Algae and smaller foraminifers of the eponymous Khuff Formation (Saudi Arabia) principally comprise Permocalculus, biseriamminids, hemigordiids and lagenids. Due to the end-Capitanian crisis (Late/Middle Permian boundary) and the regional palaeoecology, fusulinids are rare and only represented by Nankinella sp. and Eostaffella? sp. Palaeofusulinids are completely lacking. New age data shows that these foraminifers correspond to the complete Lopingian (Late Permian) as indicated by several species of Paradagmarita. The position of the Triassic/Permian Boundary is approximately characterised, but requires more accurate studies. Forty-three taxa were identified, mostly in open nomenclature. One new species is described: Glomospirella? linae n. sp. The foraminiferal assemblage is correlated with several associations in Iran, Turkey, Transcaucasia and south China.
Permian to Triassic sedimentary rocks of the Khuff Formation are exposed in central Saudi Arabia along a North-South belt, some 1,200 km long (Figure 1). The Khuff Formation rests everywhere upon a major unconformity (the Pre-Khuff Unconformity - PKU) over Lower Palaeozoic or Proterozoic shield rocks (Powers, 1968; Powers et al., 1966). The Khuff Formation is conformably overlain by the clayey and evaporitic Lower Triassic (‘Scythian’) Sudair Shale Formation. Le Nindre et al. (1990a, b) published the first synthesis of field work and extensive systematic geological mapping, conducted in the 1980s by the Saudi Deputy Ministry of Mineral Resources (DMMR) and the French Geological Survey (BRGM), including lithostratigraphy, biostratigraphy and palaeoenvironmental reconstructions. More recent compilations from central Saudi Arabia, were interpreted in terms of sequence stratigraphy by Alsharhan and Nairn (1995), Al-Aswad (1997) and Sharland et al. (2001). Sharland et al. (2001) considered the Khuff Formation as part of a tectonic megasequence dominated by thick platform and ramp carbonate, with subordinate evaporitic and clastic deposits.
A complete revision, including new field acquisitions and compilation of local studies for oil exploration (Senalp and Al-Duaiji, 1995, 2001), was prepared by Vaslet et al. (2005). It includes a reassessment of the biostratigraphy based on foraminifers and algae (Vachard et al., 2002, 2003), ostracods (Crasquin-Soleau et al., 2004, 2006) and palaeofloras (Broutin et al., 2002; Berthelin et al., 2006). Vaslet et al. (2005) divided the central Saudi Arabian outcrops of the Khuff Formation (some 200 m thick) into five members, from oldest to youngest: Ash Shiqqah, Huqayl, Duhaysan, Midhnab and Khartam members (Figure 2). The aim of this paper is to build a precise biostratigraphy and systematically describe the foraminifers of the Khuff Formation in Saudi Arabia. Types and figured specimens are housed in the BRGM (Orléans, France) collection.
LITHOSTRATIGRAPHY, BIOSTRATIGRAPHY AND SEQUENCE STRATIGRAPHY OF THE KHUFF FORMATION
The Ash Shiqqah Member (nearly equivalent to the obsolete Unayzah member of the Khuff Formation of Delfour et al., 1982) consists of terrigenous sediments with secondary clayey dolomite and local evaporite in the upper part of the unit. The palaeoenvironments range from transitional to continental and supratidal. The Unayzah Flora (Hill and El-Khayal, 1983; El-Khayal and Wagner, 1985; Broutin et al., 1995), formerly described in the lower part of the Khuff Formation, is now attributed to the underlying Unayzah Formation (Vaslet et al., 2005). Rare benthic smaller foraminifers occur locally in the upper part of the Ash Shiqqah Member, indicating a possible Middle Permian age (Midian) for this lowest unit of the Khuff Formation (Vachard et al., 2002; Vaslet et al., 2005).
The Huqayl Member is subdivided into two sequential units containing calcarenite, gypsiferous claystone, dolomite, and solution breccias related to subsurface evaporites. This marine transgressive unit is tentatively assigned a Late Permian Wuchiapingian (Dzhulfian) age according to its benthic foraminiferal content (Vachard et al., 2002; Vaslet et al., 2005). The Duhaysan Member is the first true calcareous subtidal to littoral unit of the Khuff Formation (Le Nindre et al., 1990b), and interpreted as the transgressive unit of the above Midhnab Member (Vaslet et al., 2005). The Duhaysan Member has yielded benthic foraminifers, nautiloid embryos, and abundant bactritids. A Late Permian Wuchiapingian to Changhsingian age is tentatively assigned to the Duhaysan Member (Vaslet et al., 2005).
The Midhnab Member displays a succession ranging from marine fossiliferous limestones at the base, toward gypsiferous and dolomitic rocks deposited in restricted palaeoenvironments, in the upper part. The lower part of the Midhnab Member is dated by benthic foraminifers as Late Permian Changhsingian (Dorashamian) by Vachard et al. (2002). Locally, in northern central Saudi Arabia, the topmost part of the Midhnab Member presents continental facies including lacustrine limestone, sandstone channels and claystone in meandering river systems and swamps. These facies contain drifted wood and plant remains (Hill and El-Khayal, 1983; Vaslet et al., 1985b; Le Nindre et al., 1990b; Vaslet et al., 2005). Recent descriptions of the Midhnab Flora indicate a Late Permian mixed flora including Cathaysian, Euramerian and Gondwanan plant remains (Broutin et al., 1995, 2002; Berthelin, 2002; Berthelin et al., 2006).
The Khartam Member, the uppermost mainly carbonate unit of the Khuff Formation is subdivided into two marine units characterised by littoral to tidal and intertidal palaeoenvironments. The Lower Khartam Member consists in claystone, dolomite and sands, deposited in supratidal to tidal palaeoenvironments. The Upper Khartam Member is an oolitic, peloidal and bioclastic limestone locally dolomitised, deposited in littoral to tidal and intertidal palaeoenvironments. The Lower Khartam Member yielded rare benthic foraminifers possibly dated as late Late Permian (late Changhsingian) by Vachard et al. (2002), while the Upper Khartam Member, consisting principally of reworked dasycladacean algae thalli Aciculella? sp., is characterised by the appearance of Spirorbis phlyctaena Brönniman and Zaninetti, a serpulid that is particularly abundant in the Early Triassic rocks in Neo-Tethyan areas.
According to Vaslet et al. (2005), the Khuff Formation consists of four main Depositional Sequences (DS PKh, DS PKm, DS PKk and DS TrS, see Figure 2). The last Depostional Sequence starts with the Khuff Formation and continues in the overlying Sudair Shale Formation. The DS PKh (named after Permian-Khuff-Huqayl) includes the Ash Shiqqah and the Huqayl members. Its basal Sequence Boundary (SB) corresponds to the Pre-Khuff Unconformity (PKU) and it contains the first Late Permian flooding event over central Saudi Arabian outcrop areas (MFI PKh). This flooding interval is located in the basal part of the Huqayl Member and is followed by the regressive evaporitic palaeoenvironments of the Huqayl Member (Le Nindre et al., 1990b; Vaslet et al., 2005).
The DS PKm (named after Permian-Khuff-Midhnab) starts with the deposition of subtidal to littoral Duhaysan Member above an erosive surface at the top of DS PKh, and ends with the regressive supratidal to continental deposits of the upper part of the Midhnab Member. A maximum flooding interval (MFI PKm) is clearly located in the outcrops at the base of the Midhnab Member, with abundant marine fauna including cephalopods and brachiopods (Angiolini et al., 2006; Chirat et al., 2006; Vaslet et al., 2005).
The DS PKk corresponds to the Lower Khartam Member (Permian-Khuff-Khartam), and represents the terminal Late Permian Depositional Sequence in the outcrops of central Saudi Arabia. The basal SB is marked by a return to marine subtidal conditions after the continental break at the end of DS PKm. It contains a maximum flooding interval (MFI PKk) and manifested by marine fauna, including abundant Permian ostracods (Crasquin-Soleau et al., 2005, 2006), bactritids and locally cephalopods (Chirat et al., 2006).
The DS TrS (named after the Sudair Shale Formation) starts with the littoral, tidal to intertidal deposits of the Early Triassic Upper Khartam Member of the Khuff Formation, and ends with the closed-basin, clayey to evaporitic rocks of the Lower Triassic Sudair Shale Formation (Le Nindre et al., 1990b; Vaslet et al., 2005).
Foraminifers were first recorded in the Khuff Formation of Saudi Arabia by Powers et al. (1966), and also reported by Powers (1968), who recognised a Late Permian fauna, while formally defining the unit. In the 1980s, the Saudi Arabian and French geological surveys more precisely positioned the biostratigraphy deduced from Late Permian foraminifer assemblages into the framework of the geological mapping programme of central Saudi Arabia. Foraminifers collected from these field campaigns, were identified by D. Vachard (1982-1983, quoted inLe Nindre et al., 1990b), M. Lys (in Le Nindre et al., 1990b), and A. Roux and C. Gigot (in Delfour et al., 1982). These results were reported and summarised in the explanatory notes of the Saudi Arabian geoscience maps (Delfour et al., 1982; Vaslet et al., 1983, 1985a, 1985b, 1987; Kellogg et al., 1985; Manivit et al., 1985a, 1985b, 1986; Robelin et al., 1994), and the biostratigraphic data and conclusions compiled by Le Nindre et al. (1990a, b). The present micropalaeontological study only investigates the collections made from samples taken in the field by BRGM geologists. More recent references to foraminifers of the Khuff Formation in Saudi Arabia were made by Okla (1992) while describing algae, and by Al-Aswad (1997) who placed the biostratigraphical data into stratigraphic cycles. The poorly diversified but numerous algae were studied by Rezak (1959), Roux (1991) and Okla (1992).
The Khuff Formation of Saudi Arabia was principally attributed to the Colaniella parva biozone (despite the absence of the eponymous zonal fossil), i.e. late Dzhulfian-Dorashamian (Manivit et al., 1986; Broutin et al., 1995). Guides among palaeofusulinids (e.g. Palaeofusulina, Codonofusiella) were never mentioned. The presence of “Colaniella parva ? (Colani)” was primarily indicated by Powers et al. (1966) and Powers (1968), but no advanced species of Colaniella was found during our study. The presence of Dzhulfian-aged rocks (in a strict sense), and the location of the Triassic/Permian Boundary within the Iranian equivalent of the Khuff Formation were established by Rosen (1979) in the Iranian part of the Gulf and Fars, and more recently by Virgone et al. (2002) in the Zagros and southern Pars. Another foraminiferal guide-fossil of the latest Permian (= Dorashamian = Changhsingian) (Figure 3), Paradagmarita, was first mentioned by Manivit et al. (1985b, 1986), but apparently neglected.
Alsharhan and Nairn (1995) assigned the Khuff Formation to the “Middle to Late Permian”, but they did not mention any foraminifers. Al-Aswad (1997) listed microfossils including the algae Mizzia and Permocalculus, several species of Globivalvulina and some genera of lagenids. This author assigned the Huqayl and Duhaysan members to a Kazanian age (Figure 4), and the Midhnab and Khartam members to the Tatarian (see Figure 3, for correlation of these names of stages).
Sharland et al. (2001) assigned to the Khuff Formation an age from Ufimian/Kazanian to Early Triassic, and emphasized the following Maximum Flooding Surfaces (MFS): P20 (252.5 Ma) within the Khuff D, P30 (250 Ma) within the Khuff C, P40 (249 Ma) within the Khuff B, and Tr 10 (248 Ma) in the lower part of the Khuff A. Vaslet et al. (2005) redefined maximum flooding intervals from central Saudi Arabian outcrops, reassigning the MFS Tr10 of Sharland et al. (2001) to a Late Permian (late Changhsingian = late Dorashamian) age and discussed new possible correlation with regional MFS as described by Virgone et al. (2002) in the Zagros and southern Pars in Iran.
ASSEMBLAGES OF MICROFOSSILS
Ash Shiqqah Member
Although relatively diversified, the microfacies are generally devoid of microfossils. These microfacies of the Ash Shiqqah Member are tidalitic micrites locally sandy and peloidal with extraclasts of stromatolites and calcretes; dolomicrites, eolian ferruginous sandstones, and calcareous or siliceous sandstones. The rare microfossils are Permocalculus sp., Globivalvulina sp. and hemigordiids.
The assemblage is characterised by the algae Permocalculus digitatusElliott, 1955 (Plates 1.1, 1.3 and 1.4), P. solidus (Pia, 1937) (Plate 1.5), P. plumosusElliott, 1955 (Plate 1.6 and 1.7?) and Mizzia velebitanaSchubert, 1909 (Plate 1.2); and the foraminifers Earlandia? spp. (Plates 2.1–2.4, 2.13), Nankinella spp.
(Plates 2.15 and 2.20), Globivalvulina cf. graecaReichel, 1946, Cornuspira sp. (Plates 2.9 and 2.10), Hemigordiellina regularis (Lipina, 1949) (Plates 2.6 and 2.8), Pseudomidiella cf. labensis Pronina-Nestell inPronina-Nestell and Nestell, 2001 (Plates 5.1 and 5.2), Hemigordius schlumbergeri (Howchin, 1895) (Plate 2.11), Neodiscus aff. qinglongensisWang, 1976, Agathammina sp., Nodosinelloides aff. concinna (Potievskaya, 1962) (Plate 6.13), N. shikhanica (Lipina, 1949), Polarisella elabugae (Cherdyntsev, 1914) (Plate 6.10), P.? hoae (Trifonova, 1967), Geinitzina sp., Pachyphloia spp. and Ichtyofrondina? sp. (Plate 6.30).
The organic content is composed of algae Permocalculus digitatus (Plate 1.4), P. tenellus (Pia, 1937), P. cf. plumosus (Plate 1.7); of calcispheres Radiosphaera cf. basilicaReitlinger, 1957 (Plates 2.5 and 2.7); and foraminifers Earlandia? sp., Globivalvulina graeca (Plate 3.1), Septaglobivalvulina? decrouezae (Köylüoglu and Altiner, 1989) (Plates 3.5 and 3.6), Glomospirella? sp. (Plate 4.2), Hemigordius baoqingensis Wang inZhao et al., 1981 (Plates 5.4–5.6, 5.11), Neodiscus aff. qinglongensisWang, 1976 (Plates 5.8–5.10), Pseudomidiella cf. labensis, Agathammina sp., Graecodiscus cf. kotlyarae Pronina-Nestell in Pronina-Nestell and Nestell, 2001 (Plate 2.19), Rectostipulina sp. (Plate 6.8), Nodosinelloides shikhanica (Plate 6.16), Polarisella elabugae, Geinitzina sp., Pachyphloia sp., “Lingulina” cf. semivelataCherdyntsev, 1914 (Plate 6.33), Robuloides lensReichel, 1946 (Plate 6.9) and Colaniella cf. minutaOkimura, 1988 (Plates 6.6 and 6.7).
This third assemblage is constituted by the algae Permocalculus digitatus (Plate 1.1 and 1.3) and P. plumosus (Plate 1.6) and the foraminifers Earlandia? sp., Eostaffella? sp. (Plate 2.14), Nankinella spp., Globivalvulina aff. graeca (Plate 3.13), Dagmarita? shahrezaensisMohtat-Aghai and Vachard, 2003 (Plate 3.9), Paradagmarita n. sp. (Plate 3.8), Neodiscus aff. qinglongensis (Plates 5.3–5.7), Cornuspira sp., Glomospirella? sp. (Plate 4.1), G.? linae n. sp. (Plates 4.3–4.15, 4.17), Palaeonubecularia n. sp. (Plate 2.18), Hemigordius schlumbergeri, Neohemigordius ex gr. zaninettiae (Altiner, 1978) (Plate 2.12), Multidiscus spp., Agathammina spp. (Plate 2.21), Nodosinelloides shikhanica, Polarisella elabugae (Plates 6.3–6.5, 6.20–6.22, 6.32), P. cf. elabugae (Plates 6.23–6.24, 6.31), Protonodosaria cf. proceraeformisGerke, 1959 (Plates 6.18 and 6.35), Geinitzina sp. (Plate 6.14), Pachyphloia spp. (Plate 6.34) and Cryptoseptida? sp. (Plate 6.29) and Colaniella? sp. The gastropod Bellerophon sp. (Plate 7.3) is relatively frequent.
The Khartam Member contains, at its base and in its lower part, a diversified assemblage composed of Permocalculus sp. (remains), Nankinella? sp., Globivalvulina cf. graeca, G. vonderschmittiReichel, 1946 (Plate 3.2), Dagmarita? shahrezaensis (Plates 3.10 and 3.15?), Paradagmarita flabelliformisZaninetti, Altiner and Çatal, 1981 (Plates 3.3 and 3.7), P. monodi Lys inLys and Marcoux, 1978 (Plate 3.11), P. “sp. 1” (Plates 3.4, 3.12 and 3.14), Glomospirella? linae n. sp. (Plate 4.16), Multidiscus spp. (Plate 2.16 and 2.17), Nodosinelloides cf. concinna (Plate 6.1), N. n. sp.? (Plate 6.17), Polarisella elabugae (Plate 6.2), P.? hoae (Plates 6.11 and 6.19), Protonodosaria cf. proceraeformis (Plate 6.15) and Pachyphloia spp. (Plates 6.25–6.28).
The Ash Shiqqah Member can be dated as late Midian (Figure 5) for two reasons: (1) the continuity with the early Dzhulfian Huqayl Member; (2) the lateral correlation with beds containing schwagerinoid fusulinids (e.g. Vaslet et al., 2005), for example Monodiexodina spp.
The stratigraphical range of Monodiexodina is discussed by Ueno (2003), Ehiro and Misaki (2004) and Pronina-Nestell (personal communication, 2005), but our observations in Afghanistan, Thailand and Timor (Vachard, unpublished) confirm the dating assigned by Kotlyar et al. (1989), i.e. late Midian, correlated with the Lepidolina zone, just before the disappearance of giant keriothecal fusulinids (Sheng, 1992; Jin et al., 1994).
The Huqayl Member is dated as early Dzhulfian by (1) the continuity with the late Dzhulfian Duhaysan Member; and (2) the absence of schwagerinoids, despite the relative diversity of the microfauna. The local marker seems to be Pseudomidiella cf. labensis, which appears at the base of the Huqayl Member, and is still present in the Duhaysan Member.
The Duhaysan Member is dated as late Dzhulfian by (1) the superposition to the Huqayl Member; and (2) the presence of very primitive Colaniella, considered as late Dzhulfian (e.g. Vachard et al., 2002). The earlier presence of Colaniella in the late Midian-early Dzhulfian (Chediya and Davydov, 1982; Jenny-Deshusses and Baud, 1989) was invalidated by the new dating of the beds by means of associated dunbarulid fusulinids Shindella (Kotlyar et al., 1999b, and Pronina-Nestell, personal communication, 2003). Contrary to age determination of Kotlyar et al. (1984) or Vachard et al. (2003), these beds are not late Dzhulfian but late Dorashamian in age.
The presence of Graecodiscus confirms an interval from Midian to Dorashamian (Vachard et al., 1993a; Pronina-Nestell and Nestell, 2001). Septoglobivalvulina? decrouezae is also Murgabian to Dorashamian in age, and only mentioned in the Wuchiapingian of Iran by Mohtat-Aghai and Vachard (in press).
The Midhnab Member is dated Changhsinghian (= Dorashamian) by (1) the superposition to the Duhaysan Member; (2) the presence of Paradagmarita; and (3) the presence of Glomospirella? linae n. sp.
The stratigraphical range of Paradagmarita is discussed: late Dzhulfian-early Dorashamian (Zaninetti et al., 1981; Altiner, 1981), latest Dzhulfian (Pronina 1988a, 1999), latest Dzhulfian-Dorashamian (Altiner and Özgül, 2001), Dorashamian (Vachard et al., 2002) or Changhsingian (Groves and Altiner, 2005). Gaillot (work in progress) considers a late Dzhulfian-Dorashamian distribution for P. monodi and several unpublished species, whereas he considers P. flabelliformis and P.“sp. 1” as Dorashamian fossils. Consequently, the Midhnab Member is controversially dated, i.e., late Dzhulfian or Dorashamian. It is assigned here to Changhsingian, since it contains Glomospirella? linae n. sp. synonym of “G. spirillinoides” considered as early Changhsingian in South China by Zhao et al. (1981).
The Lower Khartam Member is typically Changhsingian in age due to the specific diversification of Paradagmarita and the presence of P. flabelliformis and P. “sp. 1”. This unit probably belongs to the late Changhsingian since the Midhnab Member can be early Changhsingian, due to: (1) to the appearance of Glomospirella? linae; and (2) to the latest Permian character of the Lower Khartam Member.
The earliest fossiliferous Triassic levels in the Tethyan region with Rectocornuspira kahloriBrönnimann, Zaninetti and Bozorgnia, 1972 and Cyclogyra? mahajeriBrönnimann, Zaninetti and Bozorgnia, 1972, are identified in many areas, e.g. central Alborz (Iran), Abadeh region (Iran), northern Italy, Taurus (Turkey). They are related to stromatolitic or thrombolitic microbialites (Brönnimann et al., 1972; Altiner et al., 1979; Marcoux and Baud, 1986; Heydari et al., 2000, 2003; Crasquin-Soleau et al., 2002; Ünal et al., 2003), generally associated with a particular cement of calcite crystals (Heydari and Hassanzadeh, 2003). Until today, they have not been identified in Saudi Arabia.
The Triassic/Permian Boundary could be positioned between the Lower and Upper Khartam members (Figure 5). The basal part of the Upper Khartam Member only yields in the studied samples, a Polarisella? hoae (formerly Dentalina hoi) in a microfacies that is probably Triassic, according to the criteria of Ünal et al. (2003). The Upper Khartam Member contains the Early Triassic annelid Spirorbis phlyctaena (Manivit et al., 1985a, b, 1986; Vaslet et al., 1985b; Broutin et al., 1995). The biostratigraphical subdivisions are summarised in Figure 5.
SYSTEMATIC PALAEONTOLOGY: ALGAE
Genus: PermocalculusElliott, 1955
Type species: Gymnocodium gracilePia, 1937
Permocalculusspp. (Plates 1.1, 1.3–1.7)
Remarks: Contrary to Rezak (1959), Roux (1991) or Okla (1992), only Permocalculus was observed in our material. The genera Gymnocodium and Succodium are apparently lacking. The populations are relatively homogenous, and the different morphotypes indicated here, for example P. digitatus and P. solidus, could correspond to stages of growth of P. gracilis (Pia, 1937) as indicated by Roux (1991). Consequently our material should be reduced to P. ex gr. gracilis, moreover true P. gracilis is absent in Saudi Arabia. The alga attributed to this species by Delfour et al. (1982, fig. 15) is probably P. solidus.
Apparently, several genera or subgenera can correspond to morphotypes or stages of preservation of Permocalculus: DzhulfanellaKorde, 1965, SuccodiumKonishi, 1955, SiamporidiumEndo, 1969, TauridiumGüvenç, 1966, PyrulitesMu, 1981, NanjinophycusMu and Riding, 1983. The Gymnocodium of Delfour et al. (1982) and Okla (1992) are probably P. digitatus, and the Succodium of Okla (1992) seems to correspond to fertile (i.e. with conceptacles) specimens of P. solidus. Gymnocodium inhabits more marine environments than Permocalculus (Vachard et al., 2003).
Genus: MizziaSchubert, 1909
Type species: Mizzia velebitanaSchubert, 1909 (Plate 1.2)
1909 Mizzia velebitana n. gen. n. sp. - Schubert, pl. 16, figs. 8–12.
1992 Mizzia velebitana Schubert - Okla, pl. 45, figs. 7–10.
2000 Mizzia velebitana Schubert - Granier and Grgasovic, p. 102–107 (no illustration) (with synonymy).
Remarks: Very rare specimens are found, especially in the Huqayl Member. According to Okla (1992), the Mizzia are more abundant in the Duhaysan Member. Our material does not confirm this fact. “Mizzia” bramkampiRezak, 1959 was not observed in our samples. It is more probably Gyroporella or Eogoniolina. Consequently, the name “MizziaSchubert 1909, emend. Rezak, 1959” (Granier and Grgasovic, 2000) is considered here as superfluous and therefore only Mizzia Schubert was adopted.
Dimensions: Outer diameter = 0.630–0.930 mm, inner diameter = 0.430–0.670 mm, wall thickness = 0.100–0.130 mm, pore diameter = 0.040–0.120 mm.
Occurrence: The species and the genus are classically believed to be limited to the Permian, from Asselian to Dorashamian.
SYSTEMATIC PALAEONTOLOGY: “CALCISPHERES”
Genus: RadiosphaeraReitlinger, 1957,
Type species Radiosphaera basilicaReitlinger, 1957
cf. 1957 Radiosphaera basilica n. sp. Reitlinger, p. 775, fig. 1b.
Remarks: A unique level contains incertae sedis which look like Devonian Radiosphaera. This genus disappears in the Early Carboniferous (Mississippian). Its unquestionable presence in the Late Permian would correspond to a Lazarus effect. The morphological similarity is great with R. basilica, but the dimensions are smaller (inner diameter of R. basilica = 0.140–0.170 mm). By their size and their irregular external walls, these specimens differ from the Permian “calcispheres” of Cirilli et al. (1998).
Dimensions: Outer diameter = 0.090–0.150 mm, inner diameter = 0.045–0.080 mm, wall thickness = 0.020–0.060 mm.
Occurrence: Devonian-Mississippian, cosmopolitan. Discovered in the Duhaysan Member (late Dzhulfian) of Saudi Arabia.
SYSTEMATIC PALAEONTOLOGY: FORAMINIFERS
Genus: EarlandiaPlummer, 1930,
Type species: Earlandia perparvaPlummer, 1930
Remarks: Despite the relatively large size, the proloculus is never visible, but the morphological similarities (e.g. size and type of wall) are great with the primitive foraminifer Earlandia. Some other authors (e.g. Cirilli et al., 1998), assign the taxon to Aeolisaccus Elliott. The correct status of Aeolisaccus is not well established, it may be cyanobacteria (Chuvashov et al., 1987) or incertae sedis (Elliott, 1958). Many Aeolisaccus of the literature seem to be assigned to Earlandia, but proloculi are always lacking. Therefore, we use the name Earlandia?. A species of median size of Earlandia? in Saudi Arabia is comparable with the Viséan Earlandia minorRauzer-Chernousova, 1948a (Plate 2, figs. 1, 2: Length = 0.500–0.770 mm, Diameter = 0.115–0.130 mm, wall thickness = 0.020 mm). Another one is comparable with “Aeolisaccus” amplimuralisPantic, 1972 (Plate 2, figs. 3, 13: Diameter = 0.140–0.160 mm, wall thickness = 0.040–0.050 mm), and the last group, with “Aeolisaccus” tintinniformisMisik, 1971 (Plate 2, fig. 4: Diameter = 0.060 mm, wall thickness = 0.007 mm).
Occurrence:Earlandia is Mississippian-Pennsylvanian. “Aeolisaccus” is Middle Pennsylvanian?-Permian-Jurassic (e.g. Vachard and Montenat, 1981).
Genus: EostaffellaRauzer-Chernousova, 1948b,
Type species: Eostaffella parastruveiRauzer-Chernousova, 1948b
Eostaffella?sp. (Plate 2.14)
Remarks: Only one specimen similar to Eostaffella or Mediocris was found. These two genera disappear respectively in the Early Permian (= Cisuralian) and the Early Carboniferous (= Mississippian). According to Gaillot (unpublished data) several Eostaffellidae are also present in the Lopingian of Iran.
Dimensions: Diameter = 0.360 mm, Width = 0.180 mm, W/D = 0.50, proloculus diameter = 0.030 mm, number of whorls = 4.
Occurrence: Midhnab Member (early? Changhsingian) from Saudi Arabia.
? 1983 Nankinella minor Sheng - Vaslet et al., p. 18 (no illustration, not seen).
2005 Nankinella minor Sheng - Vaslet et al., p. 117 (no illustration).
2005 Nankinella sp. - Vaslet et al., p. 115 (fig. 34), 117.
Remarks: The material is poor and somewhat badly preserved. Furthermore, the specimens possibly suffered dwarfism and/or immaturity due to unfavourable environments. They are often abraded (corresponding to the early diagenetic types called Hayasakina). Therefore, the taxon is retained in open nomenclature. The last specimen of Nankinella (not illustrated here) from the Lower Khartam Member (sample VD 80-45) from the Al Quway’iyah Section (Figure 2) are more elongate (Diameter up to 1.500 mm and Width/Diameter = 0.40) and are tentatively assigned to Nankinella, because of apparently more curved septa.
Dimensions: Diameter = 0.720–0.950 mm, Width = 0.440 mm, W/D = 0.60, number of whorls = 5–6. Occurrence: Middle Pennsylvanian to Latest Permian, cosmopolitan.
Genus: GlobivalvulinaSchubert, 1921,
Type species: Valvulina bulloidesBrady, 1876
1946 Globivalvulina graeca n. sp. Reichel, p. 550–553, fig. in text 36, 38, pl. 19, figs. 15–17.
p. 1970 Globivalvulina graeca Reichel - Canuti et al., fig. 14. 3, 4, 6 (non fig. 14. 1 = Septoglobivalvulina gracilis).
1979 Globivalvulina graeca Reichel - Nguyen, p. 96, pl. 9, figs. 6–9 (with synonymy).
1981 Globivalvulina graeca Reichel - Altiner, p. 277–278, pl. 36, figs. 14–17 (with synonymy).
1984 Globivalvulina gracea [sic] Reichel - Altiner, pl. 1, figs. 13, 14.
v. 1985 Globivalvulina graeca Reichel - Vaslet et al., p. 16 (no illustration).
1986 Globivalvulina graeca Reichel - Marcoux and Baud, fig. 2 (no illustration).
1987 Globivalvulina graeca Reichel - Noé, p. 107, 108, pl. 30, fig. 4.
1989 Globivalvulina graeca Reichel - Köylüoglu and Altiner, pl. 7, figs. 8?, 9, 10.
1990 Globivalvulina graeca Reichel - Lin et al., p. 163, pl. 11, figs. 17–21 (with synonymy).
? 1995 Globivalvulina cf. graeca Reichel - Berczki-Makk et al., pl. 8, fig. 4. 1996 Globivalvulina graeca Reichel - Leven and Okay, pl. 8, fig. 16.
1999a Globivalvulina graeca Reichel - Kotlyar et al., p. 304, 307, 309, 311 (no illustration). 2005 Globivalvulina graeca Reichel - Vaslet et al., p. 117 (no illustration).
Remarks: Species of median size, characterised by the small number of whorls (1 to 1.5), the weakly depressed apertural face, and a possible increasing of height in the two last chambers. A population of the Midhnab Member (Khuff, Wadi Maghib Section, Figure 2) (Plate 3, fig. 13) is larger (Diameter up to 0.630 mm) with less chambers, and the last chamber is more quadratic. It is attributed to G. aff. graeca.
Dimensions: Diameter = 0.320–0.470 mm, proloculus = 0.050–0.060 mm, height of last chamber = 0.160–0.200 mm, wall thickness = 0.010–0.020 mm.
Occurrence: Middle-Late Permian, Tethyan including New Zealand (Vachard and Ferrière, 1991). Discovered in the Dzhulfian and perhaps the Dorashamian (with G. aff. graeca) of Saudi Arabia.
Globivalvulina vonderschmittiReichel, 1946 (Plate 3.2)
1946 Globivalvulina vonderschmitti n. sp. Reichel, p. 556, fig. 37a-e.
1980 Globivalvulina vonderschmitti Reichel - Vachard, pl. 29, fig. 17.
1981 Globivalvulina vonderschmitti Reichel - Altiner, p. 286–287, pl. 36, figs 10, 11, 12?, 13 (with synonymy).
1984 Globivalvulina vonderschmitti Reichel - Altiner, pl. 1, fig. 12.
1985 Globivalvulina vonderschmitti Reichel - Trifonova, pl. 1, figs. 9, 10.
v. 1985 Globivalvulina cf. vonderschmitti Reichel - Vaslet et al., p. 16 (no illustration).
1987 Globivalvulina vonderschmitti Reichel - Noé, p. 107, 108, pl. 30, fig. 5.
non 1987 Globivalvulina ex gr. G. vonderschmitti Reichel - Panzanelli-Fratoni et al., pl. 6, figs. 9–12 (probably Charliella).
1988 Globivalvulina vonderschmitti Reichel - Okimura, fig. 3. 5.
1989 Globivalvulina vonderschmitti Reichel - Köylüoglu and Altiner, pl. 7, figs. 5–7.
1989 Globivalvulina vonderschmitti Reichel - Kotlyar et al., pl. 2, fig. 27.
1991 Globivalvulina cf. vonderschmitti Reichel - Vachard and Ferrière, pl. 1, fig. 9.
1993b Globivalvulina vonderschmitti Reichel - Vachard et al., p. 97 (no illustration).
1999a Globivalvulina vonderschmitti Reichel - Kotlyar et al., p. 309, 312 (no illustration).
non v. 2002 Globivalvulina vonderschmitti Reichel - Vachard et al., pl. 1, fig. 16 (probably Charliella).
2003 Globivalvulina vonderschmitti Reichel - Ünal et al. pl. 1, fig. 13.
2004 Globivalvulina vonderschmitti Reichel - Kobayashi, fig. 6. 51.
2005 Globivalvulina vonderschmitti Reichel - Vaslet et al., p. 117 (no illustration; not seen).
Remarks: Only one specimen found; characterised by its large dimensions, its circular apertural face and its high and wide chambers.
Dimensions: Diameter = 0.790 mm, height of last chamber = 0.280 mm, wall thickness = 0.033 mm.
Occurrence: Murgabian?-Midian-Dorashamian of the Palaeo-Tethys. Discovered in the Lower Khartam Member (late? Dorashamian) from Saudi Arabia.
Remarks:Septoglobivalvulina differs from microgranular-walled Globivalvulina by the subspherical test and the inflated chambers, and differs from Paraglobivalvulina by the type of microstructure of the wall, and a less spherical test. The species G. decrouezae, which differs from typical Globivalvulina, is tentatively assigned here to Septoglobivalvulina. Some specimens of G. globosa Wang in Zhao et al. (1981) might be synonyms of G. decrouezae, with a priority of the first name.
p. 1970 Globivalvulina graeca Reichel - Canuti et al., fig. 14.1 (non fig. 14. 3, 4, 6 correctly interpreted). p. 1970 Globivalvulina sp. - Canuti et al., fig. 14. 2, 5.
?1981 Globivalvulina globosa n. sp. Wang in Zhao et al., p. 48 (in Chinese), p. 75-76 (in English), pl. 2, figs. 8-9. v. 1985 Paraglobivalvulina mira Reitlinger - Vaslet et al., p. 16 (no illustration). 1989 Globivalvulina sp. Kotlyar et al., pl. 3, fig. 21.
1989 Globivalvulina decrouezae n. sp. Köylüoglu and Altiner, p. 479-481, text-fig. 8 A-H, J-K, pl. 7, fig. 14.
?1990 Globivalvulina globosa Wang - Lin et al., p. 162, pl. 11, figs. 26-29.
1991 Paraglobivalvulina? sp. - Vachard and Ferrière, p. 2, fig. 2.
v. 1993 Globivalvulina or Paraglobivalvulina? - Fontaine et al., fig. 5E.
v. 1993 Paraglobivalvulina mira Reitlinger - Fontaine et al., fig. 6F.
v. 1994 Paraglobivalvulinoides? - Fontaine et al., pl. 47, fig. 7.
1998 Globivalvulina decrouezae Köylüoglu and Altiner - Altiner and Özkan-Altiner, pl. 3, fig. 23.
p. 2004 Globivalvulina globosa Wang - Zhang and Hong, p. 70, pl. 1, figs. 24-26.
v. 2005 Septoglobivalvulina decrouezae (Köylüoglu and Altiner) - Mohtat-Aghai and Vachard, pl. 2, fig. 17.
Remarks: Small, subspherical, biseriate test, coiled in a trochoid spire incompletely involute. Interseptal chamberlets are small. Wall very thin, microgranular. Only two specimens, both illustrated here.
Dimensions: Larger diameter = 0.340–0.360 mm, smaller diameter = 0.230–0.275 mm, wall thickness = 0.007–0.015 mm.
Occurrence: Murgabian-Dorashamian of southern Turkey (e.g. Köylüoglu and Altiner, 1989; Ünal et al., 2003; Canuti et al., 1970), Thailand and Malaysia (Fontaine et al., 1993, 1994). Late Midian of Transcaucasia (Kotlyar et al., 1989). Midian of New Zealand (Vachard and Ferrière, 1991), Dorashamian of Greece (Altiner and Özkan-Altiner, 1998). Changhsingian of South China (Lin et al. 1990). Wuchiapingian of Iran (Mohtat-Aghai and Vachard, 2005). Discovered in the Duhaysan Member (late Dzhulfian) from Saudi Arabia.
Genus: DagmaritaReitlinger, 1965,
Type species : Dagmarita chanakchiensisReitlinger, 1965
Remarks: This Dagmarita is devoid of a horn-like expansion. This character is of generic rather than of specific significance. Therefore, the taxon is denominated here D.? sharezaensis.
Dimensions: Height = 0.620–0.800 mm, Width = 0.230–0.450 mm, whorls = 4–5 pairs.
Occurrence: Dzhulfian-Dorashamian of Iran, Himalaya, Transcaucasia, South China and Malaysia. Discovered in the Midhnab and Lower Khartam members (Dorashamian) of Saudi Arabia.
Genus: Paradagmarita Lys in Lys and Marcoux, 1978,
Type species: Paradagmarita monodi Lys in Lys and Marcoux, 1978
1985b Paradagmarita flabelliformis Zaninetti, Altiner and Çatal - Manivit et al., fig. 5 (no illustration; not seen; mentioned in the Duhaysan Member).
1986 Paradagmarita flabelliformis Zaninetti, Altiner and Çatal - Manivit et al., p. 14 (no illustration; not seen; mentioned in the Huqayl and Duhaysan Members).
1989 Paradagmarita flabelliformis Zaninetti, Altiner and Çatal - Köylüoglu and Altiner, pl. 6, fig. 9.
1999 Paradagmarita flabelliformis Zaninetti, Altiner and Çatal - Pronina, tabl. 2 p. 189 (no illustration).
2005 Paradagmarita flabelliformis Zaninetti, Altiner and Çatal - Vaslet et al., p. 117 (no illustration; not seen; mentioned in the Huqayl and Duhaysan members).
Remarks: Test biseriate, secondarily coiled in an involute spire. Juvenarium with a small distortion. Adult chambers enlarged in the equatorial plane giving the characteristic flabelliform shape.
Dimensions: Height = 0.535 mm, Width = 0.195 mm.
Occurrence: Latest Dzhulfian-Dorashamian of Turkey. Discovered in the Lower Khartam Member (late? Dorashamian) of Saudi Arabia.
Paradagmarita n. sp. (Plate 3.8)
? 1988a No legend - Pronina, pl. 2, fig. 14.
? 1989 Paradagmarita sp. - Pronina, pl. 1, fig. 14.
v. 2005 Paradagmarita sp. -Vaslet et al. p. 77, 115 (fig. 34), 117, 118 (fig. 35), 127 (no illustration).
Remark: This new species differs from P. flabelliformis by the wider and more trochoid uncoiled part, and the shorter initial coiled part.
Dimensions: Height = 0.700 mm, Width = 0.560 mm, proloculus diameter = 0.030 mm, height of last chamber = 0.100 mm, 7 chambers in the arcuate part, 10? in the rectilinear biseriate part.
Occurrence: Only one section in the Midhnab Member (early? Dorashamian) from Saudi Arabia.
1981 Paradagmarita monodi Lys in Lys and Marcoux - Altiner, p. 295, 296, pl. 38, figs. 1–15.
1984 Paradagmarita monodi Lys in Lys and Marcoux - Altiner, pl. 1, figs. 1, 2. non 1985 Paradagmarita monodi Lys in Lys and Marcoux - Okimura et al., pl. 1, fig. 16 (probably a teratogenic Globivalvulina).
1986 Paradagmarita monodi Lys in Lys and Marcoux - Marcoux and Baud, figs. 2–4 (no illustration).
1986 Paradagmarita monodi Lys in Lys and Marcoux - Manivit et al., p. 14 (no illustration; not seen; mentioned in the Duhaysan Member).
1988 Paradagmarita monodi Lys in Lys and Marcoux - Loeblich and Tappan, pl. 230, fig. 6.
1996 Paradagmarita monodi Lys in Lys and Marcoux - Rauzer-Chernousova et al., p. 72, pl.18, fig. 14.
v. 2005 Paradagmarita monodi Lys in Lys and Marcoux - Vaslet et al., p. 117 (no illustration; not seen; mentioned in the Duhaysan Member).
Remarks: Wall simple, microgranular. Small specimen, which is considered here as a juvenile of P. monodi.
Dimensions: Height = 0.330 mm, Width = 0.185 mm, number of whorls = 3 pairs.
Occurrence: Taurus (Turkey), Zagros (Iran), Dzhulfian-Dorashamian. The specimen of Oman, not illustrated (Montenat et al., 1977), is probably misinterpreted because its association with late Midian Shanita is theoretically impossible (Gaillot and Vachard, unpublished data).
Paradagmarita“sp. 1” (Plates 3.4, 3.12, 3.14)
Remarks:P. sp. 1 differs from P. flabelliformis by the thicker septa often microsparitised. The specimens are here peripherically abraded.
Dimensions: Maximal length of a fragment = 0.500 mm, septa thickness = 0.040–0.055 mm.
Occurrence: Dorashamian of Iran (Gaillot, unpublished data). Discovered in the Lower Khartam Member (late? Dorashamian) of Saudi Arabia.
Genus: CornuspiraSchultze, 1854,
Type species: Orbis foliaceus Philippi, 1844 (see Loeblich and Tappan, 1988)
Remarks: Contrary to Pronina (1999), Cornuspira, a porcelaneous foraminifer, is not a synonym of Pseudoammodiscus, with a microgranular test. They belong to two different orders, respectively Miliolida and Fusulinida of the subclass Foraminiferea.
Remarks: Test small, biumbilicate, with relatively numerous and narrow whorls.
Dimensions: Diameter = 0.160–0.190 mm, Width = 0.060 mm, w/D = 0.38, proloculus diameter = 0.075 mm, whorls = 4.5–5.
Occurrence: Early Pennsylvanian (Bashkirian) to Recent.
Genus: Hemigordiellina Marie in Deleau and Marie, 1961 emend. Vachard and Beckary, 1991 (= GlomospiraRzehak, 1885 of the authors)
Type species: Glomospira diversaCushman and Waters, 1930
Hemigordiellina regularis(Lipina, 1949) (Plate 2.6 and 2.8)
1978 Glomospira regularis Lipina - Lin, p. 11, pl. 1, fig. 8.
1990 Glomospira regularis Lipina - Lin et al., p. 119, pl. 1, figs. 36–38 (with synonymy).
? 2001b Pseudoagathammina regularis (Lipina) - Vachard and Krainer, pl. 4, figs. 16, 17, 19-21.
v. 2005 “Glomospira” - Vaslet et al., p. 115, fig. 34 (no illustration).
Remarks: Porcelaneous test, streptospirally but relatively regularly coiled.
Dimensions: Diameter = 0.240–0.265 mm, proloculus = 0.050 mm, height of the last whorl of the tube = 0.050 mm, wall thickness = 0.007 mm.
Genus: ?GlomospirellaPlummer, 1945,
Type species: Glomospira umbilicataCushman and Waters, 1927
Remarks: Some undeterminated specimens having the coiling of Glomospirella are present, but Glomospirella is theoretically agglutinated and not porcelaneous. The misinterpreted Pseudovidalina spp. and Raphconilia? sp. of Kobayashi (2002: figs. 9.1–7) might belong to Glomospirella?. Moreover, true Pseudovidalina and Raphconilia, which are synonyms (Pinard and Mamet, 1998; Vachard and Krainer, 2001a), differ from this genus.
Occurrence: Duhaysan and Midhnab members of Saudi Arabia (Dzhulfian-Dorashamian).
Glomospirella? linae Vachard and Gaillot n. sp. (Plates 4.3–4.17)
1978 Glomospirella spirillinoides Lipina - Lin, p. 12, pl. 1, figs. 11–12.
1981 Glomospirella spirillinoides (Grozdilova and Glebovskaya) - Zhao et al., pl. 1, figs. 8, 9.
v. 2005 “Glomospirella spirillinoides” (Grozdilova and Glebovskaya) - Vaslet et al., p. 77, 115 (fig. 34), 117, 118 (fig. 35), 127 (no illustration).
Etymology: Dedicated to Mrs. Jiaxing Lin, eminent specialist of the Chinese Permian microfauna. Type level: Lower part of the Midhnab Member, early? Dorashamian from Saudi Arabia. Type locality: Al Huwwah Section, Saudi Arabia (Figure 2). Holotype:Plate 4, fig. 6 (Sample VD 80-84. Photo 11).
Diagnosis: The species is characterised by its relative large size, compared to the small irregular stage. The cross section of the tube is hemi-circular to bean-shaped. The wall is relatively thick.
Description: Test discoidal, generally deeply biumbilicate. The initial irregularly coiled initial part is reduced to 1–2 oscillating whorls; the diameter of the initial part does not exceed 1/5 of the test diameter. The adult second part is constituted by 4–5 aligned to planispiral whorls. The width and height of deuteroloculus increases slowly. The transverse section of this deuteroloculus is hemicircular to bean-shaped. Proloculus is spherical and relatively large, but megalo- and microspheric specimens exist. Wall porcelaneous relatively thick. Some well-preserved specimens have their primary porcelaneous (amber-coloured), becoming black (Plates 4.11, 4.14 and 4.15).
Dimensions: Diameter = (0.350–0.435) 0.520–1.180 mm, Width = (0.100–0.110) 0.120–0.220 mm, W/D = 0.16–0.21 (0.22–0.26), number of whorls = (4–4.5) 5.5–7, proloculus diameter = 0.020–0.050 mm, height of last whorl = (0.040) 0.060–0.100 mm, wall thickness = (0.005) 0.010–0.030 (0.050) mm.
Comparisons: Differs from all other Glomospirella (in a broad sense) described in the literature by the weak initial deviation and the numerous aligned whorls; differs from Brunsia spirillinoides (Grozdilova and Glebovskaya) by the wall porcelaneous and not microgranular (i.e., by the membership to the order Miliolida and not to the order Fusulinida). Differs from Glomospirella? spp. mentioned above, by its dimensions, and the number of whorls compared to the size.
Occurrence: Dorashamian (Midhnab and Lower Khartam members) of Saudi Arabia. Changhsinghian of South China.
Genus: PalaeonubeculariaReitlinger, 1950,
Type species: Palaeonubecularia fluxaReitlinger, 1950
Palaeonubecularia n. sp. (Plate 2.18)
Remark: A sessile species with a long, irregularly coiled deuteroloculus. The dimensions of the deuteroloculus increase gradually in height and width during the growth. The most closely packed first whorls form a juvenarium. The porcelaneous wall is often recrystallised in anhydrite.
Palaeonubecularia n. sp. differs from P. rustica Reitlinger, by the coiled juvenarium, different size parameters and the Late Permian age. This species builds some small foraminiferal boundstones in Iran (Gaillot, unpublished data).
Dimensions: Maximal length of the test = 0.675 mm, maximal length of juvenarium = 0.265 mm, maximal height of the deuteroloculus = 0.150 mm, maximal thickness of whorl = 0.085 mm.
Occurrence: Latest Midian-Dzhulfian of Iran (Gaillot, work in progress). Discovered in the Midhnab Member (early? Dorashamian) of Saudi Arabia.
Genus: HemigordiusSchubert, 1909,
Type species: Cornuspira schlumbergeriHowchin, 1895
1990 Hemigordius schulumbergeri (sic) (Howchin) - Lin et al., p. 212, pl. 24, fig. 22.
2000 Hemigordius schlumbergeri (Howchin) – Groves, pl. 3, figs. 21-28.
2001a Hemigordius schlumbergeri (Howchin) – Vachard and Krainer, pl. 4, fig. 8.
2001b Hemigordius schlumbergeri (Howchin) - Vachard and Krainer, pl. 9, figs. 8, figs. 18, 19, 25.
2003 Hemigordius schlumbergeri (Howchin) - Altiner et al., fig. 4 in text.
2003 Hemigordius schlumbergeri (Howchin) - Shang et al., p. 380, pl. 2, figs. 2–5.
Remarks: Test discoidal, sides almost parallel, 1–2 streptospiral initial whorls followed by 3–4 aligned whorls, last whorl often semi-evolute.
Dimensions: Diameter = 0.330 mm, Width = 0.095 mm, W/D = 0.29, proloculus = 0.030 mm.
Occurrence: From Late Pennsylvanian (Pinard and Mamet, 1998) to Middle Permian of Russia (Pronina, 1996), and South China (Lin et al., 1990). Rare in late Changhsingian of South China (Shang et al., 2003).
v. 2005 Hemigordius baoqingensis Wang - Vaslet et al., p. 77, 115 (fig. 3), 116, 118 (fig. 35), 127 (no illustration).
Remarks: Test of medium size, slightly asymmetrical and flattened on one side. Deuteroloculus with oscillating coiling. Wall recrystallised in white microsparite.
Dimensions: Diameter = 0.500–0.550 mm, Width = 0.170–0.220 mm, proloculus diameter = 0.040 mm, number of whorls = 4–5.
Genus: NeodiscusMiklukho-Maclay, 1953,
Type species: Neodiscus milliloidesMiklukho-Maclay, 1953
Remarks:Neodiscus is similar to Hemigordius, i.e., constituted by a tube, without septa or pseudosepta, enrolled streptospirally at the juvenile stage, oscillating, aligned or sigmoid at the adult stage and involute or semi-evolute. Hemigordius is compressed and discoidal, whereas Neodiscus is inflated.
aff. 1976 Neodiscus qinglongensis n. sp. Wang, p. 191–192, pl. 1, figs. 11a–d.
? v. 1978 Hemigordius sp. 1. Zaninetti et al., pl. 85, figs. 26–31.
aff. 1990 Hemigordius qinglongensis (Wang) - Lin et al., p. 214, pl. 25, figs. 14, 15.
? p. 1998 Hemigordius sp. - Cirilli et al., pl. 2, figs. 17, 18 (not figs. 15, 19 = true Hemigordius).
? 2003 Hemigordius? sp. - Ünal et al., pl. 1, fig. 46.
Remarks: Test thickly lenticular, involute, of median size. Spherical proloculus passing into tubular undivided deuteroloculus coiled in two planes. Pseudo-septa absent or very faint. 3–4 initial whorls (glomus) are streptospirally coiled; the following 1–2 whorls are slightly oscillating to aligned. Aperture terminal simple. Wall porcelaneous relatively thick, usually recrystallised in white microsparite.
Dimensions: Diameter = 0.340–0.800 mm, Width = 0.200–0.480 mm, W/D = 0.60–0.65, proloculus = 0.050–0.060 mm (for specimens of 0.460–0.800 mm).
Comparison: Differs from N. qinglongensis by its less oscillating final whorls. In Saudi Arabia, several populations exist: the Huqayl Member specimens are smaller than the Midhnab Member specimens
(Plate 5.3 and 5.7).
Occurrence: Wuchiapingian-Changhsingian type of South China. Questionable in the Dorashamian of Turkey, Iran and Italy (see list of synonymy). Discovered from the Huqayl to the Midhnab members (Dzhulfian-early? Dorashamian).
Genus: Pseudomidiella Pronina-Nestell inPronina-Nestell and Nestell, 2001,
Type species: Pseudomidiella labensis Pronina-Nestell inPronina-Nestell and Nestell, 2001
Remarks: With the exception of the type material and the population from New Zealand described by Vachard and Ferrière (1991), almost all the Baisalina of the literature are assignable to Pseudomidiella. Consequently its range is Midian (see in particular the “Baisalina” of Gargouri and Vachard, 1988) to Dorashamian in the Palaeo-Tethys and the Neo-Tethys.
v. 1985 Baisalina - Vaslet et al., p. 16 (no illustration).
cf. 2001 Pseudomidiella labensis n. sp. Pronina-Nestell in Pronina-Nestell and Nestell, p. 213, 214, pl. 1, figs. 18-21 (with synonymy).
v. 2005 Pseudomidiella cf. labensis Pronina-Nestell in Pronina-Nestell and Nestell - Vaslet et al., p. 77, 116, 118 (fig. 35), 127 (no illustration).
Remarks: Test of median size, ovoid in longitudinal section and round in transverse section. The coiling is sigmoidal, with 1–1.5 aligned last whorl. Proloculus spherical small; deuteroloculus with rare pseudo-septa. Wall porcelaneous, black. Aperture terminal simple.
Our specimens differ from typical specimens by the type of coiling and the shorter septa.
Dimensions: Diameter = 0.430–0.570 mm, Width = 0.280–0.300 mm, W/D = 0.60–0.70, whorls = 5–6,
proloculus = 0.030–0.045 mm.
Occurrence: The typical specimens have been found in the Himalaya, northern Italy and NW Caucasus. Discovered in the Huqayl Member, and rare in the Duhaysan Member (Dzhulfian) of Saudi Arabia.
Genus: NeohemigordiusWang and Sun, 1973,
Type species: Neohemigordius maopingensisWang and Sun, 1973
1978 Hemigordius zaninettiae n. sp. Altiner, p. 28, pl. 1, figs. 7–14.
1981 Hemigordius changxingensis n. sp. Wang in Zhao et al., p. 47, 73, pl. 1, fig. 16.
1988b Hemigordius (Midiella) zaninettiae Altiner - Pronina, fig. 2. 19, 20.
? 1989 Hemigordius (Midiella) zaninettiae Altiner - Kotlyar et al. tabl. 1 p. 32 (no illustration).
1990 Hemigordius changxingensis Wang - Lin et al., p. 212, pl. 24, fig. 36.
1996 Hemigordius zaninettiae Altiner - Leven and Okay, pl. 9, figs. 22, 23.
1998 Hemigordius zaninettiae Altiner - Altiner and Özkan-Altiner, pl. 4, fig. 17.
1999a Midiella zaninettiae (Altiner) - Kotlyar et al., p. 309 (no illustration).
2001 Midiella zaninettiae (Altiner) - Pronina-Nestell in Pronina-Nestell and Nestell, pl. 1, fig. 17. 2003 Hemigordius zaninettiae Altiner - Altiner et al., p. 207, fig. 6 in text. 2003 Hemigordius zaninettiae Altiner - Ünal et al., pl. 1, fig. 47.
Remarks: This is a small species of the group zaninettiae, apparently very similar to “Hemigordius” changxingensis. It is lenticular, characterised by its regularly oscillating coiling. Because of the inflated test and the streptospirally coiling followed by a planispiral coiling, this species is attributed here to Neohemigordius, equivalent to Midiella (see Vachard and Ferrière, 1991).
Dimensions: Diameter = 0.285 mm, Width = 0.115 mm, W/D = 0.40, number of whorls = 7.
Occurrence: Midian of New Zealand. Dorashamian of Turkey and Greece (see list of synonymy). Late Changhsingian of South China and NW Caucasus. Discovered in the Midhnab Member (early? Dorashamian) of Saudi Arabia.
Genus: MultidiscusMiklukho-Maclay, 1953,
Type species: Nummulostegina padangensisLange, 1925
Remarks: Several species are probably present, but each one represented by one or two specimens.
Dimensions: Specimen fig. 16: Diameter = 0.360 mm, Width = 0.145 mm, W/D = 0.40, number of whorls = 5? Specimen fig. 17: Diameter = 0.385 mm, Width = 0.180 mm, W/D = 0.47, number of whorls = 4.5.
Occurrence: Midian-Dorashamian of Tethys. Discovered in the Midhnab and Lower Khartam members (Dorashamian) of Saudi Arabia.
Genus: AgathamminaNeumayr, 1887,
Type species: Serpula pusilla Geinitz inGeinitz and Gutbier, 1848
Agathamminaspp. (Plate 2.21)
Remarks: The quinqueloculine regular coiling of the tubular undivided deuteroloculus is a characteristic feature. The material is too poor (four sections) to discriminate a species. The coiling seems to be similar to A. pusilla (Geinitz), but the wall is especially thin.
Dimensions: Diameter = 0.650 mm, Width = 0.170 mm, number of whorls = 6–7.
Occurrence: The FAD is poorly known; the genus is relatively common from Midian to Dorashamian. The LAD seems to be Rhaetian in the Austrian Alps (Salaj et al., 1983) but this reference is discussed (Pronina-Nestell, personal communication, 2005). Discovered in the Dzhulfian and Dorashamian (Huqayl and Midhnab members) from Saudi Arabia.
Genus: Graecodiscus Vachard inVachard, Clift and Decrouez, 1993a,
Type species: Graecodiscus teresae Vachard inVachard, Clift and Decrouez, 1993a
?1995 Agathammina multa Pronina - Berczi-Makk et al., pl. 13, fig. 8.
cf. 2001 Graecodiscus kotlyarae n. sp. Pronina-Nestell in Pronina-Nestell and Nestell, p. 214, pl. 1, fig. 23. v. 2005 Graecodiscus cf. kotlyarae Pronina-Nestell in Pronina-Nestell and Nestell - Vaslet et al., p. 77, 115 (fig. 34), 116, 117, 118 (fig. 35), 127 (no illustration).
Remarks: Test relatively large, inflated and prominent in the central part, discoidal in the extremities. Proloculus large. 4 initial agathamminoid whorls are followed by 2–3 aligned whorls. The last half whorl is semi-involute. This taxon differs from typical G. kotlyarae by the type of coiling.
Dimensions: Diameter = 0.460–0.850 mm, Width = 0.200–0.240 to 0.370 mm, number of whorls = 4–6.5, proloculus = 0.065–0.085 mm, height of last whorl = 0.100–0.115 mm, wall thickness = 0.007–0.020 mm.
Occurrence: Late Changhsingian of Caucasus. Discovered in the Duhaysan Member (late Dzhulfian) of Saudi Arabia.
Genus: RectostipulinaJenny-Deshusses, 1985,
Type species: Rectostipulina quadrataJenny-Deshusses, 1985
Rectostipulinasp. (Plate 6.8)
Remarks: Only one transverse section measuring: outer diameter = 0.075 mm, inner diameter = 0.035 mm.
Occurrence:Rectostipulina is Midian-Dorashamian in age in Palaeo-Tethys and Neo-Tethys. Pronina (1995) considers its FAD as Dzhulfian. Discovered in the Duhaysan Member (late Dzhulfian) of Saudi Arabia.
Genus: NodosinelloidesMamet and Pinard, 1992,
Type species: Nodosinelloides potievskayaeMamet and Pinard, 1996 (for Nodosaria gracilisPotievskaya, 1962 preoccupied)
1990 Nodosaria shikhanica Lipina - Lin et al., p. 226, 227, pl. 28, fig. 13 (with synonymy).
Remarks: Acute test with hemispherical chambers increasing in height, but not in width. Smooth surfaces of the wall and the septa. N. shikhanica is often considered synonymous with N. netschajewi (Cherdyntsev, 1914), but this species is poorly defined, and needs revision. We use the name N. shikhanica well defined by Lipina (1949).
Dimensions: Height = 0.400 mm, Width = 0.070 mm, number of chambers = 8.
Occurrence: Mainly, latest Pennsylvanian-earliest Permian, but ranges to the Dorashamian of South China (Lin et al., 1990). Discovered in the Dzhulfian-Dorashamian of Saudi Arabia (Huqayl Duhaysan, Midhnab and Lower Khartam members).
cf. 1962 Nodosaria concinna n. sp. Potievskaya, p. 70, 71, pl. 5, figs. 16, 17. cf. 1990 Nodosaria concinna Potievskaya - Lin et al., p. 222, pl. 27, fig. 18.
cf. 2004 Nodosaria concinna Potievskaya - Zhang and Hong, p. 21, pl. 2, fig. 20 (with synonymy). v. 2005 Nodosaria dzhulfensis Reitlinger - Vaslet et al., p. 77, 115 (fig. 34), 118 (fig. 35) (no illustration).
Remarks: Test rectilinear, gently tapering. Arched chambers increasing slowly in height and width. Smooth surfaces of the wall and the septa. Although this species was described in the Early Permian, the section of Saudi Arabia seems to be relatively similar to N. concinna. It was confused with N. dzhulfensis Reitlinger by Vachard in Vaslet et al. (2005).
Dimensions: Height = 0.280 mm, Width = 0.100 mm, number of chambers = 6, outer proloculus diameter = 0.050 mm.
Remarks: Similar in size to N. cf. concinna, this unique specimen differs by more inflated chambers and more depressed sutures.
Dimensions: Height = 0.250 mm, Width = 0.090 mm, number of chambers = 5?
Occurrence: Discovered in the Huqayl Member (early Dzhulfian) of Saudi Arabia.
Nodosinelloidesn. sp.? (Plate 6.17)
Remarks: Very small species very acute with constrictions and consequently deep sutures between each chamber.
Dimensions: Height = 0.160 mm, Width = 0.040 mm, number of chambers = 6 or 7.
Occurrence: Discovered in the Lower Khartam Member (late? Dorashamian) of Saudi Arabia.
Genus PolarisellaMamet and Pinard, 1992,
Type species Polarisella blindensisMamet and Pinard, 1992
1953 Nodosaria ex gr. elabugae Tscherdynzew (sic) - Rauzer-Chernousova, p. 77, fig. 32 in text.
? 1970 Frondicularia sp. - Canuti et al., fig. 12. 10 (cf. our Pl. 6, fig. 21).
p. 1970 Frondicularia cf. woodwardi Howchin - Canuti et al., fig. 12. 11 (non fig. 12. 12 = N. hoae).
1978 Nodosaria armeniensis Efimova - Lys and Marcoux, pl. 1, fig. 15.
? 1982 Nodosaria djulfensis (sic) Reitlinger - Delfour et al., fig. 16.
? 1983 Nodosaria dzhulfensis Reitlinger - Vaslet et al., p. 18 (no illustration).
1983 Nodosaria armeniensis Efimova - Jenny-Deshusses, pl. 19, fig. 1.
? 1983 Nodosaria ordinata Trifonova - Salaj et al., p. 118–119, pl. 80, figs. 9, 14, pl. 144, fig. 10. ? 1983 Frondicularia woodwardi Howchin - Salaj et al., pl. 82, figs. 2–7, 9–13.
1984 Nodosaria armeniensis Efimova - Kotlyar et al., pl. 6, fig. 13.
1985 Nodosaria armeniansis (sic) Efimova - Trifonova, pl. 3, fig. 7.
1986 Nodosaria armeniensis Efimova - Marcoux and Baud, figs. 2, 4 (no illustration). ? 1989 Nodosaria armeniensis Efimova - Kotlyar et al., table 1, p. 33 (no illustration). 1992 Polarisella elabugae (Cherdyntsev) - Mamet and Pinard, p. 377 (no illustration). 1996 Nodosaria elabugae Tcherdynzev (sic) - Pronina, p. 251, pl. 3, fig. 14.
? 1996 Cryptoseptida? sp. - Kobayashi, fig. 5. 24–25.
1998 Nodosaria elabugae Tcherdyncev (sic) - Pronina, p. 167 (no illustration).
Remarks: Test acute, gently tapering, with bridge-shaped septa, occasionally with rugosities protruding in the following chamber (Plate 6, fig. 22). Last chamber often hemispherical (Plate 6, figs. 3, 22).
The species seems to be relatively similar to several Permian and Triassic taxa: N. armeniensis, N. ordinata and F. woodwardi auctorum.
Dimensions: Height = 0.115–0.350 mm, Width = 0.050–0.090 mm, number of chambers = 4–8.
Occurrence: Permian of former USSR. Dzhulfian of Turkey and Transcaucasia. Possible presence in the Triassic. Discovered in the Dzhulfian-Dorashamian of Saudi Arabia (Huqayl, Duhaysan, Midhnab and Lower Khartam members).
Remarks: Similar to P. elabugae, but with much larger size. This morphotype is limited to the Midhnab Member in Saudi Arabia. This species is probably new (as confirmed by G. Pronina-Nestell, personal communication, 2005) but our material is too poor to allow a specific description.
Dimensions: Height = 0.385–0.450 mm, Width = 0.080–0.110 mm, number of chambers = 7?–8?
Occurrence: Discovered in the Midhnab Member (early? Dorashamian) of Saudi Arabia.
1974 Dentalina hoi Trifonova - Efimova, p. 71 (no illustration).
1975 Dentalina hoi Trifonova - Styk, p. 518–519, pl. 36, fig. 14 (with synonymy).
p. 1970 Frondicularia cf. woodwardi Howchin - Canuti et al., fig. 12. 12 (non fig. 12. 11 = N. elabugae). 1982 Nodosaria armeniensis Efimova - Delfour et al., fig. 16.
1982 Nodosaria armeniensis Efimova - Vaslet et al., p. 18 (no illustration, not seen).
v. 2005 “Dentalina” hoi - Vaslet et al., p. 77, 115 (fig. 34), 116, 117, 118 (fig. 35), 127 (no illustration).
Remarks: Test sometimes arcuate (Plate 6.11). Chambers subquadratic to pear-shaped. Wall relatively thick and chamber reduced to a conical or trapezoidal space. The characters of Dentalina are very subjective in this species, hence it is questionably attributed here to Polarisella because of the thick wall and the shape of the chambers. The rare specimens of Saudi Arabia are incomplete or immature. Adult paratypes of P.? hoae measure: Height = 0.350–0.450 mm, width = 0.070–0.100 mm for 8–9 or usually 6–7 whorls (Trifonova, 1967). Dedicated to Mrs. Ho, the Latin genitive must be hoae not hoi.
Dimensions: Height = 0.265–0.315 mm, Width = 0.030–0.070 mm, number of chambers = 5–6.
Occurrence: Late Permian of Hazro (Turkey). Triassic of central Europe (e.g. Salaj et al., 1983). Discovered from the Huqayl Member to the Upper Khartam Member (Dzhulfian-Early Triassic) of Saudi Arabia.
Genus: ProtonodosariaGerke, 1959,
Type species: Protonodosaria proceraformisGerke, 1959
cf. 1961 Protonodosaria proceraformis Gerke - Gerke, p. 163, pl. 20, figs. 12, 13, pl. 21, figs. 8–10.
cf. 1979 Protonodosaria proceraformis Gerke - Nguyen, p. 92, pl. 6, figs. 13–17.
cf. 1988 Protonodosaria proceraformis Gerke - Loeblich and Tappan, pl. 434, figs. 28, 29.
cf. 1999 Protonodosaria proceraformis Gerke - Pronina, p. 183, 184 (no illustration).
Remarks: Medium-sized taxon (one of the largest lagenids in Saudi Arabia), with chambers increasing moderately in height and width. Wall relatively thick. Many characters are common with P. proceraeformis, but the apertures were not observed.
Dimensions: Height = 0.840 mm, Width = 0.270 mm, number of chambers = 6.
Occurrence: Permian of Russia. Midian of Cambodia and Malaysia. Dorashamian of Saudi Arabia (Midhnab and Lower Khartam members).
Genus: GeinitzinaSpandel, 1901,
Type species: Geinitzina postcarbonicaSpandel, 1901
Geinitzinaspp. (Plate 6.14)
Remarks: According to the taxonomic criteria of the literature, several “species” could be present, but the group Geinitzina postcarbonica, illustrated here, is the most frequent. A possible Geinitzina “reperta” Bykova is mentioned by Delfour et al. (1982, fig. 16). In fact, the taxon reperta belongs to the Devonian genus Eogeinitzina, and the Permian specimens attributed to Geinitzina “reperta” must be renamed.
Dimensions: Height = 0.200 mm, Width = 0.150 mm, number of chambers = 7.
Genus: PachyphloiaLange, 1925,
Type species: Pachyphloia ovataLange, 1925
Remarks: Several species differing principally by the size and the general morphology (compare Plate 6.26 specimen with height = 0.320 mm, and Plate 6.34 specimen with height = 1.120 mm).
Occurrence: The FAD of Pachyphloia is discussed as probably middle Permian according to Pinard and Mamet (1998), late Early Permian, Chihsian (Lin et al., 1990), Artinskian (Groves, 2000), or may be early Early Permian (Sakmarian) (Vachard and Krainer, 2001b). The last downhole appearance (LAD) is Dorashamian (e.g. Lin et al., 1990; Pronina-Nestell and Nestell, 2001; Wang and Ueno, 2003; Shang et al., 2003).
Genus: ?CryptoseptidaSellier de Civrieux and Dessauvagie, 1965,
Type species: Cryptoseptida anatoliensisSellier de Civrieux and Dessauvagie, 1965
Remarks:Cryptoseptida is a poorly known genus. It is interpreted here as a morphological variation of Pachyphloia with deeply arched chambers. Hence, a small gap is visible between the successive whorls (compare Plate 6, fig. 29 and Plate 6, fig. 26).
Cryptoseptida?sp. (Plate 6.29)
p. 1970 Robuloides sp. - Canuti et al., fig. 12. 3 (not figs. 6, 8 = Pachyphloia). v. 1985 Cryptoseptida sp. - Vaslet et al., p. 16 (no illustration).
Remarks: Due to the absence of illustrated axial section of Cryptoseptida anatoliensis, specific identification is impossible.
Dimensions: Height = 0.430 mm, Width = 0.150 mm, number of chambers = 6?
Occurrence: Late Permian of Turkey. Dzhulfian of Transcaucasia (Kotlyar et al., 1989). Cryptoseptida? sp. is mentioned by Wang and Ueno (2003) in the Changhsingian of South China, and Cryptoseptida sp. in the Changhsingian of Japan by Kobayashi (2002). This form of Kobayashi (2002, fig. 9. 40–41) corresponds to our definition of Cryptoseptida. Another Cryptoseptida? sp. is cited in the Midian of Japan, but its illustration (Kobayashi, 2001, pl. 2, fig. 14) corresponds more probably to the genus Pseudolangella. The Triassic (Anisian) Cryptoseptida? sp. of Kobayashi (1996) is more similar to Nodosinelloides elabugae. Cryptoseptida is synonymised with Pachyphloides by Berczi-Makk (1996), and might range in this case to Jurassic, but we do not agree with this synonymy. Discovered in the Midhnab Member (early? Dorashamian) of Saudi Arabia.
Genus: RobuloidesReichel, 1946,
Type species: Robuloides lensReichel, 1946
1946 Robuloides lens n. sp. Reichel, p. 536, fig. 21–26 in text, pl. 19, figs. 6, 7.
1981 Robuloides lens Reichel - Vachard and Montenat, pl. 15, fig. 6. 1981 Robuloides lens Reichel - Altiner, pl. 42, figs. 8–14.
non 1983 Robuloides aff. lens Reichel - Salaj et al., p. 125, pl. 81, fig. 4.
1984 Robuloides aff. lens Reichel - Kotlyar et al., pl. 6, fig. 18.
1985 Robuloides lens Reichel - Pasini, pl. 61, fig. 12. 1985 Colaniella sp. - Trifonova, pl. 4, fig. 7.
v. 1985 Robuloides lens Reichel - Vaslet et al., p. 16 (no illustration).
1988 Robuloides lens Reichel - Loeblich and Tappan, pl. 437, figs. 6, 7.
1989 Robuloides lens Reichel - Köylüoglu and Altiner, pl. 10, figs. 16, 17.
1990 Robuloides lens Reichel - Lin et al., p. 193, pl. 19, figs. 19–21 (with synonymy).
1991 Robuloides lens Reichel - Baud et al., fig. 3 in text p. 193 (no illustration).
1991 Robuloides lens Reichel - Grant et al., tabl. 2 (no illustration).
1993b Robuloides lens Reichel - Vachard et al., p. 97 (no illustration).
1995 Robuloides lens Reichel - Berczi-Makk et al., pl. 9, fig. 5.
1996 Robuloides lens Reichel - Leven and Okay, pl. 9, figs. 29, 30.
1998 Robuloides lens Reichel - Altiner and Özcan-Altiner, pl. 4, fig. 14.
2003 Robuloides lens Reichel - Ünal et al., pl. 1, fig. 45.
2003 Robuloides lens Reichel - Wang and Ueno, table 1 (no illustration).
? 2005 Robuloides lens Reichel - Vaslet et al., p. 120 (no illustration; not seen).
Remarks: Small species, with lozenge-shaped outline, subcarinate, with partly diagenetically? recrystallised wall in the chamber (3 specimens in the same sample).
Dimensions: Diameter = 0.160 mm, Width = 0.090 mm, W/D = 0.56, number of whorls = 2.5 or 3. Occurrence: Late Permian of Tethys. Discovered in the Duhaysan Member (late Dzhulfian) of Saudi Arabia.
Genus: ?lchtyofrondina Vachard in Vachard and Ferrière, 1991,
Type species: Ichtyolaria latilimbataSellier de Civrieux and Dessauvagie, 1965
Ichtyofrondina?sp. (Plate 6.30)
Remarks: Large enveloping chevron-shaped chambers. The unique section is deformed? and also looks like Partisania or Nodosinvolutaria. An Ichtyofrondina latilimbata (de Civrieux and Dessauvagie, 1965), under the name Ichtyolaria, was illustrated by Delfour et al. (1982, fig. 16).
Dimensions: Height = 0.420 mm, Width = 0.250 mm, number of chambers = 7?
Occurrence: Middle-Late Permian of Tethys. Discovered in the Huqayl Member (early Dzhulfian) of Saudi Arabia.
Genus: Lingulinad’Orbigny, 1826,
Type species: Lingulina carinatad’Orbigny, 1826
cf. 1996 Lingulina semivelata Cherdyntsev - Pronina, p. 251, pl. 3, fig. 21.
cf. 1998 Lingulina semivelata Tcerdyncev (sic) - Pronina, p. 168 (no illustration).
Remarks: Subtriangular or horseshoe-shaped chambers. Septa and wall of the same thickness.
Dimensions: Height = 0.770 mm, Width = 0.130 mm, number of chambers = 7, proloculus = 0.070 mm.
Occurrence: Middle Permian of Russia and Zechstein of Baltic region (Pronina-Nestell, personal communication, 2005). Discovered in the Duhaysan Member (late Dzhulfian) of Saudi Arabia.
Genus: ColaniellaLikharev, 1939,
Type species: Pyramis parvaColani, 1924
cf. 1988 Colaniella minuta n. sp. Okimura, p. 719, 721, figs. 6. 1–6. 4.
cf. 1990 Colaniella minuta Okimura - Nakazawa, fig. 1. 10 (no illustration).
v. 2005 Colaniella sp. - Vaslet et al., p. 77, 115 (fig. 34), 116, 117, 118 (fig. 35), 127 (no illustration).
Remarks: Two very small and very primitive specimens presenting some similarities with Colaniella minuta. The genus Colaniella is absent in Zagros, Taurus, Arabia or represented by primitive forms, for instance C. bozkiriÇatal and Dager, 1974 in Turkey. The advanced Colaniella of other Neo-Tethyan areas: Alborz, Transcaucasia, Oman Mountains, Himalaya, Salt Range, central Afghanistan, are unknown in the Taurus-Zagros-Arabia Province.
Dimensions: Height = 0.115–0.130 mm, Width = 0.070–0.085 mm, number of chambers = 4.
Occurrence: Late Permian of Salt Range (horizons 3 and 4 of the Zaluch I section). Discovered in the Duhaysan Member (late Dzhulfian) of Saudi Arabia.
Metazoan fragments are rare except in the maximum flooding deposits or the middle ramps where they are represented by crinoids and bryozoa (Plates 7.1 and 7.2). In the biologically restricted environments (e.g. intertidal, inner ramp), appear the classical biumbilicate gastropods Bellerophon sp. (Plate 7.3). The annelid Spirorbis phlyctaena is the main marker of the Early Triassic (Plates 7.4–7.7).
PALAEOCOLOGY AND PALAEOBIOGEOGRAPHY
The majority of the outcropping Khuff Formation beds include sabkha deposits. More marine deposits are wackestones with bryozoa, or wackestones with lagenids. Due to the evaporitic conditions that prevailed in the inner ramp, the bioherms with calcisponges and corals, the fusulinids, and the biodiversity of the small foraminifers decreased. Consequently the list of fossils in Saudi Arabia is poorer than the list of Greece (Vachard et al., 2003), northwest Caucasus (Pronina-Nestell and Nestell, 2001), or South China (Zhao et al., 1981; Lin et al., 1990; Wang and Ueno, 2003; Zhang and Hong, 2004).
Due to the palaeoecological constraints, there is a good correlation between Saudi Arabia, southern Turkey (Altiner, 1981; Ünal et al., 2003) and southern Iran (Baghbani, 1993; Partoazar, 1995; Gaillot, work in progress). The Khuff Formation is relatively uniform in age, but diachronisms may exist. For example, in the Oman Mountains, a hiatus in the Late Permian occurs between the Shanita levels and the base of the Triassic. Moreover, the presence of: (1) Palaeofusulina sp. and Colaniella ex gr. parva (Colani) in the Musandam Peninsula (Qamar Jbel; Pillevuit, 1993, p. 47); (2) of Colaniella (Glennie et al., 1974, figs. 4.4.2-4.4.3), probably C. parva in the Oman Mountains; and (3) of Robustopachyphloia sp. in Batain Plain (Hauser et al., 2000) confirm the presence of marine Lopingian in Oman.
Turkey, the Zagros Mountains, Saudi Arabia, and parts of Oman form a biogeographic province (Figure 6) dominated by Paradagmarita (Sengör et al., 1988; Altiner et al., 2000; Gaillot and Vachard, work in progress). The stratigraphy of several parts of Oman (for example Al Jabal al Akhdar) are different. The presence of Paradagmarita is questionable, and a hiatus seems to exist between the latest Midian Shanita zone and the Triassic transgression (Vachard et al., 2002). Nevertheless, in many works the Akhdar Group is dated Guadalupian to Lopingian (e.g. Weidlich and Bernecker, 2003). Paradagmarita is rare but present in the other provinces of the world (Altiner et al., 2000; Gaillot, work in progress).
The sea of Taurus-Zagros-Saudi Arabia is considered to have been confined due to some barriers located between the Abadeh region and Zagros in Iran (Vachard et al., 2002). The open sea, rich in fusulinids, was located in the Abadeh area where the biostratigraphical scale with fusulinids is accurately established (Kobayashi and Ishii, 2003a, b). The Abadeh platform might have been located at the margin of the Neo-Tethys Ocean. During the late Permian the Alborz may have constituted another barrier, because of its paucity in Palaeofusulina and advanced Colaniella. In contrast, the Oman Mountains were directly connected with the Neo-Tethys Ocean and contain Colaniella and Palaeofusulina, as well as Transcaucasia (for the palaeogeographical distribution of Palaeofusulina, see Kobayashi, 1999).
Pseudomidiella cf. labensis and Graecodiscus cf. kotlyarae are relatively similar to species of the northwest Caucasus. Compared with this area, Lasiodiscus, advanced Permian lagenids, true Dagmarita, Paraglobivalvulina, Abadehella, Neoendothyra, Urushtenella, Paraglobivalvulinoides are lacking. Colaniella is very poorly represented. Palaeofusulinids are completely absent. This absence is probably related with provincial characters, because they are also missing in Zagros and Turkey, but relatively dispersed in Palaeo-Tethys and Neo-Tethys (Gaillot and Vachard, work in progress). Tetrataxis and Climacammina are also lacking although they are relatively tolerant.
The relationships between the Cimmerian continent (e.g. Sengör, 1979, Kobayashi, 1999; Ueno, 2003 = Extragondwan Domain of Vachard, 1980), the Peri-Gondwanan border in Iran and Turkey, and South China can be easily explained by a model of Pangea B (Morel and Irving, 1981; Erwin, 1992; Besse et al., 1998; Crasquin-Soleau et al., 2001, 2004, 2006).
Regarding the problem of the Triassic/Permian Boundary, several taxa have been cited as displaying a stratigraphic range that transgresses this event and include Cornuspira, Agathammina, Hemigordiellina (= Glomospira of the authors), Glomospirella?, Nodosinelloides and Cryptoseptida?. They are at the origin of all the lineages of Miliolina and Lagenina/Rotaliina of the Mesozoic-Cenozoic. Earlandia? is here the only survivor of the Fusulinida. These taxa are more numerous according to Salaj et al. (1983) and include Rhizammina, Hyperammina, Ammodiscus, “Glomospira”, Glomospirella, Turritellella, Tolypammina, Reophax, Lituotuba, Ammobaculites, Spiroplectammina, Textularia, Caligella, Earlandia, Earlandinita, Nodosinella, Pachyphloia, Geinitzina, Multiseptida, Palaeonubecularia, Tetrataxis, Endothyra, Neoendothyra, Endothyranella, Paraendothyra, Rectoseptaglomospiranella, Haplophragmella, Haplophragmina, Klubovella, Endothyranopsis, Glyphostomella, Agathammina, Calcitornella, Arenovidalina, Nodosaria, Rectoglandulina, Dentalina, Robuloides, Lingulina, Frondicularia, Protonodosaria, Ichtyolaria, Pseudoglandulina and Permodiscus.
We do not agree with the complete list of Salaj; moreover it remarkably indicates that the survivors of the Permian are more numerous in the Triassic, an observation that contradicts the published compilations for these species (e.g. Erwin, 1992; Rampino and Adler, 1998; Benton and Twitchett, 2003).
Due to the abundance and biodiversity of Paradagmarita, Taurus, Zagros, Oman and Saudi Arabia constitute a bioprovince limited by the Neo-Tethys, the Palaeo-Tethys, the seaway to Madagascar and Karoo and the African and Arabian continent (Figure 6). Connections existed with Afghanistan and other sectors of the Cimmerian blocks (according to the geographical distribution of Paradagmarita; Gaillot and Vachard, work in progress). In Afghanistan, a species of the group P. monodi is a nomen nudum: P. dubreuilli (Vachard, 1980), and is known as far as Japan (Iwai-Kanyo area; Chichibu Terrane), with Paradagmarita sp. (Kobayashi, 1997, pl. 4, fig. 19: 2004, fig. 6.47–6.50). Paradagmarita is also known in the Alborz and Djulfa area (northern Iran) (Okimura et al., 1985; Partoazar, 1995). This palaeogeography existed at least since the latest Midian, with the Shanita province (e.g. Sengör et al., 1988; Ueno, 2003).
The base of the Khuff Formation appears to be diachronous, because it is dated as early Wordian in the Haushi-Huqf area (southeastern Oman) (Angiolini et al., 1998, 2004). This part of the Khuff seems to be coeval with the Saiq Formation of the Oman Mountains, because the Khuff Formation is not Lopingian in Al Jabal al Akhdar but limited to the Midian (Vachard et al., 2002). In fact the conodont assemblage of the Haushi-Huqf is similar to that of the Amb Formation of the Salt Range (Angiolini et al., 1998), attributed to the early Midian by Vachard et al. (2002). In this case, the diachronism of the base of Khuff Formation might to be limited to the Midian.
Conversely, the Wordian age attributed to the major Neo-Tethys transgression (Angiolini et al., 2003) is probably underestimated; this transgression (Figure 6) is more probably Capitanian (Midian) in age. According to the fusulinoid assemblages, the Wordian-Murgabian ages deduced for the ancient classifications of Leven used in the Perigondwan (e.g. Lys, 1988) are in fact Capitanian (Midian), due to the re-establishment of the importance of Neoschwagerina schuberti Kochansky-Devidé, N. craticulifera (Schwager), N. margaritae Deprat, compared with the FAD of Yabeina, Lepidolina, Dunbarula, Sumatrina, Codonofusiella and Reichelina (e.g. Leven, 2003).
The recently proposed Deev Jahi model of Heydari and Hassanzadeh (2003), indicating that gas hydrates are the main cause of biological crisis on Earth, is particularly interesting to verify in the Province Neo-Tethys-Taurides-Saudi Arabia-Oman, since the mass extinction started in the ocean (Neo-Tethys) and spread landwards to Saudi Arabia.
The Khuff Formation in the outcrops of central Saudi Arabia is composed of five members, which are assigned to the late Guadalupian Epoch (late Middle Permian, late Capitanian or late Midian Stage), Lopingian Epoch (Late Permian, Wuchiapingian-Changhsingian or Dzhulfian-Dorashamian stages) and Early Triassic ages. Palaeobiogeographically, the regions of the Taurides, Zagros, Oman and Saudi Arabia constitute a bioprovince limited by the Neo-Tethys, the Palaeo-Tethys, the Seaway to Madagascar and Karoo and the Afro-Arabian landmass. Communication existed with Afghanistan and other sectors of the Cimmerian blocks (according to the geographical repartition of Paradagmarita), and as far as Japan.
Algae are represented only by Permocalculus and Mizzia. Smaller foraminifers are numerous, but the fusulinoids are restricted to the primitive genera Eostaffella? and Nankinella. Foraminiferal biomarkers are generally lacking, for example Pseudodunbarula, Paradunbarula, Palaeofusulina or advanced Colaniella. The local markers are similar to those of Taurides and Zagros: Paradagmarita and Septoglobivalvulina?.
A Lazarus effect is observed with the genus Radiosphaera, known respectively in the Late Devonian-Early Carboniferous and in the Late Permian. Earlandia? is diversified. It is confined in hypersaline to normal environments, and can be confused with Aeolisaccus.
The biseriamminids/globivalvulinids are relatively diversified, especially the genus Paradagmarita, with four taxa. The cornuspirids and hemigordiids are also abundant. The hemigordiopsids Hemigordiopsis, Lysites and Kamurana are absent. A new species Glomospirella? linae can be locally common, and can characterise secondarily the local Dorashamian. The lagenids are principally represented by species of genera Nodosinelloides and Polarisella, mainly with P. elabugae and P.? hoae. Taphonomically, the general abrasion of the tests of Nankinella and Paradagmarita “sp. 1” is noticeable.
(1) aff. (Latin affinis) means “with only some relations with”.
(2) cf. (Latin confer) means “almost similar to”.
(3) ex gr. (Latin ex gregae, literally: out of the flock) means “only related with a group of species not to a peculiar species” or in the case of forams in thin sections “with a type of section common to several species”).
(4) p. for Latin pars or partim; to mean in part when only a part of the figure correspond to our understanding of the taxon.
(5) sp. means “indeterminate species”. Consequently, we can denominate a species in the decreasing order of certainty: Hemigordiellina regularis, Hemigordiellina cf. regularis, Hemigordiellina aff. regularis, Hemigordiellina ex gr. regularis or finally, Hemigordiellina sp.
(6) spp. (Latin abbreviation, with the double letter at the end which is the mark of plural), means “several indeterminate species”.
(7) v. for Latin video or vidimus; to mean that the present authors have seen the material in question.
(8) non v. is non video/vidimus; the material was not seen by us.
(9) ? to mean questionable identification.
(10) FAD means ‘First Appearance Datum’ and LAD means ‘Last Appearance Downhole’.
(11) Parentheses (at the author name and date of creation) correspond to a nomenclatural change at the generic level; i.e., Glomospira regularisLipina, 1949 becomes Hemigordiellina regularis (Lipina, 1949).
The authors thank G. Pronina-Nestell and M. Nestell (Arlington University), G. Wyn Hughes (Saudi Aramco) and Michael Stephenson (British Geological Survey) for their constructive remarks. The authors also thank GeoArabia for assisting with the editing and design of the manuscript.
ABOUT THE AUTHORS
Daniel Vachard is a Researcher at the French National Centre of Scientific Research (CNRS). He is with the UMR 8014 of the University of Lille (France). His PhD thesis (1974) was a study of the Palaeozoic of southern France, and his Doctorate of Sciences at Dijon University (1980) concerned the Carboniferous-Permian biostratigraphy of Afghanistan. His work deals mainly with five topics: (1) Fusulinids; (2) Palaeozoic smaller foraminifers; (3)Palaeozoic algae sensu lato (including pseudo-algae and other incertae sedis); (4) carbonate microfacies; and (5) Carboniferous-Permian biostratigraphy and palaeobiogeography. Daniel has contributed to the development of reference scales for the Carboniferous and Permian of numerous areas (e.g. southern France, Afghanistan, Libya, Morocco, Sumatra, Carnic Alps, Oman and Mexico). His interest in the Late Permian of Saudi Arabia began with an unpublished report in 1976 for the French Geological Survey (BRGM).
Jérémie Gaillot received his MSc in Biodiversity of present-day and fossil ecosystems in 2002 from the University of Science and Technology, Lille, France. After his studies, he participated as a Junior Geologist in a synthesis project on the biostratigraphy of the Upper Dalan Formation of Iran. In June 2003, Jéfémie began his PhD on the Biostratigraphy and Palaeoecology of the Khuff Formation of the Arabian Platform at the University of Lille 1, funded by Total. The main focus of his PhD is the sequential foraminiferal biostratigraphy of epeiric platforms and the mechanisms of mass extinction at the Permian-Triassic Boundary.
Denis Vaslet is Head of the Geology and Geoinformation Division at the Bureau de Recherches Géologiques et Minières (BRGM), the French Geological Survey. He has 30 years of experience in the geology of the Middle East. From 1977 to 1979 Denis was involved in geological mapping and phosphate prospecting in Iran for the Geological Survey of Iran and the National Iranian Oil Company. From 1979 to 1991, he was responsible for the Cover Rocks mapping program in Saudi Arabia for the Saudi Arabian Deputy Ministry for Mineral Resources. Denis has been involved in the complete lithostratigraphic revision of the Phanerozoic rocks of central Saudi Arabia, for which he received his Doctorate of Sciences from the University of Paris in 1987. He is currently in charge of geological and geophysical mapping both in France and overseas, and for the production and distribution of digital geological information at BRGM. Denis remains involved in several research projects in the sedimentary geology and stratigraphy fields within the Arabian Peninsula.
Yves-Michel Le Nindre has more than 10 years of experience in the geological mapping of the Phanerozoic rocks of Saudi Arabia. He received his Doctorate of Sciences from the University of Paris in 1987. Yves-Michel’s dissertation was on the sedimentation and geodynamics of Central Arabia from the Permian to the Cretaceous. He is currently working with the Bureau de Recherches Géologiques et Minières on sedimentary basin analysis and modelling, particularly in hydrogeology, and is also involved in present-day littoral modelling.