Introduction

The Miocene epoch (23.03–5.33 Ma) was a time interval of global warmth, and also, continental configurations and mountain topography transitioned toward modern conditions, many faunae and florae evolved into the same taxa that exist today (Steinthorsdottir et al. 2021). During the Miocene, the Terminal Tethyan Event, and subsequently the closure of the Tethyan Seaway, which connected the proto-Mediterranean Sea with the Indian and Atlantic Oceans, blocked thermohaline exchanges between them and possibly caused a major climate change in the Middle Miocene Climatic Transition (Sun et al. 2012). Although the Tethyan Seaway was closed due to the collision of the African/Arabian and Iranian/Eurasian plates, Sadr and Schmiedl (2017) mentioned that a climate deterioration had been related to major paleogeographic changes, including the separation of Antarctica and the closure of the Tethyan Ocean, resulting in the reorganization of ocean circulation and heat transport. In other words, the closure was related to both tectonic and climatic changes (Sadr and Schmiedl 2017).

However, to date, the exact timing and the involved forcing mechanism of the final closure of the Tethyan Seaway are still debated. Sun et al. (2012) suggested that this Seaway experienced stepwise evolutionary processes changing from a partially opened seaway, restricted marine connection, intermittent connections, to permanent closure in its northwestern segment during the early and middle Miocene.

Reuter et al. (2007) mentioned the Tethyan seaway as Iranian gateways, and suggested that continuous shallowing and restriction of the seaway generated an increasingly hostile hypersaline environment and a biogeographic barrier for marine biota, during the end of the Burdigalian, by comparing Atlantic-Mediterranean and Indo-Pacific bioprovinces with the environmental conditions of the Zagros Basin.

Studied strata belong to the Zagros Basin and Makran Basin, and it is formally suggested that they are separated from each other by the Minab Fault, but the boundary is not accurate. The Minab Fault marks a sharp structural and stratigraphic divide between the Zagros Basin and the Iranian Makran (Gansser, 1955; Falcon, 1969).

Tectonically, Shearman et al. (1976) divided Makran and Zagros folded zone and mentioned that the deep structural simplicity below the plains suggests a long period of gentle subsidence in pre-Miocene times, perhaps going back to Early Mesozoic times, allowing several kilometers of sediment to be deposited, in Minab anticline, where the deep crustal fracture below the Zendan fault line (i.e., the Oman Line) could have been active in a north-south direction, during all pre-Miocene time and did not disturb this area, which was, however, compressed by the westerly Mio-Pliocene movements that are still going on (Falcon 1976).

In sum, there is almost no research and investigation about Mio-Pliocene strata in Qeshm Island, Zagros Basin (fig. 1a). Also, in the Makran Basin, this study only sampled from one section (Gushi Marl), from the Makran Unit (see Tab. 1), where the accurate ages of strata were never studied, and all formations and geological units are informal.

The Mio-Pliocene strata of Qeshm Island have been suggested as Mishan/Aghajari formations (see fig. 1a; Huber 1977; Hassani et al. 2014), and Gushi Marl strata from the Makran Unit have been mentioned as Late Miocene strata, while none of the studied outcrops have been studied before. This study aims to reveal the age and paleoenvironmental conditions of the Gushi Marl and the Mishan Formation by using isolated foraminifera samples, and also to investigate the timing possibilities of closure of the Iranian Gateway.

Fig 1a- Geological map of the investigated areas; A. Terrain map of Iran; B. Geological map of Qeshm Island, Persian Gulf (after Huber 1977), with three sampled localities: 1) Direstan outcrop near the village; 2) Kendaloo outcrop in NE of the Kendaloo harbour; 3) Stars Valley outcrop.

Fig 1b- Geological map of Minab region (after Peterson and Rudzinskas 1982), 4) Bemani outcrop.

Fig 2- Some index foraminifera of the studied sections. Including Direstan outcrop (Qeshm Island, Zagros Basin), Kendaloo outcrop (Qeshm Island, Zagros Basin), and Bemani outcrop (Minab Province, Makran Basin).

Geological Settings

The Zagros Basin is one of the most universal oil and gas basins, located in the west to south of Iran and in the north of the Arabian Plate (Alsharhan and Nairn 1997; Daneshian et al. 2016).

The geological structure of Qeshm Island consists of four main anticlines, including Souza-Zirang, Holor, Salakh, and Goveie, with axis in strike of north-east to south-west and Gavarzin with axis in strike of north-west to south-east, and the main rock formations of the island are the Salt dome of Hormoz, marl of Mishan, marl and sandstone of Aghajari, and Plio-Pleistocene rocks and Quaternary calcareous traces (Gerivani et al. 2011).

Mishan Formation (age: Early to middle Miocene) consists of low-weathering gray marl and ridge-forming ribs of shelly limestone with abundant microfauna, in the type section (Fars Province;Rahmani and Vaziri-Moghaddam 2010)

The top of the highest gray, marine marl of the Mishan Formation is defined as the basal contact of the Aghajari Formation, which transitionally overlies the Mishan Formation (James and Wynd 1965). The Marly Member conformably overlies the top of the Guri Member in Bandar Abbas (Gholamalian et al. 2023), in the Northern lands of the Strait of Hormoz.

In Qeshm Island, and at the Direstan outcrop, sandy fossiliferous limestone, which was assumed as the uppermost part of the Mishan Formation, includes almost all organisms that have been reported from the Mishan Formation in other parts of the Zagros Basin (e.g., Vega et al. 2012; Fanati Rashidi et al. 2014; Daneshian et al. 2016). In Kendaloo sandy fossiliferous limestone is very similar to Direstan’s sandy fossiliferous limestone, lithologically. The uppermost strata of the Mishan Formation on Qeshm Island consist of sandy, fossiliferous limestone. These deposits are characterized by a diverse and exceptionally well-preserved microfauna, including foraminifers, ostracods, fish teeth, early-stage bivalves, and echinoid needles (Haskouei et al. 2024b).

Palaeobiological studies of the Mishan Formation (Direstan outcrop) and Gushi marls (Makran Basin) have reported ostracods and pectinid bivalves. The ostracod fauna of the studied outcrops shows significant similarity to that of the Fatha Formation in Northern Iraq, supporting its interpretation as an intermediate bioprovince between the Mediterranean and Indo-Pacific realms.

 (Hawramy and Khalaf 2013), and were mentioned as an intermediate bioprovincial zone between Mediterranean and Indo-Pacific (Haskouei et al. 2025a). Overall, in Qeshm Island, studied strata that have been assumed as uppermost parts of Mishan Formation (e.g. Parast et al. 2020; Haskouei 2025) include organisms as bryozoans, echinoids, corals, gastropods, pectinid and oyster bivalves, crustaceans such as crabs and balanoids, and their shell fragments, while Gushi marls only bear ostreids (oyster bars) and balanoids, as macrofossils (Haskouei et al. 2025a; see fig 2).

McCall (2002) suggested Gushi Marl as Late Miocene strata of Makran accretionary prism (Tab. 1), which is dominated by grey, gypsiferous marl, mudstone and shale, with subordinate brown interbeds of friable sandstone (poorly sorted lithic arenite) and siltstone., and also, has been mentioned that the Gushi Marl crops out in the western part of the Minab and Taherui quadrangles (fig. 1b), where has an estimated thickness of 2250 m. The Kheku Sandstone overlies the Gushi Marl conformably, in Taherui quadrangles (Peterson and Rudzinskas 1982; McCall 2002).

Overall, in compare to Qeshm Island, the among of Miocene strata of Makran Basin are a very huge among of accumulation, which convinced us to show a detailed list of Miocene strata of Makran Basin (see Tab. 1). Also, it is necessary to mention that Gushi Marl is a very thickened strata and the samples of this paper have been collected from the uppermost part of the Gushi Marl strata.

Table 1: Mio-Pliocene Stratigraphic units of the Makran accretionary prism, listed with their age ranges (modified after McCall 2002)

Material and Methods

Sediment samples were collected from three locations in Qeshm Island (SE of Zagros Basin) and one location in the Minab region (SW of Makran Basin), see fig. 1b. The studied sections from Qeshm Island are Stars Valley (26°52'9.99"N; 56°7'20.42"E), Direstan (26°44'14"N; 55°56'07"E), and Kendaloo (26°41'45.84"N; 55°55'25.67"E). The access roads to all three sections are available due to the first and the last are tourist attraction places, and Direstan is a Village near the road to Qeshm Island airport. Likely, the only section of Makran basin, is named Bemani (26°55'42"N; 57°07'25"E) is accessible cause of being close to the main road of Minab County (fig. 1b). This paper examines the taxonomic description of 67 samples of foraminifera in total, 48 benthic foraminifera samples, which show identified 10 genera and 18 species, and also, 10 planktonic foraminifera samples, which show identified 6 genera and 5 species, illustrated by SEM images. All specimens are housed in the Department of Geology, Faculty of Science, University of Isfahan, Iran, under the acronym of IUMC.

Systematic Paleontology

Multiple references have been used for the systematic description of foraminifera (Luczkowska 1974; van der Zwaan 1982; Papp and Schmid 1985; Loeblich and Tappan 1988; Szczechura and Abd-Elshafy 1988; van de Poel 1992; Hauser and Grunig1993; Debenay 2012; Poole and Wade 2019; Tabita and Nathan 2019).

Abbreviations: St_ Stars Valley outcrop, K_ Kendaloo outcrop, D_ Direstan outcrop, BM_ Bemani outcrop, BF_ Benthic Foraminifera, P_ Planktonic, ML_ Miliolid.

Class GLOBOTHALAMEA Pawlowski, Holzmann, Tyszka 2013

Order ROTALIIDA Delage and Hérouard 1896

Superfamily ROTALIOIDEA Ehrenberg 1839

Family AMMONIIDAE Saidova 1981

Subfamily AMMONIINAE Saidova 1981

Genus ASTEROROTALIA Hofker 1950

Asterorotalia dentata (Parker and Jones 1865)

Fig. 4A-I

1865 Rotalia beccarii (Linnaeus) var. dentata (Parker and Jones), pp. 387-388, 422, pl. 19, fig. 18a-c.

2006 Asterorotalia dentata (Parker and Jones); Sohrabi et al. p. 18, pl. 1, fig. 1.

2012 Asterorotalia dentata (Parker and Jones); Mossadegh et al. p. 358, figs. 3(at–av).

2014 Asterorotalia dentata (Parker and Jones); Panchang and Nigam, pl. 36, fig. 12a-b.

2017 Asterorotalia dentata (Parker and Jones); Bhaumik et al. p. 440, pl. 1, fig. m.

2019 Asterorotalia dentata (Parker and Jones); Tabita and Nathan, p. 15, figs. 4.13-14.

Material: Altogether 9 specimens were collected and studied: one specimen from the Mishan Formation of the Direstan outcrop, two from the Kendaloo outcrop, two from the Stars Valley outcrop, Qeshm Island, and four specimens collected from the Gushi Marl, the Bemani outcrop, Minab County, southern Iran.

Diagnosis and Description: see Loeblich and Tappan (1988) and also Tabita and Nathan (2019).

Occurrence and Distribution: Miocene (Serravallian to Tortonian), Limestone Bed of Baripada, Mayurbhanj District, Odisha, India (Bhaumik et al. 2017). Middle Pliocene, Southeastern of Zagros Basin, Iran (Hassani and Hosseinipour 2017). Late Pleistocene, Kish Island, Persian Gulf, Iran (Mossadegh et al. 2012). Middle Holocene, core T2S3, Strait of Hormoz, Southern Iran (Hamzeh et al. 2021).

Remark: In the Zagros Basin (Iran), Hassani and Hosseinipour (2017) suggested that the species is a member of biozone Z7 (Globorotalia acostaensis abundance zone), which illustrates the Piacenzian-Zanclean boundary, mentioned by Bolli (1966) and Postuma (1971), and also correlated with the global biozone N20 (Hassani and Hosseinipour, 2017; also check figure 3 by Spezzaferri et al. 2002).

Environment: Bhaumik et al. (2017) reported the species from a shallow marine oxic environment, and also, Mossadegh et al. (2012) reported it from raised Coral reef sequences.

Asterorotalia pulchella (d'Orbigny 1839a)

Fig. 4J

1839a Rotalia (Calcarina) pulchella d’Orbigny, p. 80, pl. 5, figs. 16-18.

1930 Rotalia pulchella (d'Orbigny); Hofker, p. 37, Pl. 16, figs. 7-10.

1951 Asterorotalia pulchella (d'Orbigny); Hofker, p. 505-508, figs. 343-344.

1968 Asterorotalia pulchella (d'Orbigny); Hofker, p. 27, Pl. 8, figs. 8-10; Pl. 9, figs. 1-7.

2001 Asterorotalia pulchella (d’Orbigny); Szarek, p. 147, pl. 27, figs. 11-12.

2015 Asterorotalia pulchella (d’Orbigny); Hanagata and Nobuhara, p. 119, figs. 35.9-10.

2019 Asterorotalia pulchella (d’Orbigny); Tabita and Nathan, p. 12, figs 3.17-18.

Material: Only one specimen was collected and studied, from Gushi Marl, Bemani outcrop, Mina County, southern Iran.

Diagnosis and Description: see Hofker, (1951; 1971), Loeblich and Tappan, (1988) and also, Tabita and Nathan, (2019).

Occurrence and Distribution: Pliocene to Holocene of Cuba, Indonesia, Java, Persian Gulf as Rotalia pulchella (Loeblich and Tappan 1988, d’Orbigny 1839a). Quaternary, Pleistocene to Holocene of Kutei Basin, Indonesia (Loeblich and Tappan 1988). Recent, Bay of Jakarta, Java (Hofker 1968).

Remark: Despite mechanical abrasion, the species is identifiable, and characteristic features are still distinct.

Environment: This species was found at a depth of 18 m, always in a muddy environment (Hofker 1978), and also prefers the silty mud environment with badly aerated water (Hofker 1968).

Genus CHALLENGERELLA Billman, Hottinger, and Oesterle 1980

Challengerella bradyi (Billman et al. 1980)

Fig. 4K-M

1980 Challengerella bradyi, Billrnan, Hottinger, and Oesterle, p. 81.

1988 Challengerella bradyi (Billrnan et al.); Loeblich and Tappan, pl. 770, figs. 1-8.

2017 Challengerella bradyi (Billrnan et al.); Bhaumik et al. p. 440, pl. 1, figs. q-s.

2019 Challengerella sp., (Billrnan et al.); Amao et al. p. 940, figs. 4(g-i).

2023 Challengerella bradyi (Billrnan et al.); Shareef et al. p. 196, pl. 4, fig. 2.

Material: Only three specimens were collected and studied, from the Mishan Formation, Zagros Basin, Iran. One from Direstan, one Kendaloo and one Stars Valley outcrops, Qeshm Island.

Diagnosis and Description: see Loeblich and Tappan (1988).

Occurrence and Distribution: Langhian-Serravallian, Fatha Formation, Southern Iraq (Shareef et al. 2023). Serravallian to Tortonian, Limestone Bed of Baripada, Mayurbhanj District, Odisha, India (Bhaumik et al. 2017). Middle Holocene, core T2S3, Strait of Hormoz, Southern Iran (Hamzeh et al. 2021). Late Pleistocene, Kish Island, Persian Gulf, Iran (Mossadegh et al. 2012). Holocene, Gulf of Elat, Red Sea (Loeblich and Tappan 1988). Recent, the Persian Gulf, southern Iran (Saidova 2010).

Remark: In Zagros Basin (Iran), Hassani and Hosseinipour (2017) suggested that the species is a member of biozone Z1 (= Globigerinoides trilobus zone of Postuma 1971), which show timing Aquitanian before the boundary of early Burdigalian, coincides with the beginning of global biozone N5 (Hassani and Hosseinipour 2017; also check figure 3 by Spezzaferri et al. 2002).

Environment: Bhaumik et al. (2017) mentioned the species from a shallow marine oxic environment. Moreover, the species were described from depths of 20–80 m, in the modern Persian Gulf (Saidova 2010), and also from raised Coral reef sequences (Mossadegh et al. 2012).

Genus ROTALINOIDES Saidova 1975

Rotalinoides compressiuscula (Brady 1884)

Fig. 4N-P

1884 Rotalia papillosa var. compressiuscula Brady, p. 708, pl. 107, fig. 1.

2015 Rotalinoides compressiuscula (Brady); Hanagata and Nobuhara, p. 119, fig. 35.11-12.

2019 Rotalinoides compressiuscula (Brady), Tabita and Nathan, p. 15, figs. 4.19-2.

Material: Overall, three specimens were collected and studied from Direstan, Kendaloo, and Stars Valley outcrops, Qeshm Island, Zagros Basin, Iran.

Diagnosis and Description: See Tabita and Nathan (2019).

Occurrence and Distribution: Mio-Pliocene, Mishan Formation, south of Iran (Hassani and Hosseinipour 2017). Recent Coast of India (Tabita and Nathan 2019).

Remark: In Zagros Basin (Iran), Hassani and Hosseinipour (2017) mentioned that the species is a member of biozone Z6 (Globorotalia crassaformis range zone), which shows the Mio-Pliocene boundary, suggested by (Bolli 1966; Postuma 1971), and also illustrates a coincidence with the beginning of global biozone N19 (Hassani and Hosseinipour 2017; also check figure 3 by Spezzaferri et al. 2002).

Environment: The species mentioned from the 38 to 83 m water depth of a slightly muddy sand to muddy sand environment of the modern shelf (Anbuselvan and Nathan 2017).

Genus AMMONIA Brünnich 1772

Ammonia beccarii (Linnaeus 1758)

Fig. 5A-C

1758 Nautilus beccarii Linnaeus, p. 710, pl. 1, fig. 1.

2001 Ammonia beccarii (Linnaeus); Szarek, p. 148, pl. 26, figs. 13-15.

2003 Ammonia beccarii (Linnaeus); Javaux and Scott, p. 10, fig. 2.2-3.

2005 Ammonia beccarii (Linnaeus); Debenay, Millet and Angelidis, p. 334, pl. 2, fig. 17.

2012 Ammonia beccarii (Linnaeus), Milker and Schmiedl, p. 117, fig. 27.1-2.

2019 Ammonia beccarii (Linnaeus), Tabita and Nathan, p. 15, figs. 4.7-8.

Material: Only three specimens were collected and studied from the Mishan Formation, Zagros Basin, Iran. All are from the Direstan outcrop, Qeshm Island. Although two other outcrops on the Island bear the species, because of broken shells and lack of distinct characteristic features, we decided not to collect them.

Diagnosis and Description: See Tabita and Nathan (2019).

Occurrence and Distribution: Early to Middle Miocene (Aquitanian to Langhian), the Mishan Formation, the southeastern end of the Zagros Folded Zone, Iran (Fanati Rashidi et al. 2014). Recent (Hayward et al. 2021). Late Aquitanian to early Burdigalian, Asmari Formation, SW Zagros Basin, Iran (Roozpeykar and Moghaddam 2015).

Remark: In the present study, the species has been observed from the uppermost parts of the equal strata of the Mishan Formation, a fossiliferous limestone, associated with coral patches. Also, the species were found in limestone, marly limestone, and calcareous marl in many parts of the Khorgu section, Mishan Formation, the southeastern end of the Zagros Folded Zone (Fanati Rashidi et al. 2014).

Environment: The outer shelf environment (Fanati Rashidi et al. 2014). Modern inner shelf environment (depth 0-50 m), for A. beccarii, and as genus Ammonia, one of the two most common globally shallow-marine and estuarine foraminiferal genera (Hayward et al. 2021). Overall, A. beccarii is a euryhaline species, extensively distributed in the littoral and neritic environment (Debenay et al. 1998; Anbuselvan and Nathan 2017).

Superfamily EPONIDACEA Hofker 1951

Family EPONIDIDAE Hofker 1951

Subfamily EPONIDINAE Hofker 1951

Genus EPONIDES de Montfort 1808

Eponides repandus (Fichtel and Moll 1798)

Fig. 5E-I

1798 Nautilus repandus Fichtel and Moll, p. 35, pl. 3, figs. a-d.

1971 Eponides repandus (Fichtel and Moll); Murray: 173, pl. 72, figs. 1- 4.

1974 Eponides repandus (Fichtel and Moll); Andreieff et al. pl. 7, fig 12.

1982 Eponides repandus (Fichtel and Moll); Szczechura, pl. 14, figs. 3-4.

1988 Eponides repandus (Fichtel and Moll); Szczechura and Abd-Elshafy, pl. 11, fig. 1

1992 Eponides repandus (Fichtel and Moll); Hansen and Revets, p. 168, pl. 1, figs. 7-9.

1993 Eponides repandus (Fichtel and Moll); Hauser and Grunig, pl. 4, figs. 4-7.

2001 Eponides repandus (Fichtel and Moll); Szarek, p. 133, pl. 19, figs. 9-11.

2003 Eponides repandus (Fichtel and Moll); Javaux and Scott, p. 14, fig. 3.1-2.

2012 Eponides repandus (Fichtel and Moll); Debenay, p. 196.

2014 Eponides repandus (Fichtel and Moll); Panchang and Nigam, pl. 29, fig. 12a-c.

2019 Eponides repandus (Fichtel and Moll); Tabita and Nathan, figs 8.18-19.

Material: Altogether five specimens were collected and studied, from Zagros Basin, Iran, two from Direstan, two from Kendaloo, and one from the Stars Valley outcrop, Qeshm Island.

Diagnosis and Description: See Szczechura and Abd-Elshafy (1988) and also Loeblich and Tappan (1988). 

Occurrence and Distribution: Middle Miocene, the Gulf of Suez, Egypt (Szczechura and Abd-Elshafy 1988). Late Oligocene to early Miocene of Qom Formation, Central Iran (Nouradini et al. 2017). Holocene, Gulf of Naples, Italy (Loeblich and Tappan, 1988). Recent, the Gulf of Aqaba, Egypt (Hauser and Grunig 1993).

Environment: The species mentioned from the 38 to 83 m water depth of a slightly muddy sand to muddy sand environment of the shelf (Anbuselvan and Nathan 2017). Also, the species has been mentioned from the modern shallow-water (depth=85 m), and an almost high temperature environment, 22°C (Hofker 1978).

Eponides isabellanus (d'Orbigny 1846)

Fig. 5D

1839 Rosalina isabelleana d'Orbigny, p. 43, pl. 6, figs. 10-12.

1971 Eponides isabellanus (d'Orbigny); Herb, p. 267, pl. 1, figs. 1 and 18.

1980 Discorbis isabelleanus (d'Orbigny); Boltovskoy et al. p. 80, pl. 11, figs. 8-12.

1993 Eponides isabellanus (d'Orbigny); Hauser and Grunig, pl. 5, figs. 1-7.

Material: only one specimen was collected and studied, from the Mishan Formation, Direstan outcrop (Qeshm Island), Zagros Basin, Iran.

Diagnosis and Description: See Hauser and Grunig (1993) and also Boltovskoy et al. (1980).

Occurrence and Distribution: Recent, Drake Passage, Antarctica (Hauser and Grunig 1993). Recent, Islas Malvinas (Boltovskoy et al. 1980).

Remark: Hauser and Grunig (1993) suggested that numerous species assigned to Eponides by their authors are placed in different genera now, or several species assigned to Eponides today originally belonged to other genera. Based on this, Eponides isabellanus is an illustration of the species in the family that are different from others by the great variation in coiling and ornamentation (Hauser and Grunig 1993).

Environment: Cold water, open sea environment, Antarctica (Hauser and Grunig 1993). Inner shelf, depth 42m (Herb 1971).

Genus POROEPONIDES Cushman 1944

Poroeponides lateralis (Terquem 1878)

Fig. 5J-M

1878 Rosalina lateralis Terquem, p. 25, pl. 2, fig.11.

1964 Poroeponides lateralis (Terquem); Shchedrina in Rauzer-Tchernousova, p. 98, fig. 5(1-2).

1980 Poroeponides lateralis (Terquem); Boltovskoy et al. p. 109, pl. 25, figs. 4-7.

1994 Poroeponides lateralis (Terquem); Jones, pl. 106, figs. 2 (a-c).

2007 Poroeponides lateralis (Terquem); Talib and Farroqui, p. 25, pl. 1, fig. 18.

2012 Poroeponides lateralis (Terquem); Debenay, p. 210.

2019 Poroeponides lateralis (Terquem); Tabita and Nathan, p. 31, figs. 9(1-2).

Material: Four specimens were collected and studied from the Zagros Basin, Iran, one from Kendaloo, one from Direstan, and two from the Stars Valley outcrop, Qeshm Island.

Diagnosis and Description: See Boltovskoy et al. (1980), Loeblich and Tappan (1988), and also Jones (1994).

Occurrence and Distribution: Cosmopolitan, Pliocene to Holocene, Holocene of the West Atlantic, off Rhode Island, USA (Loeblich and Tappan 1988). Late Pliocene of the Isle of Rhodes, Antarctica (Boltovskoy et al. 1980). Recent, the Gulf of Aden (Shchedrina 1964).

Remark: Lunatic-like chambers in the species of Stars Valley (fig. 5M) are longer and narrower than) the specimens from two other outcrops, Direstan and Kendaloo (such as the Recent species studied by Shchedrina 1964).

Environment: Baccaert (1987) suggested that Poroeponides lateralis is a shallow-water species of the Peri-reefal Area of the Great Barrier Reef, in normal salinity and the depth between almost 10 m to 32 m. Also, it occurs in the mixohaline waters of Lagoa dos Patos (Boltovskoy et al. 1980).

Family ELPHIDIIDAE Galloway 1933

Subfamily ELPHIDIINAE Galloway 1933

Genus ELPHIDIUM de Montfort 1808

Elphidium fichtellianum (d'Orbigny 1846)

Fig. 6A-B

1846 Polystomella fichtelliana d'Orbigny, p.125, pI. 6, figs. 7, 8.

1951 Elphidium fichtellianum (d'Orbigny); Marks, p. 52, pl. 6, figs. 12a, b.

1976 Elphidium fichtellianum (d'Orbigny); Berggren, p. 112, pl. VI, figs. 7-8.

1982 Elphidium fichtellianum (d'Orbigny); van der Zwaan, p. 189, pl. 9, fig. 8.

1985 Elphidium fichtellianum (d'Orbigny); Papp and Schmid, p. 187, pl. 40, figs. 2-5.

1992 Elphidium fichtellianum (d'Orbigny); van de Poel, p. 9, pl. 1, fig. 4.

2000 Elphidium fichtellianum (d'Orbigny); Baggley, p. 1087, pl. 2, fig. 5.

2013 Elphidium chtellianum/complanatum (d'Orbigny); Jones, p. 283, pl. 72, figs. 5-8.

2019 Elphidium fichtellianum (d'Orbigny); Amao et al. p. 940, figs. 4(d-e).

Material: 2 specimens were collected and studied from Kendaloo and Direstan, Qeshm Island, Zagros Basin, Iran.

Diagnosis and Description: Shell coiled spirally, planispiral, hyaline, with crescentic-form chambers. Sutures are depressed, curved, and ponticuli. Shell ornamentation includes hispid, pustulose, costate, and keeled on two sides of the shell. Also, see Papp and Schmid (1985).

Occurrence and Distribution: Late Burdigalian to Langhian, Asmari Formation in NW of the Zagros basin (Roozpeykar et al. 2019). Tortonian, Kasaba Formation, Turkey (Hayward et al. 1996). Late Miocene (Messinian) of the Andalusian stratotype, western Guadalquivir Basin, SW Spain (Berggren 1976). The latest Tortonian to earliest Messinian, Carboneras-Nijar Basin, SE Spain (van de Poel 1992), and also, the Mediterranean Late Miocene strata (the lowermost of Messinian), Abad Member, Turre Formation, SE Spain (Baggley 2000). Recent, the Iranian coast of the Persian Gulf (Amao et al. 2019).

Remark: Despite mechanical abrasion, the species is identifiable, and characteristic features are still distinct.

Environment: Shallow water, estimated at ~30 m water depth, sandy marls environment (Berggren 1976). The Mediterranean species, indicating Warm–Temperate conditions, warmer than the same species obtaining at present (Jones 2013). Benthic species characteristic of deep open marine (van de Poel 1992), and also, common in pre-concentration basins of natural salt works of the Mediterranean coastal modern environments (Zaninetti 1982; van de Poel 1992)

Elphidium crispum (Linnaeus 1758)

Fig. 6C-E

1758 Nautilus crispus Linnaeus, p. 709, pl. 19, fig.1d.

1964 Elphidium crispum (Linnaeus); Voloshinova and Kuznetsova in Rauzer-Tchernousova, p. 151, pl. I, figs. 1(a-b).

1985 Elphidium crispum (Linnaeus); Papp and Schmid, p. 187, pl. 40, figs. 6-10.

2000 Elphidium crispum (Linnaeus); Baggley, p. 1087, pl. 2, fig. 4.

2001 Elphidium crispum (Linnaeus); Szarek, p.150, pl. 28, fig. 3.

2006 Elphidium crispum (Linnaeus); Oflaz, p. 235, pl. 11, fig. 11.

2007 Elphidium crispum (Linnaeus); Talib and Farroqui, p. 21, pl.1, fig. 25a-b.

2012 Elphidium crispum (Linnaeus); Debenay, p. 216.

2012 Elphidium crispum (Linnaeus); Milker and Schmiedl, p. 120, fig. 27.13-14.

2013 Elphidium crispum (Linnaeus); Jones, p. 283, pl. 72, figs. 1-2.

2013 Elphidium crispum (Linnaeus); Hewaidy et al., p. 43, pl. 11, figs. 12(a-b).

2019 Elphidium crispum (Linnaeus); Tabita and Nathan, p. 21, fig. 6.14

Material: Overall, three specimens were collected and studied, from the Mishan Formation, Zagros Basin, Iran, each of them are from Direstan, Kendaloo, and Stars Valley outcrops, Qeshm Island.

Diagnosis and Description: See Hayward et al. (1997), and also, Papp and Schmid (1985).

Occurrence and Distribution: Cosmopolitan, Burdigalian to Langhian of the Asmari Formation in NW of the Zagros basin (Roozpeykar et al. 2019). Tortonian of Kasaba Formation, Turkey (Hayward et al. 1996). The topmost part of the Miocene succession of the Rosetta Formation, Nile Delta area, Egypt (Hewaidy et al. 2013). The Mediterranean Late Miocene strata (the lowermost of Messinian), Abad Member, Turre Formation, SE Spain (Baggley 2000). Miocene to Recent (Jones 1994).

Remark: The species has been mentioned as an indicator of Cool-Temperate aspect (slight cooling), Slindon formation, Boxgrove, Britain (Jones 2013). Also, Boltovskoy et al. (1980) suggested that Elphidium crispum is one of the species of higher latitudes, which are significantly affected by temperature changes.

Environment: According to Jones (1994), neritic to middle bathyal (to 355 fathoms), and Farooqui (1990) mentioned as a widely recorded and commonly occurring species in shallow turbulent water of different parts of the modern environments, which can tolerate a wide range of salinity and temperature fluctuation. Also, the species have been mentioned from shallow backreef modern environments with intertidal to subtidal conditions, depth shallower than 10 m, eventual salinity fluctuations or normal salinity, also from barrier reef system (intertidal to slightly subtidal), and from peri-reefal area of Great Barrier Reef, in normal salinity and the depth between almost 10 m to 32 m (Boltovskoy et al. 1980).

Elphidium craticulatum (Fichtel and Moll 1798)

Fig. 6F-G

1798 Nautilus craticulatus Fichtel and Moll, p. 51, pl. 5, figs. h-k.

1987 Elphidium craticulatum (Fichtel and Moll); Baccaert, p. 252; pl. 102, fig. 8; pl. 103, figs 1a, b.

1994 Cellanthus craticulatum (Fichtel and Moll); Loeblich and Tappan, p. 167; pl. 380, figs 1, 2, 7-10.

1997 Elphidium craticulatum (Fichtel and Moll); Hayward et al., p. 73; pl. 7, figs 5-12.

2007 Elphidium craticulatum (Fichtel and Moll); Talib and Farroqui, p. 21, pl. 1, fig. 24 a-b.

2010 Elphidium craticulatum (Fichtel and Moll); Sohrabi-Mollayousefi and Sahba, p. 965, pl. 1, fig. 5.

2012 Elphidium craticulatum (Fichtel and Moll); Debenay, p. 219.

2014 Elphidium craticulatum (Fichtel and Moll); Panchang and Nigam, pl. 38, fig. 5a-b.

2019 Elphidium craticulatum (Fichtel and Moll); Tabita and Nathan, p. 21, fig. 6(13).

2023 Elphidium craticulatum (Fichtel and Moll); Shareef et al. p. 196, pl. 4, figs. 10 and 10A.

Material: Two specimens were collected and studied from the Mishan Formation, in Stars Valley and Direstan outcrops, Qeshm Island.

Diagnosis and Description: See Debenay (2012), and also, Tabita and Nathan (2019).

Occurrence and Distribution: Langhian-Serravallian, Fatha Formation, Southern Iraq (Shareef et al. 2023). Holocene of Indonesia (Loeblich and Tappan 1988). Recent, the Great Barrier Reef of Australia (Baccaert 1987), and New Caledonia, Southwestern Pacific (Debenay 2012), also, Persian Gulf (Sohrabi-Mollayousefi and Sahba 2010).

Remark: have been reported from the Persian Gulf as among some species, which indicate abnormal tests and are capable of tolerating elevated levels of nutrients; the average temperature of the Persian Gulf is 23.5 °C (Sohrabi-Mollayousefi and Sahba 2010).

Environment: a shallow-water species of the Peri-reefal Area of Great Barrier Reef, in normal salinity and the depth between almost 10 m to 32 m (Baccaert 1987), typically associated with coral reefs and hardgrounds (Sen Gupta 1999; Haunold et al. 1997; Parker 2009; Parker and Gischler 2015). Also, it has been mentioned as a limited value environmental parameter, which belongs to the highly complex modern environments, with an enormous variety of ecological niches and of environmental parameters that are acting in this wide carbonated lagoon with reefs, deep depressions, and various continental inputs; depth from 1 to 30 m (Debenay 2012). Modern carbonate ramp (Parker and Gischler 2015).

Elphidium asiaticum (Polski 1959)

Fig. 6H

1959 Elphidium discoidale (d’Orbigny) var. asiaticum; Polski, p. 585, pl. 78, fig. 2a-b.

2000 Elphidium asiaticum (Polski,); Kim and Kucera, p. 1073, fig. 4(a-b).

2015 Cibrononion asiaticum (Polski); Lei et al. p. 250, pl. 1, fig. 4a-e.

2019 Elphidium asiaticum (Polski,); Tabita and Nathan, p. 21, fig. 6(8).

Material: Only one specimen was collected and studied, from Gushi Marl, Bemani outcrop, Minab Province, southern Iran.

Diagnosis and Description: See Tabita and Nathan (2019).

Occurrence and Distribution: Recent, East Coast of India (Tabita and Nathan 2019). Middle Holocene (Wu et al. 2020). Latest Quaternary, Core DH1-4, the Yellow Sea, central Korea (Kim and Kucera 2000).

Remark: Despite abrasion, the identity of the species based on characteristic features is admissible.

Environment: Liu et al. (2018) suggested the species as shallow marine species strata, which are typical of inner- to middle-shelf environments (Wang et al. 1985; Zhou et al. 1996). Also, it has been mentioned from the tidal flats (Kim and Kucera 2000), and from stormy intertidal (Wu et al. 2020), shallow water, and medium-depth water (Li et al. 2015).

Elphidium advenum macelliforme (McCulloch 1981)

Fig. 6I-J

1981 Elphidium macelliforme McCulloch, p. 119, p1.40, fig. 1.

1993 Elphidium macelliforme (McCulloch); Albani and Yassini, p.28, figs.65,66.

1997 Elphidium advenum macelliforme (McCulloch); Hayward et al. p. 130, pl. 5, figs 6-12

Material: only two specimens were collected and studied, from Gushi Marl, Bemani outcrop, Minab Province, and from Direstan outcrop, Qeshm Island, Persian Gulf, southern Iran.

Diagnosis and Description: See Hayward et al. (1997).

Occurrence and Distribution: Early Oligocene to Pliocene (Hayward et al. 1997).

Environment: See Hayward et al. (1997).

Elphidium advenum maorium (Hayward et al. 1997)

Fig. 6K-M

1952 Elphidium advenum (Cushman); Finlay in Marples, p.61 and p. 62.

1974 Elphidium advenum (Cushman); Collins, p. 4l.

1979 Elphidium advenum (Cushman); Hayward and Buzas, p.52, p1.12, fig. 157.

1990b Elphidium advenum var. depressulum (Cushma); Hayward, p.96.

1984 Elphidium advenum var. depressulum (Cushma); Hayward et al. p.162.

1945 Elphidium sp. cf. simplex (Cushman); Parr, p. 216, pl. 11, fig.8.

1966a Elphidium charlollense (Vella); Kenneti, p.61, p1.8, figs. 123, 124.

1966b Elphidium charlollense (Vella); Kenneti, p.206-207.

1997 Elphidium advenum maorium Hayward n.ssp., pl. 1, fig. 7; pl. 4, figs. 11-16; pl. 5, figs. 1-5.

Material: three specimens were collected and studied, from Stars Valley, Kendaloo, Qeshm Island, and also from Gushi Marl, Bemani outcrop, Minab Province, southern Iran.

Diagnosis and Description: See Hayward et al. (1997).

Occurrence and Distribution: Late Eocene to Recent (for more, see Hayward et al. 1997).

Environment: See Hayward et al. (1997).

Elphidium advenum limbatum (Chapman, 1907)

Fig. 6N-Q

1907 Polystomella macellum var. limbatum Chapman, p.142, pl. 10, figs.9a-b.

1922 Polystomella striatopunctata (Fichtel and Moll); Heron-Allen and Earland, p.229.

1933 Elphidium advenum (Cushman) var. depressulum Cushman, p.51, fig. 4 a,b.

1939 Elphidium macellum var. limbatum (Chapman); Cushman, p.52, pl. l4, fig.5.

1958 Elphidium limbatum (Chapman); Collins, p.421;

1961 Elphidium cf. advenum (Cushman); Hornibrook, p.129.

1965 Elphidium aff. advenum? (Cushman); Kustanowich, p. 53,61.

1990b Elphidium advenum (Cushman); Hayward, p.96.

1996 Elphidium advenum (Cushman); Hayward et al. pl. 2, figs.6-7.

Material: four specimens were collected and studied, two from the Mishan formation in Direstan and Stars Valley outcrops, and two from the Kendaloo outcrop, Qeshm Island, southern Iran.

Diagnosis and Description: See Hayward et al. (1997).

Occurrence and Distribution: Late Eocene to Recent (Hayward et al. 1997).

Environment: See Hayward et al. (1997).

Superfamily NONIONOIDEA Schultze 1854

Family NONIONIDAE Schultze 1854

Subfamily NONIONINAE Schultze 1854

Genus NONION de Montfort 1808

Nonion fabum (Fichtel and Moll 1798)

Fig. 6R-S

1798 Nautilus faba Fichtel and Moll, p. 103, pl. 19, figs. a-c.

2010 Nonion fabum (Fichtel and Moll); Margreth, p. 123, pl. 36, fig. 2a-c.

2012 Nonion fabum (Fichtel and Moll); Milker and Schmiedl, p. 112, fig. 25.22-24.

2014 Nonion fabum (Fichtel and Moll); Panchang and Nigam, pl. 33, fig. 11a-b.

2019 Nonion fabum (Fichtel and Moll); Tabita and Nathan, p. 21, fig. 6(20).

2023 Nonion fabum (Fichtel and Moll); Shareef et al. p. 198, pl. 5, fig. 8.

Material: Only two specimens were collected and studied, from Stars Valley and Direstan, Qeshm Island, Persian Gulf, southern Iran.

Diagnosis and Description: See Loeblich and Tappan (1988) and Tabita and Nathan (2019).

Occurrence and Distribution: Burdigalian to Langhian, Asmari Formation, Zagros Basin, West of Iran (Roozpeykar et al. 2019). Serravallian-Langhian, Fatha Formation, Iraq (Shareef et al. 2023). Recent, the Persian Gulf (Maghsoudlou et al. 2021).

Environment: In a relatively deep basin, consistent with the outer shelf-slope environment (Murray 1991; Schmiedl et al. 2003; Roozpeykar et al. 2019).

 Superfamily BOLIVINITOIDEA Cushman 1927

Family BOLIVINITIDAE Cushman 1927

Subfamily BOLIVINITINAE Cushman 1927

Genus BOLIVINA d’Orbigny 1839

Bolivina spathulata (Williamson 1858) dentellata type

Fig. 7A

1858 Textularia variabilis (Williamson) var. spathulata Williamson; p. 76, pl. 6, figs. 164-165.

1979 Brizalina spathulata (Williamson); Hageman, p. 89, pl. 2, figs. 5a, b.

1955 Bolivina dentellata (Williamson); Tavani, p. 144, pl. 1, figs. 1-3.

1982 Bolivina spathulata (Williamson); van der Zwaan, pl. 2, figs. 1-5; text-fig. 59

1992 Bolivina dentellata (Williamson); Tavani, p. 11, pl. 2, fig. 10.

2014 Brizalina spathulata (Williamson); Nabavi et al. p. 25, pl. 2, fig. d.

Material: Only one specimen was collected and studied, from Gushi Marl, Bemani outcrop, Minab Province, southern Iran.

Diagnosis and Description: See van der Zwaan (1982).

Occurrence and Distribution: Cosmopolitan, the late early Messinian (van de Poel 1992). Burdigalian to Langhian, Asmari Formation, Lurestan province, NW Zagros basin, Iran (Roozpeykar et al. 2019). Langhian to Serravallian, carbonate strata, Siwa Oasis, Egypt (Abdel-Fattah et al. 2013). Middle Holocene, core T2S3, Strait of Hormoz, Southern Iran (Hamzeh et al. 2021). Recent, Britain (Williamson 1858).

Environment: The abundance patterns of B. spathulata suggest that it was probably a mud-dweller with a wide environmental range; it was tolerant to high salinities and to oxygen deficiency. Possibly, it preferred a high-nutrient environment. It is a low-oxygen marine species, characteristic of the shelf edge and upper slope (Murray 1973; Pujos 1976; Jorissen 1988; van de Poel 1992), and has also, have been reported from warm waters of the tropical to subtropical zone in carbonate platform (Roozpeykar et al. 2019). In modern environments, from different depths between ~120 m and ~250 m or 20-40 m (Debenay 2012). Furthermore, it has been described as a deep infaunal and dysoxic species occurring in high organic flux environments (Bernhard and Gupta 1999; Das et al. 2017), with more salinity conditions (Eris et al. 2011; Suokhrie et al. 2021).

Suborder GLOBIGERININA Delage and Hérouard 1896

Superfamily GLOBIGERINOIDEA Carpenter 1862

Family GLOBIGERINIDAE Carpenter, Parker and Jones 1862

Subfamily GLOBIGERININAE Carpenter, Parker and Jones 1862

Genus GLOBIGERINA d’Orbigny 1826

Globigerina bulloides (d’Orbigny 1826)

Figs. 7B

1826 Globigerina bulloides d’Orbigny, p. 277, no.1.

1982 Globigerina bulloides (d’Orbigny); Brasier, p. 155, fig. 5.14 (a-c).

1983 Globigerina bulloides (d’Orbigny); Kennet and Srinivasan, p. 36, pl. 6, figs. 4-6.

1985 Globigerina bulloides (d’Orbigny); Papp and Schmid, p. 215, pl. 54, figs. 1-6.

2003 Globigerina bulloides (d’Orbigny); Hanagata, p. 322, pl. 9, fig. 3.

2005 Globigerina bulloides (d’Orbigny); Narayan et al. p. 151, pl. 6, figs. 6-8.

2010 Globigerina bulloides (d’Orbigny); Ovechkina, Bylinskaya and Uken, p. 239, fig. 5G-5I.

2019 Globigerina bulloides (d’Orbigny); Tabita and Nathan, p. 47, figs. 13(4-5).

Material: two samples were collected and studied from Kendaloo and Stars Valley, southern Iran.

Diagnosis and Description: See Tabita and Nathan (2019), and also Narayan et al. (2005).

Occurrence and Distribution: Early to Middle Miocene (Aqutanian to Langhian), Mishan Formation in the Khorgu and Khamir anticlines, Bandar Abbas, southern Iran (Fanati Rashidi et al. 2014). Late Burdigalian to Langhian of the Asmari Formation in NW of the Zagros basin (Roozpeykar et al. 2019). Middle Miocene to Holocene, NW Pacific and Canadian Arctic (Narayan et al. 2005). Late Pliocene, East Antarctica Quilty (2010).

Remark: Steinthorsdottir et al. (2021) mentioned that strengthening of the South Asian monsoon at ∼8 Ma was interpreted from an increase in the foraminifer Globigerina bulloides, which thrives on upwelling of deep nutrient-rich waters caused today by summer monsoon winds driving surface waters offshore (Kroon et al. 1991; Prell et al. 1992), but in Kendaloo and Stars Valley outcrops, only two G. bulloides and G. cf. bulloides were found.

Environment: According to evidence presented by Be and Hutson (1977), G. bulloides has an optimum temperature range of 13.4°±7.8°C, and also, it lives in depths of less than 50 m (Chaproniere 1992).

Genus TRILOBATUS Spezzaferri et al. 2015

Trilobatus (Globigerinoides) trilobus (Reuss 1850)

Fig. 7D

1850 Globigerina triloba Reuss, 374, pl. 47, fig. 11a–e.

1957 Globigerinoides triloba triloba (Reuss); Bolli, p. 112, pl. 25, fig. 2a–c.

1960 Globigerinoides triloba triloba (Reuss); Jenkins, p. 353, pl. 2, fig. 5a–c.

1966 Globigerinoides trilobus trilobus (Reuss); Jenkins, p. 9, pl. 2, fig. 8a–c.

1967 Globigerinoides quadrilobatus trilobus (Reuss); Closs, p. 340, pl. 1, fig. 22.

1975 Globigerinoides quadrilobatus trilobus (Reuss); Srinivasan, p. 139, pl. 2, fig. 7.

1983 Globigerinoides triloba (Reuss); Kennett and Srinivasan, p. 62, pl. 13, figs 1–3.

1994 Globigerinoides trilobus (Reuss); Loeblich and Tappan, p. 107, pl. 206, figs 1–6.

2012 Globigerinoides trilobus (Reuss); Rögl, p. 181, pl. 1, figs 1–7.

2018 Trilobatus trilobus (Reuss); Spezzaferri, Olsson, and Hemleben, p. 300–302, pl. 9.14, figs. 1–21.

2019 Trilobatus trilobus (Reuss); Poole and Wade, p. 1998, figs. 6A–O; p. 2014, fig. 16E; p. 2015, figs. 17A, E.

Material: Only one sample was collected and studied from Gushi marl, Minab Province, Makran Basin, southern Iran.

Diagnosis and Description: See Poole and Wade (2019).

Occurrence and Distribution: Late Burdigalian, Zagros Basin (Roozpeykar et al. 2019). Middle Miocene (early Langhian to early Badenian), between 15.2 and 14.8 Myr. of Vienna Basin, Austria (Harzhauser et al. 2020). The middle Miocene to lowermost Pliocene, Mishan Formation, Bandar Abbas (Hassani and Hosseinipour 2017). Langhian to Serravallian, in the lower part of the marly member, Mishan Formation, southern Iran (Gholamalian et al. 2015; 2020).

Remark: In Zagros Basin (Iran), Hassani and Hosseinipour (2017) suggested that the species is a member of biozone Z1 (= Globigerinoides trilobus zone of Postuma 1971), which shows timing Aqutanian before the boundary of early Burdigalian, which coincides with the beginning of global biozone N5 (Hassani and Hosseinipour 2017; also check figure 3 by Spezzaferri et al. 2002).

Environment: common in subtropical/ tropical assemblages worldwide throughout their long stratigraphical ranges from the latest Oligocene/early Miocene to Recent (Poole and Wade 2019), and also have been reported from outer neritic to upper bathyal depositional environments with high nutrient content and connection to the open sea (Harzhauser et al. 2020).

Genus PRAEORBULINA Olsson 1964

Praeorbulina transitoria (Blow 1956)

Fig. 7E

1956 Globigerinoides transitoria Blow, p.65 text-figs. 2.12-13.

1971 Praeorbulina transitoria (Blow); Postuma, p. 378-379

1985 Praeorbulina transitoria (Blow); Iaccarino, p. 298.

1985 Praeorbulina transitoria (Blow); Bolli and Saunders, p.200, figs. 23.6, 7, 9, 12.

1988 Praeorbulina transitoria (Blow); Rateb 1988, p.18, pl.3, figs. 8(a-b).

2013 Praeorbulina transitoria (Blow); Hewaidy et al. p. 41, pl. 9, figs. 3(a-b).

Material: Only one sample was collected and studied from Stars Valley, Qeshm Island.

Diagnosis and Description: See Hewaidy et al. (2013).

Occurrence and Distribution: Late Burdigalian to Langhian, Guri Member, Mishan Formation, Zagros Basin, South Iran (Daneshian et al. 2016). Late Burdigalian to Langhian, the lower Miocene Qantara Formation and middle Miocene Sidi Salem Formation, Nile Delta area, Egypt (Hewaidy et al. 2013). Correlated with the boundary of two global biozones N8 to N9, in Langian (for more, see chart 6.2 in BouDagher-Fadel 2015).

Environment: open shelf environments (Rahmani and Vaziri-Moghaddam 2010).

Order TEXTULARIIDA Delage and Hérouard 1896

Suborder TEXTULARIINA Delage and Hérouard1896

Superfamily TEXTULARIOIDEA Ehrenberg 1838

Family TEXTULARIIDAE Ehrenberg 1838

Subfamily TEXTULARIINAE Ehrenberg 1838

Genus TEXTULARIA Defrance 1824

Textularia agglutinans (d’Orbigny 1839b)

Figs. 7F-H

1839b Textularia agglutinans d’Orbigny; p. 144, pl. 1, figs. 17-18, 32-34.

1935 Textularia agglutinans (d’Orbigny); Keijzer, p. 128, 132, fig. 25 a-g.

2003 Textularia agglutinans (d’Orbigny); Javaux and Scott, p. 22, fig. 5(8-9).

2007 Textularia agglutinans (d’Orbigny); Talib and Farroqui, p. 18, pl. 1, fig. 1.

2012 Textularia agglutinans (d’Orbigny); Debenay, p. 95.

2012 Textularia agglutinans (d’Orbigny); Milker and Schmiedl, p. 38, fig. 10.15-16.

2015 Textularia agglutinans (d’Orbigny); Nouradini et al., p. 69, figs. 9-11.

2015 Textularia agglutinans (d’Orbigny); Hanagata and Nobuhara, p. 18, figs. 7(3-4).

2017 Textularia agglutinans (d’Orbigny); Anbuselvan and Nathan, fig. 8(3, 3a).

2019 Textularia agglutinans (d’Orbigny); Tabita and Nathan, p. 52, fig. 15(9).

Material: Three specimens were collected and studied, from Mishan Formation, Zagros Basin, Iran, each of the are from Direstan, Kendaloo, and Stars Valley outcrops, Qeshm Island.

Diagnosis and Description: See Tabita and Nathan (2019), Mohan et al. (2018), and also Boltovskoy et al. (1980).

Occurrence and Distribution: Cosmopolitan, has been reported from numerous locations worldwide (Markado et al. 2015). In Iran: Lower Miocene, Qom Formation, Central Iran (Nouradini et al. 2015). Recent, the Bay of Bengal, east coast of India (Anbuselvan and Nathan 2017).

Remark: Nouradini et al. (2015, 2017) mentioned the species in oxic, down to 50 m, carbonate sand littoral inner–middle shelf paleoenvironment of the NE margin of the Tethyan Seaway in the Qom fore arc in Central Iran.

Environment: The widespread species from the Indian to the West Pacific Oceans, including the modern Bay of Jakarta, Java (Hofker 1968). Also, has been mentioned from the modern shallow-water environment (depth: 85 m), and almost high temperature waters (temperature 22°C) of North coast Ambon Island, Indonesia (Hofker 1978), and have been reported from sandy to slightly muddy sand environment, restricted to 30-80 m of water depth, in shallower inner shelf (Anbuselvan and Nathan 2017). Baccaert (1987) suggested it as a shallow-water species of the Peri-reefal Area, in the Great Barrier Reef, in normal salinity, and the depth between 10 m to 32 m.

Class TUBOTHALAMEA Pawlowski, Holzman and Tyszka 2013

Order MILIOLIDA Delage and Hérouard 1896

Suborder MILIOLINA Delage and Hérouard 1896

Superfamily MILIOLOIDEA Ehrenberg 1839

Family HAUERINIDAE Schwager 1876

Subfamily MILIOLINELLINAE Vella 1957

Genus TRILOCULINA d’Orbigny 1826

Triloculina trigonula (Lamarck 1804)

Figs. 7K-M

1804 Miliolites trigonula Lamarck, p. 351, pl. 17(15), fig. 4.

1826 Triloculina trigonula (Lamarck); d’Orbigny, p. 299, pl. 16,

1858 Miliolina trigonula (Lamarck); Williamson, p. 83, pl. 7, figs. 180-182.

1960 Miliolina trigonula (Lamarck); Barker, pl. 3, figs. 15-16.

1884 Miliolina trigonula (Lamarck); Brady, p. 164, pl. 3, figs. 15, 16 (not fig. 14).

1989 Miliolina trigonula (Lamarck); Hermelin, p. 39, pl. 3, fig. 8.

1994 Miliolina trigonula (Lamarck); Jones, p. 20, pl. 3, figs. 15-16.

2006 Triloculina trigonula (Lamarck); Oflaz, p. 177, pl. 4, fig. 4.

2007 Triloculina trigonula (Lamarck); Talib and Farroqui, p. 19, pl. 1, fig. 16a-b.

2010 Triloculina trigonula (Lamarck); Margreth, p. 103, pl. 11, fig. 2a-c.

2012 Triloculina trigonula (Lamarck); Debenay, p. 138.

2013 Triloculina trigonula (Lamarck); Holbourn et al. p. 566, figs, 1-2.

2014 Triloculina trigonula (Lamarck); Panchang and Nigam, pl. 13, fig. 7a-b.

2019 Triloculina trigonula (Lamarck); Tabita and Nathan, p. 70, figs. 21(1).

Material: three samples were collected and studied, two from Kendaloo and one from Direstan outcrop, Qeshm Island, Zagros Basin, southern Iran.

Diagnosis and Description: See Tabita and Nathan (2019).

Occurrence and Distribution: Worldwide, Early Eocene to Recent (Jones 1994; Holbourn et al. 2013). Late Oligocene (Chattian), Asmari Formation, Qeshm Island, Persian Gulf, southern Iran (Sajadi and Fanati Rashidi 2019). Chattian to Aqutanian, Asmari Formation, SW Zagros Basin, Iran (Roozpeykar and Moghaddam 2015).  And also, the Al-Kharrar Lagoon (modern environment), eastern Red Sea coast, Saudi Arabia (Al-Dubai et al. 2017).

Environment: Lower neritic to bathyal (Holbourn et al. 2013). Shoreline to 100 fathoms (Jones 1994). Reported from the lower bathyal zone in the eastern Pacific Ocean (see Hermelin 1989). Also, it has been mentioned that it shows positive relationships with pH, deep muddy substrates enriched in organic matter, and is negatively correlated with high temperature and salinity (Al-Dubai et al. 2017).

Triloculina terquemiana (Brady 1884)

Figs. 7N-O

1884 Miliolina terquemiana Brady, p. 166, pl. 114, fig. 1a-b.

1999 Triloculina terquemiana (Brady); Nigam and Khare, p. 292, pl. 3, fig. 7.

2007 Triloculina terquemiana (Brady); Talib and Farroqui, p. 18, pl. 1, fig. 15a-b.

2012 Triloculina terquemiana (Brady); Debenay, p.138.

2012 Triloculina terquemiana (Brady); Mossadegh et al., p. 354, figs. 11(r-t).

2019 Triloculina terquemiana (Brady); Tabita and Nathan, p. 67, figs. 20(16-17).

Material: Two samples were collected and studied, from Direstan and Stars Valley outcrops, Qeshm Island, Zagros Basin, southern Iran.

Diagnosis and Description: See Debenay (2012).

Occurrence and Distribution: Late Oligocene (Chattian), Qom Formation, Central Iran (Nouradini et al. 2017). Late Pleistocene, from raised Coral reef sequences, Kish Island, Persian Gulf, Iran (Mossadegh et al. 2012). Recent, Coastal areas of the Indian Ocean (Farooqui1990), and also, the Southwestern Pacific (Debenay 2012).

Remark: Nouradini et al. (2017) reported the species from the NE margin of the Tethyan Seaway in the Qom fore arc in Central Iran.

Environment: have been mentioned from oxic, 0–50 m, muddy epifaunal of inner shelf of paleoenvironment (Nouradini et al. 2017), also, from shore sand (Brady 1884), sandy beaches and coastal modern environments (see Farooqui1990). In the Ligurian Sea (Italian part of the Mediterranean Sea) the Miliolidae occur in large numbers from 20 to 90 m (Giunta 1955; Luczkowska 1974).

Triloculina tricarinata (d’Orbigny 1826)

Fig. 7P

1826 Triloculina tricarinata d’Orbigny, p. 299, pl. 1, fig. 8.

2010 Triloculina trigonula (Lamarck); Margreth, p. 103, pl. 11, fig. 2a-c.

2012 Triloculina tricarinata (d’Orbigny); Debenay, p. 138.

2012 Triloculina tricarinata (d’Orbigny); Milker and Schmiedl, p. 68, fig. 17.23-24.

Material: only one sample was collected and studied, from the Kendaloo outcrop, Qeshm Island, Zagros Basin, southern Iran.

Diagnosis and Description: See Debenay (2012).

Occurrence and Distribution: Chattian to Aquitanian, Asmari Formation, SW Zagros Basin, Iran (Roozpeykar and Moghaddam 2015). Early to Middle Miocene (Aquitanian to Langhian), Mishan Formation, southern Iran (Fanati Rashidi et al. 2014). Langhian to Serravallian, in the lower part of the marly member, Mishan Formation, southern Iran (Gholamalian et al. 2015; 2020). Recent, the Persian Gulf (Sohrabi-Mollayousefi and Sahba 2010).

Environment: T. tricarinata suggests a particular numerous occurrence in down to 20 m (Giunta 1955; Luczkowska 1974). Living in the Persian Gulf under conditions of seasonal temperature and salinity, the average temperature is 23.5 °C, average dissolved oxygen exceeded 6.1 mg/lit (Sohrabi-Mollayousefi and Sahba 2010). Also, it has been reported from the low-energy and the shelf lagoon deposition environment of Mishan Formation (Fanati Rashidi et al. 2014).

Subfamily HAUERININAE Schwager 1876

Genus QUINQUELOCULINA d’Orbigny 1826

Quinqueloculina bogdanowiczi (Serova 1955)

Figs. 7Q-S

1955 Miliolina bogdanowiczi Serova, p. 309, pl. 4, figs. 1-3.

1961 Quinqueloculina bogdanowiczi (Serova); Didkovsky, p. 22, pl. 1, fig. 4.

1961 Quinqueloculina brevia (Didkovsky); Ibidem, p. 45, pl. 9, fig. 2.

1974 Quinqueloculina bogdanowiczi (Serova); Luczkowska, p. 170, pl. V, figs. 3(a-c), 4(a-c); p. 44, text-fig. 10.

2014 Quinqueloculina bogdanowiczi (Serova); Filipescu et al. p. 71, fig. 5(7).

Material: three samples were collected and studied from Direstan and Stars Valley outcrops, Qeshm Island, also from Gushi marl, Minab Province, Iran.

Diagnosis and Description: See Luczkowska (1974).

Occurrence and Distribution: Serravallian (the early Sarmatian), Pannonian Basin, Romania (Filipescu et al. 2014). Upper Tortonian and also the sea of the lower Sarmatian, Poland, and Lower and Upper Tortonian, the southwestern border of the Russian Platform, and Lower Tortonian, the West Ukraine (Luczkowska 1974).

Remark: The reporting of the species from Ukraine, the southwestern border of the Russian Platform, the Pannonian Basin, and the Sarmatian Sea in central Europe, suggests that it may be an Eastern-Central Paratethyan species.

Environment: Paleogeographically, reported from a sandy environment (Luczkowska 1974), and marginal to shallow marine, with fairly high energy and fluctuating salinity (Filipescu et al. 2014).

Evidence of the Connection

The micropaleontological assemblages recovered from Qeshm Island (Zagros Basin) and Bemani (Makran Basin) provide some evidence for evaluating the status of the Iranian Gateway during the Late Miocene–Early Pliocene, including the co-occurrence of taxa such as Quinqueloculina bogdanowiczi, previously restricted to the Paratethys (Harzhauser and Piller 2007; Reuter et al. 2009), together with tropical planktonic markers including Praeorbulina transitoria and Globigerinoides trilobus (Poole and Wade 2019), suggests intermittent marine connections between the Indo-Pacific and the proto-Mediterranean realms. Of planktonic foraminifera paleotemperature markers (Spezzaferri et al. 2002; Holcová and Zágoršek 2008), Globigerina bulloides is a marker of cold temperature, and Globigerinoides (Trilobatus) trilobus is a marker of warm temperature (Poole and Wade 2019). In Qeshm Island, the presence of Globigerina bulloides, a cold-water indicator (Spezzaferri et al. 2002) and other taxa like Elphidium crispum and Challengerella bradyi, alongside warm-water taxa including Triloculina group and Eponides group, further supports the interpretation of episodic faunal exchange through narrow seaways (fig. 3), what is conclusionable in Minab region based on the co-occurrence of Asterorotalia group and Elphidium advenum group with G. trilobus and Bolivina spathulata (see table 2). These assemblages correlate with global planktonic biozones N8–N9 and N19–N20 (Boboye and Adeleye 2009), indicating that the Iranian Gateway remained at least partially open until the Messinian–Zanclean interval. Thus, the new data strengthen existing models of episodic marine incursions (“tongues”) advancing westward into Mesopotamia (Harzhauser et al. 2007; Piller et al. 2024). 

Table 2: The environmental indicator foraminifera in this study are briefly listed below to provide an overview of intermittent marine connections between the Indo-Pacific and the proto-Mediterranean realms.

* Collected from in Qeshm Island and Minab region, both.

Table 3: Occurrences of studied foraminifera at the nearest parts of the world to all four outcrops in the present study, and also correlation of three global biozones of N5, N19, and N20 with studied foraminifera (for more see “Distribution and Occurrence” sections of every sample in this manuscript). Note:  K= Kendaloo, B= Bemani, St= Stars Valley, D=Direstan, MM= marly member, GM=Guri Member, Q= Qeshm Island (Kendaloo, Direstan, Stars Valley), As = Asmari Formation, K= Kasaba Formation, W=worldwide, Fm.=Formation, E= Egypt, M= Mediterranean, IO= Indian Ocean, SWP= South West of Pacific, NWP= North West of Pacific, EC=Eastern-Central, SH= Strait of Hormoz, PG=Persian Gulf, CI=Central Iran, GA= Gulf of Aden, RS= Red Sea, DK= Drake Passage, Antarctica, (*) = all four outcrops.

Fig 3. Geographic map of the Paleo-Basins, where the species in the present study have been reported. These Paleo-Basins may show probable connections between numerous parts of Paratethys (from Europe to Qom Basin), Mediterranean/Proto-Mediterranean Basin (Egypt, Spain, and Italy), to the nearest Basins around the studied area (Iraq and other parts of the Zagros basin), and the studied area itself, all during the Miocene (after Sun et al. 2021).

Discussion

From nearest area of the southern Iran, Shareef et al. (2023) reported some of the same species including Ammonia beccarii, Challengerella bradyi, Elphidium advenum, E. craticulatum, E. crispum, Nonion fabum, from Fatha Formation of Iraq (NW of Persian Gulf and subsequently all studied areas), and suggested that the taxa probably show presence of a thin seaway connection, which had been remained open (the connection between Mediterranean Sea and Indo-Pacific Oceans), during the time of Middle-Late Miocene, which had been in direct contact with the open sea.

In Central Iran (see fig. 3), Nouradini et al. (2014) reported the shallow and warm water environment strata of Qom Formation, at the Burdigalian, including some foraminiferal species such as Textularia agglutinans, Elphidium fichtellianum, Triloculina tricarinata, Triloculina trigonula, Globigerinoides trilobus, which are palaeobiogeographically located in the marginal seaways (Qom Basin), the connection between the West, Central Paratethys and Indo-Pacific Ocean during the Lower Miocene. Also, the planktonic foraminiferal fauna, including Praeorbulina transitoria, has been mentioned as a typical Tethyan indicator of tropical-subtropical climatic conditions (Hani and Abawi 2005). On the other hand, Barbieri and Vaiani (2018) suggested that the species Elphidium advenum lives in coastal areas and is recorded in low percentages in the Mediterranean lagoons (Albani and Serandrei Barbero 1990; Hohenegger et al. 1993). Also, Safari et al. (2020), reconstructed the depositional sequences of the Qom Basin and suggested that the Basin were influenced primarily by local tectonics, while global sea level changes had a greater impact on the southern Tethyan seaway and Paratethys basins, which may show the depositional basins of the Tethyan seaway (southern Tethyan seaway, Paratethys Basin and Qom Basin) were probably related during the Burdigalian to Langhian and early Serravallian.

Paleoecologically, the majority of Elphidium species indicate lagoonal environments with brackish to marine water (Rao and Rao 1974; Shareef et al. 2023), which occurs in the Direstan outcrop. On the other hand, of the 7 studied Elphidium species, 6 samples are from Direstan.

In case of paleoenvironmental suggestions corresponding to the morphology of the species, Edwards (1982) reported Eponides repandus from very coarse sandy gravels of the eastern Shiant bank, North Minch Channel, Scotland, and mentioned that the species displays a secondary thickening of shell material on the exterior surface which serves to strengthen it against damage through current buffeting occurring in the high energy environment. Not only Eponides, but all vagrant benthonic forms reported by Edwards (1982), such as miliolids and Elphidium species, are suggested as the species occur in association with high-energy environments, where such free-living forms are generally thick-shelled to withstand vigorous current activity. By paying attention to this suggestion, it is necessary to mention that the Eponides species samples from Stars Valley have more fragile shells than the samples of Kendaloo and Direstan.

Moreover, Edwards (1982) suggested that open umbilical apertures like Ammonia and Elphidium are adaptive in littoral habitats of varying salinity but may be less important in some sublittoral and marsh habitats. However, Ammonia beccarii is the most euryhaline among living trochospiral Rotaliina and occurs in various forms, some called species, that dwell in different habitats.

From the nearest area to the present studied sections, Nouradini et al. (2015) mentioned some genera like Elphidium as an oxic indicator and Bolivina as dyoxic indicators of the Qom Basin paleoenvironment, and suggested that the frequency of bryozoans, brachiopods, echinoids, and bivalves, associated with the presence of widespread euryhaline taxa (e.g., Elphidium), indicates a normal marine salinity dominated. Also, low diversity, high abundance, and small test size of foraminifers suggest a stable environment.

Overall, the foraminiferal fauna, including Triloculina, Elphidium, Challengerella, Bolivina, Ammonia, Textularia, Nonion, and Asterorotalia show the shelf edge to upper slope association characteristics (Saidova 2010; Murray 2006; Hamzeh et al. 2021).

Miliolids-dominated benthic foraminiferal assemblages are a key indicator of restricted lagoonal environments, characterized by decreased circulation and often hyposaline, hypersaline, or low-oxygen conditions. While miliolids can occur in normal-salinity settings like sand shoals, their dominance is widely used as diagnostic evidence for lagoon restriction (Brasier 1975a; 1975b; Brandano et al. 2008; Sajadi and Fanati Rashidi 2019).

Nouradini et al. (2017) suggested that miliolids such as Triloculina and Quinqueloculina are euryhaline (Murray 1991) and able to adapt to various salinities.

In sum, the ecological distribution of studied foraminifera from Kendaloo, Direstan, Stars Valley, and Bemani outcrops is illustrated based on BouDagher -Fadel (2018) in fig. 9 (a to d).

Conclusion

In all outcrops from Qeshm Island (fig. 8a-c), the planktic foraminifera species show a low diversity, and it may reflect a restricted environment or almost restricted, where connection with the open marine must be limited, topographically, but not because of high energy of water. Miliolids samples of all four outcrops cannot help us to prove the presence or absence of any kind of barriers and/or connections to the open marine environment, because Triloculina and Quinqueloculina genera are euryhaline and able to adapt to various salinities. The most dominant foraminiferal species in Bemani (Gushi marl) is Asterorotalia dentata, while in Qeshm Island outcrops, a variety of genera and species of benthic foraminifera occurred, which refer to shallow marine oxic environments. In all four outcrops, planktonic foraminifera are rare, and from all studied samples the most common non-planktonic species is Textularia agglutinans, which shows the normal salinity conditions, and the depth between almost 10 m to 32 m. The foraminiferal fauna in Qeshm Island outcrops represent the stable, oxic environments in normal marine salinity and colder waters than modern environments, with fairly high energy.

The foraminiferal fauna of Gushi marl (Bemani outcrop) indicates oxic, shallow, connected to both open sea and silty mud environments, which may refer to the shelf edge to upper slope, while the temperature is warmer than modern environments of the tropical to subtropical zones in present-day environments (fig. 8d).

In Bemani, the presence of Trilobatus (Globigerinoides) trilobus, Bolivina spathulate (dentellata), Asterorotalia pulchella, Asterorotalia dentata (see table 3), shows the probable age range, referring to the latest Miocene (Messinian) to early-middle Pliocene (Piacenzian-Zanclean boundary), for Gushi marl (in Makran Basin), which is possible to have a correlation with the time between global biozones N19-N20.

The foraminiferal species of Qeshm Island outcrops represent the age of the Langhian-Serravallian boundary for white sandy limestone of Mishan Formation at the Stars Valley outcrop, while uppermost strata of Mishan Formation (sandy fossiliferous limestones Direstan) show the age of late Serravallian to Mio-Pliocene boundary, in Qeshm Island, which is a possibility to indicate a correlation with the time between global biozones of N8-N9 and N19-N20.

The presence of Quinqueloculina bogdanowiczi in Qeshm Island outcrops, which have only been reported from Central Eastern Europe (Paratethys) during Serravallian to Tortonian, probably refer to the assumption of presence of a narrow temporarily connection between the Iranian Gateway and the Iraqi Basin (Fatha Formation), to the eastern parts of Indo-Pacific Ocean and western coastal proto-Mediterranean, and also show a connection between the marginal seaways of eastern Paratethys (Qom Basin) and the Iraqi Basin (Fatha Formation) to the Central Paratethys, and proto-Mediterranean Sea (see fig 3). Not only the presence of foraminifera, but also the presence of the rare reported taxa from the Basins mentioned above can be more convincing evidence of the assumption. The reported taxa including rare species of Ostreids Ostrea callifera (Lamarck) from Miocene of Europe (Berning 2020), Qom Formation and (Fuchs 1879) Gushi Marl of Makran Basin (Haskouei 2025). Also, the study is focused on the foraminiferal species, including Asterorotalia dentata, A. pulchella, Triloculina tricarinata, T. terquemiana, T. trigonula, Trilobatus (Globigerinoides) trilobus, Globigerina bulloides, Quinqueloculina bogdanowiczi, Textularia agglutinans, Elphidium crispum, E. craticulatum, E. asiaticum, E. advenum limbatum, E. advenum macelliforme, E. advenum maorium, Poroeponides lateralis, Eponides repandus, Eponides isabellanus, Rotalinoides compressiuscula, Challengerella bradyi, Ammonia beccarii, Bolivina spathulata, which almost all of them are reported for the first time from the studied outcrops.

Acknowledgments

Hereby, we acknowledge the Vice Chancellor for Research and Technology of the University of Isfahan. As well, the authors appreciate Department of Geology to the scientific and logistical supports.