass sections andShakardara Well-1, the occurrence of R. sindensistogether with R. sahni in association with M. juliettae,L. haimei, C. rajkae and F. alavensis suggests an earlierfirst appearance within the SBZ3 Biozone (Figs 3, 5).Tosquella et al. (1998) found R. sindensis restricted tothe SBZ3 Biozone in the Pyrenean Basin, France. InOman, India and Pakistan, R. sindensis has beenreported from the Late Paleocene to earliest Eocene(Nagappa 1959a; Butt 1991; Racey 1995).

Ferrandez-Canadell (2002) described S. tibetica andO. punjabensis as endemic to the south-central Asianregion. In our sections, we have recorded these generaassociated with LBFs characteristic of the SBZ3 andSBZ4 biozones, for example in the Shakardara Well-1,Kotal Pass and Zranda sections (Figs 3, 5). This is inagreement with the findings of Ferrandez-Canadell

(2002), who reported these LBFs from the uppermostHangu, Lockhart and lower Patala formations in theSalt Range area of the Upper Indus Basin, whichaccording to Afzal et al. (2009) can be regarded asassignable to biozones SBZ3 and SBZ4.

Discocyclina ranikotensis first appears within theSBZ4 Biozone in our successions (Fig. 3). In Pakistanand India, other studies have also indicated that D.ranikotensis ranges from the Late Paleocene to EarlyEocene (e.g. Nagappa 1959b; Samanta 1969; Butt1991; Weiss 1993; Akhtar & Butt 1999, 2000). Thespecies appears to be endemic to the Indian region(east Tethys) as it has not been reported from westTethys carbonate platforms, where different lineagesof discocyclinids developed during the Late Paleocene(e.g. Samanta 1969; Adams 1970; Serra-Kiel et al.1998; Less et al. 2007).

Early Eocene LBFs

According to Serra-Kiel et al. (1998), A. dandotica isrestricted to the SBZ5 Biozone in the Tethyan realm.However, Tosquella et al. (1998) demonstrated thatA. dandotica extends to the SBZ6 Biozone. Ranikotha-lia nuttalli ranges from the SBZ5 to SBZ6 biozones inTethyan platform successions (e.g. Serra-Kiel et al.1998). Scheibner & Speijer (2009) recorded R. nuttalliwithin the SBZ4 Biozone in Egypt. In our sections, wehave not found R. nuttalli in the Paleocene (Figs 3, 510), which is in agreement with previous LBF studiesin Pakistan and India (e.g. Jauhri 1998; Jauhri & Agar-wal 2001; Jauhri et al. 2006; Afzal et al. 2009). How-ever, R. nuttalli ranges from the SBZ5 to SBZ6biozones in India, for example in the Ladakh (Mathuret al. 2009) and Assam-Shillong areas (Jauhri 1998;Jauhri & Agarwal 2001; Jauhri et al. 2006; Tewari et al.2010). Therefore, the last occurrence of A. dandoticatogether with R. nuttalli in Indus Basin successions iscorrelated to the upper boundary of the SBZ5 6 bio-zonal interval (Figs 57, 10).

Miscellanea miscella has been reported fromAfghanistan (Kaever 1970), northeastern Turkey (Si-rel 1997), Iran (Rahaghi 1983) and India (e.g. Jauhri1998; Jauhri & Agarwal 2001; Jauhri et al. 2006) fromhorizons equivalent to the SBZ5 Biozone. In parts ofthe Lower Indus Basin of Pakistan (e.g. in the Sindand Balochistan provinces), M. miscella occurs inassociation with LBFs characteristic of the SBZ5Biozone (e.g. Hottinger 1971; Hottinger et al. 1998;Akhtar & Butt 2000). However, in India, M. miscellahas been reported ranging up to the SBZ6 Biozone(e.g. Mathur et al. 2009; Tewari et al. 2010). In theHanna Lake and Zranda sections, the disappearanceof M. miscella together with A. dandotica and R. nut-talli suggest its upper biostratigraphic limit is within


the SBZ6 Biozone (Figs 6, 7). In the Muree Brewerysection, the occurrence of M. miscella in the interreg-num interval between the last occurrences of LBFs

typical of the SBZ5 6 biozones (e.g. A. dandotica, R.nuttalli etc.) and first appearance of Nummulites atac-icus and N. globulus ( = SBZ8 Biozone) suggests that

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its biostratigraphic range may extend into BiozoneSBZ7 (Fig. 6).

Serra-Kiel et al. (1998) showed that R. sindensisranges up to the SBZ5 Biozone in Tethyan succes-sions. Racey (1995) reported R. sahni associated withA. dandotica ( = Biozone SBZ5) in Oman. In theZranda section, the last occurrences of R. sindensisand R. sahni were recorded within the SBZ5 6 bioz-ones (Fig. 5).

In our sections and based on previous studies ofLBFs in the Indus Basin, D. dispansa is associated withLBFs of the earliest Eocene (e.g. Butt 1991; Akhtar &Butt 1999, 2000) (Figs 57). However, LBF studies inwest Tethys indicate that D. dispansa ranges frombiozones SBZ7 to SBZ20 (e.g. Serra-Kiel et al. 1998;Less et al. 2007; Less & Kovacs 2009).

Evolution of Early Palaeogene LBFsin the east Tethys Ocean

The evolution of LBFs during the Early Palaeogenewas characterized primarily by a long-term reorgani-zation of faunal communities following the KT crisis(which caused extinction of 83% of LBFs, Bou-Dagher-Fadel 2008). Here, based on biostratigraphicanalysis, we investigate LBF long-term evolutionarypatterns in east Tethys.

Biozones SBZ1SBZ2 are represented by the fewsurviving LBF lineages from the KT crisis and domi-nated by taxa with a small test and no dimorphism(Hottinger 1998, 2001) (Fig. 10). This phase is repre-sented in the Indus Basin by only a few genera ofsmaller rotaliids (Rotalia and Laffitteina) and miliolids(Idalina) (Fig. 4). Lockhartia, Bangiana, Miscellaneaand Ornatononion are absent in east Tethys (India andPakistan) (e.g. Adams 1970; Kureshy 1978; Jauhri1998; Jauhri & Agarwal 2001; Jauhri et al. 2006; Greenet al. 2008; Afzal et al. 2009; Mathur et al. 2009; Te-wari et al. 2010), although they are commonlyreported from this interval in west Tethys (e.g. Serra-Kiel et al. 1998). Potential explanations for this

difference could either be inhospitable environmentsin the east Tethys region as deposition of pure carbon-ate did not commence fully till the SBZ3 Biozone (e.g.Jauhri 1998; Jauhri & Agarwal 2001; Jauhri et al. 2006;Green et al. 2008; Afzal et al. 2009; Mathur et al.2009; Tewari et al. 2010), or an inability to reach thearea from the region of origin of the taxa.

The SBZ3SBZ4 biozones are characterized byincreasing generic diversity and reflect a time whenLBFs began experimenting with new morphologies(Hottinger 2001; Scheibner & Speijer 2008a) (Fig. 10).This interval in the Indus Basin commenced with theappearance of many new taxonomic lineages includ-ing genera of rotaliids (Lockhartia and Kathina), mil-iolids (Triloculina and Quinqueloculina), pellatispirids(Miscellanea), alveolinids (Glomalveolina), coskinoli-nids (Coskinon), spirocyclinids (Saudia), dictyoconids(Dictyoconus and Fallotella), nummulitids (Ranikotha-lia, Assilina and Operculina), discocyclinids (Discocy-clina) and lepidorbitoids (Daviesina, Orbitosiphon andSetia) (Figs 35). This faunal pattern is consistentwith other parts of Tethys (Hottinger 1997, 1998;Jauhri 1998; Serra-Kiel et al. 1998; Jauhri & Agarwal2001; Jauhri et al. 2006; Green et al. 2008; Scheibner& Speijer 2008a, 2009; Zamagni et al. 2008; Mathuret al. 2009; Tewari et al. 2010).

In biozones SBZ5SBZ6, a general reorganizationin LBF communities is recorded with a diversificationof species, marking the recovery of LBF k-strategists(characterized by long life and low reproductivepotential) (Hottinger 1997, 2001; Fig. 10). The latestPaleocene (Biozone SBZ4) miscellanids and raniko-thalids are replaced by Early Eocene alveolinids andnummulitids, which come to dominate LBF assem-blages in the western Tethyan realm at the PEboundary (e.g. Scheibner et al. 2005; Scheibner &Speijer 2008a), the so-called larger foraminifer-turnover (LFT, Orue-Etxebarria et al. 2001).

However, in the Indus Basin, LBF assemblages ofthe lowest Eocene (biozones SBZ5 6) are still domi-nated by Ranikothalia and Miscellanea, while newLBFs that first emerged within this time interval

Fig. 8. Biostratigraphically important larger benthic foraminifera from the PaleoceneEarly Eocene (SBZ1SBZ5 6), Indus Basin, Pakistan.All microphotographs taken in plane-polarized light. Sample numbers with ZRD are from Zranda section (Fig. 5); KL from Kotal Pass sec-tion (Fig. 3); PD from Mughal Kot section (Fig. 4); 4472.24 from Shakardara Well-1 (Fig. 3). AC, Laffitteina bibensis Marie, SBZ1SBZ2: A,axial section, PD-16; B, off-centre axial section, PD-8; C, equatorial section, PD-8. D, Lockhartia conditi (Nuttall), SBZ3SBZ5 6, axial sec-tion, KL53. E, Lockhartia conica Smout, SBZ3SBZ4, axial section, 4472.24. F, Lockhartia haimei (Davies), SBZ3SBZ5 6, axial section, ZRD-33. G, Kathina selveri Smout, SBZ3SBZ4, axial section, 4468.61. H, Laffitteina erki (Sirel), SBZ3SBZ4, near axial section, ZRD-21. IJ, Mis-cellanea juliettae Leppig, SBZ3, megalospheric form: I, equatorial section, PD-1; J, axial section, ZRD-28. KL, Miscellanea meandrina (Car-ter), SBZ4: K, microspheric form, equatorial section, ZRD-20; L, megalospheric form, equatorial section, ZRD-20. M, N, Miscellanea miscella(DArchaic & Haime), SBZ4, megalospheric form: M, equatorial section, ZRD-13; N, near axial section, ZRD-8. O, Glomalveolina primaevaReichel, SBZ3SBZ4, non-centred axial section, ZRD-36. P, Ranikothalia sahni Davies, SBZ3SBZ5 6, megalospheric form, axial section,ZRD-4. Q, Ranikothalia sindensis (Davies), SBZ3SBZ5 6, microspheric form, axial section, ZRD-4. R, Spiroloculina sp., axial section, ZRD-12. S, Idalina sinjarica Grimsdale, SBZ2SBZ5 6, axial section, 4471.89. T, Glomalveolina telemetensis Hottinger, SBZ3SBZ4, axial section,ZRD-9. U, Dictyoconus sp., SBZ3SBZ5 6, axial section, ZRD-34. V, Coskinon rajkae Hottinger & Drobne, SBZ3, axial section, ZRD39. W,Fallotella alavensis Mangin, SBZ3, ZRD-39. X, Keramosphaera cf. iranica Rahaghi, SBZ3SBZ4? near equatorial section, ZRD-26. SBZ-shallowbenthic zones.


elsewhere (e.g. Assilina, Alveolina and Discocyclina) areless important and Nummulites are absent (Figs 57,10). Later, in the Early Eocene there was a gradualdiversification of Discocyclina, Operculina and Assilinaspecies and an appearance of new forms including

large Orbitoclypeus, Opertorbitolites and Somalina,while Ranikothalia disappeared and Miscellaneabecame less important by the end of the SBZ5 6 bio-zonal interval. Similar LBF assemblages have beenrecorded in other parts of east Tethys (especially India

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and Pakistan) (e.g. Nagappa 1959b; Hottinger 1971;Jauhri 1998; Akhtar & Butt 2000; Jauhri & Agarwal2001; Jauhri et al. 2006; Scheibner & Speijer 2008a;Tewari et al. 2010). Such LBF assemblages in easternTethys thus differ profoundly from west Tethys.

The SBZ7SBZ8 biozonal interval is characterizedby foraminiferal size increase and further diversifica-tion of successful LBF lineages in Tethys (Hottinger2001) (Fig. 10). In the Indus Basin, Miscellanea disap-pear in the SBZ7 Biozone while Assilina, Discocyclina,Operculina and Alveolina start to diversify and show aclear gradual size increase and dimorphism. The firstNummulites appear in the SBZ8 Biozone and rapidlybecome an important component, while Alveolina,Discocyclina, Assilina and Operculina continue to showdiversification with the evolution of new lineages, forexample Actinocyclina (Figs 6, 7, 8 and 10). Theseforaminiferan assemblages are comparable with thoseof west Tethys and they occur in carbonate platform-building quantity.

LBFs and India-Asia Collision at thePE boundary

The leading edge of the Indo-Pakistan plate under-went subduction and orogenic processes at about55 Ma (the PaleoceneEocene boundary, Khan & Sri-vastava 2006; Najman 2006; Copley et al. 2010)(Fig. 11). Significant stratigraphic changes in parts ofthe basin have been recorded at the PE boundaryinterval producing unconformities, intermittent shal-low- and deep-marine sediments and in some places adramatic shift from marine to continental deposits,the latter containing endemic land mammals in partsof the Indus Basin (see Afzal et al. 2009 and referencestherein) (Fig. 2). This may have been caused by com-pression, uplift and erosion associated with India-Asiacollision (Afzal et al. 2009; Fig. 2).

Plate tectonic movements control the spatial distri-bution and variability of suitable shallow marine habi-tats for benthic foraminifera (Renema et al. 2008).Subject to global climatic constraints, tectonic events

will modulate ocean circulation, resulting in changesin surface water characteristics as well as altering con-nectivity between populations (Renema et al. 2008).The early collision of India with Asia may have gener-ated biogeographical barriers between east (India andPakistan) and west (Europe, North Africa) Tethys,preventing migration of certain LBFs (e.g. species ofNummulites and Alveolina) between these two prov-inces (Fig. 11). The biogeographical patterns of mod-ern LBFs can be interrogated using moleculartechniques, but in fossil populations morphologicalcharacteristics are used to detect biogeographical bar-riers such as intervening eutrophic waters or landmasses (e.g. Langer & Hottinger 2000; Renema et al.2008). Within the tropical subtropical belt, present-day LBFs can migrate relatively easily along marineshelves and even between islands separated by deepwater provided that these are not far apart (Adams1989). The terminal Paleoceneearliest Eocene palaeo-geographical reconstruction by Smith et al. (1994)shows northern India at low northerly latitudes, witha jettisoned Seychelles block then sitting midwaybetween the Indian sub-continent and Madagascar(e.g. Todal & Edholm 1998; Fig. 11A). Ali & Aitchison(2008) suggested that India may have possessed bothsoutherly and northerly connections with other conti-nents (Africa in the south and Asia in the north) pos-sibly forming a route for terrestrial and freshwaterfaunal migration (Fig. 11A). The biological recordalso indicates that Asian land mammals were presenton the Indian sub-continent in the Early Eocene (e.g.Clyde et al. 2003; Rose et al. 2006; Sahni 2006), aswere certain freshwater gastropods (Kohler & Glaubr-echt 2007). This suggests that India-Asia collision atthe PE boundary had probably created land bridgeswith other continents which could have acted as barri-ers for the migration of shallow marine biota (includ-ing LBFs) occupying marine platforms surroundingthe Indian continent (Fig. 11A).

However, although India-Pakistan may have hadconnections with southerly and northerly landmasses,there is no evidence for a westerly placed landmassthat may have restricted LBF migration from western

Fig. 9. Biostratigraphically significant larger benthic foraminifera from the Early Eocene (SBZ5SBZ8 biozones), Indus Basin, Pakistan. Allmicrophotographs taken in plane-polarized light. Sample numbers with ZRD are from Zranda section (Fig. 5); BD from Muree Brewery sec-tion (Fig. 6); HLD from Hanna Lake section (Fig. 7). A, Opertorbitolites sp., SBZ5 6, axial section, ZRD-1. B, Ranikothalia nuttalli (Davies),SBZ5 6, axial section, HLD-3. C, D, Assilina dandotica Davies, SBZ5 6: C, near axial section, HLD-3; D, axial section, HLD-6. E, Somalinasp., SBZ5 6, axial section, BD-3. F, G, Discocyclina dispansa (Sowerby), SBZ5 6SBZ8: F, microspheric form, axial section, BD-15; G,embryon of megalospheric form, equatorial section, HLD-14. H, Discocyclina sella (DArchiac), SBZ5 6SBZ8, microspheric form, axial sec-tion, BD-11. I, Discocyclina ranikotensis Davies, SBZ4SBZ7, microspheric form, axial section, ZRD-2. J, Orbitoclypeus sp., SBZ5 6SBZ8,embryon of megalospheric form, axial section, HLD-13. K, Orbitoclypeus sp., SBZ5 6SBZ8, embryon of megalospheric form, equatorial sec-tion, HLD-13. L, Actinocyclina sp., SBZ8, axial section, HLD-14. M, Alveolina sp., SBZ5 6SBZ8, off-centred axial section, BD-15. N, Alveoli-na sp., SBZ5 6SBZ8, off-centred equatorial section, ZRD-2. O, Glomalveolina ?lepidula (Schwager), SBZ5 6SBZ8, off-centred axial section,BD-5. P, Nummulites sp., SBZ8, axial section, BD-14. Q, R, Nummulites atacicus Leymerie, SBZ8, megalospheric form, axial section, BD-13.S, T, Nummulites globulus Leymerie, SBZ8, megalospheric form, HLD-19; S, axial section; T, off-centred equatorial section. SBZ-shallow ben-thic zones.


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Tethys (see Ali & Aitchison 2008 and referencestherein for palaeogeography). Thus, in the absence ofa land barrier, LBF migration may have been restrictedby an oceanographic barrier, although the nature ofthis is uncertain (Fig. 11A). Similar oceanographicbarriers separate elements of the modern Mediterra-nean biota (e.g. some barnacles and nearshore fish)from those of the Atlantic (Springer 1982; Pannacciul-li et al. 1997). Biogeographical isolation may also haveled to lower diversity in east Tethys, promoting lowercompetition for the same habitat space and enhancingthe survival of existing lineages (e.g. Miscellanea andRanikothalia) (Leppig 1988) during the earliest Eocene(biozones SBZ5 6). The persistence of Miscellaneaand Ranikothalia during the earliest Eocene interval ineastern Tethys indicates that inhospitable lithofaciestypes are not the reason for the absence of Nummulitesas both groups of LBFs occupy the same types of habi-tat (Hottinger 1997).

The different LBF assemblages of east and westTethys during the earliest Eocene are succeeded by acosmopolitan fauna in the SBZ8 Biozone, suggestingthat the continued northward drift of the Indian plate(e.g. Copley et al. 2010) may have collapsed biogeo-graphical barriers between the two Tethyan regions.

Response of LBF assemblages to thePETM in eastern Tethys

As well as the long-term evolutionary changes to thePalaeogene foraminiferal assemblages (Hottinger1997, 2001, 2006), a short-term climatic perturbation

(PETM CIE) at the PE boundary (Zachos et al.2003) had a major impact on shallow marine biota(LBFs and coral-reefs) (Scheibner et al. 2005; Scheib-ner & Speijer 2008a,b) (Fig. 10). This climate pertur-bation caused the extinction of 25% of LBFs(BouDagher-Fadel 2008).

During the PETM, sea surface temperatures rose by5C in low latitudes and 8C in mid to high latitudes(Zachos et al. 2003; Tripati & Elderfield 2004), andhumidity and increased rainfall resulted in increasedterrestrial runoff from areas bordering Tethys (Bolle &Adatte 2001). There was also enhanced upwelling ofdeep water onto the shelf (Speijer & Wagner 2002).This led to a transient expansion of the trophicresource continuum (i.e. the quantity of nutrients insea water) resulting in eutrophic conditions on thecontinental shelves bordering the Tethys Ocean (Spei-jer et al. 1997). Black shale deposition in Tethys (Spei-jer & Wagner 2002; Gavrilov et al. 2003), the collapseof the oligotrophic Gavelinella beccariiformis benthicforaminifera (Speijer et al. 1997), and widespreadApectodinium blooms (Crouch et al. 2003; Crouch &Brinkhuis 2005) have been linked with increasedproductivity related to the PETM.

Larger bebthic foraminiferas are considered k-strat-egists (Hottinger 1983, 1998) and are, therefore, thefirst to disappear during a breakdown or interruptionof stable oligotrophic conditions (Hottinger 1983).According to Scheibner et al. (2005) and Scheibner &Speijer (2008a), the PETM represents such an inter-ruption causing the disappearance of typical latestPaleocene k-strategist LBFs (e.g. Miscellanea, Raniko-thalia) from carbonate platforms in western Tethys,

A B

Fig. 11. Palaeogeography and collision history of the Indian sub-continent. A, earliest Eocene (SBZ5 6 biozones) reconstruction (based onSmith et al. 1994 with data from Gaetani & Garzanti 1991; Gingerich et al. 1997). Dark-grey dashed line with arrow heads show possibleroutes (through land masses) for rodent mammal migration between India and other continents. These land masses likely acted as geographi-cal barriers for LBFs to reach shallow marine habitats of the Indian subcontinent. A postulated oceanographic barrier (possibly larger dis-tances and or eutrophic waters) is shown as a grey band between east and west Tethys, which may also have prevented LBFs migrating fromwest to east. B, rate of convergence (black line) between India and Asia from 75 Ma (after Copley et al. 2010). The grey bands show the tim-ings of notable events in the history of collision. The collision is thought to have begun at the western edge of the plate boundary during PEboundary times (Rowley 1996; Copley et al. 2010).


while more moderate k-strategists like glomalveolinidsthrived without apparent disruption through the PEinterval. Later, oligotrophic conditions were re-estab-lished in the basins and on the platforms during theearliest Eocene and LBFs recovered, with the radiationof Alveolina and Nummulites (LFT of Orue-Etxebarriaet al. 2001).

However, in eastern Tethys, LBFs such as Miscella-nea and Ranikothalia dominated the earliest Eocenecarbonate platforms while Alveolina was less impor-tant and Nummulites absent (e.g. Nagappa 1959b;Hottinger 1971; Jauhri 1998; Akhtar & Butt 2000; Ja-uhri & Agarwal 2001; Jauhri et al. 2006; Scheibner &Speijer 2008a). Other parts of Tethys also show thepersistence of Miscellanea and Ranikothalia beyondthe PE boundary, but as a minor component in asso-ciation with Alveolina and Nummulites-dominatedassemblages. For example, Miscellanea has beenreported together with Alveolina vredenburgi ( = Bioz-one SBZ5) in Turkey (Hottinger 1960; Sirel 1997),Afghanistan (Kaever 1970), Iran (Rahaghi 1983) andIraq (Hottinger 1960) and Ranikothalia together withDiscocyclina in the early Ilerdian of the Northern Cal-careous Alps (Moussavian 1984). Recently, in SWSlovenia, Zamagni (2009) reported Ranikothalia andMiscellanea in the SBZ5 Biozone and Ranikothalia upto Biozone SBZ7, together with small-sized Alveolinaand Nummulites.

These data from Tethys suggest that LBFs like Mis-cellanea and Ranikothalia were not significantlyaffected by the PETM and that they rapidly adapted tonew climatic and environmental regimes, being mostsuccessful in east Tethys. Direct ecological comparisonof these LBFs with modern forms is not possible asthey became extinct during the Early Eocene. How-ever, palaeoecological studies based on carbonate mi-crofacies types and palaeoecological investigations ofother fauna associated with Miscellanea and Raniko-thalia show that they were k-strategists and, as mostk-strategists carry symbionts, lived within the upperphotic zone in carbonate shelf environments (Pignatti1994; Hottinger 1997). Recent studies of living LBFshave demonstrated that taxa with symbionts show ataxon-specific response to environmental changesincluding nutrient and turbidity stress (Langer &Hottinger 2000; Hallock et al. 2003; Renema 2008).Hallock (1987, 1988) described the community struc-ture of symbiont-bearing benthic foraminifera as afunction of nutrient flux, directly influencing euphoticzone depth. These studies predict that species that arehighly specialized to particular levels of light availabil-ity can respond to some loss of water transparency (afunction of plankton densities and fluvial influx ofdissolved organic matter and suspended sediments)by partially compressing their depth ranges. However,

if water transparency is reduced too much, most willprobably disappear, especially if less specialized speciesare present (Hallock 1987). Furthermore, Renema &Troelstra (2001) studied LBFs in the tropical Sper-monde Shelf (SW Sulawesi, Indonesia) and demon-strated that species show a greater flexibility in theirhabitat preferences; not all taxa react in the same wayto the described environmental parameters. Environ-mental factors, and possibly also interaction withother species, affect the distribution of LBFs (Hohe-negger 2004) and thereby, local survivorship orrecruitment (Newberry et al. 1996; Nekola & White1999).

The persistence of Miscellanea and Ranikothalia ineast Tethys during the earliest Eocene (biozonesSBZ5 6) may suggest their flexibility in palaeoecologi-cal adaptation under stressful conditions, controlledby local palaeoecological conditions and eco-spacecompetition with other species. The lack of Nummu-lites and scarcity of Alveolina during the earliest Eocenemay have facilitated the survival of these LBFs in eastTethys. In other parts of Tethys, where typical Paleo-cene LBFs were completely replaced in the Eocene byNummulites, Assilina and Alveolina (e.g. Orue-Etxe-barria et al. 2001; Scheibner et al. 2005; Scheibner &Speijer 2008a, 2009; Pujalte et al. 2009b), k-strategistsappear to have been supplanted (Leppig 1988) duringa rapid adaptive radiation of these genera.

Based on the discussion presented here, we postu-late that the LFT was not an instantaneous and syn-chronous event across the whole of the Tethyanregion caused only by the PETM (as suggested byOrue-Etxebarria et al. 2001; Scheibner et al. 2005;Scheibner & Speijer 2008a, 2009; Pujalte et al. 2009b)but was a gradual process primarily driven by long-term evolutionary patterns of LBFs (Hottinger 2001).Other factors, like local palaeoecological conditions,eco-space competition between taxa, and local tecton-ics may have played an important role in the timingand extent of the LFT in Tethyan carbonate platforms.

Conclusions

The main conclusions of our study are as follows:

1. The LBFs of the Indus Basin provide a biostratig-raphy through the Paleocene to Early Eocene inter-val (SBZ1SBZ8 biozones). The first appearance ofAlveolina, together with A. dandotica, R. nuttalliand D. dispansa, marks the earliest Eocene.

2. Stable isotope data for d13Ccarb show more positivevalues for the Late Paleocene (+3.4& to +3.0&)and lower values (+2.7& to +1.6&) for the earli-est Eocene.


3. The absence of Nummulites and scarcity of Alveoli-na in eastern Tethys during the earliest Eocenemay have been caused by the early collision ofIndia with Asia, creating biogeographical barriersbetween west and east Tethys. This may have low-ered LBF diversity in east Tethys and enhanced thechance of survival of typical Late Paleocene Miscel-lanea and Ranikothalia into the Eocene.

4. The PETM event did not affect LBFs such as Mis-cellanea and Ranikothalia severely and uniformly,and thus the long-term evolutionary patterns toge-ther with local palaeoecological conditions, eco-space competition between taxa, and localtectonics have played an important role in the tim-ing and extent of the larger foraminifer-turnover.

Acknowledgements. We thank the National Centre of Excellencein Geology (NCEG), University of Peshawar, Pakistan, the Univer-sity of Leicester, UK and the British Geological Survey palaeocli-mates programme for funds and laboratory facilities. We aregrateful to Dr Abdul Salam and Aimal Khan Kasi of the Centre ofExcellence in Mineralogy and Dr Akhtar M. Kassi and M. Mohi-bullah of the Geology Department, University of Balochistan, Paki-stan for literature and for facilitating field studies in the Quettadistrict during 2008. Antonino Briguglio from the Department ofPaleontology, University of Vienna, Austria is thanked for his com-ments on identification of Nummulites. We thank ChristianScheibner (Bremen University) and Willem Renema (NationaalNatuurhistorisch Museum Naturalis, Netherlands) for their con-structive reviews.

References

Adams, C.G. 1970: A reconsideration of the East Indian letter clas-sification of the Tertiary. Bulletin of the British Museum (NaturalHistory) Geology 19, 1137.

Adams, C.G. 1989: Foraminifera as indicators of geological events.Proceedings of the Geologists Association 100, 297311.

Afzal, J., Williams, M. & Aldridge, R.J. 2009: Revised stratigraphyof the lower Cenozoic succession of the Greater Indus Basin inPakistan. Journal of Micropalaeontology 28, 723.

Aguirre, J., Baceta, J.I. & Braga, J.C. 2007: Recovery of marineprimary producers after the Cretaceous-Tertiary mass extinc-tion: Paleocene calcareous red algae from the Iberian Penin-sula. Palaegeography, Palaeoclimatology, Palaeoecology 249,393411.

Akhtar, M. & Butt, A.A. 1999: Lower Tertiary biostratigraphy ofthe Kala Chitta Range, northern Pakistan. Revue de Paleobiologie,Geneve 18, 123146.

Akhtar, M. & Butt, A.A. 2000: Significance of Miscellanea miscella(D Archiac & Haime) in the Early Palaeogene stratigraphy ofPakistan. Revue de Paleobiologie, Geneve 19, 123135.

Ali, J.R. & Aitchison, J.C. 2008: Gondwana to Asia: plate tectonics,paleogeography and the biological connectivity of the Indiansub-continent from the Middle Jurassic through latest Eocene(16635 Ma). Earth-Science Reviews 88, 145166.

Aubry, M.P. & Ouda, K. 2003: Introduction to the Upper Paleo-cene-Lower Eocene of the Upper Nile Valley. In Ouda K., AubryM.P. (eds): Part 1. Stratigraphy, iiiv. Micropaleontology Press,New York.

Aubry, M.P., Berggren, W.A., Cramer, B., Dupuis, C., Kent, D.V.,Ouda, K., Schmitz, B. & Steurbaut, E. 1999: Paleocene Eoceneboundary sections in Egypt. 1st International Conference on theGeology of Africa, Assiut, Egypt , 111.

Aubry, M.P., Ouda, K., Dupuis, C., Berggren, W.A., Van Couver-ing, J.A. & the Members of the Working Group on the Paleo-cene Eocene Boundary 2007: Global Standard Stratotype-section and Point (GSSP) for the base of the Eocene Series in theDababiya section (Egypt). Episodes 30, 271286.

Beck, R.A., Burbank, D.W., Sercombe, W.J., Riley, G.W., Barndt,J.K., Berry, J.R., Afzal, J., Khan, A.M., Jurgen, H., Metje, J., Che-ema, A., Shafique, N.A., Lawrence, R.D. & Khan, M.A. 1995:Stratigraphic evidence for an early collision between northwestIndia and Asia. Nature 373, 5558.

Berggren, W.A. & Pearson, P.N. 2005: A revised tropical to sub-tropical Palaeogene planktonic foraminiferal zonation. Journal ofForaminiferal Research 35, 279298.

Berggren, W.A., Kent, D.V., Swisher, C.C. & Aubry, M.P. 1995: Arevised Cenozoic geochronology and chronostratigraphy. InBerggren W.A., Kent D.V., Aubry M.P., Hardenbol J.A. (eds):Geochronology, Time Scales and Global Stratigraphic Correlations:A Unified Temporal Framework for an Historical Geology, 129212. Society of Economic Paleontologists and Mineralogists,Houston, TX, USA, Special Volume 54.

Bignot, G. & Neumann, M. 1991: Les grands foraminife`res duCretace Terminal et du Palaeogene du Nord-Ouest Europeen:recensement et extensions chronologiques. Bulletin de la Infor-mation Geologique de Bass 28, 1329.

Bolle, M.P. & Adatte, T. 2001: Paleoceneearly Eocene climaticevolution in the Tethyan realm: clay mineral evidence. ClayMinerals 36, 249261.

BouDagher-Fadel, M.K. 2008: The Cenozoic larger foraminifera:the Palaeogene. In BouDagher-Fadel M.K. (ed.): Evolution andGeological Significance of Larger Benthic Foraminifera (Develop-ments in Palaeontology and Stratigraphy, 21), 297418. Elsevier,Amsterdam.

Bowen, G.J., Koch, P.K., Gingerich, P.D., Norris, R.D., Bains, S. &Corfield, R.M. 2001: Refined isotope stratigraphy across the con-tinental PaleoceneEocene boundary on Polecat Bench in theNorthern Bighorn Basin. In Gingerich P.D. (ed.): PaleoceneEocene Stratigraphy and Biotic Change in the Bighorn and ClarksFork Basins, Wyoming, 33. University of Michigan Papers onPaleontology, Michigan.

Butt, A.A. 1991: Ranikothalia sindensis Zone in Late Paleocene bio-stratigraphy. Micropaleontology 37, 7785.

Butterlin, J. & Fourcade, E. 1989: Extension stratigraphique etdistribution ge`ographique du genre Lockhartia Davies, 1932(foraminife`re, rotaliidae). Revue de Micropaleontologie 31, 225242.

Clyde, W.C., Khan, I.H. & Gingerich, P.D. 2003: Stratigraphicresponse and mammalian dispersal during initial IndiaAsia col-lision: evidence from the Ghazij Formation, Balochistan, Paki-stan. Geological Society of America Bulletin 31, 10971100.

Copley, A., Avouac, J.P. & Royer, J.Y. 2010: The India-Asia colli-sion and the Cenozoic slowdown of the Indian plate; implica-tions for the forces driving plate motions. Journal of GeophysicalResearch 115, B03410.

Crouch, E.M. & Brinkhuis, H. 2005: Environmental change acrossthe PaleoceneEocene transition from eastern New Zealand: amarine palynological approach. Marine Micropaleontology 56,138160.

Crouch, E.M., Dickens, G.R., Brinkhuis, H., Aubry, M.P.,Hollis, C.J., Rogers, K.M. & Visscher, H. 2003: The Apec-todinum acme and terrestrial discharge during the Paleo-cene Eocene Thermal Maximum: new palynological,geochemical and calcareous nannoplankton observations atTawanui, New Zealand. Palaeogeography, Palaeoclimatology,Palaeoecology 194, 117.

Drobne, K., Cosovic, V. & Robinson, E. 2002: Larger miliolids ofthe Late Cretaceous and Palaeogene seen through space andtime. Geologija 45, 359366.

Dupuis, C., Aubry, M.P., Steurbaut, E., Berggren, W.A., Ouda, K.,Magioncalda, R., Cramer, B.S., Kent, D.V., Speijer, R.P. &Heilmann-Clausen, C. 2003: The Dababiya Quarry Section:Lithostratigraphy, clay mineralogy, geochemistry and paleontol-ogy. Micropaleontology 49, 4159.


Eames, F.E. 1952: A contribution to the study of the Eocene in wes-tern Pakistan and western India; Part A. The geology of standardsections in the western Punjab and in the Kohat District. Quar-terly Journal of the Geological Society of London 107, 159171.

Ferrandez-Canadell, C. 2002: New Paleocene orbitoidiform forami-nifera from the Punjab Salt Range, Pakistan. Journal of Forami-niferal Research 32, 121.

Gaetani, M. & Garzanti, E. 1991: Multicyclic history of the north-ern India continental margin (northwestern Himalaya). TheAmerican Association of Petroleum Geologists Bulletin 75, 14271446.

Gavrilov, Y.O., Shcherbinina, E.A. & Oberhansli, H. 2003: Paleo-ceneEocene boundary events in the northeastern Peri-Tethys.In Wing S.L., Gingerich P.D., Schmitz B., Thomas E. (eds):Causes and Consequences of Globally Warm Climates in the EarlyPalaeogene, 147168. The Geological Society of America,Boulder, CO, USA.

Gingerich, P.D., Abbas, S.G. & Arif, M. 1997: Early Eocene Ouetta-cyon parachai (Condylarthra) from the Ghazij Formation of Ba-luchistan (Pakistan): oldest Cenozoic land mammal from SouthAsia. Journal of Vertebrate Paleontology 17, 629637.

Green, O.R., Searle, M.P., Corfield, R.I. & Corfield, R.M. 2008:Cretaceous-Tertiary Carbonate platform evolution and the ageof the IndiaAsia collision along the Ladakh Himalaya (North-west India). The Journal of Geology 116, 331353.

Hallock, P. 1987: Fluctuation in the trophic resource continuum: afactor in global diversity cycles? Paleoceanography 2, 457471.

Hallock, P. 1988: Interoceanic differences in Foraminifera withsymbiotic algae: a result of nutrient supplies? Proceedings SixthInternational Coral Reef Symposium Townsville, Australia 3, 251255.

Hallock, P., Lidz, B.H., Cockey-Burkhard, E.M. & Donnelly, K.B.2003: Foraminifera as bioindicators in coral reef assessment andmonitoring: The FORAM Index. Environmental Monitoring andAssessment 81, 221238.

Hodges, K.V. 2000: Tectonics of the Himalayan and southern Tibetfrom two perspectives. Geological Society of America Bulletin 112,324350.

Hohenegger, J. 2004: Depth coenoclines and environmental con-siderations of western Pacific larger foraminifera. Journal ofForaminiferal Research 34, 933.

Hottinger, L. 1960: Recherches sur les Alveolines du Palece`ne et delEoce`ne. Schweizerische Palaeontologische Abhandlungen 75 76,1243.

Hottinger, L. 1971: Larger foraminifera common to Mediterraneanand Indian Paleocene and Eocene formations. Annals of the Geo-logical Institute, Hungary 54, 145151.

Hottinger, L. 1983: Processes determining the distribution of largerforaminifera in space and time. Utrecht Micropaleontological Bul-letins 30, 239253.

Hottinger, L. 1997: Shallow benthic foraminiferal assemblages assignals for depth of their deposition and their limitations. Bulle-tin de la Societe Geologique de France 168, 491505.

Hottinger, L. 1998: Shallow benthic foraminifera at the PaleoceneEocene boundary. Strata Serie 19, 6164.

Hottinger, L. 2001: Learning from the past? In Box E., Pignatti S.(eds): Volume IV: The Living World. Part Two, 449477. Aca-demic Press, San Diego.

Hottinger, L. 2006: Illustrated glossary of terms used in foraminif-eral research. Carnets de Ge`ologie Memoir 2, 1126.

Hottinger, L. & Drobne, K. 1988: Tertiary Alveolinids: problemslinked to the conception of species. Revue Palaeobiologie (Ben-thos 86) 2, 665681.

Hottinger, L., Lehmann, R. & Schaub, H. 1964: Donnees actu-elles sur la biostratigraphie de Nummulitique Mediterraneen.Memoires du Bureau de Recherches geologiques et minie`res 28,611652.

Hottinger, L., Sameeni, J. & Butt, A.A. 1998: Emendation of Alveo-lina vredenburgi Davies & Pinfold 1937 from the Surghar Range,Pakistan. Dela Sazu, Ljubljana 34, 155163.

Inan, N., Tasli, K. & Inan, S. 2005: Laffitteina from the Maastrich-tian-Paleocene shallow marine carbonate successions of the

Eastern Pontides (NE Turkey): biozonation and microfacies.Journal of Asian Earth Sciences 25, 367378.

Jauhri, A.K. 1998: Miscellanea Pfender, 1935 (Foraminiferida) fromthe South Shillong region, N.E. India. Journal of the Palaeonto-logical Society of India 43, 7383.

Jauhri, A.K. & Agarwal, K.K. 2001: Early Palaeogene in the southShillong Plateau, NE India: local biostratigraphic signals of glo-bal tectonic and oceanic changes. Palaeogeography, Palaeoclima-tology, Palaeoecology 168, 187203.

Jauhri, A.K., Misra, P.K., Kishore, S. & Singh, S.K. 2006: Largerforaminiferal and calcareous algal facies in the Lakadong Forma-tion of the South Shillong Plateau, NE India. Journal of the Pal-aeontological Society of India 51, 5161.

Kaever, M. 1970: Die alttertiaren Grossforaminiferen Sudost-Afghanistans unter besonderer Berucksichtigung der Num-mulitiden-Morphologie, Taxonomie und Biostratigraphie.Munsterische Forschungen zur Geologie und Palaontologie, Mun-ster, Heft 16 17, 1400.

Kassi, A.M., Kelling, G., Kasi, A.K., Umar, M. & Khan, A.S. 2009:Contrasting Late CretaceousPaleocene lithostratigraphic suc-cessions across the Bibai Thrust, western Sulaiman Fold-ThrustBelt, Pakistan: Their significance in deciphering the early-colli-sional history of the NW Indian Plate margin. Journal of AsianEarth Sciences 35, 435444.

Khan, A.M. & Srivastava, S.K. 2006: The paleogeographic signifi-cance of Aquilapollenites occurrences in Pakistan. Journal ofAsian Earth Sciences 28, 251258.

Khan, S.R., Khan, M.A., Nawaz, R. & Karim, T. 1990: Strati-graphic control for the age of Peshawar plain magmatism,northern Pakistan. Geological Bulletin, University of Peshawar23, 253263.

Kiessling, W. 2002: Secular variations in the Phanerozoic reef eco-system. In Kiessling W., Flugel E., Golonka J. (eds): PhanerozoicReef Patterns, 625690. Society for Sedimentary Geology, Tulsa.

Kohler, F. & Glaubrecht, M. 2007: Out of Asia and into India: onthe molecular phylogeny and biogeography of the endemicfreshwater gastropod Paracrostoma Cossmann, 1900 (Caenogas-tropoda: Pachychilidae). Biological Journal of the Linnean Society91, 627651.

Kump, L.R. 2001: Chill taken out of the tropics. Nature 413, 470471.

Kureshy, A.A. 1978: Tertiary larger foraminiferal zones of Pakistan.Revista espanola de micropaleontologa 10, 467483.

Langer, M.R. & Hottinger, L. 2000: Biogeography of selectedlarger foraminifera. Micropaleontology 46, 105126.

Leppig, U. 1988: Structural analysis and taxonomic revision of Mis-cellanea, Paleocene larger Foraminifera. Eclogae Geologicae Hel-vetiae, Basel 81, 689721.

Less, Gy. & Kovacs, L.O. 2009: Typological versus morphometricseparation of orthophragminid species in single samples a casestudy from Horsarrieu (upper Ypresian, SW Aquitaine, France).Revue de Micropaleontologie 52, 267288.

Less, Gy., Ozcan, E., Baldi-Beke, M. & Kollanyi, K. 2007: Thanetianand early Ypresian Orthophragmines (Foraminifera: Discocy-clinidae and Orbitoclypeidae) from the Central Western Tethys(Turkey, Italy and Bulgaria) and their revised taxonomy andbiostratigraphy. Rivista Italiana di Paleontologia e Stratigrafia113, 419448.

Magioncalda, R., Dupuis, C., Smith, T., Steurbaut, E. & Gingerich,P.D. 2004: PaleoceneEocene carbon isotope excursion inorganic carbon and pedogenic carbonate: direct comparison in acontinental stratigraphic section. Geology 32, 553556.

Martini, E. 1971: Standard Cenozoic and Quaternary calcareousnannoplankton zonation. Proceedings of the Second InternationalConference on Planktonic Microfossils Rome 2, 739777.

Mathur, N.S., Juyal, K.P. & Kumar, K. 2009: Larger foraminiferalbiostratigraphy of lower Palaeogene successions of ZanskarTethyan and Indus-Tsangpo Suture Zones, Ladakh, India in thelight of additional data. Himalayan Geology 30, 4568.

Moussavian, E. 1984: Die Gosau- und Alttertiar-Geroller der An-gerberg-Schichten (Hoheres Oligozan, Unterinntal, NordlicheKalkalpen). Facies 10, 186.


Nagappa, Y. 1959a: Note on operculinoids Hanzawa 1935. Palae-ontology 2, 2123.

Nagappa, Y. 1959b: Foraminiferal biostratigraphy of the Creta-ceous-Eocene succession in the India-Pakistan, Burma region.Micropaleontology 5, 145192.

Najman, Y. 2006: The detrital record of orogenesis: a review ofapproaches and techniques used in the Himalayan sedimentarybasins. Earth Science Review 74, 172.

Nekola, J.C. & White, P.S. 1999: The distance decay of simi-larity in biogeography and ecology. Journal of Biogeography26, 867878.

Newberry, D., Cambell, E.F.J., Proctor, J. & Still, M.J. 1996: Pri-mary lowland dipterocarp forest at Danum Valley, Sabah,Malaysia. Species composition and patterns in the understory.Vegetatio 122, 193220.

Orue-Etxebarria, X., Pujalte, V., Bernaola, G., Apellaniz, E., Baceta,J.I., Payros, A., Nunez-Betelu, K., Serra-Kiel, J. & Tosquella, J.2001: Did the Late Paleocene Thermal Maximum affect the evo-lution of larger foraminifers? Evidence from calcareous planktonof the Campo Section (Pyrenees, Spain). Marine Micropaleontol-ogy 41, 4571.

Ozgen, N. & Akyazi, M. 2001: The benthic foraminifera of Thane-tian of Padriciano (Italy), Sopada (Slovenia) and Western Pont-ids (Turkey). Bulletin of Earth Sciences Application and ResearchCentre of Hacettepe University 23, 8798.

Ozgen-Erdem, N., Inan, N., Ajyazi, M. & Tunoglu, C. 2005:Benthonic foraminiferal assemblages and microfacies analysisof PaleoceneEocene carbonate rocks in the Kastamonuregion, Northern Turkey. Journal of Asian Earth Sciences 25,403417.

Pannacciulli, F.G., Bishop, J.D.D. & Hawkins, S.J. 1997: Geneticstructure of populations of two species of Chthamalus (Crusta-cea: Cirripedia) in the north-east Atlantic and Mediterranean.Marine Biology 128, 7382.

Pearson, P.N., Ditchfield, P.W., Singano, J., Harcourt-Brown, K.G.,Nicholas, C.J., Olsson, R.K., Shackleton, N.J. & Hall, M.A. 2001:Warm tropical sea surface temperatures in the Late Cretaceousand Eocene epochs. Nature 413, 481487.

Pignatti, J.S. 1994: Paleobiogeography of Palaeogene larger forami-nifera from the Mediterranean Tethys to the Western Pacificusing parsimony analysis: a preliminary attempt. Bollettino DellaSocieta` Paleontologica Italiana 2, 243252.

Pignatti, J., Di Carlo, M., Benedetti, A., Bottino, C., Briguglio, A.,Falconi, M., Matteucci, R., Perugini, G. & Ragusa, M. 2008: SBZ2-6 larger foraminiferal assemblages from the Apulian and Pre-Apulian Domains. Atti del Museo civico di storia naturale di Trie-ste 53, 131146.

Pujalte, V., Baceta, J.I., Schmitz, B., Orue-Etxebarria, X., Payros,A., Bernaola, G., Apellaniz, E., Caballero, F., Robador, A., Serra-Kiel, J. & Tosquella, J. 2009a: Redefinition of Ilerdian Stage(early Eocene). Geologica Acta 7, 177194.

Pujalte, V., Schmitz, B., Baceta, J.I., Orue-Etxebarria, X., Bernaola,G., Dinares-Turell, J., Payros, A., Apellaniz, E. & Caballero, F.2009b: Correlation of the ThanetianIlerdian turnover of largerforaminifera and the Paleocene- Eocene thermal maximum:confirming evidence from the Campo area (Pyrenees, Spain).Geologica Acta 7, 161175.

Racey, A. 1995: Lithostratigraphy and larger foraminiferal (num-mulitid) biostratigraphy of the Tertiary of Northern Oman.Micropaleontology 41, 1123.

Rahaghi, A. 1983: Stratigraphy and faunal assemblage of Paleo-cene-Lower Eocene in Iran. National Iranian Oil Company, Geo-logical Laboratories, Teheran 10, 173.

Renema, W. 2008: Habitat selective factors influencing thedistribution of large benthic Foraminifera assemblages over theKepualauan Seribu. Marine Micropaleontology 68, 286298.

Renema, W. & Troelstra, S.R. 2001: Larger foraminiferadistribution on a mesotrophic carbonate shelf in SW Sulawesi(Indonesia). Palaeogeography, Palaeoclimatology, Palaeoecology175, 125147.

Renema, W., Bellwood, D.R., Braga, J.C., Bromfield, K., Hall, R.,Johnson, K.G., Lunt, P., Meyer, C.P., McMonagle, L.B., Morley,R.J., ODea, A., Todd, J.A., Wesselingh, F.P., Wilson, M.E. &

Pandolfi, J.M. 2008: Hopping hotspots: global shifts in marinebiodiversity. Science 321, 654657.

Rose, K.D., Smith, T., Rana, R.S., Sahni, A., Singh, H., Missiaen, P.& Folie, A. 2006: Early Eocene (Ypresian) continental vertebrateassemblage from India, with description of a new anthracobunid(mammalia, tethytheria). Journal of Vertebrate Paleontology 26,219225.

Rowley, D.B. 1996: Age of initiation of collision between India andAsia: a review of stratigraphic data. Earth and Planetary ScienceLetters 145, 113.

Sahni, A. 2006: Biotic response to the India-Asia collision. Palaeon-tographica Abteilung A-Stuttgart 278, 1526.

Samanta, B.K. 1969: Taxonomy and stratigraphy of the Indian spe-cies of Discocyclina (foraminifera). Geological Magazine 106,115129.

Sameeni, S.J. & Butt, A.A. 2004: Alveolinid biostratigraphy of theSalt Range succession, Northern Pakistan. Revue de Paleobiologie,Geneve 23, 505527.

Schaub, H. 1981: Nummulites et Assilines de la Tethys paleoge`ne.Taxonomie, phylogene`se et biostratigraphie. Schweizerische Pal-aeontologische Abhandlungen 104 105 106, 1238.

Scheibner, C. & Speijer, R.P. 2008a: Late Paleocene-early EoceneTethyan carbonate platform evolution: a response to long- andshort-term paleoclimatic change. Earth-science reviews 90, 71102.

Scheibner, C. & Speijer, R.P. 2008b: Decline of coral reefs dur-ing Late Paleocene to Early Eocene global warming eEarth3, 1926.

Scheibner, C. & Speijer, R.P. 2009: Recalibration of the TethyanShallow-Benthic Zonation Across the PaleoceneEocene Bound-ary; the Egyptian Record. Geologica Acta 7, 195214.

Scheibner, C., Speijer, R.P. & Marzouk, A.M. 2005: Larger forami-niferal turnover during the Paleocene Eocene thermal maxi-mum and paleoclimatic control on the evolution of platformecosystems. Geology 33, 493496.

Schmitz, B., Asaro, F., Molina, E., Monechi, S., von Salis, K. &Speijer, R.P. 1997: High-resolution iridium, d13C, d18O, forami-nifera and nannofossil profiles across the latest Paleocene ben-thic extinction event at Zumaya, Spain. Palaeogeography,Palaeoclimatology, Palaeoecology 133, 4968.

Schmitz, B., Pujalte, V. & Nunez-Betelu, K. 2001: Climateand sealevel perturbations during the Initial Eocene Ther-mal Maximum: evidence from siliciclastic units in the Bas-que Basin (Ermua, Zumaia and Trabakua Pass), northernSpain. Palaeogeography, Palaeoecology, Palaeoclimatology 165,299320.

Serra-Kiel, J., Hottinger, L., Caus, E., Drobne, K., Fernandez, C.,Jauhri, A.K., Less, G., Pavlovec, R., Pignatti, J., Samso, J.M.,Schaub, H., Sirel, E., Strougo, A., Tambareau, Y., Tosquella, Y.& Zakrevskaya, E. 1998: Larger foraminiferal biostratigraphy ofthe Tethyan Paleocene and Eocene. Bulletin de la Societe Geologi-que de France 169, 281299.

Shah, S.M.I. 1977: Stratigraphy of Pakistan. Memoirs of the Geologi-cal Survey of Pakistan 12, 1137.

Sirel, E. 1997: The species of Miscellanea PFENDER, 1935 (Forami-niferida) in the Thanetian-Ilerdian sediments of Turkey. Revuede Paleobiologie, Gene`ve 16, 7799.

Sirel, E. 1998: Foraminiferal description and biostratigraphy of thePaleocene-Lower Eocene shallow-water limestones and discus-sion on the CretaceousTertiary boundary in Turkey. GeneralDirectorate of the Mineral Research and Exploration, MonographySeries 2, 1117.

Smith, A.G., Smith, D.G. & Funnell, B.M. 1994: Atlas of Mesozoicand Cenozoic Coastlines, 99 pp. Cambridge University Press,Cambridge.

Smout, A.H. 1954: Lower Tertiary foraminifera of the QatarPeninsula. Bulletin of the British Museum (Natural History)xx, 180.

Speijer, R.P. & Wagner, R. 2002: Sea-level changes and black shalesassociated with the late Paleocene Thermal Maximum; organic-geochemical and micropaleontologic evidence from the south-ern Tethyan margin (Egypt-Israel). In Koeberl C., MacLeodK.G. (eds): Catastrophic Events & Mass Extinctions: Impacts and


Beyond, 533549. Geological Society of America Special Paper,Boulder, CO, USA.

Speijer, R.P., Schmitz, B. & van der Zwaan, G.J. 1997: Benthic fora-miniferal extinction and repopulation in response to latestPaleocene Tethyan anoxia. Geology 25, 683686.

Springer, V.G. 1982: Pacific plate biogeography with special refer-ence to shore fishes. Smithsonian Contributions to Zoology 367,1182.

Tewari, V.C., Kumar, K., Lokho, K. & Siddaiah, N.S. 2010: Laka-dong limestone: PaleoceneEocene boundary carbonates sedi-mentation in Meghalaya, northeastern India. Current Science 98,8895.

Thiry, M., Milnes, A.R., Rayot, V. & Simon-Coincon, R. 2006:Interpretation of paleoweathering features and successive silicifi-cations in the Tertiary regolith of inland Australia. Journal of theGeological Society 163, 723736.

Todal, A. & Edholm, O. 1998: Continental margin off westernIndia and Deccan large igneous province. Marine GeophysicalResearch 20, 273291.

Tosquella, J., Serra-Kiel, J., Ferra`ndez-Canadell, C. & Samso, J.M.1998: Las biozonas de nummultidos del Paleoceno SuperiorEoceno Inferior de la Cuenca Pirenaica. Acta Geologica Hispanica31, 2336.

Tripati, A. & Elderfield, H. 2004: Abrupt hydrographic changes inthe equatorial Pacific and subtropical Atlantic from foraminif-eral Mg Ca indicate greenhouse origin for the Thermal Maxi-mum at the PaleoceneEocene boundary. GeochemistryGeophysics Geosystems 5, Q02006.

Tripati, A.K., Delaney, M.L., Zachos, J.C., Anderson, L.D., Kelly,D.C. & Elderfield, H. 2003: Tropical sea-surface temperaturereconstruction for the early Palaeogene using Mg Ca ratios ofplanktonic foraminifera. Paleoceanography 18, 1101.

Weiss, W. 1993: Age assignments of larger foraminiferal assem-blages of Maastrichtian to Eocene age in northern Pakistan.Zitteliana 20, 223252.

White, M.R. 1992: On species identification in the foraminiferalgenus Alveolina (Late Paleocene-Middle Eocene). Journal ofForaminiferal Research 22, 5270.

Zachos, J.C. 1994: Evolution of early Cenozoic marine tempera-tures. Paleoceanography 9, 353387.

Zachos, J., Pagani, M., Sloan, L., Thomas, E. & Billups, K. 2001:Trends, rhythms, and aberrations in global climate 65 Ma topresent. Science 292, 686693.

Zachos, J.C., Wara, M.W., Bohaty, S., Delaney, M.L., Petrizzo,M.R., Brill, A., Bralower, T.J. & Premoli-Silva, I. 2003: A tran-sient rise in tropical sea surface temperature during the Paleo-ceneEocene Thermal Maximum. Science 302, 15511554.

Zamagni, J. 2009: Responses of a Shallow-Water Ecosystem to theEarly Palaeogene Greenhouse Environmental Conditions. Unpub-lished PhD thesis, University of Potsdam, Potsdam.

Zamagni, J., Mutti, M. & Kosir, A. 2008: Evolution of shallow ben-thic communities during the Late Paleocene-earliest Eocenetransition in the Northern Tethys (SW Slovenia). Facies 54,2543.


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Afzal Et Al. 2011, Lethaia, Vol. 44