species from the Lower Cretaceous carbonate facies

Arab J Geosci (2017) 10:31 DOI 10.1007/s12517-016-2787-x

ORIGINAL PAPER

Salpingoporella (dasycladalean algae) species from the Lower Cretaceous carbonate facies of Kopet Dagh basin (NE Iran) Morteza Taherpour Khalil Abad 1

Received: 11 July 2015 / Accepted: 5 December 2016 # Saudi Society for Geosciences 2017

Abstract Salpingoporella species from algal bearing of Barremian-Aptian limestones in the Kopet Dagh basin (NE of Iran) are described. Different species (S. cemi, S. hasi, S. heraldica, S. hispanica, S. milovanovici, S. muehlmbergi, S. parapiriniae, S. piriniae, S. cf. biokovensis, S. steinhauseri, S. polygonalis) are investigated from different biometrical aspects such as depositional environments and biogeographical distribution as well as their systematic palaeontology from two formations (the Tirgan and Sarcheshmeh formations) in nine stratigraphic sections. Keywords Salpingoporella . Barremian-Aptian . Tirgan . Sarcheshmeh . Kopet Dagh . NE Iran

Introduction and stratigraphic framework The Kopet Dagh (or Koppeh Dagh, Kopet Dag) mountain range represents a NE-trending, about 650-km long and about 200-km wide, active fold belt at the border between Iran and Turkmenistan, east of the Caspian Sea. This sedimentary basin is located in the northeast of Iran and south of Turkmenistan (54° 00, 61° 14´E and 36° 00, 38° 16´N) as an intracontinental basin (Fig. 1). It was formed on a Hercynian metamorphosed basement, at the SW margin of the Turan Platform. The belt consists of about 10 km of Mesozoic and Tertiary sediments, mostly carbonates (e.g. Berberian and King 1981; AfsharHarb 1994; Golonka 2007; Taherpour Khalil Abad et al. * Morteza Taherpour Khalil Abad [email protected] 1

Young Researchers and Elite Club, Mashhad Branch, Islamic Azad University, Mashhad, Iran

2013). The formations start from the Shemshak to the Khan Giran Formation from bottom to up (Fig. 2). These sediments were deposited in a marginal sea of the northern Tethys Ocean, one of the so-called Peri-Tethyan basins (Brunet and Cloething 2003). Activities for discovery of oil and gas in Jurassic-Cretaceous sediments of Kopet Dagh range led to greater geological research in this part of the country. The Cretaceous sequence in the Kopet Dagh range studied in detail by geologists of the National Iranian Oil Company, Geological Survey of Iran and other geologists appears to be more complete than in other parts of northern, central and eastern Iran. It includes also marine deposits of Early Cretaceous age, which elsewhere are missing except, perhaps, at the few places (south of Kerman, western and eastern Alborz), where Calpionella limestone of Tithonian–? Berriasian age has been observed (modified after Stocklin and Setudehnia 1991). In this sequence, the Lower Cretaceous strata are developed to a thickness of approximately 2300 m (in the western Kopet Dagh). From these formations, two are selected and investigated for studying the Salpingoporella species assemblages. These two formations are selected from the Lower Cretaceous strata in this sedimentary basin as follows: (1) The Tirgan Formation: this formation is named from its exposure in the Tirgan Valley in the central part of Kopet Dagh by geologists of the National Iranian Oil Company (Afshar-Harb 1969, 1970). It applies to a feature-forming unit of massively bedded, oolithic and organodetrital limestone occurring through the Kopet Dagh ranges. The thickness of this formation is measured from 50 m and less in the eastern Kopet Dagh to 700 m in the western part (Afshar-Harb 1969). It overlies the Shurijeh Formation and underlies the Sarcheshmeh Formation; both contacts are conformable but a transitional

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Fig. 1 The location map of Kopet Dagh sedimentary basin and studied stratigraphic sections (Modified from Liberis and Menby 1999; Afshar-Harb 1994)

interfingering between the Tirgan and the Shurijeh exists in southeastern Kopet Dagh. The Tirgan Formation is essentially Barremian-Early Aptian but in some sections is dating as early Early Cretaceous (such as in Ali Abad stratigraphic section, see Carević et al. 2013) locally extending as high as the Aptian. Except for rare occurrences of Calpionella limestone, the Tirgan Formation is the only marine deposit of proven Early Cretaceous age in North of Iran (Stocklin and Setudehnia 1991).

(2) The Sarcheshmeh Formation: this formation is named from its exposure in the Sarcheshmeh village in the north of Bojnourd city, west of Kopet Dagh by Bozorgniain an unpublished National Iranian Oil Company report (Afshar-Harb 1969, 1970). This formation is contained of a sequence of grey marls and pencil-shales with subordinate Orbitolina-bearing limestone intercalations, developed as a conformable cover of the Tirgan Formation throughout the Kopet Dagh ranges. Lateral interfingering

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Fig. 2 The position of Salpingoporella-bearing limestones in the studied stratigraphic sections

between the upper part of the Tirgan and the lower part of the Sarcheshmeh has been observed in some places. The Sarcheshmeh Formation is everywhere overlain by the Sanganeh Formation, the contact being marked by a persistent key bed of BCoquina^ limestone that contains Late Aptian ammonites (such as Deshayesites sp. cf. D. planus, see Raisossadat 2004) and is taken as top of the Sarcheshmeh. The thickness increases from 100 m in the southeastern most Kopet Dagh to more than 500 m in the central part of the Kopet Dagh (Stocklin and Setudehnia 1991).

Stratigraphic sections (from lithology, palaeontology and age determination points of view) In order to conduct stratigraphy and paleontological investigations, nine stratigraphic sections are selected from the Tirgan and Sarcheshmeh formations which from the east to the west are as follows (Fig. 3):

1. Abderaz stratigraphic section: this stratigraphic section is situated in about 100 km far from Mashhad city along the Mashhad-Sarakhs road and is accessible from the subsidiary road to the Bazangan Lake. The Abderaz stratigraphic section is located at 36° 14´N and 60° 29´E. The Tirgan Formation is measured 45 m in this section. –

Lithology, biota assemblages and age: the Tirgan Formation in this section is mainly composed of medium to thick-bedded sandy limestone, thin to thickbedded limestone and shale. These strata are contained of benthic foraminifera (?Balkhania balkhanica, Charentia cuvillieri, Dictyoconus pachymarginalis, Dictyoconus sp., Cyclamminid, Isteriloculina elliptica, Isteriloculina alimanensis, Lenticulina sp., Iraqia simplex, Melathokerion valseriensis, Montseciella arabica, Mayncina bulgarica, Nautiloculina oolithica, Orbitolina sp., Palorbitolina lenticularis, Praechrysalidina sp., Pseudocyclammina lituus, Glomospira watersi, Rummanoloculina pseudominima, Nezzazata isabellae and Textularid) and calcareous algae

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sp., Scythiolina sp., Textularid, Vercorsella arenata, Vercorsella sp.) and calcareous algae (Boueina sp., ?Juraella bifurcate, ?Supilulimaella sp., Acicularia sp., Arabicodium sp., Deloffrella sp. cf. D. quercifoliipora, Halimeda sp., Marinellalugeoni, Montiella ? elitzae, Kopetdagaria sphaerica, N e o m e r i s c re t a c e a , P e r m o c a l c u l u s s p . , Pseudoactinoporella fragilis, Rajkaella laskervi, Salpingoporella sp., Thaumatoporella sp.). Based on the mentioned assemblage, Late BarremianEarly Aptian is assigned to the Tirgan Formation in this stratigraphic section. 3. Qheshgheh stratigraphic section: this stratigraphic section is situated in about 55 km far from Mashhad city along the Mashhad-Kalat-e-Naderi road. This section is located at 36° 39´N and 60° 3´E, near Qheshqheh village. The Tirgan Formation is measured 84 m in this section. –

Fig. 3 Distribution of Salpingoporella species during the Early Cretaceous (Modified from Carras et al. 2006)

(Boueina sp., Coptocampylodon sp., Boueina hochstetteri, Halimeda sp., Kopetdagaria sphaerica, Marinella lugeoni, Montiella ? elitzae, Neomeris cretacea, Permocalculus sp., Permocalculus innopinatus, Salpingoporella sp., Salpingoporella hasi). Based on the mentioned assemblage, Late Barremian-Early Aptian is assigned to the Tirgan Formation in this stratigraphic section. 2. Zavin stratigraphic section: this stratigraphic section is situated in about 87 km far from Mashhad city along the Mashhad-Kalat-e-Naderi road. This section is located at 36° 44´N and 59° 56´E, northwest of Zavin village. The Tirgan Formation is measured 132 m in this section. –

Lithology, biota assemblages and age: the Tirgan Formation in this section is mainly composed of thin- to medium-bedded sandy limestone, siltstone, thin- to thick-bedded limestone and shale. These strata are contained of benthic foraminifera (?Haplophragmoides sp., Balkhania balkhanica, Charentia cuvillieri, Debarina hahounerensis, Dictyoconus pachymarginalis, Dictyoconus sp., Isteriloculina elliptica, Lenticulina sp., Marsonella sp., Melathokerion valseriensis, Nautiloculina bronnimanni, Nautiloculina oolithica, Orbitolina sp., Palorbitolina lenticularis, Pseudocyclammina lituus, Rummanoloculina pseudominima, Sabaudia

Lithology, biota assemblages and age: the Tirgan Formation in this section is mainly composed of thin-bedded sandy limestone, siltstone, medium- to thick-bedded limestone and shale. These strata are contained of benthic foraminifera (Balkhania balkhanica, Charentia cuvillieri, Commaliama sp., Cuneolina sp. cf. C. pavonia, Dictyoconus pachymarginalis, Dictyoconus sp., Gaudryna sp., Iraqia simplex, Isteriloculina elliptica, Lenticulina sp., Mayncina bulgarica, Melathokerion valseriensis, Montseciella arabica, Nautiloculina bronnimanni, Nautiloculina oolithica, Nezzazata isabellae, Orbitolina sp., Palorbitolina lenticularis, Praeorbitolina cormyi, Pseudocyclammina lituus, Quinqueloculina robusta, Rummanoloculina pseudominima, Scythiolina sp., Simplorbitolina ? manasi, Textularid, Torinosuella peneropliformis, Trocholina sp. cf. T. odukpaniensis, Vercorsella arenata, Vercorsella sp.) and calcareous algae (Acicularia sp., Arabicodium sp., Boueina h o c h s t e t t e r i , B o u e i n a s p . , H a li m e d a s p . , Kopetdagaria sphaerica, Lithocodium aggregatum, Marinella lugeoni, Montiella ? elitzae, Neomeris cretacea, Permocalculus sp., Pseudoactinoporella fragilis, Salpingoporella cemi, Salpingoporella muehlbergii, Salpingoporella sp., Terquemella sp., Thaumatoporella sp.). Based on the mentioned assemblage, Late Barremian-Early Aptian is assigned to the Tirgan Formation in this stratigraphic section.

4. Chenaran stratigraphic section: this stratigraphic section is situated in about 25 km far from Chenaran city and is accessible via Mashhad-Chenaranmain road. This section is located at 36° 49´N and 59° 14´E, northeast of

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Glomospira sp. cf. G.watersi, Iraqia simplex, Isteriloculina alimanensis, Lenticulina sp., Isteriloculina elliptica, Lituolids, Mayncina bulgarica, Melathokerion valseriensis, Mesorbitolina sp., Nautiloculina sp., Neotrocholina aptiensis, Nezzazata isabellae, Nezzazatinella sp. cf. N. picardi, Montseciella arabica, Orbitolina sp., Palorbitolina lenticularis, Praechrysalidina sp., Praeorbitolina cormyi, Pseudocyclammina lituus, Pseudocyclammina sp., Rummanoloculina ponticuli, Sabaudia sp. cf. S. minuta, Trocholina sp. cf. T. odukpaniensis, Vercorsella sp., Vercorsella sp. cf. V.arenata) and calcareous algae (Acicularia sp., Boueina sp., Boueina sp. cf. B. hochstetteri, Cayeuxia sp., Clypeina sp., Clypeina sp. cf. C. solkani, Conradella bakalovae, Cylindroporella sp., Halimedacea, Juraella bifurcata, Lithocodium aggregatum, Kopetdagaria sphaerica, Marinella lugeoni, Neomeris sp. cf. N. cretacea, Montiella ? elitzae, Permocalculus sp., Praturlonella sp. cf. P. n e r a e , P s e u d o a c t i n o p o re l l a ? i r a n i c a , Salpingoporella sp. aff. S. hasi, Salpingoporella muehlbergii, Salpingoporella sp., Supilulimaella sp. cf. S. tuberifera, Terquemella sp., Thaumatoporella sp.). Based on the mentioned assemblage, Late Barremian-Early Aptian is assigned to the Tirgan Formation in this stratigraphic section (for more detail, see Carević et al. 2013; Taherpour Khalil Abad et al. 2013).

Chenaran township. The Tirgan Formation is measured 117.6 m in this section. –

Lithology, biota assemblages and age: the Tirgan Formation in this section is mainly composed of thin- to thick-bedded sandy limestone, thin- to thick-bedded limestone and shale. These strata are contained of benthic foraminifera (?Anchispirocyclina lusitanica, Bellorusiella sp., Charentia cuvillieri, ?Choffatella decipiens, Commaliama sp., Debarina hahounerensis, Dictyoconus pachymarginalis, Dictyoconus sp., Gaudryna sp., Glomospira watersi, Valserina b ro n i m a n n i , I s t e r i l o c u l i n a a l i m a n e n s i s , Isteriloculina elliptica, Lenticulina sp., Mayncina bulgarica, Melathokerion valseriensis, Montseciella arabica, Nautiloculina bronnimanni, Nautiloculina oolithica, Nezzazata isabellae, Orbitolina sp., Palorbitolina lenticularis, Pseudocyclammina lituus, Quinqueloculina robusta, Scythiolina sp., Textularid, Triloculina sp., Trocholina sp. cf. T. odukpaniensis, Vercorsella arenata, Vercorsella sp.) and calcareous algae (Acicularia sp., Boueina hochstetteri, Boueina sp., Clypeina sp., Coptocampylodon sp., Deloffrella sp. cf. D. quercifoliipora, Halimeda sp., Halimedacea, Kopetdagaria sphaerica, Marinella lugeoni, Montiella ? elitzae, Neomeris cretacea, Permocalculus sp., Pseudoactinoporella fragilis, Rajkaella laskervi, Salpingoporella muehlbergii, S a l p i n g o p o r e l l a s p . , Te r q u e m e l l a s p . , Thaumatoporella sp.). Based on the mentioned assemblage, Late Barremian-Early Aptian is assigned to the Tirgan Formation in this stratigraphic section.

5. Sisab stratigraphic section: this stratigraphic section is situated in about 35 km far from Bojnourd city and is accessible via Mashhad-Bojnourd main road, from the subsidiary road to the Esfidan village. This section is located at 38° 19´N and 57° 35´E, southeast of Bojnourd city. The Tirgan Formation is measured 215.5 m in this section. –

Lithology, biota assemblages and age: the Tirgan Formation in this section is mainly composed of thin-bedded sandy limestone, thin- to thick-bedded limestone and shales (contain of rework debris and well-preserved echinid specimens which belong to Toxasteridae). These strata are contained of benthic foraminifera (Ammobaculites sp., Balkhania balkhanica, Charentia cuvillieri, Cyclammina sp., Cuneolina sp., Cuneolina pavonia, Debarinaha hounerensis, Derventina filipescui, Dictyoconus p a c h y m a rg i n a l i s , E v e r t i c y c l a m m i n a s p . , Everticyclammina hedbergi, Gaudryna sp.,

6. Ali Abad stratigraphic section: this stratigraphic section is situated in about 28 km far from Bojnourd city along the Mashhad-Bojnourd road. This section is located at 37° 26´N and 57° 13´E, northeast of Bojnourd city. The Tirgan Formation is measured 195.30 m in this section. –

Lithology, biota assemblages and age: the Tirgan Formation in this section is mainly composed of sandstone, thin- to thick-bedded limestone and shale. These strata are contained of benthic foraminifera (Anchispirocyclina lusitanica, Ammobaculites sp., Charentia cuvillieri, Comaliamma sp., Debarina sp. cf. D. hahounerensis, Everticyclammina hedbergi, Gaudryina sp., Glomospira sp. cf. G. watersi, Isteriloculina elliptica, Istriloculina sp., Lenticulina sp., Marsonella sp., Mayncina b u l g a r i c a , N a u t i l o c u l i n a b ro e n n i m a n n i , Pseudocyclammina lituus, Rumanoloculina pseudominima, Trocholina elongata, Trocholina odukpaniensis, Trocholina sp.) and calcareous algae (Acicularia sp., Boueina sp., Cayeuxia sp., Clypeina sp., Clypeina sp. cf. C. parasolkani, Halimedacea, Juraella bifurcate, Kopetdagaria sphaerica,

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Linoporella ? sp., Marinella lugeoni, Neomeris sp., Permocalculus sp., Rajkaella sp. cf. R. laskarevi, Salpingoporella sp., Suppiluliumaella sp. cf. S. tuberifera, Thaumatoporella sp.). Based on the mentioned assemblage, Tithonian-Valanginian is assigned to the Tirgan Formation in this stratigraphic section.

at 37° 33´N and 57° 11´E, west of Bojnourd city. The Tirgan Formation is measured 330 m in this section. –

7. Arkan stratigraphic section: this stratigraphic section is situated in about 10 km far from Bojnourd city and is accessible via Bojnourd-Esfarayen main road, from the subsidiary road to the Arkan village. This section is located at 38° 19´N and 57° 35´E, southwest of Bojnourd city. The Tirgan Formation is measured 215.5 m in this section. –

Lithology, biota assemblages and age: the Tirgan Formation in this section is mainly composed of thin-bedded sandy limestone, thin- to thick-bedded limestone, marl (contain of echinid specimens which belong to Toxasteridae, see Taherpour Khalil Abad et al. 2011) and shale. These strata are contained of benthic foraminifera (Balkhania balkhanica, Choffatella decipiens, Debarina hahounerensis, Dobrogelina sp. cf. D. ovidi, Dictyoconus pachymarginalis, Everticyclammina hedbergi, Iraqia simplex, Mayncina bulgarica, Melathokerion sp. cf. M. valseriensis, Mesorbitolina sp., Mesorbitolina lotzei, Nautiloculina oolithica, Neotrocholina sp., Orbitolinopsis gr. killiani/cuvillieri, Orbitolinopsis praesimplex, Palorbitolina lenticularis, Paracoskinolina sunnilandensis, Praeorbitolina sp., Pseudochrysalidina sp., Pseudocyclammina sp., Trocholina sp. cf. T. odukpaniensis, Vercorsella sp., Vercorsella arenata, Actinoporella podolica, Clypeina gigantean, Clypeina solkani, Coptocampylodon lineolatus, Cylindroporella elliptica, Ethelia sp. cf. E. alba, Holosporella ? sp., Juraella bifurcata, Montiella ? elitzae, Praturlonella dalmatica, Praturlonella sp. cf. P. nerae, P s e u d o a c t i n o p o re l l a s p . c f . P. f r a g i l i s , Salpingoporella sp. cf. S. cemi, Salpingoporella hasi, Salpingoporella hispanica, Salpingoporella sp. cf. S. muehlbergii, Salpingoporella sp. aff. S. muehlbergii, Salpingoporella parapiriniae, Steinmaniporella ? parsica). Based on the mentioned assemblage, Late Barremian-Early Aptian is assigned to the Tirgan Formation in this stratigraphic section (for more detail, see Carević et al. 2013).

8. Akher Dagh stratigraphic section: this stratigraphic section is situated in about 29 km far from Bojnourd city along the Bojnourd-Gonbad road. This section is located

Lithology, biota assemblages and age: the Tirgan Formation in this section is mainly composed of sandstone, thin- to thick-bedded limestone and shale. These strata are contained of benthic foraminifera (?Valserina bronnimanni, Ammobaculites sp., Bellorusiella sp., Charentia cuvillieri, Choffatella decipiens, Commaliama sp., Cuneolina pavonia, Cyclamminid, Debarina hahounerensis, Derventina f i l i p e s c u i, D i c t y oc o nu s pa c hy m a rgi na l i s , Dictyoconus sp., Dobrogelina sp. cf. D. ovidi, Everticyclammina hedbergi, Gaudryna sp., Glomospira watersi, Haplophragmoides sp., Iraqia simplex, Isteriloculina alimanensis, Isteriloculina elliptica, Lenticulina sp., Marsonella sp., Mayncina bulgarica, Melathokerion valseriensis, Mesorbitolina sp., Montseciella arabica, Nautiloculina oolithica, Neotrocholina aptiensis, Nezzazata isabellae, Orbitolina kurdica, Orbitolina sp., Palorbitolina lenticularis, Praechrysalidina sp., Praeorbitolina cormyi, Pseudocyclammina lituus, Quinqueloculina robusta, Rummanoloculina pseudominima, Sabaudia sp., Scythiolina sp., Spiroloculina cretacea, Textularid, Thaumatoporella sp., Triloculina sp., Valvulammina picardi, Vercorsella arenata, Vercorsella sp.) and calcareous algae (Acicularia sp., Boueina sp., Cayeuxia sp., Clypeinasp. cf. C. solkani, Clypeina sp., Coptocampylodon lineolatus, Coptocampylodon sp., Halimeda sp., Lithocodium aggregatum, Marinella lugeoni, Montiella ? elitzae, Neomeris cretacea, Permocalculus sp., Praturlonella nerae, Pseudoactinoporella fragilis, Salpingoporella cemi, Salpingoporella hasi, Salpingoporella istriana, Salpingoporella muehlbergii, Salpingoporella sp., Terquemella sp., Triploporella sp. cf. T. bacilliformis). Based on the mentioned assemblage, Early Aptian is assigned to the Tirgan Formation in this stratigraphic section.

9. Gelian stratigraphic section: this stratigraphic section is situated in about 30 km far from Shirvan city. This section is located at 37° 15´N and 57° 55´E, south of Shirvan city. The Tirgan Formation is measured 112 m in this section. –

Lithology, biota assemblages and age: the Sarcheshmeh Formation in the studied section is mainly composed of marl, shale with some intercalations of limestones. The shaley beds are rich in ammonites (family Deshayesitidae). According to the identified microfossils (mainly Foraminifera) such as Ammodiscus penaensis, Gavelinella

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barremina, Globigerina infracretacea, Haplophragmoides sp., Lenticulina munsteri, Nautiliculina oolitica, Orbitolina discoidea, O. kurdika and O. lenticulina, the Sarcheshmeh Formation is interpreted to be of Aptian age (Kalantari 1969; Raisossadat and Moussavi-Harami 1993). Next to the introduced fauna by Raisossadat and Moussavi-Harami (1993), there are rich fauna and flora assemblages in this formation. Some of the fauna are as follows: Palorbitolina lenticularis, Praeorbitolina comyi, Dictyoconus pachymarginalis, Charentia cuvillieri, Marsonella sp. etc. and a rich-Salpingoporella assemblage such as Salpingoporella muehlbergi, S. sp. cf. S. hasi, S. heraldica, S. polygonalis etc. (see paleontological part of the present paper). The inferred age for these strata is assigned to Middle Aptian (Gargasian).

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Geological Survey of Iran and Geosciences Research Center, NE territory (M. Taherpour Khalil Abad collection). Palaeontology Ordo Dasycladales Pascher 1931. Familia Triploporellaceae (Pia 1920) nom. transl. Tribus Salpingoporelleae Bassoullet et al. 1979 Sub-tribus Salpingoporellinae Bassoullet et al. 1979 Genus Salpingoporella Pia, in Trauth, Pia 1918, emend. Diagnostic features Cylindrical thallus, euspondyle. Only primary laterals in closed whorls, phloiophorous, making a cortex. Reproduction is cladosporous (for more detail, see Barattolo: BMicropaleontology of algae from Permian to Eocene^ or Carras et al. 2006). Diagnostic features Triassic, Late Jurassic-Cretaceous, Paleocene?. Salpingoporella cemi Radoičić 1975 Pl. 3, Figs. 1–9

Material and method

Holotype

Detailed palaeontology, biometrical aspects (Fig. 4) and sedimentological investigations have been carried out on eight stratigraphic sections of the Tirgan Formation (Late BarremianEarly Aptian = Bedoulian) and one stratigraphic section of the Sarcheshmeh Formation (Middle Aptian = Gargasian) in northeast Iran, Kopet Dagh sedimentary basin, supported by the analysis of 573 thin sections with emphasis on the BSalpingoporella^ species which are the main constituents of Late Barremian-Middle Aptian carbonate sediments of the Kopet Dagh basin. The thin sections are housed in the

1968 Pianella cemi n. sp.- Radoičić, Pls. 9–11; outer Dinarides; late Late Barremian-early Early Aptian [nom. nud., Bsyntypes^] (Fig. 6). Selected citations 1970 Pianella aff. cemi- Conrad, Pl. 9, Figs. 2, 3; Spain; Barremian. 1974 Pianella aff. cemi- Radoičić, Pl. 8, Fig. 4; Herzegovina; Late Barremian-Early Aptian.

Fig. 4 Salpingoporella biometry elements (Left ones modified from Conrad et al. 2008)

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1999 Salpingoporella cemi- Yilmaz, Pl. 2, Figs. 31, 32; Fele area, Western Taurides, Turkey, Hauterivian-Barremian. 2006 Salpingoporella cemi- Carras et al., Pl. 2, Figs. 1–5; Trgaj, Podgorica area, Montenegro; Late HauterivianBarremian. 2008 Salpingoporella cf. cemi- Michetiuc et al., Pl. III, Fig. 8; Vâlcan Mountains, Danubian Domain; Barremian-Aptian. 2008 Salpingoporella cemi- Sokač and Grgasović, Pls. 1 and 2; Pl. 3, Figs. 1–8; Dinaric karst, Croatia; Neocomian. 2010 Salpingoporella cemi- Taherpour Khalil Abad et al., Fig. 7a, b; Kopet Dagh basin, NE Iran; Late BarremianEarly Aptian. 2013 Salpingoporella cf. cemi- Carević et al., Fig. 17 w; Kopet Dagh basin, Iran; Late Barremian-Early Aptian. Diagnostic features According to Carras et al. 2006, this species is Blarge, with laterals forming quincunxes, 10-15 per verticil, horizontal or slightly tilted, funnel-like (first with a very short proximal narrowing, then gradually widening toward the distal part), circular in section, or very seldom somewhat polygonal in subcortical and cortical section. Massive calcareous skeleton made of colourless sparry calcite^. Some new data has been added on the previous descriptions of this species by Sokač and Grgasović (2008) as follows: cylindrical, unramified, thallus is characterized by thick walls built of mosaic calcite derived by heteroaxial transformation of, probably, primary aragonitic skeleton. Along the margins of the skeleton, tiny, short-prismatic, calcite crystals are developed, whereas the remainder of the skeleton is filled up with middle- to coarse-grained mosaic calcite cement. Skeletal walls are delineated with very thin micritic lining of constant thickness. Remarks This species seems to be less common then other species of this genus (Sokač and Grgasović 2008). The dimensions, shape of the branches and calcification pattern are similar to Likanella ? danilovae. However Likanella ? danilovae has clustered branches which emerge in various directions (Yilmaz 1999). Another similarity is to Salpingoporella ? turgida but in S. turgida thallus sometimes club-shaped, and the tubular part of the laterals is longer but in S. cemi laterals are regularly widening out after a very short proximal narrowing and to Salpingoporella ubaiydhi but in S. cemi, the thallus is larger, with larger h values and larger laterals, set in quincunxes (Carras et al. 2006).

Range in studied sections Early Aptian (Arkan, Qheshgheh and Akher Dagh stratigraphic sections). Environments Inner platform, protected (Carras et al. 2006). Shallow subtidal with slow carbonate deposition (Sokač and Grgasović 2008) in association with S. tari (Fig. 5). Biogeographical distribution Spain, SE France, Herzegovina, Albania, Romania, Greece, Turkey, Montenegro, Croatia and NE Iran. Salpingoporella hasi Conrad et al. 1977 Pl. 3, Figs. 10–19 Holotype 1977 Salpingoporella hasi n. sp.- Conrad et al., Figs. 7 and 8; Pl. 1, Figs. 1–10; Metohija (Jugoslavia) and Lisboa (Portugal); Late Albian-Middle Cenomanian (Fig. 6). Selected citations 1973 Pianella dinarica- Berthou, Pl. 15, Figs. 1, 2; Pl. 20, Fig. 1; Pl. 31, Fig. 4; Portugal, Cenomanian. 1976 Salpingoporella aff. hasi nom. nud.- Conrad and Peybernes, Fig. 13 (d, e); Eastern Pyrenees, Late BarremianEarly Bedoulian. 1999 Salpingoporella cemi- Yilmaz, Pl. 3, Figs. 38, 39; Fele area, Western Taurides, Turkey, Hauterivian-Barremian. 2006 Salpingoporella cemi- Carras et al., Pl. IV, Figs. 8–11; Dobrušte, SE of Mt. Pastrik (Mirdita Zone); AptianCenomanian, affinis in the Barremian. 2010 Salpingoporella aff. hasi- Taherpour Khalil Abad et al., Fig. 7 (j, k); Kopet Dagh basin, NE Iran, Late BarremianEarly Aptian. 2013 Salpingoporella aff. hasi- Carević et al., Fig. 17 (x); Kopet Dagh basin, NE Iran, Late Barremian-Early Aptian. 2014 Salpingoporella aff. hasi- Khodashenas et al., Fig. 4 (K); Esfiyukh Mountain, NE Iran; Barremian-Aptian.

Dimensions See Table 1.

Diagnostic features According to Carras et al. 2006, Bspecies of the genus Salpingoporella whose verticils are made up of a small number of branches, transversally flattened, and which tubular proximal part is well developed. At their distal end, the branches widen out quickly and become rectangular. The nearby branches belonging to the same verticil may touch each other or not, depending on the space available, which in turn depends from their length and from the diameter of the siphon. The calcareous envelope is made up of a simple mosaic of hyaline crystals^.

Total range Late Hauterivian-Late Barremian (Carras et al. 2006). Late Hauterivian-Early Aptian (Granier and Deloffre 1993) (Fig. 3).

Remarks This species next to Salpingoporella granieri, S. circassa and S. urladanasi belongs to the Salpingoporella group with flattened horizontal laterals which have

Percentage of specimens 9%. (See Fig. 8).

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Plate 1 1–17. Salpingoporella muehlbergii, 18–20. S. piriniae (Scales. yellow: 200 μm, white: 250 μm, blue: 500 μm)

rectangular (different in size, more or less elongated or high) cortical pattern (Dieni and Radoičić 1999). In addition, S. hasi is similar to S. genevensis, S. hispanica, S. urladanasi, S. dinarica, and S. biokovensis with rectangular ramifications in tangential sections. This species differs from S. muehlbergii

and S. katzeri in the number of branches, dimensions, type of calcification and shape of the branches, which have curved edges (Yilmaz 1999). Percentage of specimens 17%. (See Fig. 8).

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Plate 2 1–26. Salpingoporella muehlbergii (Scales. white: 250 μm, blue: 500 μm)

Dimensions See Table 1. Total range Aptian-Albian (Varol et al. 1988), Albian-Middle Cenomanian (Granier and Deloffre 1993), Aptian-Cenomanian, (affinis in the Barremian) (Carras et al. 2006) (Fig. 3). Range in studied sections Late Barremian-Early Aptian (Sisab stratigraphic section), Early Aptian (Abderza, Akher Dagh, Arkan stratigraphic sections).

Environments Low energy, inner platform. (Carras et al. 2006). Marine part of infralittoral (Conrad and Peybernès 1976) (Fig. 5).

Biogeographical distribution Portugal, Italy, Venezuela, Oman, Turkey, Jordan, Croatia and NE Iran. Salpingoporella heraldica Sokač 1996. Pl. 3, Fig. 25

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Plate 3 1–3, 6–7, 9. Salpingoporella cemi; 4, 5. S. sp. cf. cemi; 8. S. sp. aff. cemi; 10–16. S. sp. aff. hasi; 17–19. S. sp. cf. hasi; 20, 21. S. hispanica; 22. S. sp. aff. hispanica; 23. S. sp. cf. hispanica; 24. S. sp. aff. S. milovanovici; 25. S. heraldica; 26. S. sp. cf. S. biokovensis; 27. S. steinhauseri; 28, 29. S. sp. cf. polygonalis; 30, 32. S. sp. cf. S. parapiriniae; 31. S. parapiriniae; 33. S. sp. cf. S. piriniae; 34. S. piriniae (Scales. yellow: 200 μm, white: 250 μm, blue: 500 μm)

Holotype

Selected citations

1996 Salpingoporella heraldica n. sp.- Sokač, Pl. 7, Figs. 1–18; Dalmatian islands (Croatia), Barremian-Aptian transition (Fig. 6).

2006 Salpingoporella heraldica- Carras et al., Pl. IV, Figs. 12–16; Dalmatian islands (Croatia); Barremian-Aptian.

31 Table 1

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Page 12 of 21 The calculation of biometric data of determined Salpingoporella species

Dimensions (μm) Species

D

d

h

w

e

pwd

dw (y)

dw (n)

pi

S. cemi S. hasi S. heraldica S. hispanica

500–680 780–1610 # #

230–320 430–650 # #

150–170 # # 410

7–9 6–11 # #

130 # # 540

100–140 # # 340

90–220 # # 570

180 # # #

110–120 # 120 210

S. milovanovici S. muehlmbergi S. parapiriniae S. piriniae S. sp. cf. S. biokovensis S. steinhauseri S. polygonalis

1390 180–1530 580–830 370–1530 760 620 400–520

560 90–410 250–400 140–660 290 260 250–260

# 70–280 190 110–350 270 150 #

# 8–11 7- ∼ 10 5–10 # # 9

# 80–190 140 120–450 260 220 #

# 60–80 # 290 220 # #

# 90–240 # 220–330 180 100 #

# 90–230 # 330–350 # # #

# 30–70 # 120–150 120 60 #

2008 Salpingoporella heraldica- Michetiuc et al., Pl. IV, Fig. 8. Vâlcan Mountains, Danubian domain; BarremianAptian. 2014 Salpingoporella heraldica- Granier et al., Fig. 7k, Canton of Vaud, Switzerland, Late Hauterivian. Diagnostic features BLarge axial cavity and funnel-shaped, alternating ramifications, which in tangential sections appear square-shaped, producing the characteristic chequered appearance. Small to medium-sized, with a large main axis. Laterals

set in quincunxes, 11-14 per verticil, horizontal, funnel-like, regularly and uniformly widening out, squared at distal end, with a chequered pattern. Calcareous skeleton made of yellowish sparry calcite^ (Carras et al. 2006). Remarks BThe chequered pattern of the laterals is typical from the species^ (Carras et al. 2006). Also, this species is clearly distinguishable from other Salpingoporella species from those in which the distal parts of the ramifications are either longitudinally or transversely compressed. It belongs to

Fig. 5 Distributional pattern of dasyclad associations on Early Cretaceous carbonate platforms (Modified from Flugel, 2004 and Carras et al. 2006)

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Fig. 6 Schematic drawings of Salpingoporella species holotypes (After Carras et al. 2006)

the group of rather small Salpingoporrellae by its size (dimensions), the d/D relationship, and the number of branches per whorl (Sokač 1996). This species is rare in the studied sections and only is found in the Sarcheshmeh Formation. Percentage of specimens 1%. (See Fig. 8). Dimensions See Table 1. Total range Late Hauterivian-Late Barremian, possibly Early Aptian (Carras et al. 2006) (Fig. 3). Range in studied sections Late Early Aptian-?Middle Aptian. Environments Peritidal-tidal bars in the type level (Sokač 1996) (Fig. 5).

Biogeographical distribution Croatia, Italy, SE France, Switzerland and NE Iran. Salpingoporella hispanica Conrad and Grábner 1975. Pl. 3, Figs. 20–23 Holotype 1975Salpingoporella hispanica n. sp.- Conrad and Grabner, Figs. 1–7; Spanish Pyrenees, Barremian (Fig. 6).

Selected citations 1986 Apinella jaffrezoi n. gen., n. sp.- Granier et al., Fig. 8; Pl. 1, Figs. a–j; Pl. 2, Figs. a–j; Chiapas (Mexico), Kimmeridgian. 1999 Siensiporella cf. vinogradovii n. sp.- Dragastan, Pl. 5, Figs. 6–10; Moesian Platform, Romania, Berriasian. 1976 Salpingoporella cf. hispanica- Masse, Pl. 3, Figs. 5, 6; SE France, Late Barremian.

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Fig. 7 Deposited formations in the Kopet Dagh sedimentary basin during Jurassic-Paleogene (Modified after Aghanabati, 2008)

1977 Salpingoporella hispanica- Chiocchini and Mancinelli, Pl. 26, Fig. 1; Latium (Italy), Late Barremian. 1978 Salpingoporella hispanica- Bassoullet et al., Pl. 29, Figs. 11–12. 1979 Salpingoporella hispanica- Chiocchini et al., Pl. 2, Fig. 1; Latium, Middle Barremian. 1979 Salpingoporella hispanica- Peybernes and Conrad, Pl. 2, Figs. 1, 2; Hungary, Late Barremian-Bedoulian. 1981a, b Salpingoporella aff. hispanica- Bucur, Pl. 2, Figs. 3–4; Apuseni Mts. (Romania), Barremian-Aptian. 1984 Salpingoporella hispanica- Peybernes et al., Pl. 1, Fig. 5; Algeria, Barremian. 1986 Apinella hispanica- Granier et al., p. 804. 1986 Salpingoporella gr. hispanica- Radoičić, Pl. 5, Figs. 2–7; Montenegro (Yugoslavia), NeocomianBarremian.

1987 Salpingoporella gr. hispanica- Radoičić, Pl. 2, Figs. 1– 3; Dalmatian Islands, Early Aptian. 1992 Salpingoporella hispanica- Mancinelli, Pl. 3, Figs. 1–5; Aurunci Mts. (Latium, Italy), Barremian. 1993 Salpingoporella hispanica- Bodrogi et al., Pl. 2, Figs. 7– 10; Villany zone (Hungary), Berriasian. 1993 Salpingoporella jaffrezoi n. comb.- Bodrogi et al., P. 66. 1994 Apinella hispanica- Bucur, Pl. 2, Figs. 24–27; Southern Carpathians, Late Barremian-Early Aptian. 1994 Salpingoporella hispanica- Chiocchini et al., Pl. 36, Figs. 6, 7; Abruzzo (Italy), Barremian. 1994 Apinella jaffrezoi- Chiocchini et al., Pl. 38, Figs. 5–8; Abruzzo (Italy), Late Portlandian. 1994 Salpingoporella hispanica- Gielisch, Pl. 5, Fig. 8; Desfina (Greece), Late Barremian. 2006 Salpingoporella hispanica- Carras et al., Pl. V, Figs. 1– 5; Northeast Spain, Barremian.

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Fig. 8 The percentage of determined Salpingoporella species in the studied stratigraphic sections. During determined taxa, S. muehlbergii is the most abundant one which is well distributed from east to west of Kopet Dagh sedimentary basin

2010 Salpingoporella cf. hispanica- Taherpour Khalil Abad et al., Pl. 4; Kopet Dagh basin, NE Iran, Late Barremian-Early Aptian. Diagnostic features The laterals are asymmetrical in vertical section, with a tilted proximal narrowing, much larger h values and a thin calcareous skeleton. BRather small to mediumsized. Short laterals set in quincunxes, 4-5 (seldom 3) per verticil, starting with a short peduncle leaning up, then abruptly swelling, asymmetrical (swelling more pronounced towards the upper part of the thallus), first rectangular in section, then hexagonal, horizontally elongated. Thin calcareous skeleton made up of colourless sparry calcite^ (emended diagnosis by Carras et al. 2006). Remarks Salpingoporella hispanica is similar to S. biokovensis by its outer branches, but the number of branches and their dimensions clearly distinguish these species from each other (Yilmaz 1999). It is also can be confused with S. katzeri but the conical shape of the primary branches and hexagonal shapes in tangential sections distinguish S. katzeri from this species. Also is similar to S. hasi in having rectangular ramifications in tangential sections. Percentage of specimens 8%. (See Fig. 8). Dimensions See Table 1. Total range Kimmeridgian-Early Aptian (Carras et al. 2006) (Fig. 3).

Range in studied sections Late Barremian-Early Aptian (Arkan stratigraphic section). Environments Inner platform, protected (Carras et al. 2006) (Fig. 5). Biogeographical distribution Romania, Spain, France, Italy, Hungary, Algeria, Greece, Croatia and NE Iran. Salpingoporella milovanovici Radoičić 1978. Pl. 3, Fig. 24 Holotype 1978 Salpingoporella milovanovici n. sp.- Radoičić, Pl. 1, Figs. 1–7; Pl. 2, Figs. 1–6; Pl. 3, Figs. 1–10; Mt. Gorica (Mirdita zone, Kosovo), lower part of the Late Cenomanian (Fig. 6). Selected citations 1998 Salpingoporella milovanovici- Hernandez-Romano et al., Pl. 2 (A, F), 4 (A, B), Mexico, Late Cenomanian. 2000 Salpingoporella milovanovici- Sokač, Dalmatia (Croatia), Early Cretaceous [Specimens are not illustrated]. 2006 Salpingoporella milovanovici- Carras et al., Pl. VI, Figs. 9–13; Kosovo, Late Cenomanian. 2010 Salpingoporella aff. milovanovici- Taherpour Khalil Abad et al., Fig. 7 (h); Kopet Dagh basin, NE Iran, Late Barremian-Early Aptian. 2013 Salpingoporella aff. milovanovici- Carević et al.; Kopet Dagh basin, NE Iran, Late Barremian-Early Aptian.

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Diagnostic features BRather small, with laterals set in sometimes irregular vertical files, 9-16 per verticil, first horizontal, funnel-like but asymmetrical in vertical section (swelling more pronounced towards the upper part of the thallus), first narrow, and then abruptly widening out. Section of the laterals elliptical, vertically elongated in the distal part. Calcareous skeleton made of colourless sparry calcite^ (Carras et al. 2006). Remarks This species is similar to Salpingoporella donatae but S. milovanovici is smaller in general and have differently shaped branches giving differently shaped pores in tangential sections (Sokač 2000). Also is similar to S. circassa but S. circassa is distinguishable in having laterals perpendicular to the axis, horizontally elongated (Carras et al. 2006). Percentage of specimens 5%. (See Fig. 8). Dimensions See Table 1. Total range Albian?-Cenomanian (Carras et al. 2006) (Fig. 3). Range in studied sections Late Barremian-Early Aptian (Arkan stratigraphic section). Environments Open marine with normal marine salinity (Hernández-Romano et al. 1998). Inner platform (Carras et al. 2006) (Fig. 5). Biogeographical distribution Jordan, Egypt, Mexico, Turkey, Croatia and NE Iran. Salpingoporella muehlbergii (Lorenz 1902) Pia in Trauth Pia 1918. Pl. 1, Figs. 1–17 and Pl. 2, Figs. 1–26 Holotype 1902 Diplopora Muehlbergii n. sp.- Lorenz, p. 52–54 ; Figs. 3–6, 7?; Rhaetikon (Eastern Switzerland), BarremianAptian (Fig. 6). Selected citations 1999 Salpingoporella muehlbergii- Yilmaz, Pl. 3, Figs. 42, 43; Fele area, Western Taurides, Turkey, Barremian. 2006 Salpingoporella muehlbergii- Carras et al., Pl. VII, Figs. 1–4; SE France, Late Hauterivian-Late Barremian. 2008 Salpingoporella muehlbergii- Conrad et al., Pl. 2 (j–l); Southeast France, Early Barremian- Early Aptian. 2008 Salpingoporella muehlbergii- Sokač and Grgasović, Pl. VI, Figs. 9–11; Pl. VII; Dinaric karst, Barremian.

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2008 Salpingoporella muehlbergii- Michetiuc et al., Pl. IV, Fig. 3; Vâlcan Mountains, Danubian Domain; BarremianAptian. 2010 Salpingoporella muehlbergii- Taherpour Khalil Abad et al., Fig. 7 (f, g); Kopet Dagh basin, NE Iran, Late Barremian-Early Aptian. 2013 Salpingoporella aff. muehlbergii- Carević et al., Figs. 15 B, C, 17 Y, BB; Carpatho-Balkanides and Kopet Dagh basin, NE Iran; Late Barremian-Early Aptian. 2014 Salpingoporella muehlbergii- Granier et al., Fig. 7a–c; Canton of Vaud, Switzerland, Late Hauterivian-Early Barremian. Diagnostic features According to Carras et al. (2006), Blaterals set in quincunxes, 5-11 per verticil, horizontal (sometimes slightly leaning up), first with a narrow tubular proximal part, then widening out, circular, rhombic or irregular in section. Distal part of the laterals circular or slightly horizontally elongated. Questionable presence of cysts in the laterals. Calcareous skeleton made of colourless sparry calcite^. Remarks This species is similar to S. robusta but in S. muehlbergii the ramifications widen more strongly only at its distal ends (Sokač 1993). Ramifying thallus is the generic characteristics of this species (Sokač and Grgasović 2008). This species is characterized by its larger dimensions and transversally flattened branches. It is distinguished from other species by its rhombic branches, which have a transversally flattened shape. Salpingoporella dinarica, S. genevensis, S. katzeri and S. circassa have a similar branch type, but the varying dimensions between two portions of branches, and the number and shape of the branches are different to those found in these species (Yilmaz 1999). The specimen referred to Salpingoporella muehlbergii by Conrad (1970, Pl. 6, Fig. 4) actually corresponds to the basal, sterile portion (handle) of an alga that has been attributed to different genera and is presently assigned to Korkyrella texana (Carras et al. 2006). This is the most abundant species among the other species of Salpingoporella in the Kopet Dagh sedimentary basin. Its abundance is in the Sarcheshmeh Formation Percentage of specimens 52%. (See Fig. 8). Dimensions See Table 1. Total range Late Hauterivian-Early Aptian (Granier and Deloffre 1993, Carras et al. 2006), Late Aptian unconfirmed (Carras et al. 2006) (Fig. 3). Range in studied sections Late Barremian-Early Aptian (Arkan, Chenaran, Qheshqheh, Sisab stratigraphic sections), Early Aptian (Akher Dagh stratigraphic section).

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Environments Infralittoral, fresh water (Conrad and Peybernès 1976), inner to outer platform (Carras et al. 2006) (Fig. 5).

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Range in studied sections Late Barremian (Arkan stratigraphic section). Environments Lagoon (Fig. 5).

Biogeographical distribution According to Carras et al. (2006), this species next to S. annulata is cosmopolitan, with a wide longitudinal and latitudinal distribution, in northern, central and southern regions of the Tethys. Switzerland, Italy, France, Lebanon, Spain, Croatia, Hungary, Western Alps, Georgia, Romania, Oman, Portugal, Austria, Turkey, NE Iran. Salpingoporella parapiriniae Conrad et al. 2008 Pl. 3, Figs. 30–32 Holotype 2008 Salpingoporella parapiriniae n. sp.- Conrad et al., Fig. 2a–f; Southeast France, Early Barremian-Early Aptian (Fig. 6). Selected citations 1999 Salpingoporella cf. piriniae- Bucur, Pl. 1, Figs. 16, 21; Early Cretaceous, Romania. 2001 Salpingoporella cf. piriniae- Massera, Pl. 10, Photo 6, Southeast France, Bedoulian. 2010 Salpingoporella cf. parapiriniae- Taherpour Khalil Abad et al., Fig. 7e, Kopet Dagh basin, NE Iran, Late Barremian-Early Aptian. 2013 Salpingoporella cf. parapiriniae- Carević et al., [Specimen is not illustrated]; Kopet Dagh basin, NE Iran; Late Barremian-Early Aptian. 2014 Salpingoporella cf. parapiriniae- Khodashenas et al., Fig. 4 (B); Esfiyukh Mountain, NE Iran; Barremian-Aptian. Diagnostic features BThallus small, cylindrical, with laterals first forming even quincunxes, then somewhat irregularly arranged at periphery. Laterals funnel-like, horizontal or slightly tilted, circular in deep tangential section, sub-circular to irregularly polygonal at tip. Calcareous skeleton solid, made of colourless sparry calcite^ (Conrad et al. 2008). Remarks This species is similar to S. piriniae but in S. piriniae the laterals form irregular quincunxes or vertical files and the values of d are essentially smaller (Conrad et al. 2008). Percentage of specimens 2%. (See Fig. 8).

Biogeographical distribution Romania, SE France and NE Iran. Salpingoporella piriniae Carras and Radoičić 1991. Pl. 1, Figs. 18–20 and Pl. 3, Figs. 33, 34 Holotype 1991Salpingoporella piriniae n. sp.- Carras and Radoičić, Pl. 1; Figs. 1–7 (p.p.); Pl. 2, Figs. 1–17; Pl. 3, Figs. 1–9; Parnassus platform (Greece) and Southern Apennines (Italy), Early Barremian (Fig. 6). Selected citations 2006 Salpingoporella piriniae - Carras et al., Pl. VIII, Figs. 1– 6; Kefali hill, Greece; Early Barremian. 2008 Salpingoporella piriniae- Conrad et al., Fig. 2 (g–i); Southeast France, Late Barremian. 2010 Salpingoporella cf. piriniae- Taherpour Khalil Abad et al., Fig. 7 (c, d), Kopet Dagh basin, NE Iran, Late Barremian-Early Aptian. 2013 Salpingoporella cf. piriniae- Carević et al., [Specimen is not illustrated]; Kopet Dagh basin, NE Iran; Late BarremianEarly Aptian. Diagnostic features According to Carras et al. (2006), small, with laterals forming irregular quincunxes or vertical files, 10–14 per verticil, horizontal, funnel-like, circular in deep tangential section, polygonal at tip. Calcareous skeleton solid, made of colourless sparry calcite. Remarks This species is similar to S. muehlbergii but S. muehlbergii usually larger d/D ratio, w value usually smaller, spacing of the verticils (h value) proportionally larger, laterals usually circular or slightly flattened at tip (Carras et al. 2006). Percentage of specimens 2%. (See Fig. 8). Dimensions See Table 1. Total range Barremian-Early Aptian (Carras et al. 2006) (Fig. 3).

Dimensions See Table 1. Total range Barremian-Aptian (Conrad et al. 2008) (Fig. 3).

Range in studied sections Late Barremian (Arkan stratigraphic section).

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Environments Middle energy, back-reef to open lagoon (Carras et al. 2006) (Fig. 5). Biogeographical distribution Italy, Greece, Turkey, Romania, SE France and NE Iran. Salpingoporella sp. cf. S. biokovensis Sokač & Velic 1979 Pl. 3, Fig. 26 Holotype 1979 Salpingoporella biokovensis n. sp.- Sokač and Velic, Pls. 1, 2 and 3; Dalmatia (Croatia), Early Aptian (Fig. 6). Diagnostic features According to Carras et al. (2006), it is Bmedium-sized, with laterals forming vertical files (often slightly disturbed), 7-8 per verticil, horizontal, communicating with the main axis through a tiny pore, then regularly widening in the proximal half and abruptly swelling in the distal part, squared at distal end (rarely preserved). Calcareous skeleton made up of colourless sparry calcite^. Remarks Salpingoporella biokovensis differs from other species of the genus by the shape of its branches. The distal part of the branches in tangential sections, which exhibits deformed squares, clearly distinguishes the species. Salpingoporella hispanica and Salpingoporella adriatica have similar outer branches, but the number of branches and their dimensions clearly distinguish Salpingoporella biokovensis from these species (Yilmaz 1999). Percentage of specimens 1%. (See Fig. 8).

Holotype 1973 Salpingoporella steinhauseri n. sp.- Conrad et al., Fig. 7; Pl. 1, Figs. 1–4; Switzerland and SE France, Early-Middle Berriasian (Fig. 6). Selected citations 2006 Salpingoporella steinhauseri- Carras et al., Pl. X, Figs. 1–3; Mt. Mouty (SE France), Berriasian, Canton Neuchâtel, Switzerland), Early-Middle Berriasian. 2013 Salpingoporella steinhauseri- Schlagintweit and Enos, Fig. 7j, Berriasian-?Hauterivian, DSDP Sites 390 and 392, USA. 2014 Salpingoporella steinhauseri- Granier et al., Pl. 5, Figs. f–h; Middle Berriasian, Switzerland. Diagnostic features According to Carras et al. (2006), it is Bquite small, with laterals forming irregular quincunxes or vertical files, 6-11 per verticil, leaning up, starting with a rather unconspicuous constriction, then funnel-like, circular in tangential section. Calcareous skeleton made of colourless sparry calcite^. Remarks This species is similar to Salpingoporella m i l o v a n o v i c i a n d S . c i rc a s s a b u t t h e th a l l us o f S. milovanovici is usually smaller, with laterals also forming vertical files, but horizontal, more numerous and compressed. S. circassa differs from in having laterals perpendicular to the axis, horizontally elongated (Carras et al. 2006). Salpingoporella steinhauseri is an important chronostratigraphic marker of the Middle Berriasian in France and Switzerland (Conrad et al. 1973) but this is the first report on its occurrence in the Middle Aptian (Gargasian) in the Sarcheshmeh Formation (NE Iran).

Dimensions see Table 1. Percentage of specimens 1%. (See Fig. 8). Total range Late Hauterivian-Aptian (Granier and Deloffre 1993). late Late Hauterivian-Early Aptian (Carras et al. 2006) (Fig. 3). Range in studied sections Middle Aptian (Gelian stratigraphic section). Environments Open lagoonal to back-reef (Carras et al. 2006) (Fig. 5). Biogeographical distribution Croatia, Italy, Cuba, Greece, Turkey and Romania. Salpingoporella steinhauseri Conrad et al. 1973 Pl. 3, Fig. 27

Dimensions See Table 1. Total range Berriasian-Valanginian-Hauterivian? (Northern Tethyan domain), late Late Tithonian (Central Tethyan domain) (Carras et al. 2006) (Fig. 3). Range in studied sections Middle Aptian (Gelian stratigraphic section). Environments Marine, shallow water, inner platform (Carras et al. 2006) (Fig. 5). Biogeographical distribution Switzerland, SE France, Serbia, China, Turkey, Romania, Slovakia, Greece, Italy, USA and NE Iran.

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Salpingoporella polygonalis Sokač 1996. Pl. 3, Figs. 28, 29 Holotype 1996Salpingoporella polygonalis n. sp.- Sokač, Pl. 3, Figs. 3– 10; Pl. 4, Figs. 1–17; Mljet island Dalmatia, Croatia), Early Barremian (Fig. 6). Selected citations 2006 Salpingoporella polygonalis- Carras et al., Pl. VIII, Figs. 12–16; Dalmatian islands, Croatia, Early Barremian. Diagnostic features According to the original description (Sokač 1996), Bthe massive calcareous skeleton of the cylindrical thallus of the new species is composed up of light grey, very coarse-grained, and variously recrystallized calcite. The outer surface of the calcareous envelope is always damaged, partly destroyed or dissolved, and subsequently recrystallized. That the wall was originally comparatively thick is indicated by rarely preserved, outwardly projecting, thin partitions between the distal ends of the ramifications^. Remarks This species clearly differs from S. katzeri, in which the branches also swell very rapidly, often being more swollen toward the upper side, with thin partitions between the branches and the more or less pronounced hexagonal to rhombic pattern al the distal ends. According to the original description, the main axis makes 30–35% of the total diameter when the thin calcareous partitions between the distal end of the laterals are preserved. Due to the secondary destruction of the partitions, the outer diameter of the skeleton commonly appears smaller, resulting in the d/D ratio to be larger (44– 55%) (Carras et al. 2006). This species seems to be missing on the northern Tethyan Domain.

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Conclusion The Order Dasycladales is poorly introduced and described from the Barremian-Aptian strata of the Kopet Dagh sedimentary basin (NE Iran), except in Taherpour Khalil Abad et al. (2010), Bucur et al. (2013) and Carević et al. (2013). The present study is a contribution to knowledge of BarremianAptian triploporellacean algae from the Tirgan and Sarcheshmeh formations. The Salpingoporella contents of the Barremian-Aptian limestone successions of the Tirgan and Sarchehsmeh formations are documented, and some species (Salpingoporella heraldica, S. biokovensis, S. steinhauseri and S. polygonalis) have described for the first time from this sedimentary basin. New approaches in systematic description (biometrical features) are yielded from the Northern Tethyan Domain which shows differences in measuring between north and south of the Tethyan Domain. In addition, by detailed studies on the flora association, S. muehlbergii is the main instructive species between the other ones (for more data, see Fig. 8). Acknowledgements Anonymous reviewers are thanked for their helpful comments and constructive review of the manuscript.

Appendix: list of taxa Salpingoporella cemi Salpingoporella hasi Salpingoporella heraldica Salpingoporella hispanica. Salpingoporella milovanovici Salpingoporella muehlmbergi Salpingoporella parapiriniae Salpingoporella piriniae Salpingoporella sp. cf. S. biokovensis Salpingoporella steinhauseri Salpingoporella polygonalis

Percentage of specimens 2%. (See Fig. 8). Dimensions See Table 1.

References

Total range Late Hauterivian-Early Barremian (Sokač 1996), Late Hauterivian-Early Aptian (Carras et al. 2006) (Fig. 3).

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Range in studied sections Middle Aptian. Environments Shallow subtidal, inner platform (Carras et al. 2006) (Fig. 5). Biogeographical distribution France, Hungary, Romania, Italy, Greece, Montenegro and NE Iran.

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