Cretaceous Research 32 (2011) 331e356

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Cretaceous Research
journal homepage: www.elsevier.com/locate/CretRes

Palaeoecology and biostratigraphy of early Cretaceous (Aptian) calcareous

nannofossils and the d13Ccarb isotope record from NE Iran
Azam Mahanipour a, c, *, Jörg Mutterlose b, Anoshiravan L. Kani a, Mohammad Hosein Adabi a
a
Department of Geology, Faculty of Earth Sciences, Shahid Beheshti University, Tehran, Iran
b
Institut für Geologie, Mineralogie und Geophysik, Ruhr-Universität Bochum, Universitätsstrasse 150, D-44780 Bochum, Germany
c
Department of Geology, Faculty of Science, Shahid Bahonar University, 22 Bahman Bolvar, Kerman, Iran

a r t i c l e i n f o a b s t r a c t

Article history: Upper BarremianeLower Aptian sediments of the Sarcheshmeh and Sanganeh formations in the Kopet
Received 22 December 2008 Dagh area, northeast Iran were studied with regard to their calcareous nannofossil content and their d13Ccarb
Accepted in revised form 24 January 2011 signal. The sediments are composed mainly of marlstones, argillaceous limestones and limestones. Based
Available online 2 February 2011
on the occurrence of biostratigraphic index taxa, the calcareous nannofossil zones NC5, NC6 and the NC7A
Subzone were recognised. The calcareous nannofossils and the d13Ccarb data enable recognition of the early
Keywords:
Aptian Oceanic Anoxic Event 1a (OAE 1a). The deposits of the OAE 1a interval are characterised by the rarity
Calcareous nannofossils
of nannoconids and a sharp negative d13Ccarb excursion (1.36&), followed by an abrupt positive d13Ccarb
Stable isotopes
OAE 1a
excursion of 4e5&; both events have been recognised elsewhere in OAE 1a deposits in the Tethys. In the
Northern Tethys OAE 1a interval, the relative abundance of Watznaueria barnesiae/Watznaueria fossacincta is higher (more
Aptian than 40%) than that of Biscutum spp., Discorhabdus spp. and Zeugrhabdotus spp., which indicates dissolution.
Kopet Dagh Basin In the upper part of the section, the higher relative abundance of mesotrophic and oligotrophic taxa
Iran (Watznaueria spp. and nannoconids respectively) and the enhanced relative abundance of eutrophic taxa
(Biscutum spp., Discorhabdus spp., Zeugrhabdotus spp.) is indicative of an environment with slightly
increased nutrient content. The presence of warm water taxa (Rhagodiscus asper and nannoconids) and the
absence of cool water taxa (Repagulum and Crucibiscutum) suggest warm surface-water conditions.
Ó 2011 Elsevier Ltd. All rights reserved.

1. Introduction
The Kopet Dagh Basin (¼KDB) is located within the border
triangle of Iran, Turkmenistan and Afghanistan (Fig. 1a). Sediments
of Cretaceous age are well exposed in the northeastern part of Iran
in the KDB. The National Iranian Oil Company (¼NIOC) has carried
out various geological studies in this area. Afshar-Harb (1969, 1979)
worked on the petroleum geology of the Iranian part of the KDB.
Kalantari (1969), Seyed-Emami (1980), Seyed-Emami and Aryai
(1981), Seyed-Emami et al. (1984, 1994, 1996), Immel et al.
(1997), Raisossadat and Moussavi-Harami (2000) and Raisossadat
(2004, 2006) studied the biostratigraphy and sedimentology of
the Cretaceous (BerriasianeMaastrichtian). In the western part of
the basin, Upper BarremianeLower Aptian strata are well exposed
in the Takal Kuh mountain and have been analysed for calcareous
nannofossils and stable isotopes in the present study.
* Corresponding author. Department of Geology, Faculty of Science, Shahid
Bahonar University, 22 Bahman Bolvar, Kerman, Iran. Tel.: þ98 9132993410; fax:
þ98 3413222035.
E-mail address: a_mahanipour@mail.uk.ac.ir (A. Mahanipour).
The global Early Aptian Oceanic Anoxic Event 1a (OAE 1a; e.g.
Arthur et al., 1990; Jenkyns, 1999) is characterised by increased
organic carbon burial accompanied by low carbonate sedimenta-
tion (Schlanger and Jenkyns, 1976; Menegatti et al., 1998; Bralower
et al., 1999; Jenkyns, 1999; Leckie, 2002; Erba, 2004). Different
studies have shown that this event is associated with increased
marine productivity, enhanced preservation of organic matter, sea
level rise, significant changes in marine nannoflora and microfauna
(Haq et al., 1988; Erbacher et al., 1996; Jenkyns, 1999; Erba, 2004)
and abrupt carbon isotope excursions (Menegatti et al., 1998;
Bralower et al., 1999), which are indicative of significant pertur-
bations of the global carbon cycling. The onset of OAE 1a is char-
acterised by a short-lived negative d13Ccarb excursion which has
been attributed by some authors to a rapid methane release from
continental margins (Jahren and Arens, 1998; Opdyke et al., 1999;
Jahren et al., 2001; Beerling et al., 2002), while others relate it to
intensified marine volcanism that introduced light carbon into the
oceaneatmospheric system (Arthur et al., 1985; Larson, 1991;
Arthur, 2000). This negative excursion is followed by
a pronounced positive d13C excursion. The processes that caused
the positive excursion are still a matter of debate and two models

0195-6671/$ e see front matter Ó 2011 Elsevier Ltd. All rights reserved.
doi:10.1016/j.cretres.2011.01.006
332 A. Mahanipour et al. / Cretaceous Research 32 (2011) 331e356
Fig. 1. The extension of the Kopet Dagh Basin in Iran and neighbouring countries (a) and the road map of the study area (b), modified after Berberian and King (1981).
have been proposed. Some workers attribute this excursion to
increased productivity and enhanced fertility in ocean surface
waters that caused a major perturbation of the global carbon
budget (Arthur et al., 1987; Erba, 1994; Weissert et al., 1998;
Jenkyns, 1999; Leckie et al., 2002) while others (Luciani et al.,
2001; Heimhofer et al., 2006) show that the high surface-water
productivity during the OAE 1a is not supported by data from some
Tethyan localities. The positive excursion is instead explained by
the enhanced preservation of organic matter in oxygen-depleted
ocean waters (e.g. Schlanger and Jenkyns, 1976; Arthur et al., 1990;
Tyson, 1995; Erbacher et al., 2001).
Whatever the cause, these perturbations of the ocean/climate
system led to a change in calcareous nannofossil assemblages (e.g.
nannoconid crisis; Erba, 1994, 2004; Erba and Tremolada, 2004) that
marks a global event in the Early Aptian OAE 1a (Erba, 1994). Such
a change in nannofloral assemblages is recorded worldwide (Bralower
et al., 1994; Aguado et al., 1999; Bersezio et al., 2002; Erba and
Tremolada, 2004). Planktonic foraminifera (Premoli Silva et al., 1999;
Leckie et al., 2002) and radiolarians (Erbacher et al., 1996) were also
inflicted by palaeoenvironmental changes during the OAE 1a.
The main goals of this study are 1) to establish a biostratigraphic
framework for the Upper BarremianeLower Aptian sediments of
the KDB using calcareous nannofossils, 2) to see if the OAE 1a is
recorded in the northeastern part of the Tethys basin (NE Iran),
using calcareous nannofossils and stable isotopes.
2. Geological setting and lithology
The sediments of the KDB (northern part of the Tethyan Realm;
e.g. Berberian and King, 1981; Glennie, 2000) are exposed from the
Caspian Sea in the west to Turkmenistan and Afghanistan in the
east (Fig. 1a and b). The Iranian part of the basin is nowadays
located between 35 300 e38 150 E and 54 000 e61130 N (Fig. 1b).
The studied section (the Takal Kuh section) is located 70 km west of
Ashkhaneh city, near the AshkhaneheMaraveh Tapeh road at
37430 N and 56100 E (Fig. 1b).
 A. Mahanipour et al. / Cretaceous Research 32 (2011) 331e356 333

Fig. 2. Lithostratigraphy of the Sarcheshmeh and Sanganeh Formations in the Takal Kuh section. Biostratigraphic important nannofossil events are indicated. The “nannoconid
crisis” interval is shaded.
334 A. Mahanipour et al. / Cretaceous Research 32 (2011) 331e356

Fig. 3. Vertical distribution of CaCO3%, absolute abundance, diversity, Shannon index and evenness of calcareous nannofossils in the Takal Kuh section. The “nannoconid crisis”
interval is shaded.

The Cretaceous deposits of the KDB cover the complete Berria-
sianeMaastrichtian interval. The 1668 m thick sediments of the
Sarcheshmeh and Sanganeh formations were investigated in the
current study. Based on ammonite occurrences, these two forma-
tions have been assigned a late Barremianeearly Aptian age
(Raisossadat 2004, 2006). The Sarcheshmeh Formation (Upper
BarremianeLower Aptian) consists of marlstones, argillaceous
limestones and limestones, the overlying Sanganeh Formation
(Lower Aptian) consists of marls (Fig. 2).
3. Material and methods
3.1. Calcareous nannofossils
The 1668 m thick Upper BarremianeLower Aptian sedimentary
sequence of the Takal Kuh section was logged and sampled in 2007. A
total of 288 samples was collected from the section at 3e5 m
intervals in order to study the calcareous nannofossils. For
biostratigraphic investigations all of the samples were processed
Table 1
Absolute abundance, species richness, Shannon Index, evenness, preservation, CaCO3% and counting results of calcareous nannofossils in the upper part of the Takal Kuh section (samples 223e288).

Systematic Sample number: 223 224 225 226 227 228 229 230 231 232 233 234 235 236 237
a
Absolute abundance 163.4 352.8 285.1 345.4 452.6 446.4 608.4 327.9 365.9 454.3 293.5 353.2 405.3 387.9 388.8
(milion Ind./g Sed.)
Species richness 22.0 31.0 38.0 54.0 55.0 46.0 39.0 58.0 58.0 51.0 48.0 63.0 55.0 44.0 54.0
Shannon Index 1.1 2.2 2.3 2.9 2.6 2.7 2.7 2.6 2.9 2.6 2.6 2.6 2.9 2.5 2.6
Evenness 0.4 0.6 0.6 0.7 0.7 0.7 0.8 0.7 0.7 0.7 0.7 0.6 0.7 0.7 0.7
E2- E2- E2- E2-
Preservation E3 E2 E3 E2 E2 E3 E3 E2 E2 E2 E2 E2 E2 E2 E2
% calcium carbonate 8 11 9 27 58 12 20 15 18 16 15 15 15 14 15
Order Family Total number of species 322 338 363 410 377 369 363 375 404 336 354 376 358 360 380
in each sample
Eiffellithales Chiastozygaceae Bukrylithus ambiguus # 1 2 # 1 # 2 2 2 4 # 1 2 2
Chiastozygus spp. # 1 1 1 # # 2 1
Loxolithus armilla 2 # 17 6 3 # 14 1
Staurolithites crux 4 1 2 2 4 1 1 # 1 #
Staurolithites glabra # # 1 # #
Staurolithites mutterlosei # #

A. Mahanipour et al. / Cretaceous Research 32 (2011) 331e356

Staurolithites stradneri 2 1 1 1 # # 2 1 1 # # 1 #
Staurolithites sp. 4 6 7 6 5 16 1 2 6 # 5 5 9 2
Tranolithus gabalus # 1
Zeugrhabdotus clarus 1 1 1 # # # # 1 #
Zeugrhabdotus 2 1 4 # 5 7 2 3 2 5 1 4 4 1
diplogrammus
Zeugrhabdotus elegans 3 # 2
Zeugrhabdotus cf. # 1 # 3 1 # # 1 1 1
Z. embergeri
Zeugrhabdotus 1 # 1 # 1 # # 1 1 # #
embergeri
Zeugrhabdotus erectus 1 4 5 1 1 1 # 2 # 1
Zeugrhabdotus scutula 1 # # #
Zeugrhabdotus trivectis 1 # # 1 2 # # 1
Zeugrhabdotus xenotus 1 # 2 1 2 1 # 1 #
Zeugrhabdotus sp. 8 14 23 8 12 55 8 18 19 17 13 25 11 12
Eiffellithaceae Tegumentum octiformis # 1 1 1 #
Tegumentum striatum # # 2 # 1 1 # 1 2 #
Rhagodiscaceae Percivalia fenestrata 2 1 1 3 # 1 2 3 # 1 2 #
Rhagodiscus angustus 2 8 2 14 5 10 14 6 15 10 9 12 9 4 7
Rhagodiscus asper 20 35 24 84 64 13 31 50 60 29 50 58 48 44 54
Rhagodiscus dekaenelii # 1 #
Rhagodiscus gallagheri # # #
Rhagodiscus robustus # #
Rhagodiscus sp. 1 7 2 14 4 8 12 7 20 10 8 1 7 13 11
Stephanolithiales Stephanolithiaceae Rotelapillus laffittei 2 2 10 4 3 4 6 # 3 9 4 # 3 5
Stoverius achylosus
Stradnerlithus #
geometricus
Stradnerlithus
Rhombicus
Podorhabdales Axopodorhabdaceae Axopodorhabdus # 1 5 3 3 2 # 1 5 13 3 2
dietzmannii
Ethmorhabdus 3 1 2
hauterivianus
Tetrapodorhabdus 1 3 1 1 1 3 # 1 3 # 1
coptensis
Biscutaceae Biscutum constans 12 12 28 47 16 18 18 17 15 8 9 26 11 13
Discorhabdus ignotus 2 8 11 21 22 6 11 12 13 13 16 8 9 11 12

335
(continued on next page)
Table 1 (continued )

336
Systematic Sample number: 223 224 225 226 227 228 229 230 231 232 233 234 235 236 237
Cruciplacolithus haye 2 2 # 1 2 2
Sollasites sp. 1 7 # 1 3 # 2 # 1 2
Cretarhabdaceae Cretarhabdus sp. # 3 2 # 1 3 9 5 4 # 1 3 3
Cretarhabdus conicus 3 # # 2 # # 1 4 1 # 2 1 3 4
Flabellites oblongus 1 1 1 # # 2 # #
Helenea chiastia # # # # # 2 1 # 1 1 1
Pickelhaube furtiva # # 2 1
Retecapsa sp. # 4 8 2 3 1 2 1 1
Retecapsa angustiforata 1 1 # # 1
Retecapsa surirella 3 # 1 # # # # 3 # # # #
Tubodiscaceae Manivitella spp. 1 1 1 # 1 3 1 # 2 3 1 # 2 4 #
Tubodiscus burnettiae 5 3 11 9 3 11 6 4 2 8 5 6 # 3 3
Tubodiscus jurapelagicus #
Watznaueriales Watznaueriaceae Cyclagelosphaera margerelii 11 7 4 3 3 5 2 # #
Cyclagelosphaera sp. 1
Cylindralithus nudus # 1 # 1 2 4 # 2 1 1 9
Diazomatolithus lehmanii 1 2 1 6 # # # 2 # 1 1 #
Watznaueria barnesiae 247 146 146 85 129 86 77 143 117 131 128 146 89 136 138

A. Mahanipour et al. / Cretaceous Research 32 (2011) 331e356

Watznaueria biporta 1 3 2 1 3 4 5 4 2 1
Watznaueria britannica 1 # 1
Watznaueria fossacincta 18 35 64 26 14 94 41 14 19 14 21 18 30 26 27
Watznaueria ovata 5 1 4 3 14 8 3 4 4 2 3 7 3 2
Arkhangelskiales Arkhangelskiellaceae Broinsonia matalosa 2 2 6 2 # 1 2 2 # 2
Uncertain Haqius circumradiatus # # # 1 1 1 # # 2
Heterococcoliths Haqius ellipticus # # # # # # #
Holococcoliths Calyptrosphaeraceae Calculites spp. # 10 8 11 7 12 15 10 10 6 16 23 8 14 24
Nannoliths Braarudosphaeraceae Braarudosphaera africana #
Braarudosphaera
hockwoldensis
Braarudosphaera regularis
Braarudosphaera sp. 1 1 1 1
Micrantholithus obtusus # # 2 4 # 1 2 3 1 2
Micrantholithus hoschulzii # 2 1 1 # 1 # # #
Nannoconaceae N. abundance #
Nannoconus boneti #
Nannoconus bucheri # 1 # 1 # 1
N. carniolensis #
Nannoconus circularis #
Nannoconus elongatus # # 1 # 1 1
Nannoconus fragilis # #
Nannoconus grandis
Nannoconus globulus #
Nannoconus inconspicuus 1 1
Nannoconus inornatus
Nannoconus kamptneri #
Nannoconus minutus 1 # 1 5 1 1 # 1 1
Nannoconus multicadus
Nannoconus quadriangulus #
Nannoconus truitti 1 1 # # # #
Nannoconus truitti frequens
Nannoconus truitti 1 #
rectangularis
Nannoconus vocontiensis #
Nannoconus wassallii 1 # 1
Nannoconus sp. 1 1 8 3 4 2 3 2 # 2
Polycyclolithaceae Eprolithus spp. # # 1 # 2 # #
Radiolithus planus # 1 # #
 Uncertain Rucinolithus irregularis # 2 # 4 1 1 1 2 #
polycycloliths Rucinolithus sp. 1 # 1 1
Hayesites spp. 1 1 1
Microrhabdulaceae Lithraphidites carniolensis # 38 27 22 20 18 20 42 34 33 29 34 27 41 26

Systematic Sample number: 239 240 241 242 243 244 245 246 247 248 249 250 251 252 253
b
Absolute abundance 437.2 591.7 524.6 421.0 331.4 263.5 543.3 191.6 362.5 297.5 397.1 479.1 414.5 428.4 512.7
(milion Ind./g Sed.)
Species richness 58.0 50.0 52.0 55.0 45.0 49.0 60.0 51.0 61.0 59.0 59.0 58.0 64.0 55.0 56.0
Shannon Index 2.8 2.9 2.2 2.8 2.8 2.6 3.0 2.8 2.9 3.1 3.1 3.1 3.0 3.1 3.0
Evenness 0.7 0.8 0.6 0.7 0.7 0.7 0.7 0.7 0.7 0.8 0.8 0.8 0.7 0.8 0.8
E2- E2- E2- E2-
Preservation E1 E1 E2 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E1 E2
% calcium carbonate 43 48 14 37 36 37 52 14 16 22 27 47 62 42 15
Order Family Total number of species 376 383 360 377 347 358 364 363 340 352 333 342 352 339 361
in each sample
Eiffellithales Chiastozygaceae Bukrylithus ambiguus # 1 2 # 2 1 1 # 1 2 3
Chiastozygus spp. # 1 1 3 1 3 # 1
Loxolithus armilla 1 1 8 3 1 1 #

A. Mahanipour et al. / Cretaceous Research 32 (2011) 331e356

Staurolithites crux 1 1 # 1 # # # # 1
Staurolithites glabra # 1
Staurolithites mutterlosei # #
Staurolithites stradneri 1 1 1 2 # 1 # # # # # 1 #
Staurolithites sp. 1 4 1 3 4 2 5 8 4 5 4 2 4 10
Tranolithus gabalus # 1
Zeugrhabdotus clarus # 1 # # # # # 1 # #
Zeugrhabdotus diplogrammus 2 # # # # 1 4 4 # # # 4
Zeugrhabdotus elegans
Zeugrhabdotus cf. Z. embergeri 1 1 # 1 1
Zeugrhabdotus embergeri # # # # # # # # 1 # 2 1
Zeugrhabdotus erectus # 1 1 # # 2 4
Zeugrhabdotus scutula # # 1
Zeugrhabdotus trivectis # # # #
Zeugrhabdotus xenotus # # # 1 # # 1 # 1
Zeugrhabdotus sp. 16 8 15 20 29 6 6 10 13 12 17 5 16 5 11
Eiffellithaceae Tegumentum octiformis # # 1 1
Tegumentum striatum # 1 # 1 # # # # 2 1 5
Rhagodiscaceae Percivalia fenestrata 1 1 1 1 1 # # #
Rhagodiscus angustus 14 16 8 14 17 14 11 5 11 8 6 15 11 9 8
Rhagodiscus asper 90 52 34 56 34 57 40 31 43 37 43 51 67 35 44
Rhagodiscus dekaenelii # # #
Rhagodiscus gallagheri 1 # 1 1 3
Rhagodiscus robustus # # # # # 1
Rhagodiscus sp. 4 19 1 1 7 8 8 11 21 8 13 13
Stephanolithiales Stephanolithiaceae Rotelapillus laffittei 1 2 2 5 2 1 1 # 3 3 2 2 # 3
Stoverius achylosus 1
Stradnerlithus geometricus 1
Stradnerlithus rhombicus 1 1 #
Podorhabdales Axopodorhabdaceae Axopodorhabdus dietzmannii 1 1 4 # 1 1 # 1 # 3 1 2
Ethmorhabdus hauterivianus 3 1 1 1 1 1
Tetrapodorhabdus coptensis 1 1 1 1 # 3 1 # # # 1 # 4
Biscutaceae Biscutum constans 27 35 12 28 30 15 40 11 6 18 35 30 40 39 19
Discorhabdus ignotus 19 28 17 17 13 20 28 14 9 18 26 25 25 18 29
Cruciplacolithus haye 3 # 1 1 # # 1 4
Sollasites sp. 1 1 1 2 # 2 2 #
Cretarhabdaceae Cretarhabdus sp. 3 3 2 9 5 1 1 3 4 4 1 1 1
Cretarhabdus conicus # # 3 # # # 1 2 # 7 # # 1 # #
Flabellites oblongus 1 1 1 # # #

337
(continued on next page)
Table 1 (continued )

338
Systematic Sample number: 239 240 241 242 243 244 245 246 247 248 249 250 251 252 253
Helenea chiastia 2 1 3 # 2 3 1 1 3 1 1 5
Pickelhaube furtiva 1 1 1
Retecapsa sp. # 3 1 3 1 2 1 # 2 2 1
Retecapsa angustiforata 2
Retecapsa surirella # 1 # 1 1 # # # # # # #
Tubodiscaceae Manivitella spp. 1 # 1 1 1 # # # # # # #
Tubodiscus burnettiae 3 8 5 2 4 5 9 5 10 5 8 4 13 2 8
Tubodiscus jurapelagicus 1 # 1 # # #
Watznaueriales Watznaueriaceae Cyclagelosphaera margerelii 2 # # # # # 4 #
Cyclagelosphaera sp. 1
Cylindralithus nudus 3 6 # 5 1 3 5 8 2 6 5 5 3 3 1
Diazomatolithus lehmanii 3 # # 2 3 3 3 5 4 4 7 13 2
Watznaueria barnesiae 86 74 181 113 104 126 75 104 112 78 68 45 56 67 83
Watznaueria biporta 2 4 1 1 2 # 6 1 2 1 3 2
Watznaueria britannica # # # # # 3 # # # #
Watznaueria fossacincta 16 15 31 10 9 17 18 57 21 35 9 2 9 8 20
Watznaueria ovata 3 2 5 5 9 2 2 9 16 8 6 3 8 4
Arkhangelskiales Arkhangelskiellaceae Broinsonia matalosa # 2 1 3 # 1 2 1 1 # # 3

A. Mahanipour et al. / Cretaceous Research 32 (2011) 331e356

Uncertain Haqius circcumradiatus # 1 1 2 3 1 # # 1 # 1 1 2
Heterococcoliths Haqius ellipticus # # #
Holococcoliths Calyptrosphaeraceae Calculites spp. 4 3 # 3 9 8 4 6 4 9 1 3 2 3 11
Nannoliths Braarudosphaeraceae Braarudosphaera africana
Braarudosphaera # # 3 1 # # 1 # 2
hockwoldensis
Braarudosphaera regularis 1
Braarudosphaera sp. # 1 3 3 3 4 3 2 2 2
Micrantholithus obtusus 4 2 1 8 7 3 8 2 5 10 6 3 1 23 3
Micrantholithus hoschulzii # # # # 1 2 # # 1 # # 1
Nannoconaceae N. abundance # #
Nannoconus boneti 1 #
Nannoconus bucheri 2 # # # 1 2 2 2 4 4 1
N. carniolensis
Nannoconus circularis 1 # # # 2 # 1 # #
Nannoconus elongatus # # # 5 3 5 4 4 4 10 8 13 3 3
Nannoconus fragilis 2 2 1 #
Nannoconus grandis #
Nannoconus globulus 1 1 1 1 # 1
Nannoconus inconspicuus 1 1
Nannoconus inornatus #
Nannoconus kamptneri # 6 # 1 1 # 7 4
Nannoconus minutus 5 24 # 10 6 6 24 14 12 15 9 17 11 19 5
Nannoconus multicadus # #
Nannoconus quadriangulus 1
Nannoconus truitti 1 # 1 2 3 4 4 # 1 1 3 3 2 2
Nannoconus truitti frequens 1 #
Nannoconus truitti 1 #
rectangularis
Nannoconus vocontiensis # # # # 1 1
Nannoconus wassallii # 1 # # #
Nannoconus sp. 11 24 1 2 4 9 28 9 11 17 18 34 19 22 3
Polycyclolithaceae Eprolithus spp. # 2 1 # 1 3 # # # # 1
Radiolithus planus 1 1 # # # 1 1 1
Uncertain Rucinolithus irregularis 1 # 1 1 # 2 # # # 1
polycycloliths Rucinolithus sp. # 2 1 8 6 1 2 2 1 2
Hayesites spp. 1 1 # 1
Microrhabdulaceae Lithraphidites 37 27 21 22 24 37 19 28 16 20 14 19 16 14 29
carniolensis
Systematic Sample number: 254 255 256 257 258 259 260 261 262 263 264 265 266 267 268
c
Absolute abundance 543.6 429.8 426.4 496.9 326.4 178.7 232.9 384.3 156.0 697.8 700.5 740.2 921.2 812.8 821.4
(milion Ind./g Sed.)
Species richness 61.0 62.0 61.0 61.0 47.0 42.0 54.0 55.0 41.0 40.0 56.0 43.0 52.0 56.0 63.0
Shannon Index 2.9 3.1 3.0 3.0 2.8 2.8 2.9 3.0 2.2 2.8 3.1 2.4 2.5 2.8 3.1
Evenness 0.7 0.8 0.7 0.7 0.7 0.7 0.7 0.7 0.6 0.8 0.8 0.6 0.6 0.7 0.7
E2- E2- E2- E2-
Preservation E1 E1 E1 E1 E2- E1 E1 E1 E2 E2 E1 E1 E1 E1 E1
% calcium carbonate 37 37 45 41 23 29 29 22 15 15 64 22 35 31 36
Order Family Total number of species 348 332 341 350 341 314 298 338 323 335 320 349 325 333 326
in each sample
Eiffel lithales Chiastozygaceae Bukrylithus ambiguus 2 1 4 2 2 # 2 2 5 3 # 1 1
Chiastozygus spp. 1 1 # # 1 # # # #
Loxolithus armilla 2 1 2 5 1 1
Staurolithites crux 1 # 3 # 2 1 #
Staurolithites glabra # # 1 1
Staurolithites mutterlosei # # 1
Staurolithites stradneri 1 # # 1 3 1 # 1 # # 1
Staurolithites sp. 3 3 # 3 3 2 2 1 3 1 2 # 3 4

A. Mahanipour et al. / Cretaceous Research 32 (2011) 331e356

Tranolithus gabalus 1 #
Zeugrhabdotus clarus # # # 1 #
Zeugrhabdotus diplogrammus # 5 # 1 13 2 3 # # # # # # 1
Zeugrhabdotus elegans # #
Zeugrhabdotus cf. Z. embergeri # # 1 1 # 1 # 1
Zeugrhabdotus embergeri # 1 1 # # # 7 2 # # 1
Zeugrhabdotus erectus 1 1 1 1 1 1 # 2 # 1
Zeugrhabdotus scutula #
Zeugrhabdotus trivectis 1 1 # 1 #
Zeugrhabdotus xenotus 1 # # # # #
Zeugrhabdotus sp. 14 6 4 4 11 8 1 10 2 19 7 7 3 14 11
Eiffellithaceae Tegumentum octiformis # # # #
Tegumentum striatum # 2 # # 1 # 2
Rhagodiscaceae Percivalia fenestrata 1 # # 1 # 2 # # # # 1 2 1
Rhagodiscus angustus 11 15 8 8 3 1 7 12 2 11 7 4 6 14 7
Rhagodiscus asper 92 51 31 49 49 41 61 57 36 42 47 88 65 71 53
Rhagodiscus dekaenelii # 2 #
Rhagodiscus gallagheri 1 1 1 1
Rhagodiscus robustus 1 1 # # #
Rhagodiscus sp. 8 7 4 10 1 9 17 9 8 1 5 18
Stephanolithiales Stephanolithiaceae Rotelapillus laffittei 2 2 3 4 5 1 3 2 1 2 5 2 4 # 8
Stoverius achylosus 1 # 1
Stradnerlithus geometricus 2
Stradnerlithus rhombicus 5
Pod orhab dales Axopodorhabdaceae Axopodorhabdus dietzmannii 1 2 1 1 1 2 2 1 # 2 1 2 # 3
Ethmorhabdus hauterivianus 1 1 1 1 1 2
Tetrapodorhabdus coptensis 1 1 3 1 2 # 1 5 3 2 2
Biscutaceae Biscutum constans 28 40 76 61 38 10 12 27 8 32 27 19 23 29 12
Discorhabdus ignotus 21 19 33 31 50 12 11 18 6 40 32 22 16 28 17
Cruciplacolithus haye 2 # # 4 2 1
Sollasites sp. 3 1 1 # 5 1 1 # 1
Cretarhabdaceae Cretarhabdus sp. 4 2 1 5 4 1 1 # # 2 3 1 3 2
Cretarhabdus conicus # # # # 1 # # # # # # 2 2 1 1
Flabellites oblongus 1 # 1 4 3 1 #
Helenea chiastia 1 # 3 # 1 2 3 4 5 1 # 2 2
Pickelhaube furtiva 1
Retecapsa sp. # 2 2 1 2 1 # 2 1 1
Retecapsa angustiforata 1 1 1
(continued on next page)

339
 340
Table 1 (continued )

Systematic Sample number: 254 255 256 257 258 259 260 261 262 263 264 265 266 267 268
Retecapsa surirella # # # # # 1 # # # # # #
Tubodiscaceae Manivitella spp. 1 # # # # 1 1 # 1 # 2 2
Tubodiscus burnettiae # 8 7 1 10 5 8 8 3 3 5 4 2 5 3
Tubodiscus jurapelagicus # # # #
Watznaueriales Watznaueriaceae Cyclagelosphaera margerelii 1 2 # 2 # #
Cyclagelosphaera sp. 2 1
Cylindralithus nudus # 5 4 5 # 3 4 4 2 3 1 2 2 3
Diazomatolithus lehmanii # 2 4 3 # 4 # 1 # # 7 2 # 1
Watznaueria barnesiae 57 54 43 37 60 81 69 79 146 74 41 110 117 71 72
Watznaueria biporta 1 5 1 1 2 2 5 3 3 1 1 1 5
Watznaueria britannica # # 1 1 2 6 # 1 1
Watznaueria fossacincta 4 6 8 6 15 4 22 14 46 15 13 14 10 2 29
Watznaueria ovata 5 5 3 1 2 1 4 7 7 11 3 2 1 7 1
Arkh angel ski ales Arkhangelskiellaceae Broinsonia matalosa # 2 # # 4 2 5 1 2 2 9
Uncertain Haqius circcumradiatus # 1 1 # # 1 # 1 # #
Heterococcoliths Haqius ellipticus 1 #
Holococcoliths Calyptrosphaeraceae Calculites spp. 1 2 1 1 5 4 6 6 17 25 10 10 3 5 11

A. Mahanipour et al. / Cretaceous Research 32 (2011) 331e356

Nannoliths Braarudosphaeraceae Braarudosphaera africana # #
Braarudosphaera 1 1 1 1 1 10 # # # #
hockwoldensis
Braarudosphaera #
regularis
Braarudosphaera sp. 2 6 4 6 5 4 1 1 1 1
Micrantholithus obtusus 14 27 18 16 4 18 7 7 4 # # 1 2 # #
Micrantholithus hoschulzii 1 1 2 1 # 2 1 1 2 1 1
Nannoconaceae N. abundance
Nannoconus boneti # 1 1
Nannoconus bucheri 3 1 2 2 1 1 4 # 1 1 1
N. carniolensis
Nannoconus circularis 1 # # # # 1 # # #
Nannoconus elongatus 2 # 1 # 1 6 # 1 2 # # # 1
Nannoconus fragilis # 1 4
Nannoconus grandis #
Nannoconus globulus 1 # # # 2 1 #
Nannoconus inconspicuus 1 3 1 6 2
Nannoconus inornatus
Nannoconus kamptneri 3 # 3 1 # 2 # #
Nannoconus minutus 7 3 13 8 4 15 7 4 7 8 4 1 4
Nannoconus multicadus #
Nannoconus quadriangulus 1 1
Nannoconus truitti 3 2 1 3 4 # 3 # 4 1
Nannoconus truitti #
frequens
Nannoconus truitti # # # 1 #
rectangularis
Nannoconus vocontiensis 2 1 # 1 # # # #
Nannoconus wassallii 1 1 # 1 # # #
Nannoconus sp. 21 17 28 33 3 32 12 6 5 38 1 5 11 15
Polycyclolithaceae Eprolithus spp. # 1 # # # # 2 1 # # 1 #
Radiolithus planus # # # 1 #
Uncertain Rucinolithus irregularis 1 1 1 1 2 1
polycycloliths Rucinolithus sp. 1 1 # 1
Hayesites spp.
Microrhabdulaceae Lithraphidites 15 19 15 22 28 21 30 20 10 16 15 30 34 29 17
carniolensis
Systematic Sample number: 269 270 271 272 273 275 276 277 278 279 280 281 282 283
d
Absolute abundance 1234.3 843.7 1318.5 1308.1 873.7 1195.2 395.2 443.9 882.8 438.3 1028.9 805.7 1463.3 1061.1
(Milion Ind./g Sed.)
Species richness 55.0 59.0 49.0 44.0 65.0 56.0 66.0 54.0 40.0 56.0 47.0 53.0 53.0 64.0
Shannon Index 2.7 2.8 2.7 2.6 3.0 2.6 3.1 3.0 2.9 2.9 2.6 2.8 2.7 2.8
Evenness 0.7 0.7 0.7 0.7 0.7 0.6 0.8 0.8 0.8 0.7 0.7 0.7 0.7 0.7
Preservation E1 E1 E2-E1 E2-E1 E1 E1 E1 E1 E1 E1 E2-E1 E1 E2-E1 E1
% calcium carbonate 53 38 16 15 42 28 53 67 52 55 28 22 26 30
Order Family Total number of 323 335 342 321 328 364 326 325 348 321 315 343 309 316
species in each sample
Chiastozygaceae Bukrylithus ambiguus 1 3 1 4 4 3 # 1 3 3 5 2 2
Chiastozygus spp. # # 1 1 # # # # #
Loxolithus armilla # 8 1 # 2 6 1 2 1 1
Staurolithites crux # # 1 1 # # # # #
Staurolithites glabra # 1 # # # # # #
Staurolithites mutterlosei # #
Staurolithites stradneri # 2 # # 1 # # # 1 # #
Staurolithites sp. 3 9 13 10 6 3 2 3 5 5 2 3 4 8
Tranolithus gabalus # # 1

A. Mahanipour et al. / Cretaceous Research 32 (2011) 331e356

Zeugrhabdotus clarus # 1 # #
Zeugrhabdotus # # 2 # 1 1 1 # 2 # # #
diplogrammus
Zeugrhabdotus elegans # #
Zeugrhabdotus # # # #
cf. Z. embergeri
Zeugrhabdotus embergeri # 1 # 1 # 1 # # # #
Zeugrhabdotus erectus # 1 1 4 2 1 1 2 #
Zeugrhabdotus scutula #
Zeugrhabdotus trivectis # # # # 2 3
Zeugrhabdotus xenotus 1 # 2 # # # # # # # 1
Zeugrhabdotus sp. 15 14 11 58 4 4 4 7 7 6 19 32 13 19
Eiffellithaceae Tegumentum octiformis 2 # 1 # 1 #
Tegumentum striatum 1 # # 1 3 1 2 # # #
Rhagodiscaceae Percivalia fenestrata 1 # # 2 1 1
Rhagodiscus angustus # 4 11 10 2 1 13 5 10 3 5 5 4 5
Rhagodiscus asper 53 55 42 36 34 64 25 37 35 64 35 31 40 42
Rhagodiscus dekaenelii # # # #
Rhagodiscus gallagheri
Rhagodiscus robustus # # # #
Rhagodiscus sp. 2 12 11 3 9 7 8 15 24 9 7 10 5 7
Stephanolithiales Stephanolithiaceae Rotelapillus laffittei 1 2 3 3 4 4 2 1 1 5 # 3 5 2
Stoverius achylosus
Stradnerlithus geometricus 1 1
Stradnerlithus rhombicus
Podorhabdales Axopodorhabdaceae Axopodorhabdus dietzmannii 1 1 4 # 5 3 # 1 # # # # #
Ethmorhabdus hauterivianus # 2 4 2 #
Tetrapodorhabdus coptensis 1 1 # # 1 1 # # 1 # 1 1 #
Biscutaceae Biscutum constans 37 26 31 18 14 24 57 38 32 33 18 15 22 24
Discorhabdus ignotus 29 22 35 23 18 24 36 31 30 29 33 33 31 23
Cruciplacolithus haye 7 2 # 2 1 # 1 #
Sollasites sp. 2 2 2 4 2 2 2 1 5 # 1
Cretarhabdaceae Cretarhabdus sp. 2 5 5 1 3 3 5 2 # 3 1 7 3
Cretarhabdus conicus # 2 # 1 1 2 1 # # # 1 # # 2
Flabellites oblongus # # 1 2 1 # 1 1 #
Helenea chiastia 1 2 1 3 5 2 1 2 1 # 2 1 #
Pickelhaube furtiva # # #
Retecapsa sp. # 1 # 3 1 2 4 4 1
Retecapsa angustiforata #

341
(continued on next page)
 342
Table 1 (continued )

Systematic Sample number: 269 270 271 272 273 275 276 277 278 279 280 281 282 283
Retecapsa surirella # # # 1 # # # 1 # #
Tubodiscaceae Manivitella spp. # # 1 # 1 2 1 1 1 # # 1
Tubodiscus burnettiae 6 2 1 1 2 3 2 6 5 3 4 6 15 3
Tubodiscus jurapelagicus #
Watznaueriales Watznaueriaceae Cyclagelosphaera 1 # # #
margerelii
Cyclagelosphaera sp. 2 2
Cylindralithus nudus 2 # 2 # 5 1 2 9 1 4 6 1
Diazomatolithus lehmanii 1 # # # 2 # 4 5 5 3 # # # #
Watznaueria barnesiae 87 95 94 98 92 135 39 72 43 57 93 108 105 113
Watznaueria biporta 3 4 4 4 1 6 4 1 1 3 1
Watznaueria britannica 1 2 1 1 # 1
Watznaueria fossacincta 11 13 11 17 7 12 12 4 19 40 17 12 6
Watznaueria ovata 1 1 1 5 1 1 2 1 3 1 1 1 # 4
Arkhangelskiales Arkhangelskiellaceae Broinsonia matalosa # 3 5 4 # 1 1 2
Uncertain Haqius circcumradiatus # 1 # 2 # 1 1 1 1 # # #

A. Mahanipour et al. / Cretaceous Research 32 (2011) 331e356

Heterococcoliths Haqius ellipticus
Holococcoliths Calyptrosphaeraceae Calculites spp. 16 19 11 1 5 11 2 2 2 5 12 9 8 5
Nannoliths Braarudosphaeraceae Braarudosphaera africana # # # # # # #
Braarudosphaera # # # # 1 2 # 1
hockwoldensis
Braarudosphaera regularis # #
Braarudosphaera sp. # # 1 3 4 1 #
Micrantholithus obtusus 1 1 # # 1 # # 2 1 3 # # 4
Micrantholithus hoschulzii # 1 2 1 #
Nannoconaceae N. abundance # # #
Nannoconus boneti # 1
Nannoconus bucheri 1 # # 6 4 5 2 # 1 #
N. carniolensis
Nannoconus circularis # # # 1 # 1 1 # #
Nannoconus elongatus # # # 1 1 2 2 3 1 #
Nannoconus fragilis 1 # 1 # #
Nannoconus grandis 1
Nannoconus globulus 2 1 3 1 1
Nannoconus inconspicuus 1 #
Nannoconus inornatus #
Nannoconus kamptneri # # 1 2 # #
Nannoconus minutus 12 2 15 1 4 5 21 3 # 3 1 1
Nannoconus multicadus
Nannoconus quadriangulus # 1 1
Nannoconus truitti 2 2 # 3 2 2 2 # # 1
Nannoconus truitti frequens # #
Nannoconus truitti # # 3 3 # 1 # #
rectangularis
Nannoconus vocontiensis # # 1 1 #
Nannoconus wassallii # 1 2 3 1 # #
Nannoconus sp. 17 4 # # 38 6 36 25 35 24 4 7 2 7
Polycyclolithaceae Eprolithus spp. # 1 # # 1 2 # # # # # #
Radiolithus planus 1 # # #
Uncertain Rucinolithus irregularis # # 1 1 # # 1 2 1
polycycloliths Rucinolithus sp. # 3 # 1 1
Hayesites spp.
Microrhabdulaceae Lithraphidites 13 23 38 16 15 25 20 12 48 13 20 21 14 23
carniolensis
Systematic Sample number: 284 285 286 287 288
e
Absolute abundance 843.6 638.1 164.1 590.6 311.0
(milion Ind./g Sed.)
Species richness 62.0 60.0 56.0 58.0 50.0
Shannon Index 2.9 3.1 2.9 2.8 2.9
Evenness 0.7 0.7 0.7 0.7 0.7
Preservation E1 E1 E1 E2-E1 E1
% calcium carbonate 36 44 53 29 51
Order Family Total number of species 324 325 339 321 341
in each sample
Eiffellithales Chiastozygaceae Bukrylithus ambiguus 6 1 # # #
Chiastozygus spp. 1 # #
Loxolithus armilla 1 1 2
Staurolithites crux 1 #
Staurolithites glabra 1 1 #
Staurolithites mutterlosei # #
Staurolithites stradneri # # # 1
Staurolithites sp. 2 11 3 4
Tranolithus gabalus # #

A. Mahanipour et al. / Cretaceous Research 32 (2011) 331e356

Zeugrhabdotus clarus # 1 1
Zeugrhabdotus diplogrammus # 1 1 #
Zeugrhabdotus elegans
Zeugrhabdotus cf. # #
Z. embergeri
Zeugrhabdotus embergeri # 1
Zeugrhabdotus erectus 1 1 5
Zeugrhabdotus scutula #
Zeugrhabdotus trivectis # # 1 #
Zeugrhabdotus xenotus 1 # #
Zeugrhabdotus sp. 20 11 7 12 7
Eiffellithaceae Tegumentum octiformis # #
Tegumentum striatum # # 1
Rhagodiscaceae Percivalia fenestrata 1 # # 1 1
Rhagodiscus angustus 3 4 1 2 5
Rhagodiscus asper 47 48 59 52 63
Rhagodiscus dekaenelii # # #
Rhagodiscus gallagheri #
Rhagodiscus robustus #
Rhagodiscus sp. 9 4 3 7 1
Stephanolithiales Stephanolithiaceae Rotelapillus laffittei 2 6 # 1 2
Stoverius achylosus
Stradnerlithus geometricus
Stradnerlithus rhombicus
Podorhabdales Axopodorhabdaceae Axopodorhabdus dietzmannii 4 1 2 2
Ethmorhabdus hauterivianus 1 # # 1
Tetrapodorhabdus coptensis # # 4 8
Biscutaceae Biscutum constans 38 42 31 21 22
Discorhabdus ignotus 29 37 38 36 28
Cruciplacolithus haye 1 #
Sollasites sp. 1 1
Cretarhabdaceae Cretarhabdus sp. 3 2 1 4 2
Cretarhabdus conicus 1 1 5 4 2
Flabellites oblongus # #
Helenea chiastia # 5 4 3 1
Pickelhaube furtiva
Retecapsa sp. 1 1 # 1 1
Retecapsa angustiforata
Retecapsa surirella # 1 2 # #

343
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Table 1 (continued )

344
Systematic Sample number: 284 285 286 287 288
Tubodiscaceae Manivitella spp. # 2 # # 3
Tubodiscus burnettiae 6 2 # 2 6
Tubodiscus jurapelagicus #
Watznaueriales Watznaueriaceae Cyclagelosphaera margerelii # #
Cyclagelosphaera sp. 2
Cylindralithus nudus 2 7 4 3 #
Diazomatolithus lehmanii 2 1 1 # 4
Watznaueria barnesiae 84 68 72 94 67
Watznaueria biporta 3 3 1
Watznaueria britannica # # 1 1
Watznaueria fossacincta 7 9 11 12 10
Watznaueria ovata 5 2 3 5
Arkhangelskiales Arkhangelskiellaceae Broinsonia matalosa
Uncertain Haqius circumradiatus 1 # 2 1
Heterococcoliths Haqius ellipticus
Holococcoliths Calyptrosphaeraceae Calculites spp. 2 5 5 3 1
Nannoliths Braarudosphaeraceae Braarudosphaera africana # # 1
Braarudosphaera # # 1 # 2

A. Mahanipour et al. / Cretaceous Research 32 (2011) 331e356

hockwoldensis
Braarudosphaera regularis
Braarudosphaera sp. 1 5 3
Micrantholithus obtusus 1 # 3 1 1
Micrantholithus hoschulzii 1 #
Nannoconaceae N. abundance 1
Nannoconus boneti
Nannoconus bucheri # 1 #
N. carniolensis
Nannoconus circularis # 2 1
Nannoconus elongatus # 1 2 2 5
Nannoconus fragilis 2 1
Nannoconus grandis 1
Nannoconus globulus # # 2 # #
Nannoconus inconspicuus # 3 2 1 1
Nannoconus inornatus
Nannoconus kamptneri # # # #
Nannoconus minutus 10 7 8 3 16
Nannoconus multicadus
Nannoconus quadriangulus # 5 1
Nannoconus truitti 2 # 8 # 18
Nannoconus truitti frequens #
Nannoconus truitti # 2
rectangularis
Nannoconus vocontiensis # # #
Nannoconus wassallii # #
Nannoconus sp. 13 10 17 7 14
Polycyclolithaceae Eprolithus spp. 1 # # 2 1
Radiolithus planus # #
Uncertain Rucinolithus irregularis 1 3 2 1 5
polycycloliths Rucinolithus sp. # 1 # 4
Hayesites spp.
Microrhabdulaceae Lithraphidites carniolensis 12 14 22 22 22
 A. Mahanipour et al. / Cretaceous Research 32 (2011) 331e356
(Fig. 2) using the gravity settling technique (e.g. Bown and Young,
1998). Out of these, 66 samples located in the upper part of the
section (Sanganeh Formation, 1385e1668 m; samples 223e288)
showing well preserved calcareous nannofossils were processed
using the settling technique of Geisen et al. (1999) in order to
perform absolute and relative abundance counts. Preservation of
nannofossils was evaluated qualitatively by using visual criteria
concerning the degree of etching (E1eE3) and overgrowth (O1eO3)
after Roth (1973).
The microscope work was performed on an Olympus BH-2 light
microscope with a magnification 1500 for biostratigraphy and
palaeoecological studies. In order to observe biostratigraphical
index taxa, 2e4 random transverses of each gravity slide were
examined. In the 66 settling slides a minimum of 300 specimens
was counted in each sample. Bibliographic references for the
calcareous nannofossils are given in Perch-Nielsen (1985) and
Bown (1998). Based on nannofossil counts, the nannofossil absolute
abundance, the species richness and the relative abundance of
nannofossils (the number of species observed by counting at least
300 specimens), the Shannon Index and evenness were calculated
according to Shannon and Weaver (1949) using the statistical
software MVSP version 3.11 (Fig. 3; Table 1).
3.2. Calcium carbonate
Eighty samples from 755 to 1668 m (samples 130e288) were
analysed for their CaCO3 content (Fig. 4) to check the sufficiency of
calcium carbonate for stable isotope analysis (d13Ccarb, d18O) in the
samples. The CaCO3 content was measured using the “Müller-Bombe”
(Müller and Gastner, 1971). The bulk rock samples (1.0 g), which were
dried and homogenised, were dissolved with HCl (15%) in a glass
bottle equipped with a manometer. The CaCO3 content was calculated
by the relative gas pressure increase inside the glass bottle.
3.3. Stable isotopes
A total of 80 bulk rock samples covering the interval from 755 to
1668 m (samples 130e288) was measured for stable isotopes (d13C,
d18O) to see if the OAE 1a is recorded in this part of the section. The
results of the stable isotopes data (d13C, d18O) are illustrated in
Fig. 4. The measurements were performed using a Finnigan MAT
251 mass spectrometer, coupled to the Carbo Kiel device at the
Leibniz-Labor for Radiometric Dating and Stable Isotope Research
at Kiel University, Germany.
4. Results
4.1. Nannofossil preservation, diversity and abundance
Preservation of nannofossils in the Takal Kuh section is poor in
the marlstones, argillaceous limestones and limestones of the
Sarcheshmeh Formation (0e1385 m; samples 1e222). It is
moderate to poor in the first 50 m of the overlying Sanganeh
Formation (1385e1435 m; samples 223e237) and good to
moderate in the marls of the upper part of the Sanganeh Formation
(1435e1668 m; samples 238e288). Because of the poor preserva-
tion of the nannofossils, the samples from the Sarcheshmeh
Formation were not used for the palaeoecological investigation and
the palaeoecological interpretations herein were based solely on
samples 223e288 from the overlying Sanganeh Formation.
Species richness varies from 0 to 47 species per sample in the
Sarcheshmeh Formation (0e1385 m; samples 1e222), while in the
Sanganeh Formation (1385e1668 m; samples 223e288) it varies
from 21 to 66 (Fig. 3). The Shannon Index and the evenness fluc-
tuate considerably throughout the succession. In the Sanganeh
345
Formation (1385e1668 m; samples 223e288), the Shannon Index
ranges from 1.1 (sample 223) to 3.1 (sample 276), the evenness from
0.3 (sample 223) to 0.7 (sample 264) as illustrated in Fig. 3.
The absolute abundance values have only been determined in the
Sanganeh Formation (1385e1668 m; samples 223e288); they vary
between 0.2 and 1.5 billion specimens/g sediment. The highest
values were recorded from samples 263e286 (Figs. 3 and 5; Table 1).
4.2. Nannofossil biostratigraphy
Nannofossil zonal schemes for the BarremianeAptian interval
have been proposed for the Tethyan Realm by Manivit (1971),
Thierstein (1971, 1973), Sissingh (1977), Roth (1978), Perch-Nielsen
(1979) and Bralower et al. (1993, 1995). We have applied the zona-
tion of Roth (1978) modified by Bralower et al. (1993, 1995).
According to this scheme, the Takal Kuh sediments cover the
Upper BarremianeLower Aptian interval (Fig. 2). The last occurrence
(LO) of Nannoconus bermudezi (Late Barremian; Perch-Nielsen,1979)
is recorded at 450 m (sample 68), while the first occurrence (FO) of
Rucinolithus irregularis (latest Barremian; Erba, 1996) is at 575 m
(sample 92). At low latitudes, the FO of R. irregularis is the most
reliable nannofossil event and the closest to the Barremian/Aptian
boundary (Channell and Erba, 1992; Coccioni et al., 1992; Erba, 1994;
Aguado et al., 1995; Channell et al., 1995; Erba, 1996). Thus the
lowermost part of the section up to the FO of R. irregularis (0e575 m;
samples 1e92) can be assigned to the Upper Barremian Watznaueria
oblonga Zone (NC5). The FO of Rhagodiscus angustus was observed at
1300 m (sample 210). Eprolithus floralis has its FO at 1415 m (sample
230), about 115 m above the FO of R. angustus. We have used the FO
of E. floralis to define the top of the NC6 Zone as suggested by
Bralower et al. (1993) and the interval between 575 m (sample 92)
and 1415 m (sample 230) is thus assigned to the Chiastozygus lit-
terarius Zone (NC6). The interval from 1415 m up to the top of the
section is assigned to the R. angustus Zone (NC7), Subzone NC7A
(Bralower et al., 1993), which is defined as the interval between the
FO of E. floralis and the LO of Micrantholithus spp. The succession
from the FO of E. floralis to the top of the section can be included in
this subzone, because Micrantholithus spp. is present up to the top of
the section. Fig. 6 shows the biostratigraphy and lithostratigraphy of
the Takal Kuh section while the biozonation scheme for the Barre-
mianeAptian interval is illustrated in Fig. 7.
4.3. Palaeoecology
The calcareous nannofossils of the Takal Kuh section are domi-
nated by cosmopolitan taxa, together with taxa more closely linked
to the Tethyan Realm. The most common species which have been
considered for palaeoecological studies are Watznaueria barnesiae/
Watznaueria fossacincta (mean relative abundance 33%), Rhago-
discus asper (mean 14%), Nannoconus spp. (mean 7%), Biscutum
constans (mean 7%), Discorhabdus ignotus (mean 6%), Zeugrhabdotus
spp. (mean 4%) and the pentalith forms, Micrantholithus spp. and
Braarudosphaera spp. (mean 2%). The vertical variation in relative
abundances of these taxa is given in Fig. 5, with data counts pre-
sented in Table 1.
W. barnesiae (Fig. 8A) and W. fossacincta are grouped with each
other as they represent end-members of a morphological
continuum (Lees et al., 2004; Bornemann and Mutterlose, 2006).
The relative abundance of W. barnesiae/W. fossacincta varies from 12
to 82% but in about three-quarters of the samples it is less than 40%.
R. asper is the second most common species, with relative
abundances varying between 4 and 26%. The relative abundance of
R. asper does not show any significant variation throughout the
observed interval (Fig. 5).
Fig. 4. Chemo and biostratigraphy of the OAE 1a interval from the Takal Kuh section (NE Iran) together with carbonate calcium measurement. The d13Ccarb of bulk carbonate carbon
reported in per mil versus VPDB. Labels C2eC8 indicate chemostratigraphic segments. Segment C1 is not identified in the studied interval. The “nannoconid crisis” interval is shaded.
Ammonite biozones after Raisossadat (2002).
 A. Mahanipour et al. / Cretaceous Research 32 (2011) 331e356
Fig. 5. Vertical distribution of CaCO3%, absolute abundance and relative abundances of the most common taxa of calcareous nannofossils in the Takal Kuh section. The “nannoconid crisis” interval is shaded.

347
348 A. Mahanipour et al. / Cretaceous Research 32 (2011) 331e356
Fig. 6. Biostratigraphy and lithostratigraphy of the Upper BarremianeLower Aptian succession in NE Iran (KDB, Takal Kuh section). NC nannofossil zones after Roth (1978); CC
nannofossil zones after Sissingh (1977). The “nannoconid crisis” interval is shaded.
Nannoconus spp. is the third most important group, with relative
abundances varying from 0 to 23%. The nannoconids observed in the
Takal Kuh section are dominated by wide canal forms. The most
frequent species are Nannoconus bucheri (Fig. 8I and J), Nannoconus
circularis, Nannoconus elongatus (Fig. 8K), Nannoconus globulus,
Nannoconus kamptneri (Fig. 8L), Nannoconus minutus (Fig. 8M),
Nannoconus truitti (Fig. 8N), Nannoconus vocontiensis (Fig. 8O) and
Nannoconus wassallii (Fig. 8P). Nannoconids are nearly absent in the
first 50 m (1385e1435 m; samples 223e237) of the Sanganeh
Formation, with relative abundances of Nannoconus spp. that do not
exceed 1%, whereas the relative abundances increase up to 23% in
samples 223e237. Nannoconids are also present in the underlying
Sarcheshmeh Formation but this formation is not discussed here.
The relative abundance of B. constans (Fig. 9L), varies from 0 to
22%, with the highest values in samples 256, 257 and 277. The
relative abundance of D. ignotus (Fig. 9M) varies from 1 to 5%
throughout the studied interval. Higher values of D. ignotus have
been observed in the upper part of the section (samples 256e258;
samples 263e288). Zeugrhabdotus spp. is represented by small
forms with a major axis smaller than 3.5 mm (e.g. Zeugrhabdotus
erectus and Zeugrhabdotus sp.). The relative abundance of Zeugr-
habdotus spp. varies from 0 to 18.4% throughout the interval, and in
most samples is below 10%. Except for samples 272 and 229, there
is no high variation in this group.
The pentalith group, comprising Micrantholithus obtusus
(Fig. 8H), Micrantholithus hoschulzii, Braarudosphaera africana,
Braarudosphaera hockwoldensis, Braarudosphaera regularis and
Braarudosphaera sp., varies from 0 to 7% in relative abundance. In
most samples they do not exceed more than 5%. Higher values have
only been observed in samples 229, 271 and 285.
4.4. Calcium carbonate
The carbonate content of the samples from the 755 m to 1668 m
interval varies from 11 to 80% (Figs. 3 and 4). The values range
between 20 and 80% from 730 (sample 130) to 1390 m (sample
224), while from 1390 (sample 224) to 1435 m (sample 236) they
range from 9 to 27% (except for sample 227, with 58%). In the upper
part of the section (1435e1668 m; samples 236e288) the
carbonate content varies between 10 and 70%.
4.5. Bulk stable isotopes
Chemostratigraphic profiles for the upper part of the Sarch-
eshmeh Formation and the whole Sanganeh Formation in the Takal
Kuh section are presented in Fig. 4. In contrast to the d13Ccarb, the
oxygen isotope values do not show any significant fluctuations
throughout the studied interval.
The carbonate carbon isotope data between 755 m and 1668 m
(samples 130e288) fluctuate between 1.32 and 5.43&, while in the
755 m (sample 130)e1380 m (sample 220) interval, i.e. coinciding
with the Late Barremianeearliest Early Aptian time interval, d13Ccarb
varies from 1.4 to 4.10&. The value is 1.88& at 1380 m (sample 220).
This is followed by an overall negative shift at 1395 m (sample 226)
with a value of 1.36& within the Lower Aptian. This negative
excursion is recorded within the uppermost part of the C. litterarius
nannofossil zone, NC6, (Fig. 4). The d13Ccarb values then show an
abrupt positive shift to 4.27& (1415 m; sample 230) which is
recorded within the transition beds between the C. litterarius (NC6)
nannofossil zone and the NC7A Subzone of the R. angustus (NC7)
nannofossil zone (Fig. 4). Thus, a positive excursion characterises the
 A. Mahanipour et al. / Cretaceous Research 32 (2011) 331e356 349

Fig. 7. Biozonation scheme of the BarremianeAptian interval.

deposits between 1415 and 1452 m (samples 230e242), with values
ranging from 3.54 to 4.68&. Fluctuating, but predominantly high
positive d13Ccarb values (between 3.1 and 5.4&) characterise the
deposits between 1452 m and 1665 m (samples 242e287). The
upper part of the succession is characterised by a second negative
shift of 1.32& at 1668 m (sample 288) in the Lower Aptian. The
negative and subsequent positive d13Ccarb excursion identified in the
section are taken to indicate Oceanic Anoxic Event 1a (OAE 1a).
5. Discussion
5.1. Nannofossils
Preservation of calcareous nannofossils is moderate to poor in
the first 50 m of the interval (Sanganeh Formation) of the Takal Kuh
section subject to palaeoecological investigation, while in the upper
part it is good to moderate. The low values of the Shannon Index
(1.1) and the evenness (0.3) in sample 223 imply high relative
abundancies of only a few species, being caused by a poor preser-
vation. The high values of both variables in samples 264 (evenness
0.7) and 276 (Shannon Index 3.1) are explained by assemblages of
high diversity and low relative abundance, reflecting good preser-
vation and stable settings.
Absolute abundances show a moderate increase towards the
upper part of the section (from 0.2 to 1.5 billion specimens/g.). The
values vary from 0.2 to 0.6 billion specimens/g in the lower parts of
the section (up to 1542 m; sample 262), which are less than the
average absolute abundance values in the upper parts. Fluctuations
in nannofossil absolute abundances may have been caused by 1)
variable sedimentation rate (e.g. Pittet and Mattioli, 2002), 2)
changes in primary productivity of calcareous nannofossils (e.g.
Giraud et al., 2009) or, 3) variable preservation of calcareous nan-
nofossils. Since the lowest absolute abundance values are recorded
from the basal shales of the Sanganeh Formation and their
Fig. 8. Calcareous nannofossils from the Lower Aptian Sanganeh Formation of the Takal Kuh section (northeast Iran). A, Watznaueria barnesiae, sample 245. B, Watznaueria bri-
tannica, sample 276. C, Watznaueria ovata, sample 235. D, Haqius circumradiatus, sample 284. E, Calculites percenis, sample 234. F, Braarudosphaera africana, sample 283. G,
Braarudosphaera hockwoldensis, sample 288. H, Micrantholithus sp., sample 272. I, J, Nannoconus bucheri, sample 251. K, Nannoconus elongatus, sample 276. L, Nannoconus kamptneri,
sample 250. M, Nannoconus minutus, sample 269. N, Nannoconus truitti, sample 269. O, Nannoconus vocontiensis, sample 279. P, Nannoconus wassallii, sample 275. Q, Eprolithus
floralis, sample 234. R, Radiolithus planus, sample 272. S, Rucinolithus irregularis, sample 288. T, Lithraphidites carniolensis, sample 280. Scale bar: 10 mm.
Fig. 9. Calcareous nannofossils from the Lower Aptian Sanganeh Formation of the Takal Kuh section (northeast Iran). A, Bukrylithus ambiguus, sample 280. B, C, Staurolithites sp.,
sample 275. D, Zeugrhabdotus embergeri, sample 273. E, Tegumentum striatum, sample 235. F, G, Percivalia fenestratea, sample 272. H, Rhagodiscus angustus, sample 235. I, J, Rha-
godiscus asper, sample 283. K, Rotelapillus laffittei, sample 235. L, Biscutum constans, sample 235. M, Discorhabdus ignotus, sample 234. N, Cretarhabdus conicus, sample 235. O,
Flabellites oblongus, sample 275. P, Helenea chiastia, sample 279. Q, Retecapsa surirella, sample 284. R, Manivitella pemmatoidea, sample 286. S, Tubodiscus burnettiae, sample 235. T,
Diazomatolithus lehmanii, sample 269. Scale bar: 10 mm
352 A. Mahanipour et al. / Cretaceous Research 32 (2011) 331e356
overlying marls, accumulation rate and dilution by clay particles
can have had no effect on absolute abundance. As both oligotrophic
and eutrophic taxa are present in the marls of the upper parts of the
Sanganeh Formation, the effect of productivity can be ruled out as
well. Thus, all these observations suggest that the highest nanno-
fossil absolute abundance values in the upper part of the Sanganeh
Formation can be the result of better preservation.
The palaeoecological affinities of the most common taxa of the
Takal Kuh section are illustrated in Table 2. W. barnesiae/W. fossa-
cincta are the most common taxa in the section, with a mean
relative abundance of 33%. W. barnesiae/W. fossacincta have a higher
value during the OAE 1a interval than in the upper part of the
section. In the Cretaceous, assemblages enriched in this dissolu-
tion-resistant genus (higher than 40%) may have been strongly
affected by diagenesis (Roth and Bowdler, 1981; Roth and
Krumbach, 1986; Premoli Silva et al., 1989; Erba et al., 1992;
Williams and Bralower, 1995). In the first 50 m of the Sanganeh
Formation of the Takal Kuh section, the greater than 40% relative
abundance of W. barnesiae/W. fossacincta can be interpreted as
a result of diagenesis.
The second most common taxon in the Takal Kuh section is R.
asper (mean 18%); other species of Rhagodiscus are rare in this
section so they have not been considered for palaeoecological
interpretation. R. asper is thought to be an indicator of warm
surface-water temperatures (Crux, 1989; Mutterlose, 1991; Erba
et al., 1992; Bischoff and Mutterlose, 1998; Hardas and
Mutterlose, 2007). Presumed cool water indicators such as Repag-
ulum parvidentatum and Seribiscutum spp. (Wise, 1988; Mutterlose
and Wise, 1990; Bown et al., 1998; Street and Bown, 2000) and R.
angustus (Crux, 1991) are extremely rare in our samples, which
suggests that warm water conditions prevailed in surface waters
during the studied time interval.
The third group of nannofossils in the Takal Kuh section
comprises Nannoconus spp. (mean 7%). Although the palae-
oecological affinities of nannoconids are still controversial, they are
usually considered as low-palaeolatitudinal, Tethyan, neritic and
warm water taxa (Roth and Krumbach, 1986; Mutterlose, 1989,
1991, 1992; Street and Bown, 2000) that lived on shallow
plateaus with low terrigenous input and in epicontinental seas,
often associated with the pentalith forms, Micrantholithus and
Braarudosphaera (Roth and Krumbach, 1986; Mutterlose, 1989,
1991, 1992).
Nannoconids are supposed to be adapted to oligotrophic envi-
ronmental settings (Busson and Nöel, 1991; Coccioni et al., 1992;
Herrle, 2003). Tremolada et al. (2006) also suggested oligotrophic
preferences for nannoconids as they show an inverse correlation
with eutrophic taxa (such as Z. erectus, Discorhabdus rotatorius and
B. constans).
Table 2
The palaeoecological affinities of the most common nannofossils that are present at
the Takal Kuh section. 1 ¼ Roth and Krumbach, 1986; 2 ¼ Premoli Silva et al., 1989;
3 ¼ Williams and Bralower, 1995; 4 ¼ Mutterlose, 1989, 1991; 5 ¼ Street and Bown,
2000; 6 ¼ Busson and Nöel, 1991; 7 ¼ Coccioni et al., 1992; 8 ¼ Herrle, 2003;
9 ¼ Tremolada et al., 2006; 10 ¼ Kessels et al., 2003; 11 ¼ Herrle et al., 2003;
12 ¼ Erba et al., 1992; 13 ¼ Crux, 1989; 14 ¼ Mutterlose and Kessels, 2000.
Nannofossils Nutrient level Temperature
Watznaueria barnesiae/ Oligotrophic1,2,3,12
Watznaueria fossacincta
Nannoconus spp. Oligotrophic6,7,8,9 Warm surface water1,4,5
Biscutum constans Eutrophic1,10,11 Cold surface water14
Discorhabdus ignotus Eutrophic1,10,11
Small forms of Eutrophic1,10,11 Cold surface water14
Zeugrhabdotus spp.
Rhagodiscus asper Oligotrophic Warm surface water13,4,12
Pentalith group Enhance nutrient5 Warm surface water4,12
In the Takal Kuh section, nannoconids are rare in the first 50 m
of the Sanganeh Formation but are present in the higher part of the
formation. The rarity of Nannoconus spp. between 1385 m (sample
223) and 1435 m (sample 237) is the most important feature of the
section. This 50 m interval is assigned to the transition between
nannofossil zone NC6 and Subzone NC7A (Early Aptian). From 1435
to 1668 m (sample 237e288), the nannoconids reappear, domi-
nated mainly by wide canal forms. The absence of Nannoconus spp.
has been observed within the Early Aptian OAE 1a of Atlantic,
Pacific, Mexican, Italian and Spanish sections (Erba, 1994; Bralower
et al., 1994; Aguado et al., 1999; Bralower et al., 1999; Luciani et al.,
2001) and in the La Bedoule succession (SE France) (Erba 1994;
Bralower et al., 1993, 1994; Moullade et al., 1998). This interval
was described for the first time by Erba (1994) from Italian sections
and was termed the “Nannoconid Crisis”, while Herrle and
Mutterlose (2003) called it “Nannoconid Crisis I” (NCI). The
change in nannofloral assemblages and the fluctuations in abun-
dance of nannoconids (nannoconid crisis) are documented world-
wide (Erba, 1994; Bersezio et al., 2002; Erba, 2004; Erba and
Tremolada, 2004) and indicate palaeoceanographic changes and
reaction of nannofloral assemblages to the perturbation of the
ocean/climate system.
Some authors (e.g. Erba, 2004) supposed that an enhanced
primary productivity and a shift to meso- and eutrophic surface-
water conditions may have caused the nannoconid crisis during the
OAE 1a interval, while others (Wissler et al., 2003) believe that
rising pCO2 (Berner, 1991) due to enhanced volcanic activity (Larson
and Erba, 1999) and a contemporaneous biocalcification crisis are
possible reasons for the nannoconid crisis. Moreover, Wissler et al.
(2003) believe that high amount of nutrients can be considered as
a local or regional factor for the nannoconid crisis and emphasis
pCO2 and the biocalcification crisis as important factors on a global
scale.
In the OAE 1a interval of the Takal Kuh section, high relative
abundances of oligotrophic taxa (W. barnesiae/W. fossacincta) and
low relative abundances of eutrophic taxa (B. constans, D. ignotus
and Zeugrhabdotus spp.) can be observed (Fig. 5). It should be noted
that the preservational state of nannofossils is poor to moderate in
the samples corresponding to the OAE 1a interval. Furthermore, the
CaCO3 content of this part of the section is very low. Thus, the data
obtained from the OAE 1a interval of the Takal Kuh section indicate
a high concentration of CO2 in the oceaneatmosphere system as
a possible cause of the nannoconid crisis.
Relative abundances of nannoconids in the Takal Kuh section
can be correlated with high calcium carbonate content. In some
parts of the section (1385e1435 m; samples 223e237), where
nannoconids are rare or less abundant, the carbonate calcium
content is low (mean 17%). In the upper part of the section
(1435e1668 m; samples 237e288), higher nannoconid abundances
coincide with higher calcium carbonate values (mean 35%).
The other group of nannofossils consists of B. constans (mean
7%), D. ignotus (mean 6%) and Zeugrhabdotus spp. (mean 4%). B.
constans and D. ignotus have lower relative abundance values in the
OAE 1a interval than in the upper part of the section. Zeugrhabdotus
spp. (with long axis of the ellipse smaller than 3.5 mm) shows some
fluctuations in relative abundance throughout the studied interval.
Taxa like Biscutum, Discorhabdus and small varieties of Zeugrhab-
dotus are considered as delicate and susceptible to dissolution (Hill,
1975; Thierstein, 1980; Roth, 1983), so an increase in diagenetic
effects may cause a decrease in the relative abundance of these taxa
and an increase in the relative abundance of the diagenetic resis-
tant taxa such as Watznaueria (Thierstein, 1980; Roth and
Krumbach, 1986; Erba, 1992). Furthermore, Biscutum, Dis-
corhabdus and Zeugrhabdotus are considered as high nutrient
indicators (Roth and Krumbach, 1986; Kessels et al., 2003; Herrle
 A. Mahanipour et al. / Cretaceous Research 32 (2011) 331e356
et al., 2003) and show an inverse abundance relationship to
Watznaueria spp. (Street and Bown, 2000; Kessels et al., 2006),
which are considered as mesotrophic taxa (Pittet and Mattioli,
2002; Giraud et al., 2006, 2009). In the OAE 1a interval in the
Takal Kuh section, the relative abundance of W. barnesiae/W. fos-
sacincta is higher than that of B. constans and D. ignotus, which
seems to indicate etching and dissolution. In the upper part of the
section, the enhanced abundance of eutrophic taxa coincides with
a high abundance of mesotrophic and oligotrophic taxa (Fig. 5). This
can be indicative of an environment with slightly increased
nutrient content, as suggested by Kessels et al. (2006).
Pentaliths (Micrantholithus spp., Braarudosphaera spp.) are
another group that are present in the Takal Kuh section (mean 2%).
Micrantholithus is assigned a broader latitudinal range than the
nannoconids (50 Ne50 S), but has a similar neritic distribution
(Street and Bown, 2000).
5.2. Calcium carbonate and carbon isotope stratigraphy
The CaCO3 content (except for sample 227 at 1402 m) is lowest
(mean 17%) in the OAE 1a interval in the Takal Kuh section. In the
upper part of the section (1435e1668 m; samples 236e288) the
carbonate content is higher (mean 35%).
Numerous authors have demonstrated that carbon isotope
geochemistry is a powerful tool in the stratigraphy of pelagic
successions (Scholle and Arthur, 1980; Schlanger et al., 1987) as well
as their shallow water equivalents (Jenkyns, 1995; Adabi, 1997;
Ferreri et al., 1997; Heldt et al., 2008) and can be used for global
correlation. The long-term variation in isotopic composition of
carbon isotopes has been interpreted as evidence of perturbations
of the global carbon cycle (e.g. Arthur et al., 1985; Weissert et al.,
1985; Shackleton, 1987). One of the best described global carbon
isotope events occurs in the lower Aptian, which corresponds to the
Oceanic Anoxic Event 1a (OAE 1a) of Arthur et al. (1990). The
negative and the subsequent positive d13Ccarb excursion of the Early
Aptian OAE 1a represent a major perturbation of global carbon
cycling (Arthur et al., 1985), which can be used to correlate OAE 1a
on a global scale (Jenkyns, 1995; Bralower et al., 1999; Ando et al.,
2002). The negative and subsequent positive d13Ccarb excursions
recorded in the Takal Kuh section are linked to the OAE 1a as
defined by Menegatti et al. (1998) and Herrle et al. (2004).
Menegatti et al. (1998) divided the d13Ccarb curve of the Barre-
mianeAlbian into several segments (C1eC8). The (?)C2, C3, C4-C6,
C7 and C8 segments are recognised in the d13Ccarb curve of the
studied section (Fig. 4). The resolution of the d13C curve is not high
enough to clearly identify the onset of the negative excursion,
which has been assigned to the C2 segment by Menegatti et al.
(1998). A first shift towards lighter values of 1.88& can be seen,
however, at 1380 m (sample 220). This sample is thus tentatively
interpreted as the C2 segment, which is followed by a negative shift
at 1395 m (sample 226; C3) with a value of 1.36&.
The d13Ccarb values of the Takal Kuh section then show an abrupt
positive shift to 4.27& at 1415 m (sample 230), which is followed
by yet another increase to 4.68& (1452 m; sample 242) (C4eC6).
The negative and subsequent positive excursions are globally
recorded and related to the OAE 1a (Menegatti et al., 1998; Erba
et al., 1999; de Gea et al., 2003). However, in the Takal Kuh
section the interval (C4eC6) does not show a step-like positive shift
from lower to higher values as described from the western Tethys
(Cismon section; Menegatti et al., 1998). The maximum positive
d13Ccarb excursion of the section precedes the interval with rela-
tively constant values (1452e1665 m; samples 242e287) ranging
between 3.1 and 5.4& (C7). There is a pronounced negative shift to
1.32& at 1668 m (sample 288; ¼C8), similar to that in other
Tethyan successions (e.g. Menegatti et al., 1998; Weissert et al.,
353
1998; de Gea et al., 2003). The segments (?)C2, C3 and lower part
of C4eC6 coincide with the C. litterarius (NC6) nannofossil zone,
while the remaining part of the C4eC6, C7 and C8 segments are
placed in the R. angustus (NC7) nannofossil zone.
Although some minor differences exist, the trend in the d13Ccarb
values in the Takal Kuh section is consistent with other records
from the Alpine Tethys (e.g. Menegatti et al., 1998). Menegatti et al.
(1998) defined the OAE 1a chemostratigraphically as the interval
from the base of the C4 segment to the boundary between
segments C6 and C7 in the Cismon section. Heldt et al. (2008)
defined the OAE 1a as the interval from the upper part of the C2
segment to C6eC7 in the southern Tethys (Tunisia). They also
reported a sudden increase in the abundance of radiolarians and
planktonic foraminifers in this interval, indicating eutrophication of
the upper water column. Li et al. (2008) assigned segments C3
through C6 to OAE 1a. In the Takal Kuh section, the nannoconid
crisis can be observed from the (?)C2 segment up to the upper part
of the C4eC6 segments. Simultaneously, the CaCO3 content (except
for sample 227 at 1402 m) is the lowest (mean 17%) in this part of
the section. A remarkable drop in carbonate content has been
documented along with a d13C excursion in the Livello Selli (e.g.
Menegatti et al., 1998) and in other parts of the world (e.g. Bralower
et al., 1993, 1994; Wissler et al., 2003; Erba and Tremolada, 2004),
which could be considered as evidence for accelerated carbonate
dissolution by a sudden increase in carbon dioxide (Wissler et al.,
2003).
All these observations support the fact that OAE 1a is recorded
in the Takal Kuh section, which was part of the Tethys. In this
section, the OAE 1a interval ranges from the ?C2 segment to the
upper part of the C4eC6 segment (Fig. 4), which coincides with
a rarity of nannoconids and a lower calcium carbonate content.
6. Conclusions
Integrated nannofossil biostratigraphy and chemostratigraphy
of stable isotopes of the Takal Kuh section (Sarcheshmeh and
Sanganeh formations) from the Kopet Dagh area, northeast Iran
provides a high resolution stratigraphy, dating the succession as
Late BarremianeEarly Aptian. The FOs of the nannofossil zonal
markers R. irregularis and E. floralis are recorded in the section.
Accordingly, the W. oblonga Zone (NC5), C. litterarius Zone (NC6)
and the NC7A Subzone of the R. angustus (NC7) Zone are
recognised.
Palaeoecological investigations of the calcareous nannofossils in
the upper part of the section (Sanganeh Formation) document that
in the first 50 m of the interval W. barnesiae/W. fossacincta are
higher in relative abundance than B. constans, Zeugrhabdotus spp.
and D. ignotus, which indicates dissolution. Enhanced relative
abundances of the eutrophic taxa (B. constans, Zeugrhabdotus spp.
and D. ignotus) and high relative abundances of mesotrophic and
oligotrophic taxa (W. barnesiae/W. fossacincta and Nannoconus spp.
respectively) in the upper part of the section suggest a slightly
increased nutrient supply. A virtual absence of nannoconids coin-
cident with lower amounts of CaCO3 is recorded in the first 50 m of
the Sanganeh Formation (upper part of the NC6 calcareous nan-
nofossil zoneelower part of the NC7A Subzone). This may be
explained by a higher pCO2 which caused a biocalcification crisis
and the dissolution of calcium carbonate. The presence of warm
water taxa like R. asper and nannoconids as well as the virtual
absence of cool water taxa indicate warm surface waters during the
Early Aptian.
Carbon isotope data (d13Ccarb) suggest the presence of a negative
(upper part of the NC6 nannofossil zone) and a subsequent positive
excursion (at the boundary of NC6eNC7A), which are recorded
worldwide during the OAE 1a interval. The carbon isotope
354 A. Mahanipour et al. / Cretaceous Research 32 (2011) 331e356
excursion reported here coincides with the nannoconid crisis and
with sediments with low calcium carbonate contents. These
observations enable the recognition of both the presence and
impact of the OAE 1a interval in the Lower Aptian sediments of the
Kopet Dagh area in northeast Iran.
Acknowledgements
We are grateful to Liam Gallagher, Silvia Gardin and Fabienne
Giraud for having improved this paper with their suggestions and
comments. We acknowledge the editorial work of D.J. Nichols and
C.J. Wood. We would like to thank Dr. N. Raisossadat (Birjand
University, Iran) for his advice and help in the field. We are also
grateful to B. Salehipour (GeoPardazesh Petroleum Exploration
Company, Iran) for his assistance during the sampling of the
different sections during the field trip. Financial support from the
Ministry of Science, Research and Technology of Iran is greatly
appreciated. We are grateful to the Ruhr University in Bochum for
providing facilities. Prof. U. Heimhofer, Dr. P. Hardas and M. Malkoc
are thanked for their help at the Ruhr University of Bochum in
Germany. Dr. H. Legge, S. Pauly and C. Linnert improved the
manuscript by thoughtful comments.
References
Adabi, M.H., 1997. Application of carbon isotope chemostratigraphy to the Renison
dolomites, Tasmania: a Neoproterozoic age. Australian Journal of Earth Sciences
44, 767e775.
Afshar-Harb, A., 1969. A brief history of geological exploration and geology of the
Sarakhs area and the Khangiran field. Bulletin of the Iranian Petroleum Institute
37, 86e93 (in Persian).
Afshar-Harb, A., 1979. The stratigraphy, tectonics and petroleum geology of the
Kopet Dagh region, northern Iran. Unpublished Ph.D. thesis, Imperial College of
Science and Technology of London, England.
Aguado, R., Castro, J.M., Company, M., Alfonso de Gea, G., 1999. Aptian bioevents e
an integrated biostratigraphic analysis of the Almadich Formation, Inner pre-
betic Domain, SE Spain. Cretaceous Research 20, 663e683.
Aguado, R., Company, M., Sandoval, J., Tavera, J.M., 1995. Biostratigraphic events
around the BarremianeAptian boundary in the Betic Cordillera (Southern
Spain). In: Second International Symposium on Cretaceous Stage Boundaries,
Abstract Volume, p. 6.
Ando, A., Kakegawa, T., Takashima, R., Saito, T., 2002. New perspective on Aptian
carbon isotope stratigraphy: data from d13C records of terrestrial organic
matter. Geology 30, 227e230.
Arthur, M.A., 2000. Volcanic contributions to the carbon and sulfur geochemical
cycles and global change. In: Sigurdsson, H., Houghton, B., McNutt, S.R.,
Rymer, H., Stix, J. (Eds.), Encyclopedia of Volcanoes. Academic Press, pp.
1045e1056.
Arthur, M.A., Dean, W.E., Schlanger, S.O., 1985. Variations in the global carbon cycle
during the Cretaceous related to climate, volcanism, and changes in atmo-
spheric CO2. In: Sundquist, E.T., Broecker, W.S. (Eds.), The Carbon Cycle and
Atmospheric CO2: Natural Variations Archean to Present. Geophysical Mono-
graphy Series, vol. 32. AGU, Washington, D.C, pp. 504e529.
Arthur, M.A., Jenkyns, H.C., Brumsack, H.J., Schlanger, S.O., 1990. Stratigraphy,
geochemistry and paleoceanography of organic-carbon rich Cretaceous
sequences. In: Ginsburg, R.N., Beaudoin, B. (Eds.), Cretaceous Resources, Events
and Rhythms. NATO ASI Series, vol. 304. Kluwer Academic Publishers, Dor-
drecht, pp. 75e119.
Arthur, M.A., Schlanger, S.O., Jenkyns, H.C., 1987. The Cenomanian/Turonian Oceanic
Anoxic Event, II: palaeoceanographic controls on organic matter production
and preservation. In: Brooks, J., Fleet, A.J. (Eds.), Marine Petroleum Source
Rocks. Geological Society, London, Special Publications, vol. 26, pp. 401e420.
Beerling, D.J., Lomas, M.R., Gröcke, D.R., 2002. On the nature of methane gashydrate
dissociation during the Toarcian and Aptian oceanic anoxic events. American
Journal of Science 302, 28e49.
Berberian, M., King, G.C.P., 1981. Towards a paleogeography and tectonic evolution
of Iran. Canadian Journal of Earth Science 18, 210e265.
Berner, R.A., 1991. A model for atmospheric CO2 over Phanerozoic time. American
Journal of Science 291, 339e376.
Bersezio, R., Erba, E., Gorza, M., Riva, A., 2002. BerriasianeAptian black shales of the
Maiolica formation (Lombardian Basin, Southern Alps, Northern Italy): local to
global events. Palaeogeography, Palaeoclimatology, Palaeoecology 180,
253e275.
Bischoff, G., Mutterlose, J., 1998. Calcareous nannofossils of the Barremian/Aptian
boundary interval in NW Europe: biostratigraphic and palaeoecologic impli-
cations of a high resolution study. Cretaceous Research 19, 635e661.
Bornemann, A., Mutterlose, J., 2006. Size analyses of the coccolith species Biscutum
constans and Watznaueria barnesiae from the Late Albian “Niveau Breistroffer” (SE
France): taxonomic and palaeoecological implications. Geobios 39, 599e615.
Bown, P.R. (Ed.), 1998. Calcareous Nannofossil Biostratigraphy. Chapman and Hall,
London, p. 314.
Bown, P.R., 2005. Early to Mid-Cretaceous calcareous nannoplankton from the
northwest Pacific Ocean, Leg 198, Shatsky Rise. Proceedings of the Ocean
Drilling Program 198, 1e82.
Bown, P.R., Young, J.R., 1998. Techniques. In: Bown, P.R. (Ed.), Calcareous Nannofossil
Biostratigraphy. Chapman and Hall, London, pp. 16e28.
Bown, P.R., Rutledge, D., Crux, J.A., Gallagher, L.T., 1998. Lower Cretaceous. In:
Bown, P.R. (Ed.), Calcareous Nannofossil Biostratigraphy. Chapman and Hall,
Kluwer Academic Publishers, London, pp. 86e131.
Bralower, T.J., Arthur, M.A., Leckie, R.M., Sliter, W.V., Allard, D.J., Schlanger, S.O., 1994.
Timing and Paleoceanography of Oceanic Dysoxic/Anoxia in the Late Barremian
to Early Aptian (Early Cretaceous). Palaios 9, 335e369.
Bralower, T.J., CoBabe, E., Clement, B., Sliter, W.V., Osburne, C., Longoria, J., 1999. The
record of global change in mid-Cretaceous, BarremianeAlbian sections from the
Sierra Madre, northeastern Mexico. Journal of Foraminiferal Research 29,
418e437.
Bralower, T.J., Leckie, R.M., Sliter, W.V., Thierstein, H.R., 1995. An integrated Creta-
ceous microfossil biostratigraphy. SEPM Special Publication 54, 65e79.
Bralower, T.J., Sliter, W.V., Arthur, M.A., Leckie, R.M., Allard, D.J., Schlanger, S.O., 1993.
Dysoxic/anoxic episodes in the AptianeAlbian (Early Cretaceous). In:
Pringle, M., et al. (Eds.), The Mesozoic Pacific: Geology, Tectonics and Volcanism.
AGU Geophysical Monograph, vol. 77, pp. 5e37.
Busson, G., Nöel, D., 1991. Les nannoconides indicateurs environmentaux des océans
et mers épicontinentales du Jurassique terminal et du Crétacé inférieur. Oce-
anologica Acta 14, 333e356.
Channell, J.E.T., Cecca, F., Erba, E., 1995. Correlations of Hauterivian and Barremian
(Early Cretaceous) stage boundaries to polarity chrons. Earth and Planetary
Science Letters 134, 125e140.
Channell, J.E.T., Erba, E., 1992. Early Cretaceous polarity chrons CM0 to CM11
recorded in Northern Italian land sections near Brescia (Northern Italy). Earth
and Planetary Science Letters 108, 161e179.
Coccioni, R., Erba, E., Premoli Silva, I., 1992. BarremianeAptian calcareous plankton
biostratigraphy from the Gorgo a Cerbara section (Marche, central Italy) and
implications for plankton evolution. Cretaceous Research 13, 517e537.
Crux, J.A., 1989. Biostratigraphy and palaeogeographical applications of Lower
Cretaceous nannofossils from north-western Europe. In: Crux, J.A., van
Heck, S.E. (Eds.), Nannofossils and Their Applications. Ellis Horwood, Chichester,
pp. 143e211.
Crux, J.A., 1991. Albian calcareous nannofossils from the Gault Clay of Munday’s Hill
(Bedfordshire, England). Journal of Micropaleontology 10, 203e222.
de Gea, G.A., Castro, J.M., Aguado, R., Ruiz-Ortiz, P.A., Company, M., 2003. Lower
Aptian carbon isotope stratigraphy from a distal carbonate shelf setting: the Cau
section, Prebetic zone, SE Spain. Palaeogeography, Palaeoclimatology. Palae-
oecology 200, 207e219.
Erba, E., 1992. Middle Cretaceous calcareous nannofossils from the Western Pacific
(ODP Leg 129): evidence for paleoequatorial crossings. Proceedings of the
Ocean Drilling Program, Scientific Results 129, 189e201.
Erba, E., 1994. Nannofossils and superplumes: the early Aptian ‘nannoconid crisis’.
Paleoceanography 9, 483e501.
Erba, E., 1996. The Aptian stage. Bulletin de l’Institut Royal des Sciences Naturelles
de Belgique 66 (Suppl.), 31e43.
Erba, E., 2004. Calcareous nannofossils and Mesozoic oceanic anoxic events. Marine
Micropaleontology 52, 85e106.
Erba, E., Castradori, D., Guasti, G., Ripepe, M., 1992. Calcareous nannofossils and
Milankovitch cycles: the example of the Albian Gault Clay Formation (southern
England). Palaeogeography, Palaeoclimatology, Palaeoecology 93, 47e69.
Erba, E., Channell, J.E.T., Claps, M., Jones, C., Larson, R., Opdyke, B., Premoli Silva, I.,
Riva, A., Salvini, G., Torricelli, S., 1999. Integrated stratigraphy of the Cismon
APTICORE (Southern Alps, Italy): a “reference section” for the Barre-
mianeAptian interval at low latitudes. Journal of Foraminiferal Research 29,
371e392.
Erba, E., Tremolada, F., 2004. Nannofossil carbonate fluxes during the Early Creta-
ceous: phytoplankton response to nutrification episodes, atmospheric CO2 and
anoxia. Paleoceanography 19, 1e18.
Erbacher, J., Huber, B.T., Norris, R.D., Markey, M., 2001. Increased thermohaline
stratification as a possible cause for an oceanic anoxic event in the Cretaceous
period. Nature 409, 325e327.
Erbacher, J., Thurow, J., Littke, R., 1996. Evolution patterns of radiolarian and organic
matter variations: a new approach to identify sea-level changes in mid-Creta-
ceous pelagic environments. Geology 24, 499e502.
Ferreri, V., Weissert, H., D’ Argenio, B., Buonocunto, P., 1997. Carbon-isotope stra-
tigraphy: a tool for basin to carbonate platform correlation. Terra Nova 9, 57e61.
Geisen, M., Bollmann, J., Herrle, J., Mutterlose, J., Young, J., 1999. Calibration of the
random settling technique for calculation of absolute abundances of calcareous
nannoplankton. Micropaleontology 45, 437e442.
Glennie, K.W., 2000. Cretaceous tectonic evolution of Arabia eastern plate mar-
gining: a tale of two oceans, in Middle East models of Jurassic/Cretaceous
carbonate Systems. SEPM Special Publication 69, 9e20.
Giraud, F., Pittet, B., Mattioli, E., Audouin, V., 2006. Paleoenvironmental controls on
the morphology and abundance of the coccolith Watznaueria brittanica (Late
Jurassic, southern Germany). Marine Micropaleontology 60, 205e225.
 A. Mahanipour et al. / Cretaceous Research 32 (2011) 331e356
Giraud, F., Courtinat, B., Atrops, F., 2009. Spatial distribution of calcareous nanno-
fossils across the Callovian-Oxfordian transition in the French Subalpine Basin.
Marine Micropaleontology 72, 129e145.
Haq, B.U., Hardenbol, J., Vail, P.R., 1988. Mesozoic and Cenozoic chronostratigraphy
and cycles of sea-level change. Sea-level changes: an integrated approach.
SEPM Special Publication 42, 71e108.
Hardas, P., Mutterlose, J., 2007. Calcareous nannofossil assemblages of Oceanic
Anoxic Event 2 in the equatorial Atlantic: evidence of an eutrophication event.
Marine Micropaleontology 66, 52e69.
Heimhofer, U., Hochuli, P.A., Herrle, J.O., Weissert, H., 2006. Contrasting origins of
Early Cretaceous black shales in the Vocontian basin: evidence from palyno-
logical and calcareous nannofossil records. Palaeogeography, Palaeoclimatology,
Palaeoecology 235, 93e109.
Heldt, M., Bachmann, M., Lehmann, J., 2008. Microfacies, biostratigraphy, and
geochemistry of the hemipelagic BarremianeAptian in north-central Tunisia:
influence of the OAE 1a on the southern Tethys margin. Palaeogeography,
Palaeoclimatology, Palaeoecology 261, 246e260.
Herrle, J.O., Mutterlose, J., 2003. Calcareous nannofossils from the Aptian-Lower
Albian of southeast France: palaeoecological and biostratigraphic implications.
Cretaceous Research 24, 1e22.
Herrle, J.O., 2003. Reconstructing nutricline dynamics of mid-Cretaceous oceans:
evidence from calcareous nannofossils from the Niveau Paquier black shale (SE
France). Marine Micropaleontology 47, 307e321.
Herrle, J.O., Kößler, P., Friedrich, O., Erlenkeuser, H., Hemleben, C., 2004. High-
resolution carbon isotope records of the Aptian to Lower Albian from SE France
and the Mazagan Plateau (DSDP Site 545): a stratigraphic tool for paleo-
ceanographic and paleobiologic reconstruction. Earth and Planetary Science
Letters 218, 149e161.
Herrle, J.O., Pross, J., Friedrich, O., Kößler, P., Hemleben, C., 2003. Forcing mecha-
nisms for mid-Cretaceous black shale formation: evidence from the Upper
Aptian and Lower Albian of the Vocontian Basin (SE France). Palaeogeography,
Palaeoclimatology, Palaeoecology 190, 399e426.
Hill, M.E., 1975. Selective dissolution of mid-Cretaceous (Cenomanian) calcareous
nannofossils. Micropaleontology 21, 227e235.
Immel, H., Seyed-Emami, K., Afshar-Harb, A., 1997. Kreide-Ammoniten aus dem
iranischen Teil des Koppeh-Dagh (NE-Iran). Zitteliana 21, 159e190.
Jahren, A.H., Arens, N.C., 1998. Methane hydrate dissociation implicated in Aptian
OAE events. Geological Society of American Abstract Progress 30, 52.
Jahren, A.H., Arens, N.C., Sarmiento, G., Guerrero, J., Amundson, R., 2001. Terrestrial
record of methane hydrate dissociation in the Early Cretaceous. Geology 29,
159e162.
Jenkyns, H.C., 1995. Carbon-isotope stratigraphy and paleoceanographic signifi-
cance of the Lower Cretaceous shallow-water carbonates of resolution Guyot,
Mid-Pacific mountains. Proceedings of the Ocean Drilling Program Science
Results 143, 99e104.
Jenkyns, H.C., 1999. Mesozoic anoxic events and palaeoclimate. Zentralblatt für
Geologie und Paläontologie 1997 (7e9), 943e949.
Kalantari, A., 1969. Foraminifera from the Middle JurassiceCretaceous successions
of Kopet Dagh region (NE-Iran). Unpublished Ph.D. thesis, University of London,
England.
Kessels, K., Mutterlose, J., Michalzik, D., 2006. Early Cretaceous (Valanginian-Hau-
terivian) calcareous nannofossils and isotopes of the northern hemisphere:
proxies for the understanding of Cretaceous climate. Lethaia 39, 157e172.
Kessels, K., Mutterlose, J., Ruffell, A., 2003. Calcareous nannofossils from late Jurassic
sediments of the Volga Basin (Russian Platform): evidence for productivity
controlled black shale deposition. International Journal of Earth Science 92,
743e757.
Larson, R.L., 1991. Geological consequences of superplumes. Geology 19, 963e966.
Larson, R.L., Erba, E., 1999. Onset of the mid-Cretaceous greenhouse in the Barre-
mian-Aptian: igneous events and the biological, sedimentary, and geochemical
responses. Paleoceanography 14, 663e678.
Leckie, M.L., 2002. Oceanic anoxic events and plankton evolution: biotic response to
tectonic forcing during the mid-Cretaceous. Paleoceanography 17, 1029.
Leckie, R.M., Bralower, T.J., Cashman, R., 2002. Oceanic anoxic events and plankton
evolution: biotic response to tectonic forcing during the mid-Cretaceous.
Paleoceanography 17, 1e29.
Lees, J.A., Bown, P.R., Young, J.R., Riding, J.B., 2004. Evidence for annual records of
phytoplankton productivity in the Kimmeridge Clay Formation coccolith stone
bands (Upper Jurassic, Dorset, UK). Marine Micropaleontology 52, 29e49.
Li, Y.X., Bralower, T.J., Montan ez, I.P., Osleger, D.A., Arthur, M.A., Bice, D.M., Herbert, T.D.,
Erba, E., Premoli Silva, I., 2008. Towards an orbital chronology for the early Aptian
Oceanic Anoxic Event. Earth and Planetary Science Letters 271, 88e100.
Luciani, V., Cobianchi, M., Jenkyns, H.C., 2001. Biotic and geochemical responses to
anoxic events: the Aptian pelagic succession of the Gargano Promontory,
southern Italy. Geological Magazine 138, 277e298.
Manivit, H., 1971. Nannofossiles calcaires du Crétacé français
(AptieneMaestrichtien). Essai de Biozonation appuyée sur les stratotypes.
Unpublished Ph.D. thesis, Université de Paris, France.
Menegatti, A.P., Weissert, H., Brown, R.S., Tyson, R.V., Farrimond, P., Strasser, A.,
Caron, M., 1998. High-resolution d13C-stratigraphy through the early Aptian
“Livello Selli” of the Alpine Tethys. Paleoceanography 13, 530e545.
Moullade, M., Kuhnt, W., Berger, J.A., Masse, J., Tronchetti, G., 1998. Correlation of
biostratigraphic and stable isotope events in the Aptian historical stratotype of
La Bédoule (southeast France). Comptes Rendues de l’Academie des Sciences.
Paris, Science de la terre et des planètes 327, 693e698.
355
Müller, G., Gastner, M., 1971. The “Karbonat-Bombe”, a simple device for the
determination of the carbonate content in sediments, soils and other materials.
Neues Jahrbuch für Mineralogie, Monatshefte 10, 466e469.
Mutterlose, J., 1989. Temperature-controlled migration of calcareous nannofloras in
the north-west European Aptian. In: Crux, J.A., van Heck, S.E. (Eds.), Nanno-
fossils and their Applications. Proceedings of the International Nannofossil
Association Conference, London. Ellis Horwood, Chichester, pp. 122e142.
Mutterlose, J., 1991. Das Verteilungs- und Migrationsmuster des kalkigen Nanno-
planktons in der borealen Unterkreide (Valangin-Apt) NW-Deutschlands.
Palaeontographica B 221, 27e152.
Mutterlose, J., 1992. Migration and evolution patterns of floras and faunas in marine
Early Cretaceous sediments of NW Europe. Palaeogeography, Palaeoclimatology,
Palaeoecology 94, 261e282.
Mutterlose, J., Wise, S.W., 1990. Lower Cretaceous nannofossils biostratigraphy of
ODP Leg 113 Holes 692B and 693A, continental slope off east Antarctica,
Weddell Sea. Proceedings of the Ocean Drilling Program, Scientific Results 113,
325e351.
Mutterlose, J., Kessels, K., 2000. Early Cretaceous calcareous nannofossils from high
latitudes: implications for palaeobiogeography and palaeoclimate. Palae-
ogeography, Palaeoclimatology, Palaeoecology 160, 347e372.
Opdyke, B.N., Erba, E., Larson, R.L., 1999. Hot LIPs, methane, and the carbon record of
the Apticore. Eos Transactions American Geophysical Union 80, 486e487.
Perch-Nielsen, K., 1979. Calcareous nannofossils from the Cretaceous between the
North Sea and the Mediterranean. In: Wiedmann, J. (Ed.), Aspekte der Kreide
Europas. IUGS Series A, vol. 6, pp. 223e272.
Perch-Nielsen, K., 1985. Mesozoic Calcareous nannofossils. In: Bolli, H.M.,
Saunders, J.B., Perch-Nielsen, K. (Eds.), Plankton Stratigraphy. Cambridge Earth
Science Series. Cambridge University Press, pp. 329e426.
Pittet, B., Mattioli, E., 2002. The carbonate signal and calcareous nannofossil
distribution in an Upper Jurassic section (Balingen-Tieringen, Late Oxfordian,
southern Germany). Palaeogeography, Palaeoclimatology, Palaeoecology 179,
71e96.
Premoli Silva, I., Erba, E., Salvini, G., Locatelli, C., Verga, D., 1999. Biotic changes in
Cretaceous oceanic anoxic events of the Tethys. Journal of Foraminiferal
Research 29, 352e370.
Premoli Silva, I., Erba, E., Tornaghi, M., 1989. Paleoenvironmental signals and
changes in surface fertility in mid Cretaceous Corg-rich pelagic facies of the
Fucoid Marls (Central Italy). Geobios 11, 225e236.
Raisossadat, N., Moussavi-Harami, R., 2000. Lithostratigraphic and facies analyses of
the Sarcheshmeh Formation (Lower Cretaceous) in the eastern Kopet Dagh
Basin, NE Iran. Cretaceous Research 21, 507e516.
Raisossadat, S.N., 2002. Lower Cretaceous (Upper Barremian-Lower Albian)
Ammonite Faunas of the Kopet Dagh Basin, NE Iran. PhD Thesis, University
College London, England.
Raisossadat, S.N., 2004. The ammonite family Deshayesitidae in the Kopet Dagh
Basin, north-east Iran. Cretaceous Research 25, 115e136.
Raisossadat, S.N., 2006. The ammonite family Parahoplitidae in the Sanganeh
Formation of the Kopet Dagh Basin, north-eastern Iran. Cretaceous Research 27,
907e922.
Roth, P.H., 1973. Calcareous nannofossils- Leg 17, Deep Sea Drilling Project. Initial
Reports of the Deep Sea Drilling Project 17, 695e795.
Roth, P.H., 1978. Cretaceous nannoplankton biostratigraphy and oceanography of
the northwestern Atlantic Ocean. Initial Reports of the Deep Sea Drilling Project
44, 731e759.
Roth, P.H., 1983. Jurassic and Lower Cretaceous calcareous nannofossils in the
western North Atlantic (Site 534): biostratigraphy, preservation, and some
observations on biogeography and paleoceanography. In: Sheridan, R.E.,
Gradstein, F.M., et al. (Eds.), Initial Report of the Deep Sea Drilling Project, vol.
76. U.S. Govt. Printing Office, Washington, pp. 587e621.
Roth, P.H., Bowdler, J.L., 1981. Middle Cretaceous calcareous nannoplankton bioge-
ography and oceanography of the Atlantic Ocean. SEPM Special Publication 32,
517e546.
Roth, P.H., Krumbach, K.R., 1986. Middle Cretaceous calcareous nannofossil bioge-
ography and preservation in the Atlantic and Indian oceans: implications for
paleoceanography. Marine Micropaleontology 10, 235e266.
Schlanger, S.O., Arthur, M.A., Jenkyns, H.C., Scholle, P.A., 1987. The Cen-
omanianeTuronian oceanic anoxic event, I. Stratigraphy and distribution of
organic carbon-rich beds and the marine d13C excursion. In: Brooks, J., Fleet, A.J.
(Eds.), Marine Petroleum Source Rocks. Geological Society, London, Special
Publications, vol. 26, pp. 371e399.
Schlanger, S.O., Jenkyns, H.C., 1976. Cretaceous anoxic events: causes and conse-
quences. Geologie en Mijnbouw 55, 179e184.
Scholle, P., Arthur, M.A., 1980. Carbon isotopic fluctuations in pelagic limestones:
potential stratigraphic and petroleum exploration tool. American Association of
Petroleum Geology 64, 67e87.
Seyed-Emami, K.,1980. Parahoplitidae (Ammonoidea) aus dem Nordost und Zentraliran.
Neues Jahrbuch für Geologie und Paläontologie, Monatshefte 1980, 719e737.
Seyed-Emami, K., Aryai, A.A., 1981. Ammoniten aus dem unteren Cenoman von
Nordostiran (Koppeh Dagh). Mitteilungen der Bayerischen Staatssammlung für
Paläeontologie und Historische Geologie 21, 23e39.
Seyed-Emami, K., Forster, R., Mojtahedi, A., 1984. Ammoniten aus dem mittleren
Cenoman von Nordost-Iran (Koppeh-Dagh). Neues Jahrbuch für Geologie und
Paläontologie, Monatshefte, 159e172.
Seyed-Emami, K., Schairer, G., Behroozi, A., 1994. Einige ammoniten aus der Kashaf
Rud Formation (Mittlerer Jura) E Mashhad (NE-Iran). Mitteilungen der
356 A. Mahanipour et al. / Cretaceous Research 32 (2011) 331e356

Bayerischen Staatssammlung für Paläontologie und Historische Geologie 34, Haqius ellipticus (Grün in Grün and Allemann, 1975) Bown, 1992
145e158. Helenea chiastia Worsley, 1971
Seyed-Emami, K., Schairer, G., Behroozi, A., 1996. Ammoniten aus dem Oberen Bajoc Lithraphidites carniolensis Deflandre, 1963
(Mittlerer Jura) des SE-Koppeh Dagh und SE-Alborz (NE-Iran). Mitteilungen der Loxolithus armilla (Black, 1959) Noël, 1965
Bayerischen Staatssammlung für Paläeontologie und Historische Geologie 36, Manivitella pecten Black, 1973
87e106. Manivitella pemmatoidea (Deflandre, 1965) Thierstein, 1971 emend. Black, 1973
Shackleton, N.J., 1987. The carbon isotope record of the Cenozoic: history of organic M. hoschulzii (Reinhardt, 1966) Thierstein, 1971
carbon burial and of oxygen in the ocean and atmosphere. In: Brooks, J., Fleet, A. M. obtusus Stradner, 1963
(Eds.), Marine Petroleum Source Rocks. Geological Society, London, Special Nannoconus abundans Stradner and Grün, 1973
Publications, vol. 26, pp. 423e434. N. bermudezi Brönnimann, 1955
Shannon, C.E., Weaver, W., 1949. The mathematical Theory of Communication. Nannoconus boneti Treio, 1959
University of Illinois Press, 125 pp. N. bucheri Brönnimann, 1955
Sissingh, W., 1977. Biostratigraphy of Cretaceous calcareous nannoplankton. Geo- Nannoconus carniolensis Deflandre, 1962
logie Mijnbouw 56, 37e65. N. circularis Deres and Achéritéguy, 1980
Street, C., Bown, P.R., 2000. Palaeobiogeography of Early Cretaceous (Berria- N. elongatus Brönnimann, 1955
sianeBarremian) calcareous nannoplankton. Marine Micropaleontology 39, Nannoconus fragilis Deres and Achéritéguy, 1980
265e291. N. globulus Brönnimann, 1955
Thierstein, H.R., 1971. Tentative Lower Cretaceous calcareous nannoplankton Nannoconus grandis Deres and Achéritéguy, 1980
zonation. Eclogae Geologicae Helvetiae 64, 459e488. Nannoconus inconspicuus Deflandre and Deflandre-Rigaud, 1962
Thierstein, H.R., 1973. Lower Cretaceous calcareous nannoplankton biostratigraphy. Nannoconus inornatus Rutledge and Bown, 1996
Abhandlungen der Geologischen Bundesanstalt 29, 3e53. N. kamptneri Brönnimann, 1955
Thierstein, H.R., 1980. Selective dissolution of Late Cretaceous and earliest tertiary N. minutus Brönnimann, 1955
calcareous nannofossils: experimental evidence. Cretaceous Research 2, 165e176. Nannoconus multicadus Deflandre and Deflandre-Rigaud, 1960
Tremolada, F., Erba, E., Bralower, T.J., 2006. Late Barremian to early Aptian calcar- Nannoconus quadriangulus Deflandre and Deflandre, 1967
eous nannofossil paleoceanography and paleoecology from the Ocean Drilling N. truitti Brönnimann, 1955
Program Hole 641C (Galicia Margin). Cretaceous Research 27, 887e897. N. truitti frequens Deres and Achéritéguy, 1980
Tyson, R.V., 1995. Sedimentary Organic Matter. Chapman & Hall, London, UK, 615 pp. N. truitti rectangularis Deres and Achéritéguy, 1980
Weissert, H., Lini, A., Föllmi, K.B., Kuhn, O., 1998. Correlation of Early Cretaceous N. vocontiensis Deres and Achéritéguy, 1980
isotope stratigraphy and platform drowning events, a possible link? Palae- N. wassallii Brönnimann, 1955
ogeography, Palaeoclimatology, Palaeoecology 137, 189e201. Percivalia fenestrata (Worsley, 1971) Wise, 1983
Weissert, H., McKenzie, J.A., Channell, J.E.T., 1985. Natural variations in the carbon Pickelhaube furtiva (Roth, 1983) Applegate et al. in Covington and Wise, 1987
cycle during the Early Cretaceous. In: Sundquist, E.T., Broecker, W.E. (Eds.), The Radiolithus planus Stover, 1966
Carbon Cycle and Atmospheric CO2. Natural Variations Archean to Present, Retecapsa angustiforata Black, 1971
Geophysical Monograph Series, vol. 32, pp. 531e545. Retecapsa surirella (Deflandre and Fert, 1954) Grün in Grün and Allemann, 1975
Williams, J.R., Bralower, T.J., 1995. Nannofossil assemblages, fine fraction stable R. angustus (Stradner, 1963) Reinhardt, 1971
isotopes, and the paleoceanography of the Valanginian-Barremian (Early R. asper (Stradner, 1966) Reinhardt, 1967
Cretaceous) North Sea Basin. Paleoceanography 10, 815e864. Rhagodiscus dekaenelii Bergen, 1994
Wise, S.W., 1988. Mesozoic-Cenozoic history of calcareous nannofossils in the Rhagodiscus gallagheri Rutledge and Bown, 1996
region of Southern Ocean. Palaeogeography, Palaeoclimatology, Palaeoecology Rhagodiscus robustus Bown, 2005
67, 157e179. Rotelapillus laffittei (Noël, 1956) Noël, 1973
Wissler, L., Funk, H., Weissert, H., 2003. Response of Early Cretaceous carbonate R. irregularis Thierstein in Roth and Thierstein, 1972
platforms to changes in atmospheric carbon dioxide levels. Palaeogeography, Staurolithites crux (Deflandre and Fert, 1954) Caratini, 1963
Palaeoclimatology, Palaeoecology 200, 187e205. Staurolithites glabra (Jeremiah, 1996) Burnett, 1998
Staurolithites mutterlosei Crux, 1989
Appendix A. Taxonomic index Staurolithites stradneri Rood et al., 1971
Stoverius achylosus (Stover, 1966) Perch-Nielsen, 1986
Stradnerlithus geometricus (Górka, 1957) Bown and Cooper, 1989a
A full list of all taxa cited in the text, figures and table is given below. Most Stradnerlithus rhombicus (Stradner and Adamiker, 1966) Bukry, 1969
bibliographic references can be found in Perch-Nielsen (1985) and Bown (1998, 2005). Tegumentum octiformis (Köthe, 1981) Crux, 1989
Tegumentum striatum (Black, 1971) Taylor, 1978
Axopodorhabdus dietzmannii (Reinhardt, 1965) Wind and Wise, 1983 Tetrapodorhabdus coptensis Black, 1971
B. constans (Górka, 1957) Black, 1959 Tranolithus gabalus Stover, 1966 emend. Köthe, 1981
B. africana Stradner, 1961 Tubodiscus burnettiae Bown in Kennedy et al., 2000
B. hockwoldensis Black, 1973 Tubodiscus jurapelagicus (Worsley, 1971) Roth, 1973
B. regularis Black, 1973 W. barnesiae (Black, 1959) Perch-Nielsen, 1968
Broinsonia matalosa (Stover, 1966) Burnett in Gale et al., 1996 Watznaueria biporta Bukry, 1969
Bukrylithus ambiguus Black, 1971 Watznaueria britannica (Stradner, 1963) Reinhardt, 1964
Calculites percenis Jeremiah, 1996 W. fossacincta (Black, 1971) Bown, 1989
C. litterarius (Górka, 1957) Manivit, 1971 Watznaueria ovata Bukry, 1969
Cretarhabdus conicus Bramlette and Martini, 1964 Zeugrhabdotus clarus Bown, 2005
Cruciplacolithus hayi (Black, 1973) Jakubowski, 1986 Zeugrhabdotus diplogrammus (Deflandre in Deflandre and Fert, 1954) Burnett in
Cyclagelosphaera margerelii Noël, 1965 Gale et al., 1996
Cylindralithus nudus Bukry, 1969 Zeugrhabdotus elegans (Gartner, 1968) Burnett in Gale et al., 1996
Diazomatolithus lehmanii Noël, 1965 Zeugrhabdotus embergeri (Noël, 1958) Perch-Nielsen, 1984
D. ignotus (Gorka, 1957) Perch-Nielsen, 1968 Z. erectus (Deflandre in Deflandre and Fert, 1954) Reinhardt, 1965
Ethmorhabdus hauterivianus (Black, 1971) Applegate, Covington and Wise, 1987 Zeugrhabdotus scutula (Bergen, 1994) Rutledge and Bown, 1996
Flabellites oblongus (Bukry, 1969) Crux in Crux et al., 1982 Zeugrhabdotus trivectis Bergen, 1994
Haqius circumradiatus (Stover, 1966) Roth, 1978 Zeugrhabdotus xenotus (Stover, 1966) Burnett in Gale et al., 1996