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New latest Permian foraminifers from Laren (Guangxi Province, South

China): Palaeobiogeographic implications

GAILLOT, Jérémie, et al.

Abstract

Microfacies analyses performed on the latest PermianWujiaping Formation at Laren (Guangxi

Province, South China) show that the bioclasticrich limestones of Late Permian age contain a

rich and well-diversified foraminiferal fauna. This fauna is here revised in order to be

compared with time-equivalent levels of southern Iran and southern Turkey. Some new and

unexpected phylogenetic trends are highlighted among the biseriamminoids. The new or

poorly known genera Retroseptellina, Septoglobivalvulina, Paraglobivalvulinoides,

Dagmarita?, Bidagmarita nov. gen., Louisettita, Paradagmaritopsis nov. gen. and

Paradagmarita? are concerned. Nevertheless, these newly appeared biseriamminoids are

subordinate to abundant Tetrataxis and Climacammina, ultimate survivors of the families

Palaeotextulariidae and Tetrataxidae, appeared as old as the Early Carboniferous

(‘‘Mississippian''). Algae, miliolids, and nodosarioids are poorly represented. Two genera and

four species are here newly described: Globivalvulina curiosa nov. sp., Louisettita ultima nov.

sp., Bidagmarita nov. gen., Bidagmarita sinica nov. gen. nov. sp., [...]

GAILLOT, Jérémie, et al. New latest Permian foraminifers from Laren (Guangxi Province, South

China): Palaeobiogeographic implications. Geobios, 2009, vol. 42, p. 141-168

DOI : 10.1016/j.geobios.2008.02.004

Available at:

http://archive-ouverte.unige.ch/unige:2424

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Geobios Gaillot et al.

1

NEW LATEST PERMIAN FORAMINIFERS FROM LAREN (GUANGXI 1

PROVINCE, SOUTH CHINA): PALAEOBIOGEOGRAPHIC IMPLICA TIONS 2

3

NOUVEAUX FORAMINIFERES DU PERMIEN TERMINAL DE LAREN 4

(PROVINCE DE GUANGXI, CHINE DU SUD) : IMPLICATIONS 5

PALEOBIOGEOGRAPHIQUES 6

7

Jérémie Gaillota,*, Daniel Vachardb, Thomas Galfettic, Rossana Martinid 8 a TOTAL, DGEP/GSR/TG/ISS/BIO, Dpt of Biostratigraphy, CSTJF L1100, 64018 Pau, 9

France. 10 b Université de Lille 1, UFR Sciences Terre, Labo. LP3, UMR 8014 du CNRS, Bât. SN5, 11

59655 Villeneuve d’Ascq, Cédex, France. 12 c Paläontologisches Institut und Museum, Universität Zürich, Karl Schmid-Strasse 4, CH 13

8006 Zürich, Suisse. 14 d Université de Genève, Département de Géologie et Paléontologie, 13, rue des Maraîchers, 15

CH 1211 Genève 4, Suisse. 16

17

18

___________________________________________________________________________ 19

20

Abstract 21

22

Microfacies analyses performed on the latest Permian Wujiaping Formation at Laren 23

(Guangxi Province, South China) show that the bioclastic-rich limestones of Late Permian 24

age contains a rich and well-diversified foraminiferal fauna. This fauna is here revised in 25

order to be compared with time-equivalent levels of southern Iran and southern Turkey. Some 26

new and unexpected phylogenetic trends are highlighted among the biseriamminoids. The 27

new or poorly known genera Retroseptellina, Septoglobivalvulina, Paraglobivalvulinoides, 28

Dagmarita?, Bidagmarita nov. gen., Louisettita, Paradagmaritopsis nov. gen. and 29

Paradagmarita? are concerned. Nevertheless, these newly appeared biseriamminoids are 30

subordinate to abundant Tetrataxis and Climacammina, ultimate survivors of the families 31

Palaeotextulariidae and Tetrataxidae, appeared as old as the Early Carboniferous 32

(“Mississippian”). Algae, miliolids, and nodosarioids are poorly represented. Two genera and 33

four species are here newly described: Globivalvulina curiosa nov. sp., Louisettita ultima nov. 34


2

sp., Bidagmarita nov. gen., Bidagmarita sinica nov. gen. nov. sp., Paradagmaritopsis nov. 35

gen., Paradagmaritopsis kobayashii nov. gen. nov. sp. The palaeogeographic distribution of 36

these foraminifers is interpreted to be typically of Neo-Tethyan regions, ranging from 37

southern Turkey (Hazro) to South China (Laren) and up to Japan for some species (i.e., 38

Paradagmaritopsis). 39

At Laren, Late Permian strata are generally characterized by Reichelina ex gr. simplex Sheng, 40

1956. Isolated samples of packstones, collected in Tsoteng region (Guangxi Province, South 41

China), contain Sphaerulina sp. together with various smaller foraminifers and numerous 42

representatives of the new species Globivalvulina curiosa nov. sp. In this study we 43

demonstrate that the regions of Zagros (Iran), Taurus (Turkey), South China and even Japan 44

shared similar foraminiferal assemblages and represented intermittently connected 45

palaeobiogeographic provinces during Late Permian times. 46

47

Résumé 48

49

Dans la région de Laren (Province du Guangxi, Chine du Sud), des microfaciès calcaires 50

bioclastiques renferment jusqu’à la limite Permo-triasique (PTB), de riches faunes de 51

foraminifères. Elles sont révisées taxonomiquement afin d’être comparées aux éléments des 52

associations du Sud de l’Iran et du Sud de la Turquie. Des tendances évolutives 53

insoupçonnées se révèlent, surtout parmi les bisériamminoïdes, avec des genres nouveaux ou 54

peu connus : Retroseptellina, Septoglobivalvulina, Paraglobivalvulinoides, Bidagmarita nov. 55

gen., Louisettita, Paradagmaritopsis nov. gen. et Paradagmarita? Ces taxons inhabituels sont 56

cependant subordonnés à d’abondants survivants d’autres superfamilles (Palaeotextularioidea 57

and Tetrataxoidea) apparues dès le Carbonifère inférieur (“Mississippien”), tels que 58

Tetrataxis et Climacammina. Algues, miliolides, et nodosarioïdes sont peu représentés. Six 59

taxons sont décrits: Globivalvulina curiosa nov. sp., Louisettita ultima nov. sp., Bidagmarita 60

nov. gen., Bidagmarita sinica nov. gen. nov. sp., Paradagmaritopsis nov. gen., 61

Paradagmaritopsis kobayashii nov. gen. nov. sp. La répartition paléogéographique de ces 62

foraminifères est typiquement néo-thysienne, du Sud de la Turquie (Hazro) au Sud de la 63

Chine, voire jusqu’au Japon avec Paradagmaritopsis. Des échantillons isolés de packstones 64

collectés dans une autre région (Tsoteng, Province du Guangxi, Chine du Sud) contiennent 65

Sphaerulina sp. ainsi que de petits foraminifères variés et de nombreux spécimens d’une 66

espèce nouvelle Globivalvulina curiosa nov. sp. 67


3

Dans cette étude il est démontré qu’au cours du Permien supérieur, le Zagros (Iran), le Taurus 68

(Turquie), la Chine du Sud (bassin de Nanpanjiang) et même le Japon partagent des 69

associations de foraminifères similaires et que ces diverses provinces paléobiogéographiques 70

étaient périodiquement interconnectées. 71

72

Keywords: Foraminifers; Biseriamminoids; Systematics; New taxa; Late Changhsingian; 73

Upper Permian; South China 74

75

Mots clés : Foraminifères ; Bisériamminoïdes ; Systématique ; Nouveaux taxons ; 76

Changhsingien supérieur ; Permien supérieur ; Chine du Sud 77

___________________________________________________________________________ 78

79

* Corresponding author. 80

E-mail address: [email protected] (J. Gaillot) 81

82

1. Introduction 83

Traditionally, Late Permian (Lopingian) microfossils are considered as tenacious 84

survivors, between two severe biotic crises: end-Middle Permian and, then, the worst, end-85

Permian crisis, where the life has almost disappered from the earth. According to Loeblich 86

and Tappan (1987), only 15 fusuline genera in the super-families Schubertelloidea and 87

Staffelloidea persisted within Lopingian times, between the two crises. Additional 23 to 25 88

non-fusuline genera (i.e., Biseriamminoidea, Endothyroidea, Tetrataxidoidea, 89

Palaeotextulariioidea, and Lasiodiscoidea) are known from Wuchiapingian to Changhsingian 90

rocks, respectively. In this time interval, at least 35 genera of lagenids are also recognised. 91

Among them, e.g., Geinitzina, Pachyphloia, Nodosinelloides, Protonodosaria and Robuloides 92

are especially widespread. Dasycladals are very few, less than 10 genera (see compilation in 93

Granier and Grgasovic, 2000). Only bioaccumulations of Permocalculus or Gymnocodium 94

can be emphasized (e.g., Théry et al., 2007). 95

Recently, the knowledge of the Late Permian foraminifers and calcareous algae was 96

improved by different works concerning the Khuff Formation (Late Middle Permian to Early 97

Triassic) in the Middle East (Vaslet et al., 2005; Vachard et al., 2005; Insalaco et al., 2006; 98

Gaillot, 2006; Vachard et al., 2006; Gaillot and Vachard, submitted). Many new taxa are 99

illustrated and/or described within these contributions. 100


4

The Permian-Triassic Khuff Formation, our initial reference, has been defined by Steineke 101

et al. (1958) near the town of Khuff in central Saudi Arabia. This formation, about 190 m 102

thick, is essentially made of carbonates and evaporites, and less of clastics. It rests over the 103

massive sandstones of the Saq Formation upon a major unconformity (the Pre-Khuff 104

Unconformity) and is conformably overlay by the clayely and evaporitic lower Triassic 105

(“Scythian”) Sudair Shale Formation. The Khuff Formation represents a complete Lopingian 106

(Late Permian) succession (Vaslet et al. 2005; Vachard et al. 2005; Gaillot, 2006). 107

The Khuff depositional system developed on the margin of the Neo-Tethyan Ocean, 108

separating the Gondwana supercontinent from the Gondwanan terranes (Cimmerian 109

megablock). The general palaeogeographic context of this system was a marginal marine 110

shelf setting with an inner platform that was very flat, ramp-like, but with local depressions 111

(Sharland et al. 2001; Insalaco et al. 2006). 112

Comprehensive investigations recently undertaken in the Nanpanjiang Basin (South China) 113

revealed a rich and new Late Permian foraminiferal-algal association. The discovery, 114

description and revision of some new and/or stratigraphically and palaeogeographically 115

significant taxa are related, identifying the main markers for the latest Permian in South 116

China. Among the foraminifers, the group of Biseriamminoidea was the most important and 117

highly diversified, because the disappearance of the neoschwagerinid, verbeekinid and 118

schwagerinid fusulines at the end-Capitanian, 119

In South China as well as elsewhere in the Tethys, and especially in western Neo-Tethys, 120

the shallow depositional settings of the Lopingian (including Wuchiapingian and 121

Changhsingian) are generally characterized by abundant Globivalvulinidae (family including 122

Paradagmaritininae and Globivalvulininae). The representatives of this family are considered 123

as available proxies to identify semi-restricted environments. 124

During the Lopingian times, the representatives of the order Lagenida, also present in our 125

associations, have been regarded as indicative of inner to middle neritic environments 126

throughout the Tethyan domain and northern higher latitudes (Gaillot, 2006). The robust 127

biostratigraphic framework based on the detailed description of foraminifers and algae as well 128

as on the distribution of key-taxa allows us to propose new palaeobiogeographic implications 129

for the Late Permian. 130

The first aim of this work is to describe the microfacies and the microfossil content of the 131

latest Permian calcisponges-bearing reef limestones of the Wujiaping Formation from Laren 132

(Guangxi Province, South China). The second topic aims to compare the biotic content from 133

this section, with the “Khuff Formation” sensu lato time equivalents in the Middle East. This 134


5

aspect, together with the terrestrial Cathaysian flora from Iran, suggest direct connections 135

between Zagros (southern Iran) and south China during several epochs, and westward 136

movement of the South China Block. 137

138

139

2. The Nanpanjiang Basin and section location 140

141

The Yangtze carbonate platform of South China constituted a stable palaeogeographic 142

component from Late Proterozoic to the end of Middle Triassic with deposition of shallow-143

water carbonates during most of this time (Enos et al., 1998). Middle and Late Permian reef 144

limestones are widely distributed and well preserved in eastern Yunnan, southwestern 145

Guizhou and northwestern Guangxi (Sheng et al., 1985; Rigby et al., 1989 a, b; Fan et al., 146

1990). 147

The southern margin of the Yangtze platform was embayed by the Nanpanjiang Basin (see 148

Lehrmann et al., 1998 for a description of this basin) which was widely opened and extended 149

into central Guizhou and northwestern Guangxi. Dominant deep-water deposits surrounded 150

various carbonate platforms dispersed within the basin (Fig. 1). The Laren section (NW 151

Guangxi, Fengshan District), the main locality on which this study focuses, lies on the margin 152

of one of these isolated platforms. 153

The Laren profile illustrates an excellent outcrop straddling the Permo-Triassic boundary 154

(Galfetti et al., 2007). The Upper Permian beds, mainly characterized by shallow-water reef 155

limestones, are of about 20 m-thick in the measured section. They are drastically overlain by a 156

~7.5 m-thick unit of calcimicrobial limestones of earliest Triassic age (Griesbachian) (Fig. 2). 157

The very rich and biostratigraphically significant foraminiferal assemblage found within 158

the beds just before the Permian-Triassic boundary allows attributing a late Changhsingian 159

age, while the precise age of the microbial limestone immediately above the boundary is 160

indeterminable because of nearly total absence of index fossils. 161

In order to compare the foraminiferal associations of Laren with those of coeval 162

limestones, we also collected two samples in the Tsoteng area, located ~80 km south of Laren 163

(Fig. 1). At this locality the uppermost Permian Heshan Formation is mainly composed of 164

massive and thick-bedded bioclastic-rich limestones deposited in shallow-water settings. 165

166

3. Facies and microfacies 167

168


6

Late Permian rocks are generally very massive, dark-coloured and frequently accompanied 169

by centimetric to decimetric black chert concretions. At Laren these beds display an 170

extraordinary, well preserved faunal assemblage (e.g., brachiopods, gastropods, rugose corals, 171

very large calcareous algae and calcareous porifera) (Fig. 3). Geometrical orientation of 172

fossils has not been observed excepting for a sphinctozoan colony which illustrates a planar 173

setting (Fig. 3a). 174

Microfacies analysis on thin sections (Fig. 4) revealed that these skeletal, peloidal grainstones 175

are mainly associated with: rare gastropods, rare incertae sedis Tubiphytes (non Shamovella) 176

obscurus Maslov, 1956, Ramovsia? sp., Donezella sp., Radiosphaerina? sp., Asterosphaera 177

cf. pulchra Reitlinger 1957 and Asterosphaera sp., rare fusulinids Nankinella cf. inflata 178

(Colani, 1924) emend. Sheng, 1963 and Reichelina simplex Sheng, 1956, in addition to 179

abundant and diversified smaller foraminifers, especially biseriamminids. The following taxa 180

are recognized: Eotuberitina spinosa (Lys in Lys et al., 1980); Postendothyra micula (Sosnina 181

in Sosnina and Nikitina, 1977); Globivalvulina curiosa Gaillot and Vachard nov. sp.; G. 182

kantharensis Reichel, 1946; G. vonderschmitti Reichel, 1946; G. bulloides (Brady, 1876); 183

Septoglobivalvulina distensa (Wang in Zhao et al., 1981); S. cf. guangxiensis Lin, 1978; S. sp. 184

1; Dagmarita chanakchiensis Reitlinger 1965; D. altilis Wang in Zhao et al., 1981; D.? cf. 185

sharezaensis Mohtat-Aghai and Vachard, 2003; D. simplex Wang in Zhao et al., 1981; 186

Bidagmarita sinica nov. gen. nov. sp. (with pseudo-fibrous inner layer); Siphodagmarita 187

vasleti Gaillot and Vachard in Gaillot et al. (submitted); Louisettita ultima Gaillot and 188

Vachard in Gaillot et al. (submitted); Retroseptellina decrouezae (Köylüoglu and Altiner, 189

1989); Retroseptellina nitida (Lin, Li and Sun, 1990); Paraglobivalvulinoides septulifer 190

(Zaninetti and Altiner, 1981); Paradagmaritopsis kobayashii nov. gen. nov. sp. Gaillot and 191

Vachard; Paradagmaritopsis sp. and Paradagmarita sp. They are associated with relatively 192

rare nodosarioids, such as Rectostipulina sp., Nodosinelloides sp., Geinitzina sp., 193

Pseudotristix cf. solida Reitlinger, 1965, Pachyphloia pedicula Lange, 1925, P. ex gr. ovata 194

Lange, 1925, and Ichthyofrondina palmata (Wang, 1974). Miliolids are very rare as well as 195

Pseudovermiporella nipponica (Endo in Endo and Kanuma, 1954). Phylogenetically older 196

forms (first appeared as old as early to late Mississippian), such as Climacammina (C. tenuis 197

Lin, 1978) and Tetrataxis (T. lata Spandel, 1901), are relatively common (Fig. 4). This fact 198

seems to be common in the late Changhsingian (Gaillot, 2006). 199

Isolated sample T0 (Tsoteng area) is a peloidal, sandy and finely bioclastic, neosparitized 200

wackestone with Reichelina simplex, Pachyphloia sp., Dagmarita sp. and Asterosphaera sp. 201

A relatively large lithoclast of bioclastic rudstone with Sphaerulina, Nankinella, Neodiscus, 202


7

Permocalculus, Mizzia, Rectostipulina quadrata, Pachyphloia, Nodosinelloides is also present 203

in this microfacies. 204

A second isolated sample T1 (Tsoteng area) is represented by dolomitized, pyritized 205

wackestone displaying clotted structures in microfacies and yielding rare ostracodes, very rare 206

Frondina and relatively abundant Globivalvulina curiosa nov. sp. 207

208

4. Discussion 209

210

Middle and Late Permian reefs are widely distributed in north-western Guangxi Province, 211

where they are part of a carbonate platform-margin reef belt (Rigby et al. 1989 a, b). 212

According to these authors, Inozoans (unsegmented calcareous sponges with skeletons) and 213

other sponges (sphinctozoans) are major frame-builders in the Permian reefs of South China. 214

Although the biodiversity of the sphinctozoans is generally poor, smaller foraminifers are 215

highly variable taxonomically in these reefs. Biseriamminids are especially diversified with 216

the genera and subgenera Globivalvulina, Dagmarita, Septoglobivavulina, Louisettita, 217

Siphodagmarita, Paradagmarita?, Paraglobivalvulinoides and Paradagmaritopsis nov. gen. 218

This diversity was already underlined within the Chinese literature (e.g., Lin et al., 1990). 219

The late Changhsinghian age is demonstrated by the following association: Retroseptellina 220

decrouezae, R. nitida, Siphodagmarita, Paradagmaritopsis, Paradagmarita, Louisettita and 221

Paraglobivalvulinoides. Even though no marker fossils of the latest Changhsingian (e.g., 222

Gallowainella, advanced Colaniella, Geinitzinita, Robustopachyphloia, Glomomidiellopsis, 223

Paradagmarita monody) were distinguished, bloom of Robuloides (see Gaillot, 2006) was 224

encountered in this fauna. 225

The abrupt transition from latest Permian foraminifer-rich grainstones with sophisticated 226

endoskeletons, to earliest Triassic calcimicrobial limestones yielding very simple and 227

undivided foraminifers, is known from Turkey to South China (see review in Groves and 228

Altiner, 2005). Nevertheless, late Permian reefs, including the Late Changhsingian ones, are 229

very rare in these Neo- and Palaeo-Tethyan Provinces. Exceptional examples were reported 230

from Greece, South China (Flügel and Reinhardt, 1989; Weidlich, 2002; Weidlich et al., 231

2003; this study), and NW Caucasus (Théry et al., 2007) where the sphinctozoan and inozoan 232

bioconstructions are especially well-known (Rigby et al., 1989 a, b; Fan et al., 1990). 233

234

5. Taxonomy 235


8

Key of lecture: p. = pars; ? = questionable; v. = video or vidimus (from Latin, means that 236

the material was in person see by the authors). 237

238

Class ?FORAMINIFEREA 239

Family TUBERITINIDAE Miklukho-Maklay, 1958 240

Remarks: The Tuberitinidae have two stages of a cycle of life, diplospherine, free, and 241

eotuberitine, attached) (Conil et al., 1977); unknown among the truly characterized 242

Foraminifera, associated with the absence of true foraminiferal apertures, can indicate another 243

group of Protozoa or Protophyta. A relationship might exist with some calcispherids (see 244

Vachard, 1994). 245

246

Genus Eotuberitina Miklukho-Maklay, 1958 247

Type species: Eotuberitina reitlingerae Miklukho-Maklay, 1958 248

Diagnosis: Small Tuberitinidae with a flat to convex basal disc and a thin microgranular 249

wall. Groups of 2-3 specimens exist but are rare. 250

Occurrence: Silurian-Latest Permian (Vdovenko et al., 1993), cosmopolite. The presence 251

of Eotuberitina reitlingerae up to the Changhsingian was previously indicated (Lin et al., 252

1990). 253

254

Eotuberitina spinosa (Lys in Lys et al., 1980) 255

Fig. 5(7) 256

1977 Tuberitina collosa Reitlinger forma spinosa - Lys in Montenat et al., p. 244, pl. 1, 257

fig. 1 (nomen invalidum). 258

1980 Tuberitina collosa spinosa n. subsp. Lys in Lys et al., p. 81, pl. 1, fig. 11. 259

v. p. 1980 Diplosphaerina ex gr. inaequalis (Derville) - Vachard, p. 411-412, pl. 25, fig. 260

13. 261

1986 Eotuberitina reitlingerae Miklukho-Maklay - Fontaine et al., pl. 22, fig. 10. 262

1989 Altjussella grandis nov. sp. Pronina in Kotlyar et al., p. 93, pl. 3, fig. 19. 263

v. 1990 Diplosphaerina spinosa (Lys in Lys et al.) - Vachard and Miconnet, p. 302, pl. 2, 264

fig. 3. 265

1995 Tuberitina collosa spinosa Reit. (sic) - Partoazar, pl. 12, figs 5-6. 266

1996 Altjussella grandis Pronina - Pronina, pl. 1, fig. 2; pl. 2, fig. 1. 267

1999 Altjussella grandis Pronina - Kotlyar et al. p. 311 (no illustration). 268

2004 Altjussella grandis Pronina - Dronov, p. 792 (no illustration). 269


9

v. 2006 Eotuberitina spinosa (Lys in Lys et al.) - Gaillot, p. 36, pl. I.4, fig. 4, pl. I.10, fig. 270

2, pl. I.15, fig. 13, pl. I.17, fig. 14, pl. I.18, fig. 10, pl. III.3, figs 8-9, pl. III.5, fig. 7, pl. 271

VII.1, fig. 7. 272

Description: Test ampulliform with numerous small radial spines all around the periphery 273

of the hemispherical chamber. Attachment face as in the other tuberitinids. The eotuberine 274

type is common but diplospherine stages exist (e.g., Vachard, 1980, pl. 25, fig. 13; this study). 275

Wall dark, microganular, finely porous. The chamber is regularly hemispherical. The 276

attachment disk is limited to the base of the chamber. Spines are short, radial and regularly 277

spaced. 278

Discussion: This species was successively attributed to Tuberitina, Diplosphaerina and 279

Altjussella. It differs from Tuberitina by the spinose thin wall and the growth limited to a 280

single chamber, from Eotuberitina reitlingerae by the spinosity, and from Altjussella by 281

another type of wall (see the comparative illustrations of the quoted genera in Vachard, 1994, 282

fig. 8, p. 15). Based on our observations, the wall is microstructurally similar to this of 283

Eotuberitina. 284

Dimensions: Over-all diameter = 0.140-0.460 mm (0.075-0.175 mm), inner diameter = 285

0.120-0.445 mm (0.065-0.140 mm), wall thickness = 0.005-0.015 mm, height of spines = 286

0.004-0.030 mm (0.003-0.035 mm). 287

Occurrence: Midian-Lopingian. Southern Italy (Monte Facito; Vachard and Miconnet, 288

1990); Crimea (Kotlyar et al., 1999); Transcaucasia (Kotlyar et al., 1989; Pronina, 1996); NW 289

Caucasus (Pronina-Nestell and Nestell, 2001); Oman (Jebel Akhdar; Lys in Montenat et al., 290

1977); Zagros (Lys et al., 1980; this study); Alborz (Partoazar, 1995); Afghanistan (Vachard, 291

1980); Central Pamir (Dronov, 2004), and Lamayuru Block of Ladakh Himalaya (Lys et al., 292

1980); discovered in South China (this work). Therefore, E. spinosa is relatively characteristic 293

of a western to central Tethyan province, but is present in both Palaeo-Tethys and Neo-294

Tethys. 295

296

Class FORAMINIFEREA d’Orbigny, 1826 297

orth. mut. d’Eichwald, 1830 nomen translat. Lee, 1990 298

Order FUSULINIDA Fursenko, 1958 299

Suborder FUSULININA Fursenko, 1958 nomen translat. Bogush, 1985 300

Superfamily BRADYINOIDEA Reitlinger, 1950 301

nomen translat. Rauzer-Chernousova et al., 1996 (ex family) 302

Family BRADYINIDAE Reitlinger, 1950 nomen translat. Reitlinger, 1958 (ex subfamily) 303


10

Subfamily BRADYININAE Reitlinger, 1950 304

Genus Postendothyra Lin, 1984 305

Type species: Postendothyra scabra Lin, 1984. 306

Synonyms: Endothyranopsis Cummings, 1955 (pars), Bradyina von Möller, 1878 (pars). 307

Diagnosis: Small test organized as a Bradyina (planispiral, involute, nautiloid), but with 308

very finely alveolar wall and pre and post- septal lamellae faint to absent. Wall microgranular. 309

Aperture terminal simple. 310

Composition: Numerous species (but with probably many synonyms). 311

Occurrence: Midian-Changhsingian. South China (Hubei; Lin, 1984), Italy (Vachard and 312

Miconnet, 1990), Greece (Vachard et al., 1993b), Slovenia (Flügel et al., 1984), Crimea 313

(Pronina and Nestell, 1997), Cyprus (Nestell and Pronina, 1997), Turkey (Zaninetti et al., 314

1981), Afghanistan (Vachard, unpublished data), Primorye (Sosnina and Nikitina, 1977), 315

Malaysia (Fontaine et al., 1994), NW Thailand (Fontaine et al., 1993), East Thailand 316

(Fontaine et al., 1997), Cambodia (Nguyen Duc Tien, 1986b), Transcaucasia (Pronina, 1988), 317

Japan (Kobayashi, 1986 ; 1997c), Oman (Vachard et al., 2002), and Himalaya (Lys et al., 318

1980). 319

320

Postendothyra micula (Sosnina in Sosnina and Nikitina, 1977) 321

Fig. 5(3-5) 322

1977 Bradyina micula nov. sp. Sosnina in Sosnina and Nikitina, p. 46-47, pl. 1, fig. 17-18. 323

1978 Endothyranopsis guangxiensis nov. sp. Lin, p. 34, pl. 7, figs 1-3. 324

1981 Bradyina guangxiensis (Lin) - Zhao et al., pl. 1, figs 13-15. 325

p. 1986b Globivalvulina cyprica Reichel - Nguyen Duc Tien, pl. 13, figs 16?, 17-18 (non 326

figs 14-15 = Globivalvulinidae). 327

1987 Paraglobivalvulina recristallisée? - Panzanelli-Fratoni, pl. 6, fig. 9. 328

1988 Bradyina guangxiensis (Lin) - Pronina, pl. 2, figs 2-3. 329

1989 Bradyina guangxiensis (Lin) - Pronina, pl. 1, figs 2-3. 330

1989 Bradyina micula Sosnina - Kotlyar et al., pl. 1, fig. 9. 331

1990 Postendothyra guangxiensis (Lin) - Lin et al., p. 191, pl. 18, figs 21-26. 332

v. 1990 Bradyina micula Sosnina - Vachard and Miconnet, p. 303, pl. 2, fig. 6. 333

v. 1993a Bradyina micula Sosnina - Vachard et al., pl. 4, fig. 1. 334

p. v. 1994 Bradyina sp. - Fontaine et al., pl. 18, fig. 6 (no fig. 4 = Neoendothyra, nor fig. 5 335

= Endoteba). 336

1996 Bradyina? sp. - Leven and Okay, pl. 8, fig. 24; pl. 9, fig. 36. 337


11

1997 Postendothyra micula (Sosnina) - Pronina and Nestell, pl. 1, fig. 7. 338

2001 Postendothyra guangxiensis (Lin) - Pronina-Nestell and Nestell, pl. 4, fig. 15. 339

v. 2006 Postendothyra micula (Sosnina in Sosnina and Nikitina) - Gaillot, p. 41-42, pl. 340

VII.1, figs 3-5. 341

Description: We synonymize Postendothyra micula with P. guangxiensis. Our Fig. 5(3) is 342

typical of the former species, and the Fig. 5(4) from the second one. The parameters of both 343

species are similar (diameter of P. micula = 0.330-0.350 mm, diameter of P. guangxiensis = 344

0.250-0.450 mm). Measurements of our material are: Diameter = 0.360-0.440 mm, width = 345

0.260-0.280 mm, w/D = 0.59-0.76, number of whorls: 2-2.5, number of chambers at the last 346

whorl: 5, proloculus diameter = 0.020 mm, height of last chamber = 0.120-0.160 mm, wall 347

thickness = 0.006 mm. 348

Occurrence: As for the genus. 349

350

Superfamily PALAEOTEXTULARIOIDEA Galloway, 1933 351

nomen translat. Habeeb, 1979 (ex family) 352

Family PALAEOTEXTULARIIDAE Galloway, 1933 353

nomen translat. Wedekind, 1937 (ex subfamily) 354

Subfamily PALAEOTEXTULARIINAE Galloway, 1933 355

Genus Climacammina Brady in Etheridge, 1873 emend. Cummings, 1956 356

Type species: Textularia antiqua Brady in Young and Armstrong, 1871. 357

Diagnosis: Palaeotextulariinae (i.e., Palaeotextulariidae with bilayered wall) with biseriate 358

initial chambers with basal and single apertures, followed by several uniseriate and cribrate 359

chambers. 360

Occurrence: Late Brigantian-Late Changhsingian, cosmopolite since the Pennsylvanian 361

(Mamet, 1970, p. 33). 362

363

Climacammina tenuis Lin, 1978 364

Fig. 5(2) 365

1978 Climacammina tenuis nov. sp. Lin, p. 20, pl. 23, fig. 9. 366

1984 Climacammina tenuis Lin - Lin, p. 122, pl. 2, fig. 29. 367

1990 Climacammina tenuis Lin, Li and Sun, p. 144, pl. 7, figs 11-13. 368

1992 Climacammina tenuis Lin - Ueno, fig. 10: 16-17. 369


12

non 1995 Climacammina tenuis Lin - Ueno et al., fig. 3: 6 (other species, maybe C. 370

sphaerica Potievskaya). 371

v. 2006 Climacammina tenuis Lin - Gaillot, p. 55, pl. VII.1, fig. 2. 372

Description: The species is characterized by few biseriate chambers and numerous 373

uniseriate chambers, relatively wide and low with many cribrate apertures. It differs from C. 374

valvulinoides Lange, which is the species generally cited in the Changhsingian, by a smaller 375

test and from C. major Morozova sensu Bozorgnia, 1973, by more uniseriate chambers, and a 376

more tapering test. 377

Dimensions: The figured specimen of South China measures: Height = 2.200 mm, width = 378

1.000 mm, w/H ratio = 0.55, proloculus diameter = 0.160 mm, number of biseriate chambers: 379

3 pairs, number of uniseriate chambers: 8, height of last chamber = 0.240 mm, wall thickness 380

= 0.020 mm. 381

Occurrence: Early-Middle Permian of South China; Murgabian (= Wordian) of NE Japan 382

(Ueno, 1992). 383

384

Superfamily TETRATAXOIDEA Galloway, 1933 385

nomen translat. Haynes, 1981 (ex subfamily) 386

Family TETRATAXIDAE Galloway, 1933 387

nomen translat. Pokorny, 1958 (ex subfamily) 388

Synonyms: Pseudotaxidae Mamet, 1974, Valvulinellidae Loeblich and Tappan, 1984, 389

Abadehellidae Loeblich and Tappan, 1984, Endotaxidae Bogush and Brazhnikova in Rauzer-390

Chernoussova et al., 1996. 391

Diagnosis: Unique family of trochospiral Fusulinida. Conical shape, with eventual 392

diversely shaped terminal appendices. Four or five chambers per whorl. Undivided chambers 393

or more or less complex partitions. Wall unilayered dark, or bilayered with a brown inner 394

layer and a yellow outer layer. Aperture terminal umbilical. 395

Remarks: These families are synonyms, because all include rare specimens with a 396

particular type of attachments (the "microaquariums" of Vachard and Krainer, 2001). 397

Therefore, only one family of Fusulinida exhibits a trochospiral coiling during the 398

Carboniferous-Permian. 399

Occurrence: Late Tournaisian-Late Changhsingian; cosmopolite. 400

401

Genus Tetrataxis Ehrenberg, 1854 402

Type species: Tetrataxis conica Ehrenberg, 1854. 403


13

Diagnosis: Test medium to large, trochospirally coiled, with four chambers per whorl. 404

Wall bilayered, granular and pseudofibrous. 405

Remarks: An important problem is the life position of Tetrataxis: i.e., attached, not 406

attached, periodically free or attached, with or without interposition of a microaquarium 407

between the test and the substrate (Kochansky-Devidé, 1970, pl. 2, fig. 8; Toomey et al., 408

1977, fig. 8P; Poncet, 1982; Cossey and Mundy, 1990; Ueno and Igo, 1993, fig. 3. 23; 409

Mundy, 1994, text-fig. 12 p. 722; Gallagher, 1998; Vachard and Krainer 2001; Krainer et al., 410

2003a, pl. 3, fig. 38, pl. 4, fig. 17; Saïd, 2005, fig. XII.5), and the significance of the terminal 411

appendices of “Globotetrataxis” Brazhnikova in Aizenberg et al., 1983 and species like 412

Tetrataxis mira Conil and Lys, 1964 or T. fortis Bogush and Yuferev, 1976 [compare with 413

Adachi, 1985, pl. 13, figs 9-11; or Globotetrataxis sp. E (Matsusue, 1992, fig. 3. 27); 414

Tetrataxis sp. of Ueno and Igo (1993, fig. 3. 22) and Ueno et al. (1994, fig. 4. 12)]. 415

Occurrence: Late Ivorian (MFZ6) to latest Permian (Gaillot, 2006); cosmopolite. 416

417

Tetrataxis lata Spandel, 1901 418

Fig. 5(8) 419

1901 Tetrataxis conica Ehrenberg var. lata n. var. Spandel, p. 186, fig. 6 a, b. 420

1949 Tetrataxis lata Spandel - Morozova, p. 261-262, pl. 2, figs 20, 23, 28. 421

non 1961 Pseudotetrataxis conica var. lata - Deleau and Marie, p. 93, pl. 11, figs 5, 8 422

(another species). 423

1982 Tetraxis conica lata Spandel - Hao and Lin, pl. 2, fig. 7. 424

1986 Tetrataxis lata Spandel Zolotova et al. in Gorskii and Kalmykova, pl. 22, figs 10 a, b. 425

1990 Tetrataxis lata Spandel - Lin et al., p. 156, pl. 9, figs 4-5. 426

1999 Tetrataxis lata Spandel - Groves and Boardman, pl. 1, figs 7-9. 427

v. 2006 Tetrataxis ex gr. conica Ehrenberg - Insalaco et al., pl. 2, fig. 4. 428

2006c Tetrataxis sp. - Kobayashi, pl. 3, figs 15-16. 429

2006c Tetrataxis sp. - Kobayashi, pl. 1, fig. 42. 430

v. 2006 Tetrataxis lata Spandel - Gaillot, p. 57, pl. I.4, fig. 3, pl. I.5, figs 2, 14, pl. II.3, fig. 431

6, pl. III.22, fig. 4, pl. VII.1, fig. 8. 432

Discussion: The average dimensions of our specimens are: Height = 0.640 mm, diameter = 433

1.070 mm, ratio height/diameter = 0.60, number of chambers: 5, apical angle = 90, last 434

thickness = 0.70 mm. Consequently, it belongs to a great species with few whorls of the group 435

T. conica Ehrenberg, 1854 (angle of 90°, ratio of 0.50-0.60), but true T. conica is very smaller 436


14

for the same number of whorls; more precisely, according to its parameters, the specimens are 437

rather attributed to Tetrataxis lata Spandel, 1901. 438

Remarks: This species has two senior homonyms: Tetrataxis lata Potievskaya, 1958 (not 439

yet re-named) and T. lata Bogush and Juferev, 1962 (re-named T. bogushi Ueno and 440

Nakazawa, 1993). 441

Occurrence: Early Permian of Kansas, Urals, North China. Kazanian of Russia. Common 442

in the Late Changhsingian strata of South China (this work). In the Zagros (Iran), Tetrataxis 443

lata is one of the diagnostic fossil of the Wuchiapingian transgression (Gaillot, 2006). 444

445

Superfamily BISERIAMMINOIDEA Chernysheva, 1941 446

nomen translat. Marfenkova, 1991 (ex family) 447

Diagnosis: A superfamily of Fusulinida Endothyrina characterized by a sligthly 448

trochospiral to planispiral biseriate coiling, eventually uncoiled. Wall very variable, but 449

without clearly established generic criteria. Aperture single terminal basal. 450

Composition: 2 families: Biseriamminidae Chernysheva, 1941 (including Koktjubinidae 451

Marfenkova, 1991); Globivalvulinidae Reitlinger, 1950 nomen translat. herein. 452

Remarks: The limits of the genera and families and generally poorly established. 453

Biseriammina is especially poorly illustrated. 454

Occurrence: Tournaisian-latest Permian; cosmopolite. 455

456

Family GLOBIVALVULINIDAE Reitlinger, 1950 nomen translat. herein 457

(pro subfamily) 458

Subfamily GLOBIVALVULININAE Reitlinger, 1950 (sic Globivalvulinae) 459

Genus Globivalvulina Schubert, 1921 460

Type species: Valvulina bulloides Brady, 1876. 461

Diagnosis: Small, medium or large globivalvulinid test, entirely biseriate and planispiral. 462

Subglobular with lobate periphery. Valvular projection well developed. Wall black, 463

microgranular to differentiated: with a yellow pseudo-fibrous inner layer (see G. mosquensis 464

Reitlinger, 1950), Omphalotis-like (Vachard and Beckary, 1991), granular with agglutinate 465

particules: G. granulosa Reitlinger, 1950, with intermediary clear layer (“diaphanotheca” of 466

authors): G. bulloides (“Brady” of the authors). Aperture simple protected by the valvular 467

projection. 468

Remarks: Globivalvulina is composed of several groups of species according to generally 469

admitted foraminiferal criteria: wall structure, size, development of the valvulae, characters of 470


15

the chambers, and shape of the apertural face. Unfortunately, these characters appear very 471

variable among the specimens of the same population (Vachard et al., 2006). 472

Occurrence. Serpukhovian to latest Permian; cosmopolite. 473

474

Globivalvulina bulloides (Brady, 1876) 475

Fig. 6(1) 476

1876 Valvulina bulloides nov. sp. Brady, p. 89-90, pl. 4, figs 12-15. 477

1981 Globivalvulina bulloides (Brady) - Zhao et al., pl. 2, figs 4-5. 478

1990 Globivalvulina bulloides (Brady) - Lin et al., p. 86, p. 162, pl. 11, figs 3-1 (with 5 479

references in synonymy). 480

1998 Globivalvulina bulloides (Brady) - Pinard and Mamet, p. 118-119, pl. 27, figs 16-20; 481

pl. 28, figs 1-4, 5?, 6-11 (with 30 references in synonymy). 482

2005 Globivalvulina bulloides (Brady) - Brenckle, p. 47, pl. 7, fig. 18-22 (lectotype 483

designated). 484

p. 2006b Globivalvulina sp. A - Kobayashi, fig. 3.28, 30, 32-34 (non fig. 3.29, another 485

species, nor fig. 3.31, 35-36 = Retroseptellina decrouezae, see below). 486

v. 2006 Globivalvulina bulloides (Brady) - Gaillot, p. 59-60, pl. I.3, figs 1, 5, 7-8; pl. I.37, 487

figs 2-3; pl. I.43, fig. 12; pl. III.23, fig. 5; pl. IV.2, fig. 13; pl. V.3, fig. 1; pl. VI.4, figs 3, 7, 488

15. 489

v. 2006 Globivalvulina bulloides (Brady) - Vachard et al., p. 472, fig. 5. 17, fig. 6. 18-20, 490

fig. 7. 9?, 13? 491

Discussion: The type-species of the genus is proportionally medium-sized. The growth is 492

regular and the valvular projection moderate. The wall is suggested as “diaphanothecal” by 493

many authors (e.g., Pinard and Mamet, 1998; Brenckle, 2005), but according to our 494

observations, from the Bashkirian to Changhsingian times, this clear intermediary layer is not 495

always present in all the specimens of a same population. Although admitted by almost all the 496

authors, this unique criterion chosen to separate “Biseriella” and Globivalvulina must be 497

seriously re-examined. Diameter = 0.400 mm, number of whorl: 1.5, number total of 498

chambers: 5, proloculus diameter = 0.025 mm, height of last chamber = 0.115 mm, wall 499

thickness = 0.015 mm. 500

Occurrence: Early Pennsylvanian to latest Permian; cosmopolite. 501

502

Globivalvulina curiosa Gaillot and Vachard nov. sp. 503

Fig. 7(16-20) 504


16

v. 2006 Globivalvulina curiosa Gaillot and Vachard nov. sp. Gaillot, p. 63-64, pl. VII.3, 505

figs 16-20 (nomen nudum because introduced in a Ph.D.). 506

Etymology: From Latin curiosus = curious, because of the puzzling shapes of tests. 507

Type locality: Tsoteng (Guangxi, South China). 508

Type level: Late Changhsingian. Base of thrombolites, lithostratigraphically assigned to 509

the basis of Triassic (investigation of conodonts in progress). 510

Diagnosis: A Globivalvulina without any regularity neither in the coiling nor type of 511

chambers. 512

Description: (1) Proloculus: ovoid, regular in shape and location, measuring 0.011-0.033 513

mm of small axis and 0.033-0.055 mm of large axis. (2) Coiling planispiral, relatively regular 514

(Fig. 7 (16)) or with a tendency to uncoiling (Fig. 7(21)). (3) Shape of chambers: very 515

irregular (Fig. 7(16-17)), hemispherical to semi-ovoid, as in the tests affected today by marine 516

pollutions (e.g., Debenay et al., 1996; Armynot du Châtelet et al., 2004). Sutures are generally 517

deep with a lobate profile (Fig. 7(17)) or not (Fig. 7( 21)). The shape of chambers is never 518

regular; abnormal chambers are located at the end of the coiling (Fig. 7(21)), at the beginning 519

of the terminal whorl (Fig. 7(16)), or in the entire terminal whorl (Fig. 7(17)). (4) Height of 520

last chamber: relatively constant although the variations in shape: 0.155 mm (Fig. 7(16)), 521

0.244 mm (Fig. 7(17)), 0.166 mm (Fig. 7(20)) (5) Axial section (Fig. 7(20)) and oblique 522

sections (Fig. 7(18-19)) are relatively similar to that of the other Globivalvulina; nevertheless, 523

the tapering and the sutures of these sections are noticeable. (6) Endoskeleton composed of an 524

oral tongue relatively developed and incurved with a “modern style” profile (Fig. 7(20)). 525

Septa are (locally?) prolonged by small expansions thinner and more undulate (Fig. 7(17-18)). 526

Some septa are curvated backwards (Fig. 7(16, 21)). Rare calcified globules or loops are 527

visible in the last chamber of the most deformed specimen (Fig. 7(17)). (7) The wall is 528

microgranular, dark unilayered, medium-thick (0.006-0.013 mm) (theoretically these 529

specimens could be attributed to Biseriella). (8) Aperture terminal at the base of the last 530

chamber, wide and depressed (Fig. 7(20)). 531

Holotype: Fig. 7(16) (sample T1). 532

Type material: Twenty specimens in one thin-section. 533

Repository of the type: Natural History Museum, Geneva (Switzerland), holotype number 534

MHNG 07-001. 535

Comparison: According to C. Jenny-Deshusses (pers. comm. to D. Vachard. and in press) 536

“crazy globivalvulines” exist in the Lopingian of Iran (Alborz). They migth be related to this 537


17

species. All the “normal” globivalvulines of the literature are different, as well as the 538

Biseriella. 539

Remarks: Due to the Permian-Triassic transition, these forms could be related with the 540

different observed geochemical shifts (hypothesis shared with C. Jenny-Deshusses and S. 541

Richoz). 542

Occurrence: late Changhsingian? (South China, this work). 543

544

Genus Septoglobivalvulina Lin, 1978 emend. Gaillot and Vachard in Gaillot et al. 545

(submitted) 546

Type species: Septoglobivalvulina guangxiensis Lin, 1978. 547

Diagnosis: Test similar to Globivalvulina but differing by an ovoid to subspherical shape 548

and a voluminous last chamber covering entirely the preceeding whorls. Endoskeleton as in 549

Globivalvulina, i.e., reduced to an oral tongue. Wall thin dark microgranular unilayered. 550

Composition: Septoglobivalvulina guangxiensis Lin, 1978 (D = 0.820-0.920 mm), S. 551

similis Lin, Li and Sun, 1990 (D = 0.740-0.870 mm), Globivalvulina distensa Wang in Zhao 552

et al., 1981 (D = 0.500-0.700 mm); Septoglobivalvulina cf. globosa sensu Vachard et al., 553

2002 (pl. 1, figs 13-14) non Wang in Zhao et al. 1981 (D = 0.660-0.800 mm). S. sp. 1 of this 554

study (D = 0.555 mm); ?Globivalvulina cyprica sensu Nguyen Duc Tien, 1986b, pl. 6, fig. 14 555

only (non Reichel, 1946). 556

Comparison: Differs from Globivalvulina by the increasing in height and width of the last 557

chamber becoming enveloping of the preceding coiled chambers, from Paraglobivalvulina by 558

the shape not completely spherical, the thin wall and the more rudimentary endoskeleton, and 559

from Retroseptellina Gaillot and Vachard in Gaillot et al. (submitted; nomen nudum) by the 560

shape of the septa plane and not curvated backward, and the more marked increasing in height 561

of the last chamber whereas the last chambers of Retroseptellina are wider than the 562

preceeding ones. Unlike Ünal et al. (2003, pl. 1, figs 9-11), which re-assigned 563

Paraglobivalvulina gracilis Zaninetti and Altiner, 1981 to Septoglobivalvulina, we prefer 564

maintain this species in Paraglobivalvulina (Gaillot, 2006; Gaillot et al., submitted). 565

Occurrence: Early?-late Midian to Changhsingian: Oman, South China (this work), 566

Transcaucasia, Turkey (Hazro), Iran (Fars, Zagros) and Abu Dhabi. 567

568

Septoglobivalvulina cf. guangxiensis Lin, 1978 569

Fig. 7(14) 570


18

v. 2006 Septoglobivalvulina cf. guangxiensis Lin - Gaillot, p. 70-71, pl. I.4, fig. 14, pl. 571

VII.3, fig. 14. 572

v. 2006 Septoglobivalvulina cf. guangxiensis Lin - Vachard et al., fig. 9. 7. 573

Compare with: 574

1978 Septoglobivalvulina guangxiensis nov. gen. nov. sp. Lin, p. 28, pl. 5, figs 4-7. 575

1981 Septoglobivalvulina guangxiensis Lin - Zhao et al., pl. 2, fig. 10. 576

1984 Paraglobivalvulina gracilis Zaninetti and Altiner - Altiner, pl. 1, fig. 15 (the 577

chamberlets of S. gracilis are not visible). 578

1987 Paraglobivalvulina guangxiensis Lin - Loeblich and Tappan, pl. 230, figs 7-8. 579

1990 Septoglobivalvulina guangxiensis Lin - Lin et al., p. 164-165, pl. 12, figs 11-12, 13? 580

(13 might belong to Siphoglobivalvulina). 581

? 1992 Paraglobivalvulina mira Reitlinger - Berczi-Makk, pl. 4, fig. 7. 582

? p. 2005 Paraglobivalvulina globulosa (sic, probably: globosa Wang) - Hughes, pl. 2, fig. 583

21 (non figs 22-23 = ?S. distensa). 584

Comparison: Our material differs from the type material by the smaller size (Diameter = 585

0.820-0.900 mm) and thicker wall, but the chamber shape and the big proloculus are identical. 586

Diameter = 0.670 mm, width = 0.680 mm, number of whorl: 1, number of chambers: 2, 587

proloculus diameter = 0.233 mm, height of last chamber = 0.130 to 0.330 mm, wall thickness 588

= 0.020 to 0.056 mm. 589

Occurrence: Wuchiapingian-Changhsingian of South China (Lin et al., 1990, p. 95). 590

Wuchiapingian of Kuh-e Surmeh, Zagros, Iran (Gaillot, 2006). 591

592

Septoglobivalvulina distensa (Wang in Zhao et al., 1981) 593

Fig. 6(2) 594

1981 Globivalvulina distensa nov. sp. Wang in Zhao et al., p. 48 (in Chinese), 75 (in 595

English), pl. 2, figs 1-3. 596

1985 Globivalvulina sp. of G. vonderschmitti group - Okimura et al., pl. 1, fig. 10. 597

? 1988 Paraglobivalvulina? sp. - Pronina, pl. 2, figs 8-9. 598

p. 1990 Globivalvulina laxa nov. sp. Lin, Li and Sun, p. 163-164, pl. 11, fig. 36 only (non 599

figs 35, 37-38 = globivalvulinid indet.). 600

? p. 2005 Paraglobivalvulina globulosa (sic, probably: globosa Wang) - Hughes, pl. 2, figs 601

22-23 (non fig. 21 = ?S. guangxiensis). 602


19

v. 2006 Septoglobivalvulina distensa (Wang in Zhao et al.) - Gaillot, p. 71, pl. I.4, fig. 5; 603

pl. I.5, fig. 15; pl. I.7, fig. 12; pl. I.17, fig. 12; pl. I.37, fig. 9; pl. I.43, fig. 15; pl. II.8, fig. 1, 5; 604

pl. II.9, fig. 1; pl. II.31, fig. 4; pl. III.6, fig. 5; pl. III.15, fig. 14?; pl. III.16, fig. 5; pl. VI. 4, 605

figs 10-11; 16, pl. VI.5, fig. 18; pl. VI.6, fig. 21; pl. VII.2, fig. 2. 606

Description: Our specimen corresponds to the minimal dimension of the type material. 607

Diameter = 0.500 mm, number of volution: 1, number of chambers: 3 or 4, proloculus 608

diameter = 0.070 mm, height of last chamber = 0.235 mm (i.e., almost the middle of the total 609

diameter), wall thickness = 0.015 mm. 610

Occurrence: Early?-late Midian to Changhsingian: Oman, South China (this work), Turkey 611

(Hazro), Iran (Fars, Zagros), UAE. Late Changhsingian of South China (this work), and 612

?Transcaucasia. 613

614

Septoglobivalvulina sp. 1 615

Fig. 6(3) 616

v. 2006 Septoglobivalvulina sp. 1 - Gaillot, p. 71-72, pl. I.11, fig. 9; pl. I.12, figs 1-2; pl. 617

VII.2, fig. 3. 618

Description: This unique specimen has the same size than S. distensa but more whorls and 619

more chambers. Diameter = 0.555 mm, number of volutions: 2, number of chambers: 5 or 6, 620

proloculus diameter = 0.060 mm, height of last chamber = 0.360 mm (i.e., more than the 621

middle of the total diameter), wall thickness = 0.015 mm. 622

Occurrence: Early Changhsingian of Zagros (Kuh-e Surmeh) (Gaillot, 2006). Late 623

Changhsingian of South China (this work). 624

625

Genus Retroseptellina Gaillot and Vachard in Gaillot et al., submitted 626

Type species: Globivalvulina decrouezae Köylüoglu and Altiner, 1989. 627

Diagnosis: Coiling similar to Globivavulina, with few whorls and few chambers, with a 628

tendency of the increasing of the width of the chambers during the growth. Septa strongly 629

curvated backward. Wall single microgranular. 630

Composition: Globivalvulina globosa Wang in Zhao et al., 1981; G. decrouzae Köylüoglu 631

and Altiner, 1989; Paraglobivalvulina nitida Lin, Li and Sun, 1990. 632

Comparison: Differs from Globivalvulina by the shape of the septa and the chambers; 633

from Septoglobivalvulina by an increasing in width more than in height, and the strongly 634

backward curvated septa. 635


20

Occurrence: Questionable in “early Murgabian” of Thailand (Ueno and Sakagami, 1993; 636

and more probably Midian in age). Murgabian-Dzhulfian-Dorashamian of southern Turkey 637

(e.g., Canuti et al., 1970 up-dated by Gaillot et al., submitted; Koylüoglu and Altiner, 1989; 638

Ünal et al., 2003), Thailand and Malaysia (Yanagida et al., 1988; Fontaine et al., 1993; 1994). 639

Midian of Batain Plain, Oman (Vachard et al., 2002). Latest Midian-earliest Dzhulfian of 640

Transcaucasia (Kotlyar et al., 1989). Midian of New Zealand (Vachard and Ferrière, 1991). 641

Late Midian of central Japan (Kobayashi, 2006b). Dorashamian of Greece (Altiner and 642

Özkan-Altiner, 1998; Baud et al., 1991; Grant et al., 1991). Wuchiapingian-Changhsingian of 643

South China (Lin et al., 1990; (this work). Wuchiapingian of northern Italy and Iran (Mohtat-644

Aghai and Vachard, 2005). Duhaysan Member (late Dzhulfian) of Saudi Arabia (Vachard et 645

al., 2005). Changhsingian of Hungary (Théry et al., 2007). 646

647

Retroseptellina decrouezae (Köylüoglu and Altiner, 1989) 648

Fig. 6(5) 649

p. 1970 Globivalvulina graeca Reichel - Canuti et al., fig. 14.1 (non fig. 14. 3, 4, 6 650

correctly interpreted). 651

p. 1970 Globivalvulina sp. - Canuti et al., fig. 14. 2, 5. 652

1989 Globivalvulina sp. Kotlyar et al., pl. 3, fig. 21. 653

1989 Globivalvulina decrouezae nov. sp. Köylüoglu and Altiner, p. 479-481, text-fig. 8 A-654

H, J-K, pl. 7, fig. 14. 655

p. 1990 Globivalvulina globosa Wang - Lin et al., p. 162, pl. 11, figs 26-27 (non figs 28-29 656

= Globivalvulina bulloides). 657

v. 1991 Paraglobivalvulina? sp. - Vachard and Ferrière, p. 2, fig. 2. 658

v. 1993 Globivalvulina or Paraglobivalvulina? - Fontaine et al., fig. 5E. 659

v. 1993 Paraglobivalvulina mira Reitlinger - Fontaine et al., fig. 6F. 660

v. 1994 Paraglobivalvulinoides? - Fontaine et al., pl. 47, fig. 7. 661

p. 1995 Paraglobivalvulina mira Reytlinger - Berczi-Makk et al., pl. 6, figs 1-3 (non pl. 7, 662

figs 1-2 = R. nitida). 663

1998 Globivalvulina decrouezae Köylüoglu and Altiner - Altiner and Özkan-Altiner, pl. 3, 664

fig. 23. 665

2001 Paraglobivalvulina globosa (Wang) - Pronina-Nestell and Nestell, pl. 5, figs 2-3. 666

v. p. 2002 Septaglobivalvulina (sic) globosa (Wang) - Vachard et al., pl. 1, fig. 13 (non fig. 667

14 = Septoglobivalvulina distensa or gracilis). 668

2004 Globivalvulina globosa Wang - Zhang and Hong, p. 70, pl. 1, figs 24-26. 669


21

v. 2005 Septoglobivalvulina decrouezae (Köylüoglu and Altiner) - Mohtat-Aghai and 670

Vachard, pl. 2, fig. 17. 671

v. 2005 Septoglobivalvulina? decrouezae (Köylüoglu and Altiner) - Vachard et al., p. 154-672

155, pl. 3. 5-6. 673

v. 2006 Septoglobivalvulina? decrouezae (Köylüoglu and Altiner) - Insalaco et al., pl. 1, 674

fig. 20. 675

p. 2006b Globivalvulina sp. - Kobayashi, figs 3. 31, 35-36 (non figs 3. 28-30, 32-34 = truly 676

Globivalvulina spp.). 677

2006c Septoglobivalvulina? sp. - Kobayashi, pl. 2, figs 9-12. 678

v. 2006 Retroseptellina decrouezae (Köylüoglu and Altiner) - Gaillot, p. 68-69, pl. I.13, figs 679

12-14; pl. I.14, figs 1-2; pl. I.15, fig. 15; pl. I.18, figs 3-5, 9; pl. I.37, fig. 12; pl. II.8, fig. 3; 680

pl.III.2, fig. 5; pl. III.3, fig. 13; pl. III.4, fig. 13; pl. III.6, fig. 6; pl. III.7, fig. 2; pl. III.16, figs 681

16-17; pl. III.22, fig. 14; pl. VI.5, fig. 26; pl. VII.2, fig. 5. 682

v. 2006 Retroseptellina decrouezae (Köylüoglu and Altiner) - Vachard et al., fig. 9. 5. 683

Discussion: The outline is subrectangular, the number of whorls is 2 or 3. R. globosa has 684

fewer whorls, and R. nitida is more evolute. In fact, the transition from R. decrouezae to R. 685

nitida is progressive. R. decrouezae is characterized by more than 1 whorl, no increasing of 686

the last whorl, and a much larger size (> 0.500 mm). Its chambers are spacious, and the 687

backward curvature of the septa takes the form of a hook. 688

Dimensions: Diameter = 0.440 mm, width = 0.450 mm, w/D = 1.03, number of whorls: 689

1.5. The diameter of the type material of Köylüoglu and Altiner (1989) is a little larger with 690

0.520-0.640 mm. 691

Occurrence: Late Midian-Lopingian of Italy, Turkey, Thailand and Malaysia, 692

Transcaucasia, New Zealand, Greece, South China, central Iran, Saudi Arabia (see 693

compilation in Vachard et al., 2005, p. 155). Midian of Central Japan (Kobayashi, 2006b). 694

Changhsingian of Japan (Kobayashi, 2006c). Lopingian of Zagros-Fars-Abu Dhabi area. 695

Early Wuchiapingian of Hambast region (Central Iran) (Mohtat-Aghai and Vachard, 2005). 696

Hazro: Late Wuchiapingian-Early Changhsingian. 697

698

Retroseptellina nitida (Lin, Li and Sun, 1990) 699

Fig. 6(4) 700

1990 Paraglobivalvulina nitida nov. sp. Lin, Li and Sun, p. 166, pl. 12, figs 20-21. 701

p. 1995 Paraglobivalvulina mira Reitlinger - Berczi-Makk et al., pl. 7, figs 1-2 (non pl. 6, 702

figs 1-3 =). 703


22

v. 2006 Retroseptellina nitida (Lin, Li and Sun) - Gaillot, p. 84-85, pl. I.19, fig. 18; pl. II.8, 704

fig. 7; pl. VII.2, fig. 4. 705

v. 2006 Retroseptellina nitida (Lin, Li and Sun) - Vachard et al., fig. 9. 6. 706

Description: The last chambers of R. nitida are evolute. The curvature of septa is diversely 707

developed, from weak to hook-shaped. Dimensions: H = 0.500-0.580 mm, w = 0.450-0.780 708

mm, number of whorls: 2.5. The type material of Lin et al. (1990) is larger with 0.960-1.120 709

mm but for 3-4 whorls. 710

Occurrence: Lopingian of South China (Lin, Li and Sun, 1990), Hungary (Berczi-Makk et 711

al., 1995), and Zagros, Kuh-e Surmeh (Gaillot, 2006). 712

713

Genus Paraglobivalvulinoides Zaninetti and Jenny-Deshusses, 1985 714

Type species. Paraglobivalvulina? septulifera Zaninetti and Altiner, 1981. 715

Diagnosis: Test large, spherical, biserially coiled, involute, chambers wide and strongly 716

embracing. Oral chamberlets with very complex tongues. Wall microgranular, proportionally 717

thin. 718

Composition: Paraglobivalvulina? septulifera Zaninetti and Altiner, 1981, 719

Paraglobivalvulina piyasini Sakagami and Hatta, 1982, Paraglobivalvulinoides spumida Lin, 720

Li and Sun, 1990. 721

Occurrence: Latest Changhsingian, Alborz, northern Iran (Bozorgnia, 1973), Zagros, 722

southern Iran (Gaillot, 2006), ?Italy (Pasini, 1985), Greece (Vachard et al., 1993b), Himalaya 723

(Lys et al., 1980), South China (Lin et al., 1990; this work), Thailand (Sakagami and Hatta, 724

1982; Yanagida, 1988), Transcaucasia (Pronina-Nestell and Nestell, 2001), Japan (Kobayashi, 725

1997; 1999), and Malaysia (Fontaine et al., 1994). 726

727

Paraglobivalvulinoides septulifer (Zaninetti and Altiner, 1981) 728

Fig. 5(1) 729

1973 Paraglobivalvulina mira Reitlinger - Bozorgnia, p. 145, pl. 39, figs 9-11 pl. 40, figs 730

1-2. 731

1980 Paraglobivalvulina mira Reitlinger - Lys et al., pl. 3, figs 7-10. 732

1981 Paraglobivalvulina? septulifera nov. sp. Zaninetti and Altiner, p. 40-41, pl. 1, figs 733

15-19. 734

1981 Paraglobivalvulina sp. - Altiner, pl. 36, fig. 3. 735

1982 Chrysanthemina - Jiang, Yang and Zang, pl. 2, fig. 10 (nomen nudum, no 736

description). 737


23

1982 Paraglobivalvulina piyasini nov. sp. Sakagami and Hatta, p. 10, pl. 5, fig. 1. 738

1983 Paraglobivalvulina septulifera Zaninetti and Altiner - Jenny-Deshusses, pl. 23, fig. 5. 739

1985 Paraglobivalvulina septulifera Zaninetti and Altiner - Pasini, pl. 61, fig. 11. 740

? 1987 Paraglobivalvulina septulifera Zaninetti and Altiner - Noé, p. 108, pl. 30, fig. 10. 741

? 1988 Paraglobivalvulina piyasini Sakagami and Hatta - Yanagida, pl. 2, figs 28-29. 742

1990 Paraglobivalvulina? septulifera Zaninetti and Altiner - Lin et al., p. 166, pl. 12, figs 743

22-24. 744

? 1990 Paraglobivalvulina spumida nov. sp. Lin et al., p. 166-167, pl. 12, figs 25-30. 745

1993b Paraglobivalvulinoides septulifera Zaninetti and Altiner (sic without brackets) - 746

Vachard et al., pl. 8, fig. 1. 747

1993 Paraglobivalvulina sp. - Baghbani, pl. 6, fig. 5. 748

1994 Paraglobivalvulina septulifer Zaninetti and Altiner - Fontaine et al., pl. 47, fig. 6. 749

1996 Paraglobivalvulinoides septulifera (Zaninetti and Altiner) - Rauzer-Chernousova et 750

al., pl. 18, fig. 10. 751

? 1997 Paraglobivalvulina piyasini Sakagami and Hatta - Kobayashi, fig. 2. 3, pl. 4, figs 1-752

5. 753

? 1999 Paraglobivalvulina piyasini Sakagami and Hatta - Kobayashi, text-fig. 1. 11 p. 281. 754

2001 Paraglobivalvulinoides sp. - Pronina-Nestell and Nestell, pl. 5, figs 6-7. 755

? 2001 Paraglobivalvulina piyasini Sakagami and Hatta - Pronina-Nestell and Nestell, pl. 756

5, fig. 8. 757

v. 2006 Paraglobivalvulinoides sp. - Insalaco et al., pl. 2, fig. 24. 758

2006c Paraglobivalvulina sp. - Kobayashi, pl. 2, fig. 13. 759

2006c Paraglobivalvulina mira (Reitlinger) - Kobayashi, pl. 2, fig. 14. 760

v. 2006 Paraglobivalvulinoides septulifer (Zaninetti and Altiner, 1981) - Gaillot, p. 73, pl. 761

I.19, fig. 19. 762

v. 2006 Paraglobivalvulinoides septulifera (Zaninetti and Altiner, 1981) - Vachard et al., 763

fig. 9. 10. 764

Dimensions: Height = 2.300-2.860 mm, width = 2.300-2.800 mm, number of pairs of 765

chambers: 4, wall thickness = 0.050 mm. 766


768

Subfamily DAGMARITINAE Bozorgnia 1973 emend. herein 769

Synonyms: Biseriamminidae Chernysheva, 1941 (pars); Globivalvulinidae Reitlinger, 770

1950 (pars); Louisettitinae Loeblich and Tappan, 1984; Louisettitidae nomen translat. Rauzer-771


24

Chernousova et al., 1996; dagmaritin-type biseriamminids sensu Altiner (1997, text-fig. 1 p. 772

3). 773

Remarks: This subfamily corresponds to the Dagmarita-lineage and its possible ancestors. 774

The subfamily is morphologically poorly known (and/or poorly represented morphological 775

variations). The creation of Louisettininae appears, in this case, devoid of significance. 776

Composition: Sengoerina Altiner, 1999; Crescentia Ciarapica, Cirilli, Martini and 777

Zaninetti, 1986; Dagmarita Reitlinger, 1965; Siphodagmarita Gaillot and Vachard (nomen 778

nudum). Bidagmarita gen. nov.; Louisettita Altiner and Brönnimann, 1980. 779

Occurrence: Early Murgabian/Wordian to latest Changhsingian; Palaeo-Tethyan, Ne-780

Tethyan. 781

782

Genus Dagmarita Reitlinger, 1965 783

Type species: Dagmarita chanakchiensis Reitlinger, 1965. 784

Diagnosis: Test rectilinear, tapering, biseriate. Chambers hemispherical to semi-785

ellipsoidal. Wall thin, with development of lateral horn-like expansions. Wall microgranular, 786

uni- to three layered (granular or finely perforated in anatypical forms, see below: 787

Dagmarita?). 788

Composition: Many species are described in the literature, often with excessive 789

synonymies. Two groups can nevertheless be distinguished: D. chanakchiensis and D. altilis. 790

The following species were described: D. chanakchiensis Reitlinger, 1965; D. elegans 791

Sosnina in Sosnina and Nikitina, 1977; D. cuneata Sosnina in Sosnina and Nikitina, 1977; D. 792

exilis Sosnina in Sosnina and Nikitina, 1977; D. oblonga Sosnina in Sosnina and Nikitina, 793

1977; D. altilis Wang in Zhao et al., 1981; D. elongata Wang in Zhao et al., 1981; D. 794

minuscula Wang in Zhao et al., 1981; D. simplex Wang in Zhao et al., 1981; D. liantanensis 795

Hao and Lin, 1982; D. caucasica Vuks in Kotlyar et al., 1984; D. minima Lin, 1984; D. 796

shahrezaensis Mohtat Aghai and Vachard, 2003. D. shahrezaensis do not display lateral 797

expansions and might belong to another genus to be described. 798

Occurrence: Early Murgabian (Vachard, 1980) and/or early Maokouan (Lin et al., 1990) 799

to latest Changhsingian (Zhao et al., 1981; Lin et al., 1990) of Palaeo-Tethys and Neo-Tethys. 800

with the following detailed distribution from West to East: Middle Permian of Appennines 801

(Panzanelli-Fratoni et al., 1987; Vachard and Miconnet, 1990); Middle Permian of 802

Montenegro (Pantic, 1970); Late Permian of the Carnic Alps (Noé, 1987); Late Permian of 803

Hungary (Berczi-Makk, 1992); Late Permian of western Turkey (Argyriadis et al. 1976; Lys 804

and Marcoux, 1978); Late Permian of eastern Taurus (Zaninetti et al., 1981; Altiner, 1981, 805


25

1984; Koyluöglu and Altiner, 1989); Midian-Changhsingian of Hazro (Canuti et al. 1970 as 806

Palaeotextularia sp.; updated in this study); Midian-Dzhulfian of Transcaucasia (Reitlinger, 807

1965; Kotlyar et al., 1984, 1989); Late Permian of central Alborz (Bozorgnia, 1973; Jenny-808

Deshusses, 1983); Late Wuchiapingian of central Iran (Mohtat-Aghai and Vachard, 2003); 809

Lopingian of Zagros and Fars (this study); Middle Permian of central Afghanistan (Vachard 810

and Montenat, 1981); Middle Permian of Salt Range (Pakistan, Okimura, 1988); Late 811

Changhsingian of Ladakh (Himalaya, Lys et al., 1980); Maokouan- Changhsingian of South 812

China (Zhao et al., 1981; Lin et al., 1990); Middle Permian of West Thailand (Fontaine et al., 813

1988a) and Philippines (Fontaine et al., 1986); Midian-Dzhulfian of northwestern Thailand 814

(Caridroit et al., 1990, p. 342, 348); Middle Permian of Malaysia (Fontaine et al., 1988b); 815

Midian-Dzhulfian of northwestern Thailand (Caridroit et al., 1990, p. 342, 348); Middle-Late 816

Permian of Cambodia (Nguyen Duc Tien, 1979, 1986a); Late-Middle Permian of Primorye 817

(Sosnina in Sosnina and Nikitina, 1977); Late Midian-Changhsingian of Japan (Kobayashi, 818

2006c). 819

820

Dagmarita altilis Wang in Zhao et al., 1981 821

Fig. 6(19) 822

1981 Dagmarita altilis nov. sp. Wang in Zhao et al., p. 74, pl. 1, fig. 24. 823

1984 Dagmarita minima Lin, p. 112, pl. 1, figs 18-19. 824

1988 Dagmarita altilis Wang - Pronina, pl. 2, figs 10-11. 825

1989 Dagmarita altilis Wang - Pronina, pl. 1, figs 10-11. 826

1990 Dagmarita altilis Wang - Lin et al., p. 84 (no illustration). 827

v. 2006 Dagmarita altilis Wang - Insalaco et al., pl. 2, fig. 2. 828

v. 2006 Dagmarita altilis Wang in Zhao et al. - Gaillot, p. 77, pl. I.7, fig. 15 pl. I.8, figs 1-3, 829

11, 21-22, pl. I.14, figs 5-6, 11, 13¸pl. I.16, fig. 1; pl. I.9, fig. 14?, pl. II.14, figs 12-13, pl. 830

II.31, figs 5, 9, pl. III.6, figs 10-11, pl. III.15, figs 3, 8, 17, pl. III.16, figs 1, 8, pl. III.22, fig. 831

10; pl. III.24, figs 1-7; pl. III.25, figs 5-9, 15-18; pl. VI.6, fig. 2; pl. VII.2, fig. 19. 832

Description: This species differs from D. chanakchiensis by a wider test, a truncate base, 833

and wider chambers. Some specimens of D. altilis appear more or less similar to D. simplex 834

Wang in Zhao et al. by the simple wall thin dark microgranular, the basal often truncated and 835

the shape of chamber. They probably correspond to juveniles and/or deformed specimens of 836

D. altilis. 837


26

Occurrence: Early Maoukouan-Latest Changhsingian of southern China (Wang in Zhao et 838

al. 1981; Lin et al., 1990), Changhsingian of Transcaucasia (Pronina 1988; 1989), Lopingian 839

of Zagros. Lopingian of Hazro (Turkey). 840

841

Dagmarita? cf. sharezaensis Mohtat-Aghai and Vachard, 2003 842

Fig. 6(18) 843

Compare with: 844

? 1986 Dagmarita sp. - Vuks and Chediya, pl. 9, fig. 10. 845

v. 2003 Dagmarita shahrezaensis nov. sp. Mohtat-Aghai and Vachard, p. 38, 40, 42, pl. 1, 846

figs 1-14 (with 4 references in synonymy). 847

v. 2005 Dagmarita shahrezaensis Mohtat-Aghaï and Vachard - Mohtat-Aghai and 848

Vachard, p. 211, pl. 2, figs 21-22; pl. 3, fig. 5. 849

? 2005 Dagmarita chanakchiensis Reitlinger - Hughes, pl. 4, fig. 9. 850

v. ? 2005 Dagmarita? shahrezaensis Mohtat-Aghai and Vachard - Vachard et al., p. 155, 851

pl. 3, figs 9-10, 15? 852

v. 2006 Dagmarita? sharezaensis Mohtat-Aghai and Vachard - Gaillot, p. 77-78, pl. I.8, 853

fig. 6; pl. I.37, fig. 16; pl. V.3, fig. 9; pl. VI. 5, figs 1?, 19; pl. VII.2, fig. 18. 854

Comparison: This species differs from D. chanackchiensis and D. altilis by the absence of 855

hornlike expansions, but the wall and septal curvature are characteristic of Dagmarita genus. 856

This specimen is especially long and thick-walled (hence the open nomenclature). 857

Occurrence: Questionable in early Changsinghian of Saudi Arabia and Late 858

Changhsingian of Primorye. Late Wuchiapingian-?Changhsingian of central Iran, Himalaya, 859

Transcaucasia, South China and Malaysia. Late Wuchiapingian of central Iran. Late Midian-860

Changhsingian of Zagros and Hazro (Gaillot, 2006). 861

862

Genus Bidagmarita Gaillot and Vachard gen. nov. 863

Type species: Bidagmarita sinica Gaillot and Vachard nov. gen. nov. sp. 864

Etymology: From Latin bis = two times, doubly, and Dagmarita similar genus; because of 865

the double size in comparison to this latter genus and the bilayered wall. 866

Diagnosis: Dagmaritin genus characterized by a great size and a bilayered wall: 867

microgranular outer layer, pseudofibrous inner layer, and a faint curvature backward of the 868

septa. 869


27

Composition: Bidagmarita sinica nov. sp. ; ?Dagmarita chanakchiensis sensu Lys in Lys 870

et al., 1980 and sensu Fontaine et al., 1994, pl. 3, fig. 6; ?D. caucasica Vuks in Kotlyar et al., 871

1984; ?D. spp. of D. chanakchiensis group sensu Okimura et al. (1985, pl. 1, figs 7-8); D. sp. 872

sensu Fontaine et al., 1994, pl. 45, fig. 9. 873

Comparison: The new genus differs from Dagmarita by the great size and the wall, from 874

Louisettita by the wall and the absence of endoskeleton. 875

Occurrence: ?Kalabagh Member, Salt Range, Pakistan; ?Abadeh Formation, central Iran. 876

?Nikitin Suite (Late Changhsingian) of NW Caucasus. ?Late Changhsingian of Lamayuru 877

Block (Ladakh Himalaya). ?Late Changhsingian of Malaysia. Late Changhsingian of South 878

China (this work). 879

880

Bidagmarita sinica Gaillot and Vachard nov. gen. nov. sp. 881

Fig. 7(4-5) 882

v. 2006 Bidagmarita sinica Gaillot and Vachard nov. gen. nov. sp. Gaillot, p. 78-79, pl. VII.3, 883

figs 4-5 (nomen nudum in a Ph.D.). 884

v. 2006 Bidagmarita sinica Gaillot and Vachard - Vachard et al., fig. 9. 14-15 (nomen nudum, 885

no description). 886

Etymology: From Latin Sinicus = from China. 887

Type locality: Laren (Guangxi, South China). 888

Type level: Late Changhsingian. 889

Diagnosis: As for the genus. 890

Description: Test large for a Dagmaritin, obviously tapering. Proloculus spherical 891

moderately large. Sagittal axial section (Fig. 7(4)) showing a regular increasing in width and 892

height of the chambers. The pairs of septa which have the typical form of the Dagmarita at 893

the beginning: i.e., their end situated in the symmetry plane (which coincides with the axis of 894

development), are progressively distant from each other and the space without septa in the 895

center increases. Horny expansions are proportionally weak in comparison with the size of the 896

test, and a rapid examination can permit to conclude to a palaeotextulariid representative. 897

Nevertheless, the wall and the curvature of septa differ completely. Chamber subtrapezoidal 898

in shape. Septa gently curvated. A small oral tongue terminates the septa and re-inforces the 899

aperture (as in Dagmarita). Other endoskeletal elements are absent. Within the wall, the 900

microgranular layer is very thin and the pseudofibrous inner layer has three or four times 901

ticker. In frontal axial section (Fig. 7(5)), excepted the generic differences, the differences 902


28

with Dagmarita or Louisettita ultima are rather incipient. No transverse sections were 903

observed. 904

Dimensions: Height = 0.835-1.025 mm, width = 0.690 mm, test thickness = 0.465 mm, 905

proloculus diameter = 0.020 mm, number of chambers: 8 pairs, height of last chamber = 906

0.167-0.222 mm, wall thickness = 0.015 mm. 907

Comparison: No illustration in the literature is comparable. 908

Holotype: Fig. 7(4) (sample L3). 909

Type material: Rare form: 4 specimens in 3 thin-sections. 910

Repository of the type: Natural History Museum, Geneva (Switzerland), holotype number 911

MHNG 07-002. 912

Occurrence: Late Changhsingian of Laren section (South China, this work). 913

914

Genus Louisettita Altiner and Brönnimann, 1980 emend. herein 915

Type species: Louisettita elegantissima Altiner and Brönnimann, 1980. 916

Emended diagnosis: The shape is identical to Dagmarita, but some endoskeletal elements 917

appear (the “radial partitions” of the original diagnosis) that, in advanced species, form small 918

loops or equivalents to the phrenothecae of some schwagerinoid fusulinids (and not always 919

radial and perpendicular partitions as indicated in the original diagnosis), or backward 920

oriented septa. Wall microgranular, monolayered. Aperture basal simple. 921

Composition: L. elegantissima, L. ultima nov. sp., L. extraordinaria nov. sp., 922

?Paradagmarita dubreuilli Vachard, 1980 (nomen nudum) (= ?Paradagmarita sp. Pronina, 923

1988a, pl. 2, fig. 14 = Pronina, 1988, pl. 1 fig. 14). 924

Occurrence: Late Changhsingian. Turkey, Zagros, ?Afghanistan, ?Transcaucasia and 925

?South China (this work). 926

927

Louisettita ultima Gaillot and Vachard nov. sp. 928

Fig. 6(20-25) 929

1981 Dagmarita sp. - Wang in Zhao et al., pl. 1, fig. 26. 930

v. 2006 Louisettita ultima Gaillot and Vachard in Gaillot, p. 81, pl. I.14, figs 7-10; pl. I.15, 931

figs 8, 16-17; pl. I.19, fig. 14; pl. III.5, fig. 8?; pl. III.6, fig. 12; pl. VI.6, figs 9-14; pl. VI.7, 932

fig. 18; pl. VII.2, figs 20-25. 933

Etymology: From Latin ultimus = ultimate, because this species persists in the latest 934

Permian sequence prior the end Permian mass extinction. 935


29



Diagnosis: Test medium sized for the genus, wall thick unilayered, loops few visible, 938

backward curvated septa present. 939

Description: The species, medium sized for the genus, is a little larger than the type-940

species L. elegantissima, and exhibits a thicker wall. The radial partitions are visible in 941

convenient sections. The main criteria of this species is the development of a backward 942

curvature of the septa (e.g., as in Retroseptellina). Large and robust test with thick wall but 943

thin septa. No specimens were observed with regular vertical partitions. Endoskeletal 944

elements very irregularly arranged as small chamberlets: at the base of septa or laterally (Fig. 945

6 (23)), or loops (Fig. 6(20)) or curvature of septa (Fig. 6(25)). The wall of the last chambers 946

of the specimen 6(21) is finely perforated; this particularity exists also in Turkey (Altiner, 947

unpublished data). 948

Dimensions: Height (H) = 0.315-0.550 mm, width (w) = 0.145-0.220 mm, w/H = 0.40-949

0.46, proloculus diameter = 0.025 mm, number of pairs of chambers: 5-6, wall thickness = 950

0.012-0.020 mm, loop thickness = 0.002mm. 951


Type material: 28 specimens. 953

Repository of type: Natural History Museum, Geneva (Switzerland), holotype number 954

MHNG 07-003. 955

Occurrence: Changhsingian of Eastern Taurus (Hazro), Zagros (Gaillot, 2006) and South 956

China (this work). 957

958

Subfamily PARADAGMARITINAE Gaillot and Vachard (submitted) 959

2006 Paradagmaritinae Gaillot, p. 82 (nomen nudum). 960

Description: A subfamily of Globivalvulinidae (i.e., with a biserially coiled growth and a 961

microgranular wall, occasionally differenciated) characterised by an uncoiling more or less 962

developed after an initial coiling generally slightly trochospiral. 963

Composition: Paradagmarita Lys in Lys and Marcoux, 1978; Paradagmarita? (this 964

study); Paradagmaritopsis gen. nov.; Paradagmaritella nov. gen. Gaillot and Vachard 965

(nomen nudum, submitted); Paradagmacrusta nov. gen. Gaillot and Vachard (nomen nudum, 966

submitted); Paremiratella nov. gen. Gaillot and Vachard (nomen nudum, submitted). 967


30

Discussion: Differ from Globivalvulininae in the uncoiled part of the test; differ from 968

Dagmaritininae (= Louisettitinae) in the coiled part. No evidence of the phylogenetic filiation 969

from Dagmarita to Paradagmarita (as proposed by Altiner, 1997) has been observed. A 970

filiation from Globivalvulina seems to be more likely (see the specimens of Okimura et al., 971

1985, pl. 1, fig. 16, and Berczki-Makk et al., 1995, pl. 6, fig. 4). 972

Occurrence: Lopingian, frequent on both sides of western Neo-Tethys (but quoted from 973

southern Italy to Thailand), and confirmed here in Japan with Paradagmaritopsis) and 974

mentioned for the first time in South China (with Paradagmarita? sp. and Paradagmaritopsis, 975

this work). 976

977

Genus Paradagmarita Lys in Lys and Marcoux, 1978 emend. 978

Type species: Paradagmarita monodi Lys in Lys and Marcoux, 1978. 979

Emended diagnosis: Small to medium-sized globivalvulinid paradagmaritin characterized 980

by an early stage enrolled, biserial, involute, slightly trochospiral, and a later uncoiled, 981

biserial stage, relatively long. Wall dark, microgranular, relatively thin, uni- or multilayered. 982

Chambers inflated. Aperture simple, terminal, interio-marginal with a valvula. 983

Occurrence: According to Kotlyar et al. (1989, tabl. 1 n° 24), Paradagmarita appears in 984

the latest Khachikian (top of Midian), but the illustrated specimen (pl. 4, fig. 10) looks like 985

Spireitlina Vachard in Vachard and Beckary, 1991. Middle Wuchiapingian-Changhsingian. 986

Paleo-Tethyan and Neo-Tethyan (Gaillot, 2006), principally known in Turkey (Taurus), Iran 987

(Zagros), Saudi Arabia (Vachard et al., 2005) and Transcaucasia (Pronina-Nestell and Nestell, 988

2001), but mentioned from Italy to Japan. The Paradagmarita from Afghanistan described by 989

Vachard (1980) rather belong to the genus Louisettita. Forms from Thailand and Pakistan are 990

very different. The Paradagmarita of Japan (Kobayashi, 1997; 2004) belong to 991

Paradagmaritopsis, also observed in South China (this study). Rare Paradagmarita, briefly 992

described below, are present in South China. 993

994

Paradagmarita sp. 995

Fig. 6(11, 17) 996

v. 2006 Paradagmarita spp. Gaillot, p. 85-86, pl. I.6, fig. 4, pl. III.2, fig. 7, pl. VII.2, figs 997

11, 17. 998

Description: Two cross sections displaying an arcuate beginning of coiling, trocho- or 999

planispiral, followed by a plansipiral coiling with chambers enveloping the apertural face. 1000

These characters evoke the genus Paradagmarita or a more peneropliform new genus. 1001


31

Diameter = 0.300-0.355 mm, number of volution: 1-1.5, number of chambers of last whorl: 6-1002

10. 1003

Occurrence: Changhsingian of Laren (South China, this work). 1004

1005

Genus Paradagmaritopsis Gaillot and Vachard gen. nov. 1006

Type species: Paradagmaritopsis kobayashii Gaillot and Vachard nov. gen. nov. sp. 1007

Etymology: Ending opsis = similar to. 1008

Diagnosis: Coiled part very short reduced to the two first chambers, or to the curvature of 1009

the axis. Long uncoiled biseriate part. Chambers subelliptical in shape, with oblique long axis. 1010

Comparison: It differs from Paradagmarita by the very short coiled part; from Dagmarita 1011

by the absence of lateral cameral protuberances; from the Palaeotextularioidea by the type of 1012

wall and the minute size; from Globispiroplectammina Vachard, 1977 by the type of wall, and 1013

from Robustulina Gaillot and Vachard in Gaillot, 2006 by the importance of the uncoiled part. 1014

Occurrence: Late Wuchiapingian-Changhsingian of Zagros (Gaillot, 2006). 1015

Changhsingian of Japan and South China (this work). 1016

1017

Paradagmaritopsis kobayashii Gaillot and Vachard 1018

Fig. 6(6-8, 12-14, 15?, 16?) 1019

1997 Paradagmarita sp. - Kobayashi, pl. 4, fig. 19. 1020

2004 Paradagmarita sp. - Kobayashi, p. 67, fig. 6.47-6.50. 1021

v. 2006 Paradagmarita sp. 2 - Insalaco et al., pl. 2, fig. 1. 1022

v. 2006 Paradagmaritopsis kobayashii Gaillot and Vachard nov. gen. nov. sp. - Gaillot, p. 87-1023

88; pl. I.9, figs 1-9; pl. I.12, fig. 23; pl. II.15, figs 6-8, 15; pl. II.16, figs 1-2, 6-14; pl. III.15, 1024

figs 11-12; pl. III.22, fig. 13; pl. VII.2, figs 6-8, 12-15, 16? (nomen nudum; submitted). 1025

v. 2006 Paradagmaritopsis kobayashii Gaillot and Vachard - Vachard et al., figs 10. 8-10. 1026

Etymology: Dedicated to Prof. Fumio Kobayashi, for his work in Permian 1027

micropaleontology. 1028



Diagnosis: As for the genus. 1031

Description: The initial deviated part is very short composed of 2-3 pairs of chambers, 1032

small, hemispherical suceeding to the proloculus. The uncoiled biserate part is proportionally 1033

large formed of asymmetrical ogival chambers often terminated by a mamilla (holotype: Fig. 1034

6(6)). The wall is dark, thin to relatively thick (due to planes of section or local 1035


32

thickennings?). Height = 0.280-0.700 mm, width = 0-160-0.460 mm, proloculus diameter = 1036

0.040-0.050 mm, number of pairs of coiled chambers: 2-3, number of pairs of uncoiled 1037

chambers: 3-4. The specimen Fig. 6(15) is exceptionally large with H = 1.500 mm for 8 pairs 1038

of chambers, it is unique. The specimen Fig. 6(16) differs by a relatively long coitled stage of 1039

one complete whorl, nevertheless as the last chamber is similar to the chambers of the 1040

holotype (Fig. 6(6)), it is provisionally considered as forming part of the species. 1041


Repository of the type: Natural History Museum, Geneva (Switzerland) , holotype 1043

number MHNG 07-004. 1044


1046

Paradagmaritopsis sp. 1047

Fig. 6(9-10) 1048

v. 2006 Paradagmaritopsis sp. - Gaillot, p. 88, pl. VII.2, figs 9-10. 1049

Description: Two oblique sections in the Chinese material exhibit a coiling apparently 1050

diffrent of P. kobayashii. Height = 0.425-0.465 mm, width = 0.190 mm, test thickness = 0.130 1051

mm, proloculus diameter = 0.030 mm, number of chambers: 5 pairs, height of last chamber = 1052

0.110-0.130 mm, wall thickness = 0.010 mm. 1053

Occurrence: Late Changhsingian of South China (this work). 1054

1055

Suborder FUSULININA Fursenko, 1958 1056

Superfamily OZAWAINELLOIDEA Thompson and Foster, 1937 nomen translat. 1057

Solovieva, 1978 (pro family) 1058

Family REICHELINIDAE Miklukho-Maklay, 1959 nomen translat. Vachard et al., 1993a 1059

(pro Reichelininae) 1060

Genus: Reichelina Erk, 1942 1061

Type species: Reichelina cribroseptata Erk, 1942. 1062

1063

Reichelina simplex Sheng, 1956 1064

Fig. 5(6), Fig. 7(12) 1065

1956 Reichelina simplex nov. sp. Sheng, p. 290, 299, pl. 1, figs 5-6. 1066

1963 Reichelina simplex Sheng - Sheng, p. 149, pl. 1, figs 10-12. 1067

1977 Reichelina simplex Sheng - Lin et al., p. 13, pl. 2, fig. 5. 1068

1979 Reichelina simplex Sheng - Sun, pl. 1, figs 3-4. 1069


33

1984 Reichelina simplex Sheng - Sheng and Rui, p. 34, pl. 1, fig. 8. 1070

1989 Reichelina sp. - Pronina, pl. 2, fig. 5. 1071

1991 Reichelina simplex Sheng - Flügel et al., pl. 37, fig. 3. 1072

v. 2006 Reichelina simplex Sheng - Insalaco et al., pl. 2, fig. 3. 1073

v. 2006 Reichelina simplex Sheng - Gaillot, p. 44-45, pl. I.4, fig. 1, pl. I.5, fig. 12, pl. I.17, 1074

figs 2, 18; pl. I.18, fig. 2, pl. I.36, figs 1-2, 8, pl. II.3, fig. 8, pl. II.33, figs 1, 4, 15, 18; pl. III.6, 1075

fig. 13; pl. III.7, fig. 1; pl. VI.4, figs 4-5, 17-19; pl. VI.5, fig. 2; pl. VII.1, fig. 6; pl. VII.3, fig. 1076

12. 1077

Description: Our specimens correspond exactly to the smallest dimensions of the 1078

populations described by Sheng (1963), with, for the illustrated specimen of Fig. 5(6): 1079

Diameter = 0.600 mm, width = 0.150 mm, w/D = 0.25, number of whorls: probably 3, height 1080

of the last part = 0.080 mm; and for the other one (Fig. 7(12)): Diameter = 0.410 mm (only 1081

the coiled part is cut), width = 0.125 mm, w/D = 0.30, number of whorls: 3.5, proloculus 1082

diameter = 0.022 mm. 1083

Discussion: This species described in the Wuchiapingian and Changhsinghian of South 1084

China is the smallest of the adult Reichelina (we consider R. minuta Erk as a juvenile of R. 1085

cribroseptata). The latest Reichelina cited in South China is Reichelina pulchra Miklukho-1086

Maklay, 1954 (Sheng et al., 1984, text-fig. 6 p. 143). 1087

Occurrence: Late Midian of Sicily (Flügel et al., 1991), Hazro (Turkey) and Kuh-e Dena 1088

(Zagros). Late Changhsingian of Transcaucasia (Pronina, 1989). Lopingian of Kuh-e Surmeh 1089

and Fars. Not found in Abu Dhabi wells probably due to restricted environments. 1090

1091

Superfamily STAFFELLOIDEA Miklukho-Maklay, 1949 nomen translat. Solovieva, 1978 1092

Family STAFFELLIDAE Miklukho-Maklay, 1949 nomen traslat. Rauzer-Chernousova et 1093

al., 1996 pro subfamily (sic Staffellinininae) 1094

Genus Nankinella Lee, 1933 1095

Type species: Staffella discoides Lee, 1931. 1096

1097

Nankinella cf. inflata (Colani, 1924) 1098

Fig. 5(12) 1099

Compare with: 1100

1924 Fusulinella inflata nov. sp. Colani, p. 77-78, pl. 15, figs 3-5, 7-10, 13, 15; pl. 29, fig. 1101

25. 1102


34

1963 Nankinella inflata (Colani) - Sheng, p. 31-32, 155, pl. 3, figs 1-5 (with 2 references 1103

in synonymy). 1104

1977 Nankinella inflata (Colani) - Lin et al., p. 14-15, pl. 2, fig. 14. 1105

1998 Nankinella cf. inflata Colani (sic) - Zhang and Hong, p. 209, pl. 2, fig. 19 (with 5 1106


v. 2006 Nankinella cf. inflata (Colani) - Gaillot, p. 47, pl. VII.1, fig. 12. 1108

Description: Our material contains several oblique sections of a large species with 1109

relatively few whorls. They migth belong to N. inflata (Colani, 1924) sensu Sheng, 1963. 1110

Diameter = 3.000 mm, width = 2.160 mm, w/D = 0.72 for 10 whorls. Unfortunately, no axial 1111

section was found. 1112

Occurrence: Middle to Late Permian of Viet-Nam, South China, Xizang, Thailand, 1113

Cambodia, Malaysia, Greece (Chios Island). 1114

1115

Order LAGENIDA Lankester, 1885 1116

Superfamily NODOSARIOIDEA Ehrenberg, 1838 1117

Genus Pachyphloia Lange, 1925 1118

Type species: Pachyphloia ovata Lange, 1925 (see Loeblich and Tappan 1987 and not 1119

Sellier de Civrieux and Dessauvagie, 1965). 1120

Occurrence: FAD in the Sakmarian. Acme and specific diversification: Midian-Lopingian 1121

(this study in Hazro, Turkey, and Zagros, Iran). LAD Latest Permian. 1122

1123

Pachyphloia pedicula Lange, 1925 1124

Fig. 5(9) 1125

1925 Pachyphloia pediculus nov. sp. Lange, p. 232, pl. 1, fig. 23. 1126

1960 Pachyphloia pediculus Lange - Sosnina, pl. 1, fig. 6. 1127

1965 Pachyphloia “pediculus” Lange – Sellier de Civrieux and Dessauvagie, p. 114-115 1128

(no illustration). 1129

1970 Pachyphloia pedicula Lange - Pantic, pl. 10, fig. 9. 1130

1973 Pachyphloia pedicula Lange - Bozorgnia, p. 154-155, pl. 36, figs 1-2, 4. 1131

1978 Pachyphloia pedicula Lange - Lys and Marcoux, pl. 8, fig. 3 1132

1981 Pachyphloia pedicula Lange - Altiner, pl. 40, figs 19-24. 1133

1983 Pachyphloia pedicula Lange - Jenny-Deshusses, p. 106, pl. 4, fig. 1 (with 4 1134

references in synonymy. 1135

1986a Pachyphloia sp. - Nguyen Duc Tien, pl. 1, figs 19-20. 1136


35

1990 Pachyphloia pediculus Lange - Lin et al., p. 243, pl. 32, figs 7-8. 1137

1992 Pachyphloia pedicula Lange - Berczi-Makk, pl. 1, fig. 2. 1138

1995 Pachyphloia pedicula - Berczi-Makk et al., pl. 2, figs 6-8; pl. 9, figs 6?, 7? 1139

2004 Pachyphloia pediculus Lange - Zhang and Hong, p. 76, pl. 3, fig. 25 (with 2 1140


v. 2006 Pachyphloia pedicula Lange - Gaillot, p. 167, pl. I.26, fig. 7; pl. I.27, fig. 9; pl. 1142

III.27, fig. 8; pl. VI.11, fig. 2; pl. VI.12, fig. 3; pl. VI.13, fig. 3; pl. VII.1, fig. 9. 1143

Description: Test medium sized with relatively few chambers. Height = 0.750 m, width = 1144

0.560 mm, w/H = 0.75, number of chambers: 8 or 9, proloculus diameter = 0.025 mm, height 1145

of last chamber = 0.120 mm, wall thickness = 0.020 mm. 1146

Occurrence: Late Midian of Sumatra. Late Midian, Wuchiapingian and Changhsingian of 1147

Hazro (eastern Taurus, Turkey). Late Changhsingian of NW Caucasus, Hungary, Alborz, 1148

Turkey, Transcaucasia and South China. 1149

1150

Family FRONDINIDAE Gaillot and Vachard in Gaillot et al. (submitted) 1151

2006 Frondiniidae fam. nov. - Gaillot, p. 195 (nomen nudum in a Ph.D.). 1152

2006 Frondinidae fam. nov. Gaillot and Vachard in Gaillot et al., p. 158-159 (nomen 1153

nudum; submitted). 1154

Diagnosis: A family of Nodosarioidea relatively common during the Midian/Capitanian 1155

and the Late Permian. This family is characterized by its dark (microgranular?) wall and the 1156

embracing shape of the chambers. Apertures are smooth or with an internal neck. 1157

Composition: Frondina Sellier de Civrieux and Dessauvagie, 1965; Ichthyofrondina 1158

Vachard in Vachard and Ferrière, 1991 emend. 1159

Comparison: The family differs from all the other groups of Nodosarioidea by the return 1160

to a dark wall (this of the distant ancestor Earlandia; Vachard, 1994), whereas the members 1161

of the family are rigorously homeomorph of some typical Nodosarioidea such as Ichtyolaria 1162

Wedekind, 1937. The link between ancestral Nodosinelloides and Frondinidae can be 1163

represented by an unknown genus which would present the partial disappearance of the 1164

yellow external layer of the wall. 1165

Occurrence: Capitanian-Changhsingian. Palaeo-Tethys and Neo-Tethys. 1166

1167

Genus Ichthyofrondina Vachard in Vachard and Ferrière, 1991 emend. 1168

Type species: Ichthyolaria latilimbata Sellier de Civrieux and Dessauvagie, 1965. 1169


36

Synonyms: Frondina Sellier de Civrieux and Dessauvagie, 1965 (pars); Lingulina (pars); 1170

Pseudoglandulina sp.; Ichthyolaria (pars); Frondicularia Defrance, 1824 (pars). 1171

Emended diagnosis: Test uniseriate. Chambers palmate, strongly embracing, rarely 1172

evolute. Wall hyaline but apparently dark. Aperture simple, terminal, central, with or without 1173

two inner oral tongues. 1174

Composition: Ichthyolaria latilimbata Sellier de Civrieux and Dessauvagie, 1965; I. 1175

nessenensis Bozorgnia, 1973; I. primitiva Sellier de Civrieux and Dessauvagie, 1965; 1176

Frondicularia? arpaensis Pronina in Kotlyar et al., 1989; F. guangxiensis Lin, 1978?; F. 1177

simplex Lin, 1978; F. palmata Wang, 1974; Frondina parvula Pronina in Kotlyar et al., 1989; 1178

Frondina sp. 1 sensu Pronina, 1988 (pl. 2, figs 47-48), and 1989 (pl. 1, figs 23-24). 1179

Occurrence: Late Midian-Changhsinghian, Palaeo-Tethys and Neo-Tethys. (Turkey, Iran, 1180

Transcaucasia, South China, Thailand, New Zealand. Changhsingian South China, North 1181

Caucasus and Transcaucasia). 1182

1183

Ichthyofrondina palmata (Wang, 1974) 1184

Fig. 5(11) 1185

1974 Frondicularia palmata nov. sp. Wang, p. 287, pl. 149, fig. 11. 1186

1978 Frondicularia palmata Wang - Lin, p. 43, pl. 8, fig. 30. 1187

1981 Frondicularia palmata Wang - Zhao et al., pl. 3, figs 17-18. 1188

1988 Frondina palmata (Wang) - Pronina, pl. 2, figs 49-50. 1189

1989 Frondina palmata (Wang) - Pronina, pl. 2, figs 21-22. 1190

p. 1990 Frondicularia palmata Wang - Lin et al., p. 232, pl. 29, figs 12-14 (non figs 15-16 1191

= I. latilimbata). 1192

non 1997 Lunucammina palmata (Wang) - Kobayashi, pl. 3, figs 13-14 (= 1193

Nodosinelloides? sp.). 1194

non 2001 Frondina palmata (Wang) - Pronina-Nestell and Nestell, pl. 2, fig. 19 (another 1195

species of Ichtyofrondina). 1196

non 2001 Lunucammina palmata (Wang) - Kobayashi, pl. 1, figs 23-24 (= 1197

Nodosinelloides? sp., as in 1997). 1198

non 2002 Lunucammina cf. palmata (Wang) - Kobayashi, pl. 3, figs 13-14 (= Frondina or 1199

another species of Ichthyofrondina). 1200

2005 Ichthyofrondina palmata (Wang) - Groves et al., p. 25, figs 22.1-5. 1201

v. 2006 Ichthyofrondina palmata (Wang) - Gaillot, p. 160-161, pl. I.27, fig. 6, pl. VII.1, 1202

fig. 11. 1203


37

Description: Test minute. Palmate chambers, completely enveloping the preceeding ones, 1204

but with the proloculus remaining prominent. Sutures absent. I. latilimbata (Sellier de 1205

Civrieux and Dessauvagie, 1965) and its probable synonym Ichthyofrondina guangxiensis 1206

(Lin, 1978) is larger, with less enveloping chambers. Our specimens are strictly similar to the 1207

I. palmata described by Lin et al. (1990). Height = 0.300 mm, width = 0.180 mm, w/H = 0.60, 1208

number of chambers: 6, proloculus diameter = 0.020 mm, height of last chamber = 0.060 mm, 1209

wall thickness = 0.005 mm. 1210

Occurrence: Lopingian of South-China, Turkey, Zagros and Transcaucasia (Gaillot, 1211

2006). 1212

1213

6. Palaeobiogeographic implications 1214

1215

The palaeogeographic distribution of the biseriamminoids described herein is interpreted to 1216

be typically Neo-Tethyan, ranging from southern Turkey (Hazro) to South China (Laren), and 1217

even reaching Japan for some genus (i.e., Paradagmaritopsis). 1218

Kobayashi (2004) indicated the presence of Paradagmaritopsis in Japan, under the name 1219

of Paradagmarita sp., together with two important biogeographical markers: Globivalvulina 1220

sp. and Partisania sp., respectively. The recent discovery of these taxa in the Late Permian 1221

strata of the Arabian Plate (Gaillot, 2006) suggests that many taxa, which are supposed to be 1222

limited to the western Neo-Tethys, may in fact migrate towards Panthalassa (i.e., Japan). 1223

Those taxa appear to be apparently absent in other well-known Tethyan areas such as 1224

Oman, Afghanistan or Pakistan, as well as South China or Thailand. The possibilities which 1225

may explain the lack of these taxa in several regions are: (a) gaps in the geological record; (b) 1226

fossil preservation (taphonomy); (c) unfavourable depositional environments and/or substrates 1227

and (d) disappearance of the original area by subduction. The lack of data is evidently very 1228

important because it could completely avoid the palaeobiogeographical reconstructions. 1229

As put forward by various authors however, some Upper Permian biogeographic markers 1230

are useful to constrain palaeobiogeography; they are Shanita (e.g., Sengör et al., 1988; Ueno, 1231

2003), Palaeofusulina and Colaniella (e.g., Kobayashi, 1999). In addition, the distribution of 1232

Globivalvulinidae, and especially of Paradagmarita (Sengör et al., 1988; Gaillot and 1233

Vachard, 2004), re-inforces the palaeobiogeographic provinces previously proposed. They 1234

indicate that the BaoShan Block (Yunnan) and Lhassa Block (Xizang) are related to the Peri-1235

Gondwan border and/or to another ancient plate, more or less directly linked to Gondwana. 1236

According to the data provided by literature, microfaunal comparisons with foraminifers of 1237


38

Padang (Sumatra) and Salt Range, as well as of Lamayuru (close to the Indus Suture) are 1238

difficult to establish because of the differences on ages and assemblages. The North Chinese 1239

Tarim Basin and Kun-Lun Mountains are related to the Peri-Hercynian Block based on the 1240

palaeobiogeographic distribution of Viséan foraminifers and calcareous algae, as well as on 1241

the presence of Eopolydiexodina (e.g., Vachard and Bouyx, 2002). 1242

South China and Indochina Blocks are in Extra-Gondwanan (or Cimmerian: Sengör, 1979, 1243

or Extra-Hercynian new name) location. Well constrained in latitude (Ziegler et al., 1997), 1244

these two blocks are conversely not constrained in longitude and could be located more 1245

westward and almost in connection with Iran (see also the palaeomagnetic data of Besse et al., 1246

1998). A more western location of these blocks may well also explain the striking similarities 1247

between Vietnam and Greece, both studied at the beginning of the twentieth century by 1248

Deprat (1912) (see also Théry et al., 2007). 1249

Considering that the blocks of North China and Mongolia have undergone similar 1250

westward displacement, we obtain a Pangea scheme, applicable to the Carboniferous and 1251

Permian distribution of foraminifers and calcareous algae (Gaillot, 2006). 1252

In this configuration, the oceans precluding the collisional belts have been narrow oceans or 1253

sea ways, comparable to recent Red Sea. Fossils distribution emphasizes that during the Late 1254

Permian, the Zagros (Iran), Taurus (Turkey), South China and even Japan shared similar 1255

foraminiferal assemblages and represented intermittently connected palaeobiogeographic 1256

provinces. However, it is to be noted that neither Shanita nor Monodiexodina have been found 1257

in the Laren material. 1258

A second possible hypothesis could take into account the model of an ever-green-Pangea, 1259

also characterised by narrow oceans. This palaeogeographic configuration had existed from 1260

Cambrian to Early Jurassic explaining the different palaeogeographic reconstructions that 1261

followed each other all over this interval of time. The lack of oceanic bottom remnants is 1262

easily explained by the evolution through time of different Tethyan basins, which 1263

continuously originate, or vanish, from refitting and remodelling of land masses. 1264

The foraminiferal migration ways and the direction of migration are poorly understood due to 1265

the gaps indicated above and particularly due to the paucity of data in Himalaya. More likely, 1266

the two palaeo-provinces: Taurus-Zagros-Oman-Saudi Arabia-south and central Afghanistan, 1267

and western Myanmar-western Thailand-Bao Shan, could directly be connected with South 1268

China as well as with Indochina (Fig. 8). Hence, the Middle-Late Permian distribution of 1269

foraminifers and calcareous algae suggests that no major Pangea break-up, without any far 1270


39

migration of tectonic plates, occured since the end of the Early Permian, after the formation of 1271

collisional suture of Urals. 1272

1273

7. Conclusions 1274

The exhaustive study and inventory of the foraminiferal assemblage enclosed within the 1275

beds preceding the Permian-Triassic boundary in the Laren section (Guangxi Province, South 1276

China) allow us to highlight the following features: 1277

1. The Laren section illustrates an excellent outcrop straddling the Permo-Triassic 1278

boundary (Galfetti et al., 2007). 1279

2. The latest Permian is mainly characterized by shallow-water calcisponge reef 1280

limestones. The bioclastic carbonate microfacies contain a rich and well diversified 1281

foraminiferal and algae microfauna. 1282

3. The existence of new phylogenetic lineages among the biseriamminoids foraminifera is 1283

evidenced. They concern the genera Retroseptellina, Septoglobivalvulina, 1284

Paraglobivalvulinoides, Louisettita, Paradagmaritopsis nov. gen. and Paradagmarita. For 1285

South China, the main implication is that the phylogeny of the new herein described 1286

subfamily Paradagmaritinae represents the main framework for the latest Permian (Lopingian) 1287

biozonation. 1288

4. The Persian Gulf possibly acted as a radiative pole for the Paradagmaritinae. From there, 1289

a pool of initially endemic species intermittently spread towards relatively closed 1290

palaeobiogeographic domains. In South China, up to Japan, the main correlable event is 1291

represented by the lineage Globivalvulina/Paradagmarita transitional forms to 1292

Paradagmaritopsis kobayashii nov. gen. nov. sp. 1293

5. Survivors of superfamilies Palaeotextularioidea and Tetrataxoidea, have appeared since 1294

the Early Carboniferous, such as Tetrataxis and Climacammina, are relatively common up to 1295

the PTB. 1296

6. Algae, miliolids and nodosarioids are poorly represented. 1297

7. Six taxa are newly described: Globivalvulina curiosa nov. sp., Louisettita ultima nov. 1298

sp., Bidagmarita nov. gen., Bidagmarita sinica nov. gen. nov. sp., Paradagmaritopsis nov. 1299

gen., and Paradagmaritopsis kobayashii nov. gen. nov. sp. 1300

8. Paradagmarita, Paradagmaritopsis and Paradagmarita? constitute the subfamily 1301

Paradagmaritinae, which parallel evolves with the globivalvulinid families: 1302

Globivalvulininae, Dagmaritinae and Paraglobivalvulinae. 1303


40

9. The palaeogeographic distribution of the biseriamminoids is typically Neo-Tethyan, 1304

spanning from southern Turkey (Hazro) to South China (Laren); Paradagmaritopsis even 1305

attains Japan. Consequently, during the Late Permian, Zagros (Iran), Taurus (Turkey), South 1306

China and even Japan shared similar foraminiferal assemblages and represented intermittently 1307

connected palaeobiogeographic provinces. 1308

10. The migration ways and the direction of migrations are poorly understood essentially 1309

due to the paucity of data in Himalaya. Nevertheless, some key-areas such as Thailand, 1310

Burma, South China, Afghanistan, Pamir, Salt Range and Oman, should provide further data 1311

to figure out. 1312

1313

1314

Acknowledgements 1315

1316

We wish to thank the Geobios reviewers Prof. F. Kobayashi and the anonymous reviewer for 1317

constructive remarks and criticism during the review process. Prof. J. Bucur contributed for 1318

the identification of Lower Triassic algae. This work was partially supported by the Swiss 1319

National Science Foundation (RM project 200021-113816). 1320

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Vachard, D., Gaillot, J., Pille, L., Blazejowski, B. 2006. Problems on Biseriamminoidea, 1920

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Vaslet, D., Le Nindre, Y.M., Vachard, D., Broutin, J., Crasquin-Soleau, S., Berthelin, M., 1923

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Guangxi, Southwest China. Albertiana 33, 103-104. 1971

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The Changhsingian and Permian-Triassic boundary of South-China. Bulletin Nanjing 1973

Institute Geology and Paleontology, Academia Sinica 2 (4), 58-69 (in Chinese with 1974

English abstract). 1975

Ziegler, A.M., Hulver, M.L. Roeley, D.B., 1997. Permian world topography and climate. In: 1976

Martini, I.P. (Ed.) Late glacial and postenvironmental changes - Quaternary, 1977

Carboniferous-Permian and Proterozoic. Oxford Universty Press, New York, 111-1978

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Zolotova, V.P., Baryshnikov, V.V., 1980. Foraminifery kungurskogo yarusa stratotipicheskoi 1980

mestnosti (Foraminifers of the stratotype locality of Kungurian stage). In: Rauzer-1981

Chernousova, D. M., Chuvashov, B.I. (Eds.) Biostratigrafiya artinskogo i 1982

kungurskogo yarusov Urala (Biostratigraphy of the Artinkian and Kungurian stages 1983

from Urals). Akademiya Nauk SSSR, Uralskii Nauchnyi Tsentr, 72-109. 1984

1985 1986 Figure captions 1987

1988

Fig. 1. Geographic map of the Western Guangxi Province with location of the studied areas 1989

and distribution of Late Permian platforms (modified after Rigby et al. 1989b). 1990

1991

Fig. 2. Outcrop photograph of the detailed Permo-Triassic boundary at Laren section. Note 1992

the contrasting lithologies between the Late Permian calcisponge reefs of Wujiaping Fm. and 1993

the Early Triassic calcimicrobial limestones of Luolou Fm.; scale = 6 m. 1994

1995

Fig. 3. Field photographs showing latest Permian (late Changhsingian) reef limestones 1996

associated with the calcisponges bioconstruction of Laren. (a) Calcisponges in life position; 1997

scale = 5 cm. (b) Longitudinal section of the calcisponges; scale = 5 cm. (c) Transverse 1998

sections of the same calcisponges; scale = 2,5 cm. (d) Sponges of the genus Discosiphonella 1999

and silicified brachiopods; scale = 1,25 cm. 2000

2001

Fig. 4. Thin-section photomicrographs of the bioclastic wacke/packstones associated with the 2002

calcisponge reefs. (a) Neosparitized bioclastic limestone showing Neodiscus sp., 2003


61

Climacammina sp. (several sections) and Tetrataxis sp. Thin section L10b (coll. T. Galfetti); 2004

scale 500 µm. (b) Similar microfacies showing Paraglobivalvulinoides sp., Climacammina 2005

sp. and Neodiscus sp. Thin section L10b (coll. T. Galfetti); scale = 500 µm. (c) Neosparitized, 2006

slightly silicified, bioclastic and peloidal wackestone showing Climacammina sp. and 2007

Nankinella sp. Thin section L6 (collection T. Galfetti); scale = 500 µm. 2008

2009

Fig. 5. (1). Paraglobivalvulinoides septulifer (Zaninetti and Altiner, 1981). Subaxial section. 2010

Sample L6. (2). Climacammina tenuis Lin, 1978. Sagittal axial section. Sample L6. (3-5). 2011

Postendothyra micula (Sosnina in Sosnina and Nikitina, 1977). 3. Axial section. Sample L3. 2012

4. Axial section. Sample L6. 5. Transverse section. Sample L6. (6). Reichelina simplex Sheng, 2013

1956. Subaxial section. Sample L3. (7). Eotuberitina spinosa (Lys in Lys et al., 1980), 2014

Transverse section. Sample L10. (8). Tetrataxis lata Spandel, 1901. Subaxial section. Sample 2015

L3. (9). Pachyphloia pedicula Lange, 1925. Axial section. Sample L3. (10). Pseudotristix sp. 2016

Subaxial section difficult to separate from a Geinitzina but here associated in Fig. 7(3), and 2017

differing from the other true Geinitzina of the same thin section by the sises of the test and the 2018

chamber. Sample L10. (11). Ichtyofrondina palmata (Wang, 1974). Axial section. Sample L6. 2019

(12). Nankinella cf. inflata (Colani, 1924) emend. Sheng, 1963. Recrystallised axial section. 2020

Sample L10. 2021

2022

Fig. 6. (1). Globivalvulina bulloides (Brady, 1876). Transverse section. Sample L3. (2). 2023

Septoglobivalvulina distensa (Wang in Zhao et al., 1981). Transverse section. Sample L3. (3). 2024

Septoglobivalvulina sp. 1. Transverse section. Sample L10. (4). Retroseptellina nitida (Lin, Li 2025

and Sun, 1990). Subtransverse section. Sample L10. (5). Retroseptellina decrouezae 2026

(Köylüoglu and Altiner, 1989). Subtransverse section. Sample L3. (6-8, 12-14, 15?, 16?). 2027

Paradagmaritopsis kobayashii Gaillot and Vachard nov. gen. nov. sp. 6. Holotype. 2028

Transverse section. Sample L10. 7. Paratype. Axial section. Sample L3. 8. Paratype. Subaxial 2029

section. Sample L6. 12. Paratype. Axial section. Sample L10. 13. Paratype. Transverse 2030

section. Sample L3. 14. Paratype. Oblique section. Sample L10. 15. Paratype? Anatypical 2031

large subaxial section. Sample L6. 16. Paratype? Anatypical long coiled initial part. Sample 2032

L6. (9-10). Paradagmaritopsis sp. 9. Subaxial frontal section. Sample L6. 10. Sagittal axial 2033

section. Sample L6. (11, 17). Paradagmarita sp. Two subtransverse sections. Sample L10. 2034

(18). Dagmarita? cf. sharezaensis Mohtat-Aghai and Vachard, 2003. Oblique section. Sample 2035

L10. (19). Dagmarita altilis Wang in Zhao et al., 1981. Axial section. Sample L3. (20-25). 2036

Louisettita ultima Gaillot and Vachard nov. sp. 20. Holotype. Oblique section showing the 2037


62

endoskeleton. Sample L10. 21. Paratype. Oblique sagittal section; the wall of the last chamber 2038

is finely perforated; this particularity exists also in dagmaritins of Turkey (Altiner, 2039

unpublished data). Sample L10. 22. Paratype. Oblique section. Sample L6. 23. Paratype. 2040

Oblique section showing small lateral chamberlets. Sample L10. 24. Paratype. Oblique 2041

section. Sample L6. 25. Paratype. Oblique section showing the curvature of the septa and the 2042

lateral chamberlets. Sample L10. 2043

2044

Fig. 7. (1-2). Ramovsia? sp. 1. Recrystallized longitudinal section attached on a gastropod 2045

(right). Sample L3. 2. Longitudinal section with more typical wall, differing from 2046

Earlandia/Aeolisaccus by the size and the asymmetry. Sample L3. (3). Pseudotristix cf. solida 2047

Reitlinger, 1965. Transverse section. Sample L3. (4-5). Bidagmarita sinica nov. gen. nov. sp. 2048

4. Holotype. Sagittal axial section. Sample L3. 5. Paratype.Frontal axial section. Sample L3. 2049

(6, 8). Asterosphaera sp. Two sections showing the type of wall and the crenulata inner 2050

periphery characteristic of the genus according to Vachard and Clément, 1994. Sample L6. 2051

(7). Radiosphaerina? sp. The outer periphery is more irregular and the inner periphery not 2052

circular, two differences with a transverse section of a costulate nodosarioid. Sample L3. (9-2053

10). Asterosphaera cf. pulchra Reitlinger, 1957. 9. A typical tangential section. Sample L6. 2054

10. Very rare type of section showing two connected specimens. Sample L3. (11). 2055

Rectostipulina cf. quadrata Jenny-Deshusses, 1985. A very rare octogonal transverse section. 2056

Sample T0. (12). Reichelina simplex Sheng, 1956. Axial section in the coiled part. Sample T0. 2057

(13). Donezella sp. A probably new species with a large thallus without visible bifurcation, 2058

but exhibiting the typical wall. Sample L6. (14). Septoglobivalvulina cf. guangxiensis Lin, 2059

1978. Axial section. Reworked Wuchiapingian. Sample T0. (15). Sphaerulina sp. Transverse 2060

section. Sample T0. (16-21). Globivalvulina curiosa nov. sp. 16. Holotype. Transverse 2061

section. 17. Paratype. Subtransverse section. 18. Paratype. Oblique section. 19. Paratype. 2062

Oblique section. 20. Paratype. Subaxial section. 21. Paratype. Transverse section. Sample T1. 2063

2064

Fig. 8. Palaeogeographic reconstruction and location of Paradagmarita Subprovince (light 2065

grey). 1: Italy; 2: Tunisia; 3: Ex-Yugoslavia; 4: Albania; 5: Hungary; 6: Greece; 7: Cyprus; 8: 2066

Crimea; 9: Caucasus; 10: Anatolia; 11: Taurus; 12: Oman-Saudi Arabia; 13: Zagros; 14: 2067

Alborz; 15: Afghanistan; 16: Salt Range; 17: SE Pamir; 18: Himalaya; 19: Kashmir; 20: 2068

Tibet; 21: South China; 22: Thailand-Burma; 23: Malaysia; 24: Indosinia; 25: Primorye-2069

Koryak; 26: Japan; 27: “Kobayashi block” (according to Kobayashi, 2004). After Gaillot, 2070

2006, modified. 2071

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