Permian smaller foraminifers: taxonomy, biostratigraphy and biogeography

Daniel Vachard

This review has been undertaken in order to present some interpretations about the bio-stratigraphy of the smaller foraminifers belonging to four classes present during the Permian: Fusu-linata, Miliolata, Nodosariata and Textulariata. Biostratigraphic markers of these classes are principally known in the orders and superfamilies of Lasiodiscoidea, Bradyinoidea and Globival-vulinoidea (Fusulinata), Cornuspirida (Miliolata), and in the entire class Nodosariata. The class Textulariata is too little known during the Permian to play a significant biostratigraphical role: nevertheless , the appearance of the order Verneulinida is probably an important bioevent. The main genera among the lasiodiscids are Mesolasiodiscus,

Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 Permian smaller foraminifers: taxonomy, biostratigraphy and biogeography DANIEL VACHARD 1 rue des Tilleuls, 59152 Gruson, France Daniel.Vachard@univ-lille1.fr Abstract: This review has been undertaken in order to present some interpretations about the biostratigraphy of the smaller foraminifers belonging to four classes present during the Permian: Fusulinata, Miliolata, Nodosariata and Textulariata. Biostratigraphic markers of these classes are principally known in the orders and superfamilies of Lasiodiscoidea, Bradyinoidea and Globivalvulinoidea (Fusulinata), Cornuspirida (Miliolata), and in the entire class Nodosariata. The class Textulariata is too little known during the Permian to play a significant biostratigraphical role: nevertheless, the appearance of the order Verneulinida is probably an important bioevent. The main genera among the lasiodiscids are Mesolasiodiscus, Lasiodiscus, Lasiotrochus, Asselodiscus, Pseudovidalina, Xingshandiscus and Altineria; the bradyinoids Bradyina and Postendothyra; the globivalvulinoids Globivalvulina, Septoglobivalvulina, Labioglobivalvulina, Paraglobivalvulina, Sengoerina, Dagmarita, Danielita, Louisettita, Paradagmarita, Paradagmaritopsis and Paremiratella; the miliolates Rectogordius, Okimuraites, Neodiscus, Multidiscus, Hemigordiopsis, Lysites, Shanita and Glomomidiellopsis, and the tubiphytids and ellesmerellids, which might be specialized miliolate and cyanobacterium consortia, with reference to microstructures and phylogenies of these groups. The Nodosariata markers belong to Nodosinelloides, Tezaquina, Polarisella, Geinitzina, Pachyphloia, Rectoglandulina, first true Nodosaria, Langella, Pseudolangella, Calvezina, Cryptoseptida, Cylindrocolaniella, Colaniella, Frondina and Ichthyofrondina, but their lineages are too poorly understood to permit an accurate biostratigraphic use at the present time. The superfamily Geinitzinoidea is emended. Finally, palaeobiogeographical implications based on Shanita, Colaniella and Altineria are given. During the nineteenth century and the first third of the twentieth century, there were few works devoted to the smaller foraminifers, except perhaps for the Nodosariata. From 1900 to 1930, some papers were published by Spandel (1901), Cherdyntsev (1914), Lange (1925) and Likharev (1939), and in the USA by Cushman, Galloway, Waters and Harlton. In the 1940s, a fundamental contribution was provided by Reichel (1945, 1946), especially in Greece and Cyprus. This author was present at the beginning of the Swiss school, with workers Brönnimann, Zaninetti, Martini and Jenny-Deshusses. D. Altıner, trained by this school, has written since the 1980s a fundamental work about the Permian lasiodiscoid, hemigordiid and globivalvulinoid foraminifers. Other micropalaeontologists working in Turkey also provided important contributions: Güvenç, Sellier de Civrieux, Dessauvagie, Lys and Okuyucu. In the 1960s, Russian studies predominated, with A.D. and K.V. Miklukho-Maklay, Reitlinger, Sosnina, Gerke, Kireeva, and Sosipatrova. In the 1980s, Japanese teams studied the palaeontology of their country, as well as in Iran, Pakistan and Thailand. During this century, the Russian school for the Permian smaller foraminifers has been reactivated by Filimonova (2008, 2010). G. Nestell, also known as Pronina, Vuks or Pronina-Nestell, wrote several very important studies (despite her questionable foraminiferal macroclassification, from families to classes). Representatives of the French and Italian schools were productive all around the world, including Lys, Vachard, Rettori, Angiolini and Gaillot (in chronological order). Chinese colleagues, Gu, Song and Zhang, published important recent contributions, especially about the Permian –Triassic boundary foraminifers, extending the classical studies of Ho, Lin and Wang. Although considered as a subordinate biostratigraphic group, the smaller foraminifers may contribute to the biozonation of the carbonate and evaporite shallow environments with a wide distribution of the representatives of the Miliolata (e.g. Vachard et al. 2005; Gaillot & Vachard 2007; Koehrer et al. 2012 for the Khuff Formation in the Middle East; and Lucas et al. 2015; Vachard et al. 2015 for the Yeso Group of southern USA) or to deeper environments with the distribution of representatives of the Nodosariata (e.g. Gu et al. 2007; Zhang & Gu 2015) in South China; it is noteworthy that these shallowest and deepest biotopes were occupied only by foraminifers from the Permian (Vachard et al. 2010). From: Lucas, S. G. & Shen, S. Z. (eds) 2018. The Permian Timescale. Geological Society, London, Special Publications, 450, 205– 252. First published online December 8, 2016, https://doi.org/10.1144/SP450.1 # 2018 The Author(s). Published by The Geological Society of London. All rights reserved. For permissions: http://www.geolsoc.org.uk/permissions. Publishing disclaimer: www.geolsoc.org.uk/pub_ethics Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 206 D. VACHARD Previous classifications of foraminifers The current classifications of foraminifers are difficult to reconcile, both at the genus level and at the suprageneric level. However, the older classifications based on the wall microstructure have been challenged by the data and suggestions of molecular clocks (Rigaud et al. 2015b with references therein). In this biostratigraphic and practical paper, the wall microstructure is preserved as a preponderant criterion, even if the original microstructures are often difficult to identify because of the recrystallization of walls consisting of aragonite and high-Mg calcite; even low-Mg calcite transforms itself. The agglutinates are difficult to distinguish because of finely granular siliceous recrystallization. Initially, along with J. Gaillot, in trying to find a true difference between the classes Fusulinata and Textulariata, we separated the Fusulinata with calcareous agglutinates from the Textulariata with siliceous agglutinate (Gaillot & Vachard 2007; Vachard et al. 2010); however, it was observed that some representatives of the Fusulinata have a true siliceous agglutinated wall (e.g. palaeotextulariids, endothyranellins and endotebids). Currently, this subject is being examined by S. Rigaud (Rigaud et al. 2014, 2015a, b). The foraminiferal molecular clock developed notably by J. Pawlowski (e.g. Pawlowski et al. 2013) is difficult to follow because it does not correspond to available observations in sampled materials. Even if the interest of this molecular classification is to be theoretical, it is manifestly incompatible with the fossil record in Palaeozoic rocks. Some consensual syntheses have been presented in Langer et al. (1993) and Rigaud et al. (2014, 2015a, b). Despite these divergent concerns, one palaeontological observation is, nevertheless, consistent with the molecular data: the transition from an agglutinated to a calcareous wall, and vice versa, occurred several times in the foraminiferal evolution, and in independent lineages (Bowser et al. 2006; Pawlowski et al. 2013): for example, when Psammosphaera (agglutinated) give rise to miliolates (porcelaneous), as well ammodiscids (agglutinated) (Pawlowski et al. 2013), and the miliolates (porcelaneous) in turn give Miliammina (agglutinated) (Fahrni et al. 1997). Here, a proposed classification is developed according to the phylogenetic hypotheses of Vachard (1994), Gaillot & Vachard (2007), Vachard et al. (2010) and Hance et al. (2011). It differs significantly from previous systematics proposed by Mikhalevich (1980, 2013) and Loeblich & Tappan (1987, 1992) . The taxonomic rank of the foraminifera (phylum or subphylum) is still a matter of debate (see e.g. Adl et al. 2012) but, herein, the views of Cavalier-Smith (2002, 2003) are adopted. Even if macroevolution remains very questionable, the lower units of the classification, especially at the generic level, seem to be objective and can be used confidently with their FAD (first appearance datum: i.e. the oldest of the successive local appearances of a taxon) and LAD (last appearance datum: i.e. the youngest of the successive disappearances of a taxon) for the biozonation. A complete taxonomical reference list would be huge and, therefore, the reader can refer to Loeblich & Tappan (1987), Vdovenko et al. (1993), Rauzer-Chernousova et al. (1996), Pronina-Nestell & Nestell (2001), M. K. Nestell et al. (2006), Gaillot & Vachard (2007), Karavaeva & Nestell (2007), G. P. Nestell et al. (2009) and Vachard et al. (2015). Previous biostratigraphical data The recent Permian scale was established thanks to the work of Jin et al. (1997). For a long period of time, two parallel scales for the foraminifers have existed: one for the Tethyan fusulinids (Leven 1967; Leven & Gorgij 2011), and another one for the Omolon Massif (NE Siberia) nodosariates (Pronina 1999a; Karavaeva & Nestell 2007). Both are, however, difficult to apply to other areas because of: (a) the regionalism of the two groups; (b) the development of the foraminiferal classifications; (c) the progress of the correlations with the conodonts scales; (d) the problems of definition of the Leven’s stratotypes (Angiolini et al. 2015, 2016); (e) the introduction of concurrent regional stages (e.g. Hermagorian by Davydov et al. 2013); and (f ) the precise, new correlations between the different biozonations (Henderson et al. 2012). The smaller foraminifers are actually little used, but four recent types of investigations could develop their knowledge: (1) the smaller foraminifers associated with the fusulinids described by Leven (1967, 1998, 2003) are currently revised by Filimonova (2010, 2013), even if some parts of these contributions may provoke questions (see Angiolini et al. 2015, 2016); (2) the microfaunas of smaller foraminifers associated with endemic North American fusulinids (e.g. Polydiexodina) have shown that more Tethyan smaller foraminifers than expected are present in Mexico, New Mexico and Texas (Vachard et al. 1992; Nestell & Nestell 2006; Nestell et al. 2006), and needs further investigation in other states of the USA and countries in the Americas; (3) the potential of the nodosariates was re-demontrated by Karavaeva & Nestell (2007); and (4) areas, unfavourable for fusulinids because of their deposits and types of environments, have been dated and biozoned recently thanks to smaller foraminifers (Lucas et al. 2015; Vachard et al. 2015). Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 PERMIAN SMALLER FORAMINIFERS Important papers on smaller foraminifers have recently been published: relating especially to understanding the Permian–Triassic Boundary (PTB) and its extinctions in central and eastern Tethys (i.e. South China and SE Asia: Fig. 1): Altıner (1999, 2001), Groves & Altıner (2005), Groves et al. (2005), Gaillot & Vachard (2007), Gaillot et al. (2009), Altıner & Özkan-Altıner (2010), Ueno et al. (2010), Wang et al. (2010), Kobayashi (2012a, b, 2013), Song et al. (2007, 2011, 2013) and Nestell et al. (2015). The foraminiferal biostratigraphic scales are generally based on fusulinids (e.g. Leven 2003; Wilde 2006; Leven & Gorgij 2011; Henderson et al. 2012). After some attempts to establish the stratigraphical significance of smaller foraminifers, pioneering detailed biozonations have been proposed for the late Middle Permian and Late Permian (Capitanian–Lopingian) by Vachard et al. (1993a, b, 2002) and Pronina (1996). First biozonations by smaller foraminifers for the Cisuralian of North America have been published very recently (Lucas et al. 2015; Vachard et al. 2015). Proposed classification of Permian smaller foraminifers Phylum Foraminifera d’Orbigny, 1826 nom. translat. Cavalier-Smith, 2002 (subphylum) and 2003 (phylum) Class Fusulinata Gaillot & Vachard, 2007 207 The members of the class Fusulinata have a homogeneous, microgranular, primary test wall of low-Mg calcite in which crystal units are optically unordered, more or less equidimensional and are only a few micrometres in size. The class Fusulinata contains eight orders: (1) Parathuramminida Bykova in Bykova & Polenova, 1955; (2) Tuberitinida Vachard et al., 2015; (3) Earlandiida Loeblich & Tappan, 1982; (4) Archaediscida Poyarkov & Skvortsov, 1979 emend. Hance et al., 2011; (5) Pseudopalmulida Mikhalevich, 1993; (6) Tournayellida Dain in Dain & Grozdilova, 1953; (7) Endothyrida Fursenko, 1958 (including Palaeotextulariina Hohenegger & Piller, 1975); and (8) Fusulinida Fursenko, 1958. It should be noted that this latter order is the only one designated less formally as fusulines, fusulinids, fusulinoideans or fusulinaceans, and is conventionally separated in the foraminiferal treatises from the smaller foraminifers (or non-fusulines) and often even described by other authors. The Permian smaller foraminifers also include representatives of the classes Miliolata, Nodosariata and Textulariata (see later). The Permian smaller foraminifers of the class Fusulinata belong to four orders: Tuberitinida, Earlandiida, Archaediscida and Endothyrida. Regarding Archaediscida, it is noteworthy that their Permian representatives only belong to the superfamily Lasiodiscoidea. In spite of this, many genera and even species of Archaediscoidea (e.g. Permodiscus, Archaediscus, Kasakhstanodiscus: see Davydov 1988) have been mentioned from the Permian in Russia, Tajikistan Fig. 1. Location and toponymy of the main Permian areas in the world (Transcaucasus, or Transcaucasia, were the former Soviet names for a geopolitical region encompassing all of Armenia and the majority of Georgia and Azerbaijan; the classical Permian outcrops of ‘Transcaucasia’ are at this time located in Armenia and Nakhchivan, an enclave of Azerbaijan; e.g. see Théry et al. 2007, fig. 3a). Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 208 D. VACHARD and Uzbekistan. Although Pinard & Mamet (1998) have refuted this assignment and demonstrated that all these forms are, in reality, pseudovidalinids, the misinterpretations continue. For instance, an archaediscoid Permodiscus sumsariensis, which is a marker for the latest early Visean (biozone MFZ11B: Okuyucu et al. 2013), has recently been misinterpreted in the Cisuralian by Filimonova (2010). Definitively, the Archaediscoidea disappear in the lowermost early Moscovian (earliest Middle Pennsylvanian) and do not exist in the Permian. The name Fusulinata Gaillot & Vachard, 2007 is valid for designating this class: even if Fusulinata was introduced by Maslakova (1990) and made explicit in Maslakova et al. (1995), it was, in both articles, for a subclass (because the unique class recognized in both articles is Foraminifera) and, moreover, without any description and/or discussion. In contrast, the name of the subclass introduced by Maslakova (1990) has priority over Fusulinana Vachard et al., 2010 (see below), even if the spelling Fusulinana is more correct than the original Fusulinata (compared to the endings ana and ata used in the classifications of, e.g., V. Mikhalevich and G. Nestell). Subclass Afusulinana Vachard et al., 2010 Order Tuberitinida Vachard et al., 2015 Family Tuberitinidae Miklukho-Maklay, 1958 Permian genera. Tuberitina Galloway & Harlton, 1928; Eotuberitina Miklukho-Maklay, 1958 (¼Diplosphaerina Derville, 1952 part.); Mendipsia Conil & Longerstaey in Conil et al., 1980. Remarks. The Tuberitinidae are calcareous microfossils that have two stages of their cycle of life: diplospherin (free) and tuberitin (attached) (Conil et al. 1977). Diplosphaerina sensu stricto being the diplospherin stage of Eotuberitina, both genera are probably synonymous, and the former genus has priority over the second one. Despite their atypical palaeobiological characters, the tuberitinids probably belong to the foraminifers of the Afusulinana subclass due to their numerous, minute foramina and their wall microstructures, which are often dark-microgranular and hyaline-pseudofibrous, but this assignment may be discussed. The Afusulinana differ from the Parathuramminida by their biphased cycle of life, less conspicuous apertures and absence of homeomorphies with extant foraminifers (the Parathuramminida being homeomorphs of the extant and fossil genera, e.g. Thurammina Brady). According to Vachard (1994) and Vachard et al. (2015), the Tuberitinidae constitute a homogeneous group. The genus Polysphaerinella Mamet differs from this group by its bilayered test and, thus, could belong to another family or subfamily. Moreover, other phylogenetic trends, depending on distinct types of walls, exist in the subfamilies Tubeporininae Zadorozhnyi & Yuferev and Altjusellinae Zadorozhnyi & Yuferev, and the genus Tubesphaera Vachard. All these phylogenetic trends could be considered as different families in the order Tuberitinida. Biostratigraphy. The tuberitinids are not very useful for Permian biostratigraphy. An interesting thing to note is their survival after the PTB (Song et al. 2007, 2011; Okuyucu et al. 2014). Biogeography. The tuberitinids have a cosmopolitan distribution from Late Devonian to latest Permian times, and the last representatives of the earliest Triassic are currently only known in South China (Song et al. 2007, 2009, 2011) and Turkey (Okuyucu et al. 2014). Order Earlandiida Loeblich & Tappan, 1982 Superfamily Earlandioidea Loeblich & Tappan, 1982 Family Earlandiidae Cummings, 1955 emend. Vachard, 1994 Permian genera. Earlandia Plummer, 1930 (¼Aeolisaccus Elliott, 1958 ¼ Decastronema Golubic et al., 2006 (part.) ¼ Hyperammina Brady, 1878 (part.)); ?Chitralina Angiolini & Rettori, 1995; ?Giraliarella Crespin, 1958; ?Rectoformata Okuyucu, 2007. Remarks. Some tubular, undivided microfossils from the Permian-Triassic times are discussed, but are often synonymized with Earlandia. The species Aeolisaccus dunningtoni Elliott, 1958 seems to be a representative of the group Earlandia elegans, in which the proloculus would be inconspicuous and/or broken (Gaillot & Vachard 2007 and references therein). Decastronema Golubic et al., 2006, by its type species, is a junior synonym of the Cambrian –Triassic cyanobacterium Proaulopora Vologdin, 1932, but Decastronema also includes some Earlandia among its other species. However, many so-called Palaeozoic representatives of ‘Hyperammina’ (non Hyperammina Brady, 1878, which is an extant genus) generally correspond to secondary silicified Earlandia, and are interesting elements for discussing the secreted, against agglutinated, types of walls (see Vachard et al. 2010, 2015; Nestell et al. 2015; and see ‘Textulariata’ later in this section). Biostratigraphy. The family is known throughout the Palaeozoic and might survive up to the Cretaceous with the taxa Earlandia? and Giraliarella? of Arnaud-Vanneau (1980): for a discussion on this, see Vachard et al. (2010), Krainer & Vachard (2011), Hance et al. (2011) and Nestell et al. (2015). There was a widespread conviction that Earlandia was a Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 PERMIAN SMALLER FORAMINIFERS PTB survivor (e.g. Groves & Altıner 2005; Groves et al. 2005, 2007; Vachard et al. 2010; Krainer & Vachard 2011); this fact, nevertheless, has been recently disputed (Nestell et al. 2015). Chitralina and/or Giraliarella (two genera to revise and probably emendate) might be the ancestors of Rectostipulina Jenny-Deshusses, 1985 and/or Tubulastella Rigaud et al., 2015b (see below), as well as Earlandia being the ancestor of the Nodosariata for many authors (Hohenegger & Piller 1975; Vachard 1994; Altıner & Savini 1997; Groves 1997; Groves et al. 2003; Vachard et al. 2010; Krainer & Vachard 2011; Rigaud et al. 2015b). Rectoformata, known in Turkey (Okuyucu 2007) and South China (Zhang et al. 2015), might be biostratigraphically important for dating the Tethyan assemblages. Biogeography. Probably cosmopolitan during the Palaeozoic, the family seems then to be restricted to some Tethyan areas. Superfamily Caligelloidea Reitlinger in RauzerChernousova & Fursenko, 1959 nom. translat. Gaillot & Vachard, 2007 Family Insolentithecidae Loeblich & Tappan, 1986a nom. translat. Gaillot & Vachard, 2007 209 Permian genera. Lasiodiscus Reichel, 1946; Hemidiscus Schellwien, 1898 emend. Vachard & Krainer, 2001a; Lasiotrochus Reichel, 1946; Mesolasiodiscus Rauzer-Chernousova & Chermnykh, 1990. Remarks. This group has been largely neglected after the fundamental papers of Reichel (1946) and Reitlinger (1956). Recent studies of the ancestral lasiodiscids and howchiniids (Cózar et al. 2015) demonstrated the maximal complexity of the phylogenies of these groups. Consequently, the apparently well-known Permian lasiodiscids need further studies (1 & 2 in Fig. 2), peculiarly in North America where they are not yet known or are very questionable. Similarly, the Carboniferous representatives mentioned in North America (most often called Monotaxinoides) are misinterpreted miliolates (Krainer et al. 2015; Lucas et al. 2016a, b), except for ‘Monotaxinoides priscus’ sensu Groves, 1992 (although this taxon most probably belongs to Eolasiodiscus), Eolasiodiscus sp. and Mesolasiodiscus? sp. (Lucas et al. 2016b). Biostratigraphy. As the Late Silurian –Late Permian caligelloid genera are poorly known, their precise biostratigraphic contribution remains small. Insolentitheca, which is essentially a late Mississippian– Pennsylvanian genus, is rare in the Early Cisuralian of Bolivia (Mamet 1996), and in the Kungurian of southern Turkey and Tajikistan (Vachard & Moix 2013; Angiolini et al. 2016). Floritheca is only known in the Lopingian of southern Iran (Zagros, Fars), Abu Dhabi (UAE) and the southern Chichibu Terrane (SW Japan) (Gaillot & Vachard 2007). Biostratigraphy. Hemidiscus sensu stricto seems to be restricted to the Cisuralian, but it is difficult to distinguish from the Pennsylvanian genus Eolasiodiscus or Mississippian genus Hemidiscopsis Cózar in Cózar et al., 2015. Mesolasiodiscus was introduced as a Cisuralian transitional taxon between Eolasiodiscus and Lasiodiscus, but it seems to exist as early as the Middle (even Early?) Pennsylvanian: its FAD and LAD are poorly known, apparently late Asselian and Kungurian, respectively (Filimonova 2010). Similarly, the true FAD of Lasiodiscus is poorly known (many Cisuralian references to Lasiodiscus tenuis in the literature are, in reality, Hemidiscus due to the absence of umbilical fillings and sutural appendices), whereas its LAD is clearly latest Permian, perhaps even latest Changhsingian (MiklukhoMaklay 1954; Pronina-Nestell & Nestell 2001; Gu et al. 2007). Lasiotrochus sensu stricto seems to be endemic to the late Guadalupian– early Lopingian, from Greece to South China (many previous identifications are, in reality, misinterpreted Mesolasiodiscus: Vachard et al. 2002, pl. 1, figs 17 & 18; Lucas et al. 2016b; Vachard et al. unpublished data). Biogeography. The insolentithecid genera are either cosmopolitan (as Insolentitheca) or more restricted (as Floritheca). Biogeography. The Permian lasiodiscids, relatively common in the Palaeo- and Neotethys, are absent from North America. Permian genera. Insolentitheca Vachard in Bensaid et al., 1979; Floritheca Gaillot & Vachard, 2007. Remarks. This atypical group of foraminifers, ontogenetically deformed probably because of a particular infaunal life, is principally known during the Carboniferous. Rare taxa subsist in the Permian; especially in the Cisuralian. Order Archaediscida Poyarkov & Skvortsov, 1979 emend. Hance et al., 2011 Suborder Lasiodiscina Gaillot & Vachard, 2007 Superfamily Lasiodiscoidea Reitlinger in Vdovenko et al., 1993 Family Lasiodiscidae Reitlinger, 1956 emend. Gaillot & Vachard, 2007 Family Pseudovidalinidae Altıner, 1988 Permian genera. Pseudovidalina Sosnina, 1978 emend. Alipour & Vachard in Alipour et al., 2013 (¼Raphconilia Brenckle & Wahlman, 1996); Altineria Özdikmen, 2009 emend. Vachard et al., in press; Asselodiscus Mamet & Pinard, 1992; Xingshandiscus Zheng, 1986. Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 210 D. VACHARD Fig. 2. Smaller foraminifers (Fusulinata) from Trogkofel (Late Cisuralian of the Carnic Alps, Austria). Scale bar ¼ 0.100 mm, except for (2) where it is 0.250 mm. (1) Lasiodiscus tenuis Reichel, 1946. (2) Mesolasiodiscus sp. (3) & (4) Transition Asselodiscus –Pseudovidalina? sp. (5)–(11) Pseudovidalina spp. (12) – (17) Spireitlina spp. (18) – (24) Endothyra spp. Remarks. This interesting group, still poorly known during the Permian, might provide, with further studies, more accurate biomarkers for the Tethyan Permian. The Pseudovidalinidae have a double Permian history (the second episode of which was illustrated by Altıner 1988). During the first episode, during the Late Pennsylvanian –Cisuralian, except for rare Asselodiscus (Davydov et al. 2001; Alipour et al. 2013), the genus Pseudovidalina was the unique representative of the family (3–11 in Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 PERMIAN SMALLER FORAMINIFERS Fig. 2). However, during this period, Pseudovidalina displays various evolutionary stages, eventually interpretable as subgenera or genera (Raphconilia is one of these stages), as is the case for Carboniferous homeomorphous archaediscoids (see the earlier remarks about the misinterpretations of pseudovidalinids as archaediscoids and the detailed discussion in Alipour et al. 2013). Altineria (pro Angelina Altıner, 1988 pre-occupied by a Cambrian trilobite) is sometimes considered as a junior synonym of Xingshandiscus (see Pinard & Mamet 1998). This synonymy is irrelevant because Altineria is unilayered pseudofibrous, whereas the type species of Xingshandiscus exhibits a bilayered wall with a dark-microgranular inner layer and a hyaline-pseudofibrous outer layer. Currently, we are revising the taxa of this family with R. Rettori, D. Altıner and V. Gennari. Biostratigraphy. Pseudovidalina sensu lato is rare in the Gzhelian (Henderson et al. 1995; Davydov et al. 2001), relatively frequent within the Carboniferous –Permian boundary interval (¼Orenburgian ¼ Bursumian ¼ Newwellian regional substages) and subsists up to the late Changhsingian. At the end of the Cisuralian, during the Kungurian, a first evolute pseudovidalinid genus called Xingshandiscus appears. During the Middle –Late Permian boundary interval, another evolute trend is represented by Altineria. The Capitanian –Wuchiapingian biostratigraphy and periGondwanan palaeobiogeography of the Altineria lineage need further study. The possible FAD of Altineria might be early? Capitanian in Japan (Kobayashi 2006a reinterpreted), but this event most probably occurs between the LAD of Yabeina Deprat and the FAD of Nanlingella Rui & Sheng (Vachard et al. unpublished data). The type species, Altineria alpinotaurica (Altıner, 1988), is probably limited to the Capitanian –Wuchiapingian boundary interval, and, apparently, no late Wuchiapingian– Changhsingian representatives of the lineage are known. Biogeography. Genera of this family are either cosmopolitan (e.g. Asselodiscus and Pseudovidalina) or limited to the peri-Gondwanan areas (Altineria). Subclass Fusulinana Maslakova, 1990 nom. correct. Vachard et al., 2010 Order Endothyrida Fursenko, 1958 Suborder Endothyrina Bogush, 1985 Superfamily ?Endothyroidea Glaessner, 1945 Family Endothyridae Rhumbler, 1895 Permian genera. Endothyra Phillips, 1846 sensu Brady, 1876 emend. China, 1965; Endothyranella Galloway & Harlton in Galloway & Ryniker, 1930; Linendothyra Mamet & Pinard, 1992; Neoendothyra Reitlinger, 1965; Neoendothyranella 211 Nestell & Nestell, 2006; and Planoendothyra Reitlinger in Rauzer-Chernousova & Fursenko, 1959. Remarks. The Endothyridae were very diversified during the Carboniferous. They remain present during the Permian, but are rare and without biostratigraphic importance, except perhaps for some species of Neoendothyra. The name Endothyroidea Glaessner, 1945 has priority over Endothyroidea Brady, 1884 nom. translat. Loeblich & Tappan, 1981 in designating this superfamily. Biostratigraphy. Endothyra is primarily a late Tournaisian (MFZ6: Poty et al. 2006) to Permian genus. Its acme is Visean –Moscovian. Its LAD is Permian, but not yet precisely established (Early or Middle Permian?), owing to confusion in the literature with Neoendothyra (19–24 in Fig. 2). The majority of the so-called Triassic Endothyra, in reality, belong to Endotebidae (Vachard et al. 1994). Planoendothyra sensu stricto is mainly a Pennsylvanian genus, but its LAD might be Kungurian (Vachard et al. unpublished data). Endothyranella, which is essentially a Middle–Late Pennsylvanian genus, is represented by relatively atypical forms in the Cisuralian (Vachard 1980; Baryshnikov et al. 1982; Vachard & Krainer 2001b). The material of the Urals and Afghanistan displays a siliceous agglutinate wall (see ‘Textulariata’ later in this section). Neoendothyra exhibits its primitive, relatively atypical forms in the late Cisuralian (Vachard & Krainer 2001b, pl. 4, figs 6 & 15; Vachard et al. unpublished data) or the late Chihsian ¼ Kungurian–Roadian) (Lin et al. 1990); its first typical forms seem to be early Roadian (Angiolini et al. 2015), and its acme is Capitanian –Wuchiapingian and up to the PTB (Song et al. 2007; Mohtat-Aghai et al. 2009). Triassic species have been reinterpreted as belonging to Endoteba (see later). Linendothyra seems to be more common in the late Capitanian– Lopingian times (Vachard unpublished data), and Neoendothyranella is Capitanian in age. Biogeography. Endothyra is rare but cosmopolitan during the Cisuralian. Rare Permian Planoendothyra are known in Indonesia (Nguyen Duc Tien 1989a, pl. 11, fig. 1) and in the Carnic Alps (Vachard & Krainer 2001a; Vachard et al. unpublished data). Cisuralian Endothyranella are represented by relatively atypical forms in the Urals (Baryshnikov et al. 1982), Afghanistan (Vachard 1980) and Bolivia (Mamet 1996). Neoendothyra exhibits its primitive forms in South China, the Carnic Alps and the Canadian Artic (see Lin 1985; Pinard & Mamet 1998; Vachard & Krainer 2001b). The first typical forms are observed in the late Cisuralian (especially Kungurian) of the Carnic Alps (Vachard et al. unpublished data) and South China (Lin et al. 1990). The Capitanian –Wuchiapingian Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 212 D. VACHARD specimens are common on the Palaeotethyan and Panthalassan shelves: Armenia, Azerbaijan, Montenegro, Tunisia, Turkey, NW Caucasus, Iran (central Alborz), Oman, Afghanistan, Cambodia, Thailand, Malaya, Sumatra, South China, Texas, Mexico and Guatemala. Changhsingian last forms subsist in South China, Thailand, Malaysia, Primorye (in the far east of Russia) and Japan; and up to the PTB in South China and Iran. Linendothyra seems to be more common in eastern Tethys but exists in Oman (Vachard et al. 2002). Neoendothyranella is endemic to New Mexico because the specimen of Cambodia of Nguyen Duc Tien (1986a, pl. 4, fig. 2), assigned to this genus by Nestell & Nestell (2006), is most probably a Linendothyra. Biogeography. Bradyina is cosmopolitan, at least up to the early Artinskian (Blazejowski 2009). Postendothyra is Palaeotethyan, Neotethyan and Panthalassan, and is unknown in North America (see Gaillot et al. 2009). Superfamily Bradyinoidea Reitlinger, 1950 nom. translat. Rauzer-Chernousova et al., 1996 Family Bradyinidae Reitlinger, 1950 nom. translat. Reitlinger, 1958 Subfamily Bradyininae Reitlinger, 1950 Remarks. Among the palaeotextulariids, the consistent continuity in morphology is remarkable (4–7 in Fig. 3), as it is in the tetrataxids (see below). The morphologies of Climacammina and Deckerella remain remarkably constant from the beginning to the end of their range (Vachard et al. 2010). Cribrogenerina, a unique genus that appeared in the Permian, differs only by lower chambers and more numerous openings in the cribrate aperture. Permian genera. Bradyina von Möller, 1878 (¼ Bradyinelloides Mamet & Pinard, 1992); Postendothyra Lin, 1984. Remarks. There are two episodes in the history of Permian bradyinoids. At the beginning of this system, the last Bradyina (sometimes designated as Bradyinelloides) survive and are more or less cosmopolitan, whereas the terminal period is characterized by Tethyan Postendothyra, a form totally atypical because, in contrast to all other bradyinoids, it is small sized and with a simple, basal aperture. Biostratigraphy. Bradyina is a late Mississippian– Early Cisuralian cosmopolitan genus. Its last representatives are late Cisuralian in age (Baryshnikov et al. 1982; Filimonova 2010). The FAD of Postendothyra is unknown; nevertheless, this genus becomes relatively common from the Capitanian to the Changhsingian in South China (Lin 1984; Song et al. 2009, 2011; Ueno et al. 2010; Zhang et al. 2015), Italy (Vachard & Miconnet 1990), Greece (Vachard et al. 1993b), Slovenia (Flügel et al. 1984), Crimea (Pronina & Nestell 1997), Cyprus (Nestell & Pronina 1997), Turkey (Zaninetti et al. 1981), Armenia and Azerbaijan (former ‘Transcaucasia’: Pronina 1988a; Kotlyar et al. 1989), Oman (Vachard et al. 2002), Afghanistan (Vachard unpublished data), Himalaya (Lys et al. 1980) and southern Tibet (Wang et al. 2010), Malaysia (Fontaine et al. 1994), NW Thailand (Fontaine et al. 1993), East Thailand (Fontaine et al. 1997), Cambodia (Nguyen Duc Tien 1986b), Primorye (Sosnina & Nikitina 1977), and Japan (Kobayashi 1986, 1997c, 2006a, 2012a, b). Superfamily Palaeotextularioidea Galloway, 1933 nom. translat. Habeeb, 1979 Family Palaeotextulariidae Galloway, 1933 nom. translat. Wedekind, 1937 Subfamily Palaeotextulariinae Galloway, 1933 Permian genera. Palaeotextularia Schubert, 1921 emend. Galloway & Ryniker, 1930; Climacammina Brady in Etheridge, 1873 emend. Cummings, 1956; Cribrogenerina Schubert, 1908; Deckerella Cushman & Waters, 1928a. Biostratigraphy. Palaeotextularia is Middle Mississippian (late Visean: MFZ13 biozone) to latest Permian; nevertheless, after the Visean it is difficult to distinguish the adult forms of this genus from juvenile or neotenic Climacammina. The FAD of Climacammina is an important datum in the late Mississippian, but after that the morphology of Climacammina remains very stable during the Pennsylvanian and Permian times, up to the latest Changhsingian and the PTB (Gaillot & Vachard 2007; Vachard et al. 2010). Deckerella is Moscovian (Reitlinger 1950) to latest Changhsingian (Kotlyar et al. 1999; Wang et al. 2010; Ebrahim Nejad et al. 2015). Cribrogenerina is Middle – Late Permian. Biogeography. Palaeotextularia and Climacammina, which are, first, Tethyan and Uralian genera during the late Mississippian, become cosmopolitan from the Early Pennsylvanian. Deckerella is rare but cosmopolitan throughout its range. Cribrogenerina is principally an eastern Tethyan and Panthalassan genus. Superfamily Endoteboidea Vachard et al., 2013 Family Spireitlinidae Vachard et al., 2013 Permian genus. Spireitlina Vachard in Vachard & Beckary, 1991. Remarks. The Spireitlinidae, due to their stratigraphic distribution, were suggested herein as the missing link between the Palaeotextulariidae and the Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 PERMIAN SMALLER FORAMINIFERS 213 Fig. 3. Smaller foraminifers (Fusulinata and Miliolata) from Trogkofel (Late Cisuralian of the Carnic Alps, Austria). Scale bar ¼ 0.100 mm. (1) & (2) Tetrataxis sp. (3) Transition Tetrataxis– Abadehella sp. (4) & (5) Climacammina spp. (6) & (7) ?Deckerella cf. laheei Cushman & Waters, 1928a. (8)–(10) Globivalvulina spp. (11) Hedraites sp. (12) Hemigordiellina sp. (13) Hemigordius? sp. (14)–(15) Praeneodiscus sp. (16) Glomomidiella sp. (17) Hemigordius sp. Endotebidae (Vachard et al. 2012). Spireitlina, which appears during the Visean –Serpukhovian boundary interval, is relatively well represented and cosmopolitan from the Moscovian to the Artinskian, but shows a weak species diversification (12–18 in Fig. 2). Biostratigraphy. Spireitlina is rare in the latest Visean – Serpukhovian of southern France (Pille 2008; Lucas et al. 2016a) and Tian Shan (Kulagina et al. 1992, reinterpreted by Vachard et al. 2012); it is relatively common and cosmopolitan from Early Pennsylvanian (Bashkirian) to Middle Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 214 D. VACHARD Permian (Wordian) times (Vachard et al. 2012), and rare in Capitanian (Vachard & Miconnet 1990, pl. 1, fig. 18, dating reinterpreted here; Nestell & Nestell 2006; Nestell et al. 2006; Zhang et al. 2015) and Wuchiapingian (Ueno et al. 2010). Biogeography. Spireitlina is cosmopolitan, and particularly common and constant in North America (e.g. Groves & Boardman 1999; Vachard & Krainer 2001a; Nestell & Nestell 2006; Nestell et al. 2006; Lucas et al. 2016a). Family Endotebidae Vachard et al., 1994 Permian genera. Endoteba Vachard & Razgallah, 1988; Vachardella Nestell & Nestell, 2006; ?Gen. and sp. indet. A sensu Kobayashi (2001); ?Granuliferelloides? sensu Kobayashi (2001); ?‘Haplophragmina?’ sensu Nestell & Nestell, 2006 (non Reitlinger, 1950). Remarks. Being related with Palaeotextularioidea, via the Spireitlinidae, the Endotebidae might also belong to the Textulariida (Rigaud et al. 2014) (see below). As evidence of this assignment, it is noteworthy that Endoteba and Vachardella have occasionally a siliceous agglutinate (see Nestell et al. 2006, pl. 4, figs 10 –13). Endoteba probably first occurs in the Artinskian or Kungurian (Vachard et al. 2002). It is abundant in the Capitanian. Endoteba is rare in the Late Permian and earliest Triassic, but reappears and diversifies in the Middle Triassic (Vachard et al. 1994; Rettori 1995). Except for the Nodosariata, Endoteba is one of the most advanced foraminifers (because it is multichambered) that crosses over the PTB, the most severe biotic turnover in the geological history of the Earth. Finally, Triassic biseriate stages reappear among the Endotebidae with Malayspirina Vachard in Fontaine et al., 1988a (as a homeomorph of the late Tournaisian –early middle Visean genus Eotextularia Mamet). Biostratigraphy. Endoteba is questionable in questionable Artinskian olistolites (Vachard et al. 2001b), and is known from Kungurian (Angiolini et al. 2016), Roadian (Kobayashi 2012a) and Capitanian (Vachard & Razgallah 1988; Vachard & Miconnet 1990). Vachardella is only known in the late Capitanian. The taxa in open nomenclature, Haplophragmina?, Granuliferelloides? and ?Gen. and sp. indet. A, are Kungurian–Changhsingian in age. Biogeography. Endoteba, common in Tunisia (Vachard & Razgallah 1988; Ghazzay et al. 2015), is also known in Italy (Apennines: Vachard & Miconnet 1990; Sicily: Vachard et al. 2001b), Cyprus (Nestell & Pronina 1997), Turkey (Leven & Okay 1996; Moix et al. 2013; Şahin et al. 2014), Oman (Hauser et al. 2000; Vachard unpublished data), Tajikistan (South Pamirs: Angiolini et al. in press), Malaysia (Ishii et al. 1975, pl. 2, fig. 8, reinterpreted) and Japan (Kobayashi 2012a). Vachardella and ‘Haplophragmina?’ are currently endemic to Texas and New Mexico (Nestell & Nestell 2006). Granuliferelloides? and ?Gen. and sp. indet. A were found in Tethys and western Panthalassa. Superfamily Tetrataxoidea Haynes, 1981 Family Tetrataxidae Pokorny, 1958 Subfamily Tetrataxinae Galloway, 1933 Permian genera. Tetrataxis Ehrenberg, 1854 emend. Nestler, 1973; Abadehella Okimura & Ishii in Okimura et al., 1975; Polytaxis Cushman & Waters, 1928a. Remarks. Tetrataxis and Polytaxis are relatively common during the Permian, but their species repeat the Carboniferous morphologies (1 & 2 in Fig. 3) and, therefore, have no biostratigraphic value. Abadehella, with its endoskeleton, is a Permian genus that mimics the Mississippian genus Valvulinella Schubert, but there is no phylogenetical continuity between these two taxa. Biostratigraphy. Tetrataxis is Early Mississippian (late Tournaisian, MFZ6: Poty et al. 2006) to latest Permian (late Changhsingian: Gaillot 2006; Song et al. 2009; Ueno & Tsutsumi 2009). Polytaxis is Early Pennsylvanian (Bashkirian) –Late Permian (Wuchiapingian) in age. The most interesting Permian tetrataxid is Abadehella, the FAD of which is poorly known. In the Carnic Alps, we observed (Vachard et al. unpublished data) secondarily septated tetrataxids: that is Abadehella sensu lato as early as the Kungurian (3 in Fig. 3). All these tetrataxids attain the PTB, but apparently do not cross through the PTB, as the Tetrataxis of the Triassic (e.g. described by Salaj et al. 1983) obviously differ both morphologically and microstructurally. The oldest occurrence of Abadehella dates back to the Artinskian in the Carnic Alps (Vachard et al. unpublished data), the Kungurian of New Mexico (Lucas et al. 2015, fig. 33.12 & 33.13) and, possibly, to the lower Cisuralian in South China (see Lin et al. 1990). The Abadehella reported from Israel, NW Caucasus, Armenia, Azerbaijan, central Iran and SE Pamir are probably all Capitanian– Lopingian in age (Okimura & Ishii 1981; Kotlyar et al. 1984; Kobayashi 1996, 1999; Pronina-Nestell & Nestell 2001; Orlov-Labkovsky 2004). Biogeography. All tetrataxid genera seem to be cosmopolitan. Although for a long time considered as Tethyan, Abadehella is now also known to be cosmopolitan, based on its occurrence in Guerrero, Mexico (Vachard et al. 1992), and in Texas (Nestell et al. 2006) and New Mexico, in the USA (Nestell & Nestell 2006; Lucas et al. 2015). Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 PERMIAN SMALLER FORAMINIFERS Superfamily Globivalvulinoidea Hance et al., 2011 Family Globivalvulinidae Reitlinger, 1950 emend. Gaillot & Vachard, 2007 Subfamily Globivalvulininae Reitlinger, 1950 (sic Globivalvulinae) Permian genera. Globivalvulina Schubert, 1921; Charliella Altıner & Özkan-Altıner, 2001; Labioglobivalvulina Gaillot & Vachard, 2007; Retroseptellina Gaillot & Vachard, 2007; Septoglobivalvulina Lin, 1978 emend. Gaillot & Vachard, 2007. Remarks. The evolution of the globivalvulinids was slow and weak during the Pennsylvanian –Cisuralian (8 –10 in Fig. 3). Their differentiation began in the Kungurian, and accelerated during latest Capitanian–earliest Wuchiapingian time. Biostratigraphy. Globivalvulina is Late Mississippian (earliest Serpukhovian)–latest Permian (Changhsingian), with a presence to confirm in the earliest Triassic (see G. curiosa Gaillot et al., 2009; see also the double PTB event of Song et al. 2011, 2013). The Early Mississippian (latest Tournaisian) species, formerly called Globivalvulina bristolensis Reichel, 1946, is now excluded from Globivalvulina: it corresponds to another lineage and was renamed Parabiseriella bristolensis by Cózar & Somerville (2012). Charliella appears in the Wordian of South China (Zhang et al. 2015) and NW Iran (Ebrahim Nejad et al. 2015). It is mainly known in the Capitanian, but was also found in the late Wuchiapingian and early Changhsingian. Labioglobivalvulina is late Capitanian –Lopingian. Septoglobivalvulina is early?/ late Capitanian –Changhsingian. Retroseptellina is questionable in the late Roadian of Thailand (Ueno & Sakagami 1993; nevertheless, in our opinion, these levels might be Capitanian in age) and is principally Wordian –Changhsingian. Biogeography. Initially, a Palaeotethyan taxon, Globivalvulina became cosmopolitan after the late Bashkirian. Charliella is present in Turkey (NW Anatolia), Italy (Monte Facito), NW Iran (Ebrahim Nejad et al. 2015), Zagros, Fars and Abu Dhabi (Gaillot & Vachard 2007), Sumatra (as Globivalvulina cyprica sensu Nguyen Duc Tien 1986b, pl. 13, fig. 5 only), Cambodia (as Globivalvulina cyprica and Globivalvulina sp. B sensu Nguyen Duc Tien 1979, pl. 9, figs 10 –12, and 1986a, pl. 4, figs 6 & 7), central Mexico (Vachard et al. 1992 as Globivalvulina ex gr. cyprica, pl. 6, fig. 7), western Texas, USA (Nestell & Nestell 2006 as Crescentia migrantis), and Late Permian of Thailand (Vachard unpublished data). Charliella is probably present in Japan (Okimura 1972; Kobayashi 2013, fig. 7.17? & 7.18), 215 southern Tibet (Wang et al. 2010), Yunnan (Ueno et al. 2010) and Oman (Jebel Akhdar, ‘late Dzhulfian’) (Lys in Montenat et al. 1977; Vachard unpublished data). Labioglobivalvulina is found in NW Iran, Zagros and Fars (Iran), Hazro (Turkey), ?Armenia and Azerbaijan, Montenegro, Italy, Hungary, South China, central Japan, and northern Thailand (Gaillot & Vachard 2007) and Malaysia (Aw et al. 1977, pl. 43, fig. 19). Septoglobivalvulina only exists in South China, Oman, ?Armenia and Azerbaijan, Turkey (Lycian nappes, Hazro), Iran (Fars and Zagros, NW Iran), and Abu Dhabi (see Gaillot et al. 2009; Koehrer et al. 2012). Retroseptellina is widespread in the Palaeo- and Neotethys: Slovenia (Nestell et al. 2009 as Paraglobivalvulina? globosa (Wang)); Hungary (Théry et al. 2007); Greece (Baud et al. 1991; Grant et al. 1991; Altıner & Özkan-Altıner 1998); southern Turkey (Köylüoglu & Altıner 1989; Ünal et al. 2003; Gaillot & Vachard 2007; Şahin et al. 2014); northern Caucasus (Pronina-Nestell & Nestell 2001 as Paraglobivalvulina globosa); northern Italy and Iran (Mohtat-Aghai & Vachard 2005; Ebrahim Nejad et al. 2015); the Duhaysan Member of the Khuff Formation in Saudi Arabia (Vachard et al. 2005); Oman, Batain Plain (Vachard et al. 2002); Armenia and Azerbaijan (Kotlyar et al. 1989); Thailand and Malaysia (Yanagida et al. 1988; Fontaine et al. 1993, 1994); southern Tibet (Wang et al. 2010); South China (Lin et al. 1990; Song et al. 2007; Gaillot et al. 2009; Zhang et al. 2015 as Globivalvulina? sp., fig. 4QQ); New Zealand (Vachard & Ferrière 1991); and Japan (Kobayashi 2006a, 2012a, b, 2013). Subfamily Paraglobivalvulininae Gaillot & Vachard, 2007 Permian genera. Paraglobivalvulina Reitlinger, 1965; Paraglobivalvulinoides Zaninetti & JennyDeshusses, 1985; Urushtenella Pronina-Nestell in Pronina-Nestell & Nestell, 2001. Remarks. The globivalvulinin ancestors of the paraglobivalvulinins are probably Septoglobivalvulina and/or Retroseptellina. This subfamily evolves to the realization of complex, secondary, sutural apertures. Biostratigraphy. Paraglobivalvulina is Capitanian– Changhsingian; Paraglobivalvulinoides is latest Changhsingian; Urushtenella is Changhsingian. Biogeography. Paraglobivalvulina is present in Armenia and Azerbaijan, Israel, Turkey, Iran (Alborz, Zagros and Fars, NW Iran), NW Caucasus, Carnic Alps, Hungary, Greece, Cyprus, Oman, Salt Range, southern Tibet, South China, Thailand, Philippines, and Japan. Paraglobivalvulinoides was mentioned in Iran (Alborz and Zagros), ?Italy, Greece (Vachard et al. 1993b), NW Caucasus, Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 216 D. VACHARD Himalaya, South China, Thailand, Malaysia, Japan (Gaillot et al. 2009 and references therein) and SE Pamirs (Kotlyar et al. 1999). Urushtenella is endemic to NW Caucasus (Pronina-Nestell & Nestell 2001): its presence in Zagros (Gaillot & Vachard 2007), southern Tibet (Wang et al. 2010) and, especially, in the Capitanian of Japan (Kobayashi 2012a) to be confirmed. Subfamily Dagmaritinae Bozorgnia, 1973 Permian genera. Dagmarita Reitlinger, 1965; Bidagmarita Gaillot & Vachard in Gaillot et al., 2009; Crescentia Ciarapica et al., 1986; Danielita Altıner & Özkan-Altıner, 2010; Louisettita Altıner & Brönnimann, 1980 emend. Gaillot & Vachard, 2007; Sengoerina Altıner, 1999; ?Labiodagmarita Gaillot & Vachard, 2007. Remarks. The FAD of Dagmarita is discussed. However, as indicated by Altıner & Özkan-Altıner (2010), the lineage Globivalvulina cyprica– Sengoerina–Dagmarita –Danielita in Turkey suggests that the evolutionary derivations of dagmaritin genera occurred very rapidly in the Capitanian. The foraminiferal material of Zheng (1986), which is dated as Chihsian (i.e. Kungurian–early Roadian), displays some taxa that are more probably late Guadalupian or even early Lopingian in age, such as Aulacophloia Gaillot & Vachard (pl. 5, figs 22 & 23 of Zheng 1986); Dagmarita Reitlinger (pl. 5, fig. 8); Geinitzina cf. taurica Sellier de Civrieux & Dessauvagie (pl. 5, figs 9– 13c); transition Globivalvulina Schubert–Paraglobivalvulina Reitlinger (pl. 5, fig. 41); Crassiglomella Gaillot & Vachard (pl. 6, figs 3 & 4); and Multidiscus Miklukho–Maklay (pl. 6, fig. 7a, b). In this condition, four hypotheses are possible for the interpretation of the proposed age of this microfauna: (a) the Chihsian age is misinterpreted; (b) the smaller foraminifers are reworked (and biostratigraphically mixed) in calciturbidites; (c) these smaller foraminifers are contained in calcareous olistolites of different ages mixed in a tectonic mélange; and (d) some taxa appear previously in South China and then migrate to western Tethys. At the moment, there are no geological arguments allowing one of these hypotheses in particular to be selected, but it is clear that the FAD of Dagmarita is younger than the Chihsian (see below). Biostratigraphy. The FAD of unquestionable Dagmarita is late Roadian (¼early Murgabian of Vachard 1980) or early Maokouan (Lin et al. 1990); the LAD is latest Changhsingian (Zhao et al. 1981; Lin et al. 1990; Wang et al. 2010). Sengoerina is late Capitanian (Altıner 1999; Nestell & Nestell 2006; Altıner & Özkan-Altıner 2010; Zhang et al. 2015) to latest Changhsingian (Song et al. 2007, 2009, 2011). The type species of Crescentia comes from a Triassic calciturbidite of the Monte Facito (Italy), where it is associated with abundant Capitanian foraminifers and algae (Ciarapica et al. 1986; Vachard & Miconnet 1990), as for the fusulinids Neoschwagerina and Kahlerina; nevertheless, they are mixed with rarer Changhsingian taxa, such as Colaniella (JennyDeshusses et al. 2000). Consequently, owing to its probable reworked character, the exact dating of the type species of Crescentia is debatable. Rare additional specimens of this genus exist in the Changhsingian of Himalaya (Lys et al. 1980) and Zagros (Gaillot & Vachard 2007), which reinforce the ambiguous dating. Bidagmarita might be limited to the late Changhsingian. Danielita is Capitanian or Lopingian, reworked in the Late Triassic. Louisettita probably has its FAD in the late Wuchiapingian in the Persian Gulf (see Gaillot & Vachard 2007; Mohtat-Aghai et al. 2009), but is generally limited to Changhsingian. Labiodagmarita is of Lopingian age (Gaillot & Vachard 2007). Biogeography. Dagmarita is a common Palaeotethyan, Neotethyan and Panthalassan genus, with the following detailed distribution (from west to east): Tebaga, Tunisia (Vachard & Razgallah 1988; Ghazzay et al. 2015); Apennines, Italy (Panzanelli-Fratoni et al. 1987; Vachard & Miconnet 1990); Slovenia (Nestell et al. 2009); Montenegro (Pantic 1970); the Carnic Alps (Noé 1987); Hungary (Berczi-Makk 1992); Greece (Hydra: Wignall et al. 2012); Cyprus (Nestell & Pronina 1997); Turkey (western Turkey: Argyriadis et al. 1976; Lys & Marcoux 1978); eastern Taurus (Altıner 1981, 1984; Zaninetti et al. 1981; Köylüoglu & Altıner 1989); Hazro (Canuti et al. 1970 as Palaeotextularia sp.; updated in this study); Armenia and Azerbaijan (Reitlinger 1965; Kotlyar et al. 1984, 1989); Iran (central Alborz: Bozorgnia 1973; Jenny-Deshusses 1983; Mohtat-Aghai & Vachard 2003; central Iran, Abadeh: Okimura & Ishii 1981; Zagros and Fars: Gaillot & Vachard 2007; and NW Iran: Ebrahim Nejad et al. 2015); Oman (Koehrer et al. 2012); central Afghanistan (Vachard & Montenat 1981); Salt Range, Pakistan (Okimura 1988); Lamayuru, Ladakh, Himalaya (Lys et al. 1980); South China (Zhao et al. 1981; Lin et al. 1990; Gaillot et al. 2009); western Thailand (Fontaine et al. 1988c) and the Philippines (Fontaine et al. 1986); NWThailand (Caridroit et al. 1990); Malaysia (Fontaine et al. 1988b); Cambodia (Nguyen Duc Tien 1979, 1986a); Primorye (Sosnina in Sosnina & Nikitina 1977); and Japan (Kobayashi 1997c, 2006b, c, 2012a, b, 2013). Dagmarita was also mentioned in Texas, USA (Nestell et al. 2006). Sengoerina is known in Turkey, Greece (Hydra), South China and New Mexico, and doubtful in central Iran and northern Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 PERMIAN SMALLER FORAMINIFERS Afghanistan (Vachard unpublished data). Crescentia comes from Monte Facito (Italy); rare specimens also exist in Zagros (Gaillot & Vachard 2007), Himalaya (Lys et al. 1980) and Thailand (Fontaine et al. 1993); the so-called Crescentia of Texas, USA (Nestell & Nestell 2006) are misinterpreted (see above). Bidagmarita is known in the Salt Range, Pakistan, central Iran, NW Caucasus, the Lamayuru Block (Ladakh Himalaya), Malaysia and South China (Gaillot et al. 2009). Danielita is endemic to the Bursa area (Turkey). Louisettita is present in the Changhsingian of Turkey, Iran (Zagros, Julfa area), South China, and, questionably, of Afghanistan, Armenia and Azerbaijan; its FAD probably occurred in the Persian Gulf (see Gaillot & Vachard 2007; Mohtat-Aghai et al. 2009). Labiodagmarita only exists in Saudi Arabia (Vachard et al. 2005), Zagros, Fars, Abu Dhabi and Turkey (Gaillot & Vachard 2007). Subfamily Paradagmaritinae Gaillot & Vachard, 2007 Permian genera. Paradagmarita Lys in Lys & Marcoux, 1978; Paradagmacrusta Gaillot & Vachard, 2007; Paradagmaritella Gaillot & Vachard, 2007; Paradagmaritopsis Gaillot et al., 2009; Paremiratella Gaillot & Vachard, 2007. Remarks. Described by Gaillot & Vachard (2007), this family has been critically revised by Altıner & Özkan-Altıner (2010): for example, Altıner (pers. comm. March 2016) suggests that the genera Paradagmaritella, Paradagmaritopsis and Paremiratella should be included in the subfamily Dagmaritinae. However, as these taxonomic problems do not modify the biostratigraphy and biogeography of the paradagmaritinins, the previous classification is followed herein. Biostratigraphy. According to Kotlyar et al. (1989, table 1), the first representative of the subfamily appears in the latest Khachikian (interpreted by these authors as the latest Capitanian/Midian). However, this part of the Khachik Formation is, most probably, earliest Wuchiapingian in age (Leven 1998; and see below). Gaillot & Vachard (2007) indicated that the range of Paradagmarita sensu stricto is late Wuchiapingian-Changhsingian. Other genera of the subfamily have more restricted ranges: Paradagmacrusta is Changhsingian; Paradagmaritella is ?latest Capitanian– Changhsingian; Paradagmaritopsis is late Wuchiapingian–Changhsingian; and Paremiratella is Changhsingian. Biogeography. The subfamily appeared probably in Armenia or in Azerbaijan (i.e. the former Transcaucasia), but, as indicated above, the corresponding datum of Kotlyar et al. (1989) is yet to be verified. The subfamily is Palaeotethyan and Neotethyan, 217 and is principally known in Turkey (Taurus: Altıner 1984), Iran (Zagros: Gaillot & Vachard 2007; NW Iran: Ebrahim Nejad et al. 2015), Saudi Arabia (Vachard et al. 2005), Oman (Koehrer et al. 2012) and NW Caucasus (Pronina-Nestell & Nestell 2001), although it was mentioned from Italy to Japan. The so-called Paradagmarita from Afghanistan described by Vachard (1980) in reality belong to Louisettita; those from Thailand and Pakistan are very atypical; and those from Japan (Kobayashi 1997c, 2004) instead belong to Paradagmaritopsis, a genus also observed in southern China (Gaillot et al. 2009) and Oman (Koehrer et al. 2012). The other genera are more endemic: (1) Paradagmacrusta is only mentioned in Zagros and Fars (Iran); (2) Paradagmaritella, in Armenia or Azerbaijan, Fars, Zagros, Saudi Arabia, and, questionably, in NW Caucasus and Turkey; and (3) Paradagmaritopsis in Iran (Zagros and Fars), Japan and South China (Gaillot et al. 2009). Paremiratella is endemic to Zagros, Fars, Abu Dhabi and Oman (and questionable in Japan). Class Miliolata Saidova, 1981 Order Cornuspirida Mikhalevich, 1980 All Late Palaeozoic miliolates exhibit a diversely coiled, undivided, tubular chamber (11 –17 in Fig. 3 and Figs 4–6). Hence, they constitute a unique order: the Cornuspirida. Two suborders are distinguished here: the attached Nubeculariina and the free Cornuspirina. We agree with Gaillot & Vachard (2007) in speculating that all the Palaeozoic miliolates derive from a pseudolituotubellid Fusulinata at the end of the Visean, since the majority of the Pennsylvanian miliolates are calcivertellids (i.e. attached forms). However, contrary to the opinion of Gargouri & Vachard (1988), Vachard et al. (1993a) and Gaillot & Vachard (2007), who supposed that the hemigordiopsin cornuspirids gave rise to the involutinids, and in agreement with Groves & Altıner (2005), it is more likely that Triassic involutinids might have arisen from a very simple, planispiral evolute ancestor, which is either Pseudoammodiscus (consensual hypothesis), Ammodiscus sensu Nestell et al. (2015) or, most probably, Postcladella Krainer & Vachard, 2011. The name Cornuspirida Mikhalevich, 1980 has priority over Cornuspirida Vdovenko et al., 1993 and Hemigordiopsida Pronina, 1994, for designating this order of miliolates. Palaeozoic miliolates are known from the latest Visean or earliest Serpukhovian, and are common during the Permian. Nubeculariina are abundant as early as the Serpukhovian–Bashkirian. Primitive typical cornuspirids (with the typical, wellpreserved, amber-coloured test: see the discussion in Vachard et al. 2015) (see Figs 4 & 5, showing the initial aspect of the wall (Fig. 4) and its Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 218 D. VACHARD Fig. 4. Well-preserved Uralogordius (centre) and Ellesmerella (top left and bottom right) with amber-coloured wall characteristic of the extant Miliolata, to compare with the dark-microgranular wall of an earlandiid (top right). Scale bar ¼ 0.250 mm. Early Artinskian of Garnitzenbach (Carnic Alps, Austria). diagenetic transformations (Fig. 5)), Cornuspira and Hemigordiellina, are only known from the Bashkirian. More complex coilings appear with Brunsiella and then with Hemigordius in the Moscovian. The diversity is already important in the Cisuralian, but increases gradually during the Guadalupian where neodiscid and hemigordiopsid giant forms appear. The Capitanian –Lopingian is an episode of maximal diversity for the group. Suborder Nubeculariina Jones in Griffith & Henfrey, 1875 nom. translat. Mikhalevich, 1988 nomen re-translat. herein Family Calcivertellidae Reitlinger in Vdovenko et al., 1993 emend. Gaillot & Vachard, 2007 Permian genera. Calcivertella Cushman & Waters, 1928b; Ammovertella Cushman, 1928; Baryshnikovia Reitlinger in Vdovenko et al., 1993; Calcitornella Cushman & Waters, 1928b; Hedraites Henbest, 1963; Hedraites? sensu Lucas et al., 2015; Palaeonubecularia Reitlinger, 1950; Pseudoagathammina Lin et al., 1990; Pseudospira Reitlinger in Vdovenko et al., 1993; Pseudovermiporella Elliott, 1958 emend. Henbest, 1963; Tansillites Nestell & Nestell, 2006; Trepeilopsis Cushman & Waters, 1928b; ?Orthovertella Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 PERMIAN SMALLER FORAMINIFERS 219 Pennsylvanian –Cisuralian, Hedraites? sensu Lucas et al. (2015) is Kungurian, Pseudovermiporella is late Cisuralian (Artinskian or even latest Sakmarian) –latest Permian (late Changhsingian: Zhao et al. 1981; Flügel & Reinhardt 1989; Vachard et al. 2003; Mohtat-Aghai et al. 2009; Ebrahim Nejad et al. 2015), it is common in the Lopingian of southern Turkey (Hazro) and Zagros. Recently, we used the Artinskian FAD of Pseudovermiporella as a biozonal, regional marker in South China (Liu et al. in press). Fig. 5. Uralogordius sp., after diagenesis of the wall (cf. with Fig. 4); this wall became first dark, then whitish and neomicrosparitized. Scale bar ¼ 0.500 mm. Kungurian of the stratotype of the regional Bolorian Stage (Pamirs; Tajikistan). Cushman & Waters, 1928b; ?Orthovertellopsis Vachard et al., 2015. Remarks. Calcivertelloids seem to constitute the most primitive Miliolata, whereas the attachment is generally considered as a secondary trend among the foraminifers. Orthovertella and Orthovertellopsis, because of the growth of their tests, seem to have been attached during part of their life: therefore, they are questionably assigned to calcivertellids. Biostratigraphy. All these genera are long ranged, Calcivertella is Late Mississippian (Serpukhovian)–Late Permian. Calcitornella, Trepeilopsis, Ammovertella and Orthovertella are Pennsylvanian– Permian. Other genera might be more restricted in time, but they are still poorly known. Orthovertellopsis is biostratigraphically poorly known (i.e. cited as Kungurian (late Leonardian), Early Permian, Permian, Early Kazanian (Middle Permian) or ?latest Capitanian, following the areas where it is present). Tansillites is late Capitanian. Pseudoagathammina, Baryshnikovia and Pseudospira also have a questionable Permian distribution. Palaeonubecularia is Serpukhovian– Changhsingian: perhaps Triassic in Europe and Taurus (Turkey). Hedraites (11 in Fig. 3) is Late Biogeography. Calcivertella and Calcitornella are cosmopolitan. Trepeilopsis, Orthovertella and Ammovertella are more rarely cited, but are probably also cosmopolitan. Orthovertellopsis exists in New Mexico, Australia (Carnarvon Basin, Canning Basin, Tasmania, and, perhaps, the Sydney Basin), South China, the Urals (Russia) and, perhaps, Texas (Vachard et al. 2015). Pseudoagathammina was described in South China, but was recently re-found in Tunisia (Ghazzay-Souli et al. 2015). Tansillites is endemic to New Mexico. Baryshnikovia and Pseudospira are sporadically mentioned; they are, perhaps, cosmopolitan, being currently known in Donbass, the Pre-Urals, Uzbekistan and New Mexico. Palaeonubecularia is cosmopolitan. Hedraites and Hedraites? sensu Lucas et al. (2015) are common in North America and rare in Tethys. Pseudovermiporella is cosmopolitan, and is especially common during the Lopingian in Turkey and Iran (Ebrahim Nejad et al. 2015). Suborder Cornuspirina Jirovec, 1953 emend. Gaillot & Vachard, 2007 Superfamily Cornuspiroidea Mikhalevich, 1988 Family Cornuspiridae Schultze, 1854 Subfamily Cornuspirinae Schultze, 1854 Permian genera. Cornuspira Schultze, 1854 (¼Ammodiscus Reuss, 1862 (part.); ¼Pseudoammodiscus Conil & Lys in Conil & Pirlet, 1970 (part.)); Flectospira Crespin & Belford, 1957; Glomotrocholina Nikitina in Sosnina & Nikitina, 1977; Hemigordiellina Marie in Deleau & Marie, 1961 emend. Vachard in Vachard & Beckary, 1991; Hoyenella Rettori, 1994 emend. Gaillot & Vachard, 2007 (¼Glomospirella Plummer, 1945 (part.)); Pilammina sensu Vachard in Termier et al., 1977 non Pantic, 1965; Postcladella Krainer & Vachard, 2011 (¼Ammodiscus Reuss, 1862 (part.) ¼ Rectocornuspira Warthin, 1930 (part.)); Pseudohemigordius Nestell & Nestell, 2006; Streblospira Crespin & Belford, 1957. Remarks. Almost every genus of this family is discussed; especially Hemigordiellina (with Glomospira diversa Cushman & Waters, 1930 as type species) and Glomospira or Pseudoglomospira, as well as the quatuor Cornuspira, Rectocornuspira, Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 220 D. VACHARD Fig. 6. A complex specimen of Tubiphytes ex gr. obscurus Maslov, 1956 showing a well-preserved series of sagittiform (arrow-shaped) chambers. Scale bar ¼ 0.250 mm. Early Kungurian of Trogkofel (Carnic Alps, Austria). Postcladella, Pseudoammodiscus or Ammodiscus (12 & 13 in Fig. 3) (e.g. Loeblich & Tappan 1987; Vachard et al. 2010; Krainer & Vachard 2011; Nestell et al. 2015). In the Permian, the genus Cornuspira was first confused with Ammodiscus, and then with Pseudoammodiscus (e.g. Groves & Boardman 1999; Pronina, 1999a, p. 184; Nestell & Nestell 2006) or again with Ammodiscus in the Triassic (Nestell et al. 2015). Postcladella, which has distinct morphological characters compared with Ammodiscus, Cornuspira and Rectocornuspira (see the discussion in Krainer & Vachard 2011; Vachard et al. 2015), was, however, recently re-synonymized with Ammodiscus (Nestell et al. 2015): this latter interpretation requires further intensive discussion. If it is admitted that the wall microstructure has a secondary importance, Glomospira can, in turn, replace Pseudoglomospira and Hemigordiellina, despite these genera, in my opinion, corresponding to three different classes: Textulariata, Fusulinata and Miliolata. Rectocornupira is a Pennsylvanian porcelaneous genus (Warthin 1930; Loeblich & Tappan 1987). This name was erroneously applied to : (1) dark-microgranular walled, uncoiled pseudoammodiscids of the Visean –Serpukhovian boundary interval (see Vdovenko et al. 1993); and (2) to Early Triassic Postcladella kahlori (Brönnimann et al.) (see Krainer & Vachard 2011). Recently, this latter species was reconsidered as possessing an agglutinated wall and reassigned to Ammodiscus (see Nestell et al. 2015). However, because of the terminal uncoiling of Postcladella kahlori, it is to be noted that the name Rectoammodiscus Reitlinger in Vdovenko et al., 1993 would be formally more correct. Other controversial genera are Streblospira, Glomotrocholina, Pilammina? and Hoyenella. The genus Streblospira is interpreted herein as different to the Triassic marker Meandrospira, as has been done by Vdovenko et al. (1993), but contrary to Loeblich & Tappan (1964, 1987) and Kalia et al. (2000). Glomotrocholina described in the Middle– Late Permian of the Primorye has never been re-found since its creation, although it occurs in reefal environments (G. Nestell pers. comm. March 2016), which are relatively common in the Middle Permian and where the foraminifers are abundant. Glomotrocholina is therefore difficult to identify and needs a revision because it probably differs significantly from its proposed diagnosis. Eventually, Pseudohemigordius and/or Vissariotaxis? nativus Nestell & Nestell in Nestell et al., 2006 from the Capitanian of Texas might correspond to Glomotrocholina or to a homeomorph of this latter genus. Pilammina? sp., only mentioned by Vachard in Termier et al. 1977 and Gaillot & Vachard 2007, and strongly questioned by D. Altıner (pers. comm. March 2016), differs indeed from Pilammina Pantic, 1965, and is possibly only a minute species of Hemigordiellina. It is noteworthy that Palmieri (2004) proposed the new name Flectospiroides for a taxon previously called Pilammina by him, and of ‘Late Kazanian–Midian (?)’ (i.e. Middle Permian) Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 PERMIAN SMALLER FORAMINIFERS age in Australia. The genus Hoyenella sensu Gaillot & Vachard, 2007 is also discussed because: (1) it could differ from Hoyenella Rettori, 1994, the distribution of which would only be Triassic (D. Altıner pers. comm. March 2016); and (2) it is homeomorph with the microgranular genus Brunsia, the porcelaneous genus Brunsiella and the agglutinated genus Glomospirella. Owing to the current trends, there is great temptation to return to Glomospirella, but, as indicated by Vachard et al. (2015), this genus has Glomospira umbilicata Cushman and Waters for the type species, which is a very large taxon, with a diameter of up to 1.00 mm. Its age is Middle–Late Pennsylvanian, whereas other species described in Glomospirella are small to very small (because they measure generally 0.25–0.40 mm), with a second part of coiling markedly planispiral and evolute, and are often described in the Permian and the Triassic. However, it is clear that these Permian glomospirellids have a porcelaneous wall, whereas the true microstructure of Triassic Hoyenella is not well established (porcelaneous, calcitic microgranular, aragonitic microgranular or an uncharacteristic and unknown wall?). Biostratigraphy. Cornuspira is early?/late Serpukhovian (i.e. latest Mississippian) to Holocene. Postcladella is essentially an Early Triassic taxon (Krainer & Vachard 2011), but it is very rare in the late Capitanian and sporadic in the Late Permian of the Middle East (Shang et al. 2003). Hemigordiellina is Pennsylvanian and Permian; it appears as soon as the bed T in Tindouf (Morocco) (Cózar et al. 2014), which is probably late Serpukhovian in age, and earlier in Europe, if the poorly known genus Warnantella Conil & Lys in Conil et al., 1977 is synonymous of Hemigordiellina. Hoyenella has a possible FAD in the early Sakmarian (see Vachard & Krainer 2001b, pl. 5, fig. 24): its acme is Early–Late Triassic. Pilammina? is rare in the late Guadalupian of Tebaga (Tunisia) (Termier et al. 1977), and the late Wuchiapingian of Zagros (Gaillot & Vachard 2007). Glomotrocholina is Middle or Late Permian. Flectospira is Artinskian, and Streblospira is ?Artinskian–Middle Permian in age. Biogeography. Cornuspira and Hemigordiellina are probably cosmopolitan during the Permian. Postcladella is widespread in the Tethyan Early Triassic, but is very rare in the Middle East during the Permian. Hoyenella is cosmopolitan in the Permian, and is distributed, during its Triassic acme, from Western Europe to China and Japan. Pilammina? is endemic to Tunisia and Zagros. Other endemic genera are Glomotrocholina (Primorye) and Flectospira (Western Australia). The Flectospira of eastern Himalaya illustrated by Kalia et al. (2000) are misinterpreted and could correspond to every 221 other cornuspirin. Streblospira is rare out of Australia, where it was described, but was mentioned in Bolivia (Mamet 1996), Armenia –Azerbaijan (Kotlyar et al. 1984), Oman (Vachard et al. 2002) and Turkey (Wignall et al. 2012; Şahin et al. 2014). Subfamily Agathammininae Ciarapica, Cirilli & Zaninetti in Ciarapica et al., 1987 Permian genera. Agathammina Neumayr, 1887; Septagathammina Lin, 1984; Graecodiscus Vachard in Vachard et al., 1993a. Remarks. The agathamminins seem to be the group that is transitional between the Palaeozoic, atypical miliolates and the typical Mesozoic –Recent miliolinins; hence, it is difficult to believe that, as in many molecular clocks, the miliolinins derive from the very primitive, monothalamous foraminifers. Biostratigraphy. The FAD and LAD of Agathammina are poorly known (Pronina 1988b, 1999a), probably Artinskian– Kungurian and latest Changhsinghian, respectively. The genus is relatively common from the Capitanian to the Changhsingian throughout the Palaeotethys; many formerly admitted Triassic survivors are now assigned to another genus. The FAD might be Kungurian (see Vachard & Moix 2013). Septagathammina is late Chihsian – Wuchiapingian (Lin et al. 1990) or Changhsingian. Graecodiscus defined in the Lopingian might appear as early as the Roadian (Angiolini et al. 2015). Biogeography. Agathammina is traditionally cited throughout the Palaeotethys, from Tunisia to Japan. Although mentioned for a relatively short time in North America (Nestell & Nestell 2006; Nestell et al. 2006), where it can be common (G. Nestell pers. comm. March 2016). Septagathammina is known in South China (Lin et al. 1990), Sumatra, Cambodia, Armenia, Azerbaijan, Crimea, NW Caucasus, ?Hungary, Zagros, Thailand and ?Primorye (Nestell & Nestell 2006; Gaillot & Vachard 2007). Graecodiscus, described in Greece, might be also present in Saudi Arabia (Vachard et al. 2005), New Mexico (Nestell & Nestell 2006), Tajikistan (Angiolini et al. 2015) and South China (Liu et al. in press). Family Hemigordiidae Reitlinger in Vdovenko et al., 1993 Subfamily Hemigordiinae Pronina, 1994 Permian genera. Hemigordius Schubert, 1908; Arenovidalina Ho, 1959 emend. Gaillot & Vachard, 2007 (¼Multidiscus of the authors, part.); Brunsiella Reitlinger, 1950; Neodiscopsis Gaillot & Vachard, 2007; Okimuraites Reitlinger in Vdovenko et al., 1993 emend. Ebrahim Nedjad et al. 2015 (¼? Brunsispirella Gaillot & Vachard, Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 222 D. VACHARD 2007); Rectogordius Alipour & Vachard in Alipour et al., 2013 (¼Neohemigordius of the authors non Wang & Sun); ?Triadodiscus Piller, 1983. Remarks. There exist several nomenclatural problems in this family. For example, as suggested by Nestell et al. (2009), Brunsispirella could be an unnecessary junior synonym of Okimuraites: however, both taxa differ from Hemigordius (17 in Fig. 3) by their smaller, more compressed tests with many planispiral, evolute whorls. Triadodiscus either belongs to this group (partly or entirely) or to the involutinids: this depends on whether the wall remains porcelaneous or is already aragonitic in composition. Similarly, according their true microstructure, the different species of Arenovidalina probably belong to several genera. Biostratigraphy. The FAD of Hemigordius is Bashkirian. Its acme is Moscovian– Changhsingian. Rectogordius is Cisuralian (late Asselian – Artinskian) (Forke et al. 1998; Vachard & Krainer 2001a, b). Brunsiella, which appeared in the Early Pennsylvanian and is a probable ancestor of Hemigordius, was still mentioned in the Cisuralian by Groves & Boardman (1999). Okimuraites is late Capitanian–Lopingian. Arenovidalina is Late Pennsylvanian–Late Permian and Early– Late Triassic. Neodiscopsis is late Capitanian–Changhsingian. Biogeography. Hemigordius is cosmopolitan from Moscovian to Changhsingian. Rectogordius is Early Permian (Late Asselian –Artinskian) of Iran, Arctic Canada, Donets, Afghanistan and the Carnic Alps (Forke et al. 1998; Vachard & Krainer 2001a, b). Consequently, its palaeobiogeographical range seems to be located in the northern Palaeotethys, in the shelf of the Uralian Ocean and in the northern part of North America. Arenovidalina is apparently cosmopolitan, and is known especially in Greece, Italy, Turkey, Armenia –Azerbaijan, Iran (Zagros, Fars), Saudi Arabia, Oman, Sumatra, Primorye, South China and Japan. Neodiscopsis was found in Turkey (Antalya nappes and Hazro) (Şahin et al. 2014), Oman (Koehrer et al. 2012), Iran (Zagros), South China and Japan (Gaillot & Vachard 2007). Family Neodiscidae Lin, 1984 nom. translat. and emend. Gaillot & Vachard, 2007 Permian genera. Neodiscus Miklukho-Maklay, 1953 emend. Gaillot & Vachard, 2007; Crassispirella Gaillot & Vachard, 2007; Crassiglomella Gaillot & Vachard, 2007; Glomomidiella Vachard et al., 2008; Multidiscus Miklukho-Maklay, 1953 emend. Gaillot & Vachard, 2007; Neohemigordius Wang & Sun, 1973 emend. Alipour et al., 2013; Olgaorlovella Vachard et al., 2015; Praeneodiscus Vachard et al., 2015; Uralogordius Gaillot & Vachard, 2007 (¼Arenovidalina sensu Baryshnikov et al., 1982 non Ho, 1959); ?Midiella Pronina, 1988b. Remarks. This family displays a trend to the gigantism (compared with the normal size of the Miliolata); another trend is the presence of buttresses, and a differentiated wall with an outer tectum (14– 16 in Fig. 3 & Figs 4 & 5). Midiella, because of its sigmoidal coiling and its moderate size, could be transititional between the Hemigordiidae and the Neodiscidae. Biostratigraphy. Neodiscus is Middle–Late Permian. Glomomidiella is relatively rare from the Kungurian to the Wordian, and has its acme from the Capitanian to the Changhsingian. Olgaorlovella and Praeneodiscus are currently limited to the Cisuralian (?Sakmarian –Kungurian). Neohemigordius and Crassiglomella are probably Capitanian – Changhsingian. Multidiscus might be late Cisuralian, but is most probably Wordian –Changhsingian. Uralogordius is principally known in the late Cisuralian (Baryshnikov et al. 1982; Vachard et al. unpublished data), but possibly survives into the Middle Permian. Crassispirella is Lopingian, and mainly Changhsingian. Midiella is present in all of the Permian; its acme is Capitanian –Changhsingian. Biogeography. Neodiscus is present in Oman, Slovenia, Tunisia, NW Iran, central Iran, NW Caucasus, Greece, Turkey (Hazro, Himmetli), SE Pamirs, southern Tibet, South China and Japan. Glomomidiella, during its acme, was mentioned in Greece, Tunisia, Croatia, Serbia, Hungary, Italy, Armenia, Azerbaijan, Turkey, Iran, Oman, Himalaya, South China, Sumatra, Malaysia and Japan. Olgaorlovella and Praeneodiscus, described in New Mexico, might exist in Greece (as Hemigordius aff. ovatus Grozdilova sensu Vachard et al. 1993b, pl. 3, fig. 14), Japan and Bashkortostan (Russia). Neohemigordius is from South China and Japan (Kobayashi 2012a). Crassiglomella was encountered in South China, central Japan, Cambodia, Turkey (Himmetli), Iran (Julfa area, Zagros and Fars), Oman and Tunisia. Multidiscus is found in Sumatra, Italy, Slovenia, Greece, Israel, Turkey, Armenia –Azerbaijan, Saudi Arabia, Oman, Iran (Zagros, Fars), southern Tibet, South China, Japan and Texas. Uralogordius is present in the Urals, the Carnic Alps (Vachard et al. unpublished data), Armenia –Azerbaijan, northern Afghanistan, Japan and Mexico; Crassispirella in Persian Gulf (Iran), Turkey (Hazro), Saudi Arabia and South China. Midiella is distributed in the Palaeotethys and Neotethys: Turkey (Hazro), NW Iran, central Iran (Abadeh), Persian Gulf, Saudi Arabia, Oman, South China and Primorye. Family Baisalinidae Loeblich & Tappan, 1986a Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 PERMIAN SMALLER FORAMINIFERS Permian genera. Baisalina Reitlinger, 1965; Pseudobaisalina Sosnina, 1983; Nikitinella Sosnina, 1983; Pseudomidiella Pronina-Nestell in ProninaNestell & Nestell, 2001; Septigordius Gaillot & Vachard, 2007. Remarks. This family is characterized by a pseudosepation, remarkably irregularly sized and distributed compared with the other foraminifers. Some illustrations of Kobayashi (1988, pl. 2, figs 18 & 19, 2012a, pl. 3, 2013, fig. 8.38–8.42) are especially significant. Biostratigraphy. Baisalina is late Capitanian–late Changhsingian; Pseudobaisalina and Nikitinella are Wordian and/or Capitanian; Pseudomidiella is Changhsingian (Pronina-Nestell & Nestell 2001); Septigordius is late Early Permian–latest Permian. Biogeography. Baisalina is a mentioned in Armenia, Azerbaijan, Hungary, Greece, Turkey, NW and central Iran, Afghanistan, Myanmar, southern Tibet (Wang et al. 2010), South China (Zhang et al. 2015), Japan (Kobayashi 1988, 2012a, 2013), New Zealand (Vachard & Ferrière 1991) and Texas, USA (Nestell et al. 2006). Nikitinella and Pseudobaisalina are endemic to Primorye (Sosnina 1983). Pseudomidiella occurs in NW Caucasus (Pronina-Nestell & Nestell 2001), Slovenia (Nestell et al. 2009), Greece (Hydra: Vachard et al. 2008; Wignall et al. 2012). Septigordius seems to be Palaeotethyan, Neotethyan and Panthalassan: Primorye, Crimea, Cyprus, Turkey, Oman, Ladakh, South China, Japan and New Zealand (Gaillot & Vachard 2007 and references therein). Family Hemigordiopsidae Nikitina, 1969 emend. Gaillot & Vachard, 2007 Permian genera. Hemigordiopsis Reichel, 1945 (¼Gansudiscus Wang & Sun, 1973); Glomomidiellopsis Gaillot & Vachard, 2007; Kamurana Altıner & Zaninetti, 1977; Lysites Reitlinger in Vdovenko et al., 1993; Shanita Brönnimann, Whittaker & Zaninetti, 1978. Remarks. The Hemigordiopsidae is another giant family of the Miliolata during the Wordian?– Capitanian– Lopingian. It presents the first phenomenon of flosculinization (i.e. the thickenings of basal layers that reduce the height of chambers in miliolids and alveolinids) registered among the Miliolata (Gaillot & Vachard 2007; Vachard et al. 2010). Biostratigraphy. Hemigordiopsis is rare in the Wordian (Zhang et al. 2015), common in the Capitanian and the Wuchiapingian (Vachard et al. 2003), and questionable in the Changhsingian (Altıner 1984). Lysites are probably restricted in 223 the Capitanian –Wuchiapingian boudary interval. Shanita characterizes the Capitanian–Wuchiapingian boundary (Gaillot & Vachard 2007; Koehrer et al. 2012; Kolodka et al. 2012). Because of the association of Shanita with Chusenella in Oman, and Neoschwagerina in Iran, apparently preserved in situ, we consider Shanita as preferentially latest Capitanian rather than earliest Lopingian. Glomomidiellopsis is late Capitanian –Changhsingian. Kamurana is either late Capitanian or Wuchiapingian, but was mentioned in the latest Changhsingian of South China (Wignall & Hallam 1996, text fig. 4 p. 591) and southern Tibet (Wang et al. 2010). Biogeography. Hemigordiopsis with synonym Gansudiscus were unquestionably mentioned in Cyprus, Montenegro, Greece, Tunisia, Turkey, Armenia, Oman, central Iran, peninsular Thailand, South China and Japan (see discussions in Gargouri & Vachard 1988; Nestell & Pronina 1997). Lysites, even if defined in Turkey, seems to be more common in eastern Tethys (South China and Thailand). Known from Turkey and Oman to Myanmar, Thailand and Yunnan (e.g. Brönnimann et al. 1978; Sheng & He 1983; Şengör et al. 1988; Dawson et al. 1993; Jin & Yang 2004), Shanita is considered here as characteristic of these peri-Gondwanan terranes which, after the opening of the Neotethys, migrated to the Cimmeria and Sibumasu microcontinents (Nestell & Pronina 1997). Glomomidiellopsis is known in Cambodia, Primorye, Hazro (Turkey), Zagros (Iran) and Oman, and is questionable in Myanmar. Kamurana sensu stricto seems endemic of Turkey, but, because often confused with Neodiscus and/or Glomomidiellopsis, was also mentioned in Italy (Panzanelli-Fratoni et al. 1987), Armenia – Azerbaijan (Pronina 1988a), South China (Wignall & Hallam 1996) and New Zealand (Vachard & Ferrière 1991). Groups incertae sedis possibly partly related to the Miliolata Group 1: ellesmerellids Permian genera. Ellesmerella Mamet & Roux in Mamet, Roux & Nassichuk, 1987 emend. Vachard & Krainer, 2001b (eventually junior synonym of Osagia and/or Ottonosia, which designate, in reality, biopisoliths composed of Ellesmerella; e.g. see Henbest 1963; Vachard et al. 2015). Remarks. Girvanella permica Pia, 1937, the type species of Ellesmerella, was initially described as an alga or cyanobacterium. In contrast, it was proposed (Vachard & Krainer 2001b; Sanders & Krainer 2005; Vachard et al. 2015; Vachard et al. unpublished data) that it is most probably a representative of the family Nubeculariidae Jones, 1895 because of its amber-coloured well-preserved wall (Fig. 4), and because some nubeculariids can build Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 224 D. VACHARD biopisolites and bioconstructions (e.g. ‘oolithes canabinnes’, ‘microreefs’, Osagia, Ottonosia), which mimic oncoids or columnal stromatolites. Biostratigraphy. Early Cisuralian (see Vachard & Krainer 2001b). Biogeography. Probably cosmopolitan (especially in peri-Gondwanan and North American craton shelves). Group 2: tubiphytids Permian genera. Tubiphytes Maslov, 1956; Epimonella Vachard in Kolodka et al., 2012; Latitubiphytes Vachard et al., 2012; Ramovsia Kochansky-Devidé, 1973 (¼Dorudia Jenny & Jenny-Deshusses, 1978); ‘Tubiphytes’ epimonellaeformis Vachard et al., 2015. Remarks. No amber-coloured, well-preserved specimens were never encountered among the tubiphytids. However, relationships with the miliolates have been suggested for a long time (see Vachard et al. 2010 and references therein), especially for the Jurassic forms based to some similarities between the inner cavities of the tubiphytids and the shape of the chambers of some miliolates (Fig. 6). Another argument is that the first tubiphytids resemble the miliolate Palaeonubecularia. This latter genus was even proposed as a transitional form between the families Calcivertellidae and Tubiphytidae (Vachard & Krainer 2001b; Vachard et al. 2012). This transition possibly took place at the end of the Moscovian (Middle Pennsylvanian) (Vachard et al. 2012), and was probably induced by an increasingly close association (consortium) between miliolate foraminifers and cyanobacteria. Biostratigraphy. Tubiphytes has its FAD in the latest Pennsylvanian (Chuvashov et al. 1993; Vachard et al. 2012) and its LAD in the Jurassic (Crescenti 1969; Senowbari-Daryan et al. 2008). Latitubiphytes is late Moscovian–Cisuralian, ‘Tubiphytes’ epimonellaeformis is Kungurian; Epimonella is Kungurian–Capitanian; and Ramovsia is Cisuralian. Biogeography. Tubiphytes and Latitubiphytes are cosmopolitan during the Permian. ‘Tubiphytes’ epimonellaeformis is endemic of New Mexico. Epimonella has been found in Iran, southern Turkey and New Mexico. Ramovsia is only known in the Carnic Alps (Kochansky-Devidé 1973), Iran (Jenny & Jenny-Deshusses 1978) and southern Turkey (Vachard & Moix 2013). Class Nodosariata Mikhalevich, 1993 Subclass Nodosariana Mikhalevich, 1993 Order Nodosariida Calkins, 1926 The nodosariates are distributed from late early Moscovian to Recent: their genera are either cosmopolitan or endemic. The Palaeozoic Nodosariida exhibit mostly a monolamellar wall of radiate calcite, with crystal c-axes perpendicular to the surface and without secondary lamination (Loeblich & Tappan 1987). The Palaeozoic Nodosariida are divided into two superfamilies, Nodosarioidea and Geinitzinoidea emend. herein, and 10 families: Syzraniidae Vachard in Vachard & Montenat, 1981; Protonodosariidae Mamet & Pinard, 1992; Geinitzinidae Bozorgnia, 1973; Robuloididae Reiss, 1963 nom. translat. Loeblich & Tappan, 1984; Partisaniidae Loeblich & Tappan, 1984; Frondinidae Gaillot & Vachard, 2007; Colaniellidae Fursenko in RauzerChernousova & Fursenko, 1959; Nodosariidae Ehrenberg, 1838; Pachyphloiidae Loeblich & Tappan, 1984; and Ichthyolariidae Loeblich & Tappan, 1986b. The two superfamilies are distinguished by the presence of a primitive cylindrical aperture (Geinitzinoidea) or a typical stellate aperture (Nodosarioidea). The exact status of the ancient family Nodosellinidae and its possible priority on one of these names depends of the true status of its generotype Nodosinella; the revisions of Nodosinella by Cummings (1955), Foster et al. (1985) or Pinard & Mamet (1998) do not allow a decision to be made. Superfamily Geinitzinoidea Loeblich & Tappan, 1984 emend. herein Emended diagnosis. Test subcylindrical to subtriangular uniseriate, rarely coiled and lenticular, with a primitive round terminal aperture. The septa are complete in all the families, except for the Syzraniidae family (which exhibits a progressive lineage with undivided, then pseudo-septated and finally completely septated test). The round aperture is generally simple, areal and central. A wall of radial, hyaline, fibrous calcite, generally unilayered. Sometimes, the wall is bilayered with a dark-microgranular inner layer and a clear radial, hyaline-fibrous outer layer (Syzraniidae, Protonodosariidae) or, more rarely, the wall is unilayered and dark-radial (Frondinidae). Remarks. The fundamental difference is the simple (not stellate) aperture. The superfamily was called Robuloidoidea Reiss, 1963 nom. translat. Loeblich & Tappan, 1984 emend. Gaillot & Vachard, 2007, but this name is misinterpreted because the name Geinitzinoidea Bozorgnia, 1973 translated in Loeblich & Tappan (1984, p. 20) preceeds the name Robuloidoidea translated in Loeblich & Tappan (1984, pp. 32–33). Numerous homeomorphs exist between the two superfamilies, Nodosarioidea and Geinitzinoidea, and even within the same genus (e.g. Nodosaria, Dentalina and Lingulina). The oldest hyaline-fibrous forms (Syzraniidae) appear Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 PERMIAN SMALLER FORAMINIFERS in the late early Moscovian (Kashirian); the complete septation is Late Pennsylvanian in age; the acme of the group is Permian, but it also exists in the Triassic and even the Jurassic. Family Syzraniidae Vachard in Vachard & Montenat, 1981 Permian genera. Syzrania Reitlinger, 1950; Syzranella Mamet & Pinard, 1992; Amphoratheca Mamet & Pinard, 1992; Rectostipulina JennyDeshusses, 1985 emend. Rigaud et al., 2015b; Tezaquina Vachard in Vachard & Montenat, 1981 non Vachard, 1980 emend. Alipour & Vachard in Alipour et al., 2013; Vervilleina Groves in Groves & Boardman, 1999. Remarks. This family is currently unanimously accepted as the oldest family of the Nodosariata, which are consequently monophyletic (Vachard et al. 2010 and references therein). The Triassic family Tubulastellidae Rigaud et al., 2015b is homeomorphous of the Syzraniidae, but with an aragonitic wall. As indicated above, the Syzraniidae display a progressive appearance of the septation among the nodosariates (1 & 2 in Fig. 7). Biostratigraphy. Syzrania is Moscovian– Changhsingian. Amphoratheca and Syzranella are Late Pennsylvanian and Early Permian. Rectostipulina remains typically a Lopingian marker, although its divergence from Syzrania unquestionably occurs in the Capitanian (Gaillot & Vachard 2007; Song et al. 2009; Ebrahim Nejad et al. 2015; Zhang et al. 2015). It exists in the latest Changhsingian of southern Tibet (Wang et al. 2010) and in the lowermost Triassic beds in South China (Song et al. 2007), but it is absent in the coeval series of Italy (Groves et al. 2007). Tezaquina has its FAD in late or latest Gzhelian (Russia, the Canadian Arctic, the Carnic Alps, Darvas, northern Iran); its acme seems to be Asselian –Sakmarian; it is rare and/or doubtful in the other Permian stages of Russia, where these questionable representatives might belong to Biparietata (a homeomorphous genus with a multilayered wall microstructure, which is assigned here to the Protonodosariidae). Vervilleina is generally late Gzhelian –Cisuralian, but has been noted up to the latest Permian (Shang et al. 2003; Song et al. 2007). Biogeography. Syzrania is cosmopolitan from the Moscovian to the Artinskian, it is then only Tethyan. Amphoratheca and Syzranella are probably cosmopolitan, but mainly noted in North America. Rectostipulina is known in Turkey, Slovenia, Cyprus, Greece, Afghanistan, Iran (Alborz, Zagros and Julfa areas), Armenia, Azerbaijan, Saudi Arabia, Ladakh, southern Tibet, South China, western Thailand and Cambodia. Tezaquina is first distributed in Russia, the Canadian Arctic, the Carnic Alps, 225 Darvas and northern Iran. During its acme, Tezaquina is known in Donbass (Ukraine); the Perm area and Bashkortostan (Russia); northern Iran; Tezak (Afghanistan); Darvas (Tajikistan); Bolivia; and the Barents Sea, Canadian Arctic, Greenland and New Mexico. It is rare and/or doubtful in other Permian regions of Russia (the Urals, Pechora and Primorye) and, perhaps, confused with Biparietata. Vervilleina is probably cosmopolitan, but is especially known in North and South America (Groves & Boardman 1999; Wood et al. 2002), and central Palaeotethys (Groves & Boardman 1999; Filimonova 2010). Family Protonodosariidae Mamet & Pinard, 1992 emend. Gaillot & Vachard, 2007 Subfamily Protonodosariinae Gaillot & Vachard, 2007 Permian genera. Protonodosaria Gerke, 1959 non 1952 emend. Sellier de Civrieux & Dessauvagie, 1965; Biparietata Zolotova in Zolotova & Baryshnikov, 1980; Frondinodosaria Sellier de Civrieux & Dessauvagie, 1965; Nestellorella Gaillot & Vachard, 2007; Lingulonodosaria sensu K. V. Miklukho-Maklay 1960a (non Silvestri, 1903, nec sensu Sellier de Civrieux & Dessauvagie, 1965); Nodosinelloides Mamet & Pinard, 1992; Polarisella Mamet & Pinard, 1992 emend. Gaillot & Vachard, 2007; ‘Pseudoglandulina’ Cushman, 1929 (part.). Remarks. Protonodosariidae, as proposed here, encompasses the uniseriate, tapering nodosariates with more or less hemispherical chambers and an aperture that is simple, and rarely with a neck (3– 14 in Fig. 7). They differ from the Nodosariidae in the non-stellate aperture. Probably, this family is still poorly defined, with ‘mixed morphologically different genera’ (G. Nestell pers. comm. March 2016). The most questionable genus of this conventional family is Pseudoglandulina, which was used diversely among Permian taxa, either forgotten or provided with a variety of species (e.g. Filimonova 2010), and which was, in addition, interpreted as a junior synonym of Pyramidulina Fornasini (Loeblich & Tappan 1987). Biostratigraphy. As for the family, the subfamily is latest Pennsylvanian –latest Triassic in age. Protonodosaria and Nodosinelloides have a long range: Late Pennsylvanian (Kasimovian) –latest Permian/earliest Triassic; the range of the genus Polarisella, as emendated by Gaillot & Vachard (2007), is even longer, from the Cisuralian to the Middle Triassic (Anisian) and possibly up to the Jurassic of Germany. In constrast, Biparietata would be limited to the Kungurian; and Nestellorella is questionably present in the Wordian of southern Oman (Angiolini et al. 2004), and most probably the Capitanian–late Changhsingian in Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 226 D. VACHARD Fig. 7. Smaller foraminifers (Nodosariata) from Trogkofel (Late Cisuralian of the Carnic Alps, Austria). Scale bar ¼ 0.100 mm. (1) Tezaquina sp. (2) Vervilleina sp. (3)–(14) Nodosinelloides spp. (15) – (24) Geinitzina spp. (25) – (27) Pachyphloia spp. Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 PERMIAN SMALLER FORAMINIFERS 227 Armenia –Azerbaijan (Pronina 1988a; Kotlyar et al. 1989), the Capitanian in the central Pamirs (Dronov 2004), the late Capitanian in Hazro and the Lopingian of Zagros (Gaillot & Vachard 2007). Cryptoseptida were mentioned in South China (Wang & Ueno 2003), Japan (Kobayashi 1996, 2001, 2002) and Hungary (Berczi-Makk 1996). Biogeography. As for the family, the genera are partly cosmopolitan, partly Tethyan. Protonodosaria, Nodosinelloides and Polarisella are probably cosmopolitan. Nestellorella is Neotethyan in southern Oman (Angiolini et al. 2004), Armenia– Azerbaijan (Pronina 1988a; Kotlyar et al. 1989), in the Rushan –Pshart zone in the central Pamirs (Dronov 2004), Hazro (Turkey) and Zagros. Permian genus. Nodosinella Brady, 1876. Remark. Despite many attempts, the old genus Nodosinella was neither correctly revised nor refound; it perhaps resembles Protonodosaria with a differentiated and/or recrystallized wall. Subfamily Langellinae Gaillot & Vachard, 2007 Biogeography. England; all the other localities have to be revised. Permian genera. Langella Sellier de Civrieux & Dessauvagie, 1965; Pseudolangella Sellier de Civrieux & Dessauvagie, 1965; ?Cryptoseptida Sellier de Civrieux & Dessauvagie, 1965; ?Maichelina Sosnina, 1977. Remarks. Because of the reservations expressed above regarding the family, the individuality and composition of this subfamily can be discussed. Eomarginulinella and Maichelina are poorly known. Despite numerous misinterpretations of its type species C. anatoliensis Sellier de Civrieux & Dessauvagie, 1965, Cryptoseptida continues to be commonly used in the literature. It has been confused with Pachyphloides by Berczi-Makk (1996), Polarisella ex gr. elabugae by Kobayashi (1996), Pseudolangella by Kobayashi (2001, pl. 2, fig. 14) and Aulacophloia by Kobayashi (2002, fig. 9. 40, 41). We propose herein its assignment to Langellinae because of its increasing thickness of wall, the septa being thinner than the wall, the shape of the chambers and the type of aperture. Biostratigraphy. Langella, rare in the Artinskian– Kungurian (¼Longlingian of South China), has its acme in the late Chihsian and its LAD in the Changhsingian. Pseudolangella has a probably Sakmarian FAD and a Changhsingian LAD; it is questionable in the Early Permian, and has its acme in the Guadalupian (¼Middle Permian). Cryptoseptida was described in the ‘Calcaires à Bellerophon’ of Turkey: that is, a composite unit probably late Guadalupian and Lopingian in age. It was never really re-found in the Lopingian except by Kotlyar et al. (1984) in the early Wuchiapingian. Biogeography. Langella and Pseudolangella are relatively widespread in the Palaeo- and Neotethys (see Vachard 1990), from Italy to South China, and the Panthalassan in New Zealand (see the detailed distribution in Gaillot &Vachard 2007). Maichelina is endemic to Primorye. Cryptoseptida seems endemic to an area encompassing southern Turkey, Armenia and Azerbaijan (Sellier de Civrieux & Dessauvagie 1965; Kotlyar et al. 1984). Questionable ?Family Nodosinellidae Rhumbler, 1895 Biostratigraphy. Middle or Late Permian. Family Geinitzinidae Bozorgnia, 1973 Permian genera. Geinitzina Spandel, 1901 (¼ ?Neogeinitzina K.V. Miklukho-Maklay, 1954); Frondinodosaria Sellier de Civrieux & Dessauvagie, 1965; Geinitzinita Sellier de Civrieux & Dessauvagie, 1965; Gerkeina Grozdilova & Lebedeva in Sosipatrova, 1969; Howchinella Sellier de Civrieux & Dessauvagie, 1965 emend. Palmieri in Foster et al., 1985; Lunucammina Spandel, 1898 (¼?Frondicularia Defrance in d’Orbigny, 1826 sensu Sellier de Civrieux & Dessauvagie, 1965); Omoloniella Karavaeva & Nestell, 2007; Pachyphloides Sellier de Civrieux & Dessauvagie, 1965; Pseudotristix Miklukho-Maklay, 1960a (¼Multifarina Lin, 1984 ¼Tristix sensu Sellier de Civrieux & Dessauvagie non MacFadyen, 1941 (part.)); Rectoglandulina sensu Karavaeva & Nestell, 2007 non Loeblich & Tappan, 1955; Reitlingeria Pronina in Kotlyar et al., 1989; Spandelinoides Cushman & Waters, 1928c; Tauridia Sellier de Civrieux & Dessauvagie, 1965; ?Pseudonodosaria Boomgaart 1949 (partially considered as synonymous of Rectoglandulina). Remarks. Once again, the use of Recent genera for Permian taxa is questionable in this family and, except for the generotype Geinitzina, which is relatively well characterized (15–24 in Fig. 7), the other genera are discussed and/or poorly known. Pseudonodosaria is used by few authors for Permian taxa (e.g. Woszczynska 1987; Palmieri et al. 1994; Karavaeva & Nestell 2007; Filimonova 2010); it needs revision as does Rectoglandulina, which was abandoned by its originators (see Loeblich & Tappan 1987), but ‘reinstated’ recently by Karavaeva & Nestell (2007): further studies are required because the taxon described under this name by Gerke (1961) unquestionably exists and is noticeable in the Permian biostratigraphy. The genus Lunucammina remains equally mysterious, even after the long discussions of Loeblich & Tappan (1964, 1987), Sellier de Civrieux & Dessauvagie (1965) Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 228 D. VACHARD and K. V. Miklukho-Maklay (1960b). In my opinion, most species attributed to this genus in the literature belong to Frondicularia sensu Sellier de Civrieux & Dessauvagie, 1965. However, Palmieri et al. (1994) have indicated that many Russian species of Frondicularia correspond to Australian species of Howchinella and ‘Ichthyolaria’ (see below). Owing to their concomitant occurrences in Zechstein deposits, Nodosinella and Lunucammina are possibly synonymous. As for Nodosinella, the genus Tauridia remains a very enigmatic taxon (Fig. 8); moreover, it was often mispelled as Taurida (e.g. Pinard & Mamet 1998; Groves 2000), and misinterpreted by Loeblich & Tappan (1987) as a synonymous of Frondina, the wall of which differs completely. Biostratigraphy. The typical representatives of the family exist from the latest Pennsylvanian to the Triassic. Geinitzina is latest Pennsylvanian (Groves 2000, 2002) or earliest Permian (Groves & Wahlman 1997) to Changhsingian (Lin et al. 1990). The lowermost Triassic ‘Geinitzina’ recently mentioned in Italy (Groves et al. 2007) are most probably representative of Nestellorella (fig. 10.5) and Nodosaria (fig. 10.6, 14, 15). The possible synonym Neogeinitzina is limited to the Changhsingian. Geinitzinita, which is generally a Mesozoic genus, is known by two or three species in the Changhsingian (Vuks 1984; Gu et al. 2007). Pseudotristix is generally cited in the Wuchiapingian (¼Dzhulfian) from Turkey to southern China (including Multifarina, where it is indicated as Maokouan in age by Lin et al. 1990). Its FAD seems to be late Guadalupian in Sumatra, western Thailand, central Japan and Primorye, and its LAD is Changhsingian in age in Greece (Hydra), Israel, Armenia –Azerbaijan, Italy, Zagros, Cambodia, South China, North Thailand and Japan. Pachyphloides belongs to the Maokouan, Wuchiapingian? and Changhsingian. Rectoglandulina is probably homeomorphous of a taxon found in the Cretaceous and Paleocene –Holocene (?); during the Permian, it seems to be Kungurian–Changhsingian in age. The taxon ‘Pseudonodosaria’, which is generally mentioned as late Cisuralian in age, is similarly homeomorphous of a Holocene taxon; it was indicated as extending ‘beyond the P–T boundary elsewhere in South China’ (Jin et al. 2000). The range of Lunucammina ¼ Frondicularia is discussed; perhaps late Cisuralian–late Lopingian. Howchinella is Early Permian (early Cisuralian, Artinskian) –Late Permian (latest Changhsingian) (Gu et al. 2007; Karavaeva & Nestell 2007; Song et al. 2009; Zhang & Gu 2015). Gerkeina is Kungurian. Omoloniella is late Kungurian–Lopingian and perhaps Triassic. Spandelinoides is Late Pennsylvanian – Early Permian. The range of Tauridia is little known; perhaps from Cisuralian to Lopingian. Biogeography. Geinitzina is cosmopolitan; its possible synonym Neogeinitzina is, however, only Fig. 8. A typical nodosariate, Tauridia n. sp. Scale bar ¼ 0.250 mm. Late Artinskian of Zweikofel (Carnic Alps, Austria). Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 PERMIAN SMALLER FORAMINIFERS mentioned in NW Caucasus (K.V. MiklukhoMaklay 1954; Pronina-Nestell & Nestell 2001), South China (Wang 1974; Lin 1984; Lin et al. 1990; Zhang & Hong 2004) and NW Iran (Mohtat-Aghai et al. 2009). Geinitzinita is mentioned in Armenia –Azerbaijan (Vuks 1984) and South China (Gu et al. 2007). Pseudotristix is sporadically cited from Italy, Greece, Poland, Turkey, Israel, Armenia, Azerbaijan, Iran (Zagros, NW Iran), Sumatra, Cambodia, South China, North Thailand, Japan and Primorye. Pachyphloides is mentioned in NW Turkey, NW Caucasus, Zagros and South China. Rectoglandulina and Pseudonodosaria are probably cosmopolitan, but still poorly known. Lunucammina (¼Frondicularia part.) was found in Germany, Italy, Hungary, Poland, Iran (Zagros and Fars), eastern Himalaya, Australia and ?Japan (where it was probably confused with Frondina). Howchinella is probably cosmopolitan, but relatively rare everywhere, except in the Far East of Russia and Australia. Similarly, Tauridia, although poorly known, is possibly cosmopolitan, as it is currently cited in Turkey, the Carnic Alps, the Urals and New Mexico. Endemic genera are Spandelinoides (Texas, USA), Gerkeina (the Urals) and Omoloniella (NE Siberia, Russia). Family Robuloididae Reiss, 1963 nom. translat. Loeblich & Tappan, 1984 Permian genera. Robuloides Reichel, 1946; Pararobuloides Miklukho-Maklay, 1954; Eocristellaria Miklukho-Maklay, 1954; Cryptomorphina Sellier de Civrieux & Dessauvagie, 1965; Gourisina Reichel, 1946; Hubeirobuloides Lin, Li & Zheng in Lin et al., 1990; ?Calvezina Sellier de Civrieux & Dessauvagie, 1965; ?Eomarginulinella Sosnina, 1969. Remarks. This family encompasses all the planispirally coiled nodosariates of the Permian and, therefore, of the Palaeozoic. Hence, Calvezina, initially coiled, is related here to this family. Gourisina, very rarely re-found since its description, might correspond to a teratogenic form of Pararobuloides,especially the species Gourisina rossica K. V. Miklukho-Maklay, 1954. Biostratigraphy. Robuloides, misinterpreted as ‘Artinskian or younger’ (Mamet 1996), is Capitanian –latest Changhsingian (and even earliest Triassic in South China: Song et al. 2007). Pararobuloides is Wuchiapingian– Changhsingian. Hubeirobuloides is late Capitanian–late Changhsingian. The genus Eocristellaria is relatively poorly known, with an acme probably located in the Late Permian, and a possible FAD in the Middle Permian (with some early Kazanian forms described under the name of ‘Astacolus’). Calvezina, the FAD of which is probably early Middle Permian, 229 has its acme in the Capitanian; its LAD is late Changhsingian (Altıner 1984; Song et al. 2009). Its possible synonym Eomarginulinella is Middle Permian. Cryptomorphina is late Guadalupian – Changhsingian. Gourisina is only typical in the late Changhsinghian. Biogeography. Robuloides was found in Greece (Hydra, Attica), Slovenia, Hungary, Cyprus, Turkey, Israel, NW Caucasus, Armenia, Azerbaijan, Saudi Arabia, central Iran, Afghanistan, the Salt Range, SE Pamirs, southern Tibet, South China, Primorye, Japan and New Zealand; this genus is absent from the Americas (contrary to Mamet 1996). Pararobuloides is known in Greece, NW Caucasus, Turkey, Oman, southern Tibet, Thailand and Japan. Hubeirobuloides is only illustrated in South China (Lin et al. 1990), NW Caucasus (Pronina-Nestell & Nestell 2001), Crimea (Pronina & Nestell 1997) and Japan (as Pachyphloia? sp.: Kobayashi 2001). Eocristellaria probably exists in Turkey, central Iran (Abadeh: Okimura & Ishii 1981), South China (Zhang & Gu 2015) and, perhaps, in Australia (Palmieri et al. 1994). Calvezina was encountered in Turkey, Greece, Alborz, Armenia, Azerbaijan, Himalaya, Primorye, ?Italy, ?Hungary, ?Malaysia, ?Japan and ?Texas. Eomarginulinella was mentioned (often as Calvezina) in Primoye (in the far east of Russia: Sosnina 1969), Armenia–Azerbaijan (Kotlyar et al. 1989; Pronina 1996), southern Turkey (Altıner 1981), New Mexico (Nestell & Nestell 2006) and Slovenia (Nestell et al. 2009). Cryptomorphina is known in Turkey (central Taurides and Hazro), Italy (Tesero section) and Iran (Zagros; NW Iran). Gourisina is known from Greece (Reichel 1946), NW Caucasus (K.V. MiklukhoMaklay 1954) and, questionably, from central Iran (Abadeh) (Baghbani 1993). Family Partisaniidae Loeblich & Tappan, 1984 Permian genera. Partisania Sosnina, 1978 (¼Xintania Lin, 1984); Eoguttulina Cushman & Ozawa, 1930 (part.); ?Nodoinvolutaria Wang in Lin, 1978. Remarks. This poorly known family might be the ancestor of the Mesozoic polymorphinids (Rigaud et al. 2015b). Biostratigraphy. The family has been cited in Wordian, Kazanian, Capitanian, Tatarian and Lopingian. Biogeography. Palaeotethyan, Neotethyan and Panthalassan Partisania are known in Primorye, NW Thailand, South China, Philippines, Japan, Armenia, New Zealand, Oman, Cyprus, Crimea, Iran (Fars area) and New Mexico. Eoguttulina (or more probably a homeomorph of this genus) was very rarely cited in Russia (Pronina 1999a). Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 230 D. VACHARD Nodoinvolutaria is Maokouan–late Changhsingian of South China (Lin et al. 1990; Wignall & Hallam 1996; Gu et al. 2007). Family Frondinidae Gaillot & Vachard, 2007 Permian genera. Frondina Sellier de Civrieux & Dessauvagie, 1965 emend. Gaillot & Vachard 2007; Ichthyofrondina Vachard in Vachard & Ferrière, 1991 emend. Gaillot & Vachard, 2007. Remarks. This family contains two small forms that have a dark-hyaline wall, an atypical character amongst the Nodosariata. They are interesting because they are probably important for the biozonation of the late Capitanian –Lopingian. Biostratigraphy. Frondina and Ichthyofrondina are late Capitanian –Changhsinghian in age, and possibly exist up to the lowermost Triassic in Italy (Groves et al. 2007) and South China (Song et al. 2007). Furthermore, Frondina is questionably present up to the Cretaceous (Sellier de Civrieux & Dessauvagie 1965). Biogeography. Frondina and Ichthyofrondina are Palaeotethyan and Neotethyan, and both occur in Turkey, Cyprus, Greece, Tunisia, Iran, NW Caucasus, Armenia, Azerbaijan, Saudi Arabia, Oman, southern Tibet, NW Thailand, South China and Japan. Frondina has been also mentioned in New Zealand (Vachard & Ferrière 1991), Saudi Arabia (Hughes 2005) and Slovenia (Nestell et al. 2009). In North America, both genera were never correctly identified; nevertheless, in El Antimonio (Sonora, Mexico), Brunner (1979) has illustrated two specimens that resemble Frondina (pl. 3, fig. 8) and Ichthyofrondina (pl. 3, fig. 3). Family Colaniellidae Fursenko in RauzerChernousova & Fursenko, 1959 Permian genera. Colaniella Likharev, 1939 (¼Pseudocolaniella Wang, 1966; ¼Paracolaniella Wang, 1966); Cylindrocolaniella Loeblich & Tappan, 1985 (¼Wanganella Sosnina in Kiparisova et al., 1956 pre-occupied); Pseudowanganella Sosnina, 1983. Remarks. This family is well characterized by the shape of its chambers; it encompasses few genera that rapidly evolved and totally disappear. Biostratigraphy. Colaniella is latest Capitanian– late Changhsingian; Cylindrocolaniella and Pseudowanganella are Lopingian. Pseudocolaniella and Paracolaniella, which are very similar in shape to Colaniella and have the same stratigraphic distribution, are considered herein as junior synonyms of Colaniella. Biogeography. Colaniella is essentially a Palaeotethyan and Panthalassan taxon (e.g. see Ishii et al. 1975; Vuks & Chediya 1986; Okimura 1988; Shang et al. 2003; Wang et al. 2010; Kobayashi 2013; Vachard 2014); it remains rare and only represented by primitive forms in the peri-Gondwanan margin of the Neotethys (Israel, Turkey, Iran, Saudi Arabia, Oman and Tunisia). Cylindrocolaniella and Pseudowanganella are endemic of Primorye because Pseudowanganella, illustrated by Nestell & Pronina (1997, pl. 1, figs 26 & 27) in Cyprus, corresponds, in our opinion, to a more primitive taxon; the Japanese forms assigned to Wanganella (Ishii et al. 1975; Okimura 1988) most probably being true Colaniella to compare, for example, with Colaniella bozkiri Çatal & Dager, 1974 from Taurus (southern Turkey). Superfamily Nodosarioidea Norvang, 1957 nom. translat. Loeblich & Tappan, 1961 Family Nodosariidae Ehrenberg, 1838 Subfamily Nodosariinae Ehrenberg, 1838 nom. translat. Reuss, 1862 Permian genera. Nodosaria Lamarck, 1816 non 1812 (see synonyms in Loeblich & Tappan 1987; and those which are different in Karavaeva & Nestell 2007); Dalongella Gu et al., 2007; ‘Dentalina’ non? d’Orbigny, 1826; ‘Lingulina’ d’Orbigny, 1826 emend. Sellier de Civrieux & Dessauvagie, 1965. Remarks. Uniseriate, rectilinear, with hemispherical to elongate chambers and a stellate aperture. Lingulina is diversely interpreted, sometimes as a Frondina. Biostratigraphy. Nodosaria is Permian–Recent. Karavaeva & Nestell (2007) indicated a Roadian FAD for the ‘first true Dentalina-like forms’, but the second, false and/or typical specimens of ‘Dentalina’ are not well characterized and are probably homeomorphs of undeterminate genera. ‘Lingulina’ is generally late Capitanian–Changhsingian in age (Pronina, 1999b), whereas Dalongella is Changhsingian. Apparently, the taxon ‘Pseudolingulina’ spp. (sic) of Orlov-Labkovsky (2004, text-fig. 2), mentioned in the late Wuchiapingian–early Changhsingian of Israel, is a nomen nudum. Biogeography. Nodosaria is cosmopolitan. Dalongella is endemic of South China. Permian ‘Dentalina’ and ‘Lingulina’ have been cited all around the world, especially in Russia, Western Europe and China. Family Pachyphloiidae Loeblich & Tappan, 1984 Permian genera. Pachyphloia Lange, 1925; Robustopachyphloia Lin, 1980; Aulacophloia Gaillot & Vachard, 2007; Sosninella Sellier de Civrieux & Dessauvagie, 1965 nomen imperfectum. Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 PERMIAN SMALLER FORAMINIFERS Remarks. This typical Permian family is now relatively well known, except for the ancestral forms of Pachyphloia, which probably appear in the late Cisuralian (25–27 in Fig. 7). Biostratigraphy. The complete range of Pachyphloia in the literature is Early Permian –Jurassic: however, Pachyphloia has its FAD in the Kungurian (Lin et al. 1990; Pinard & Mamet 1998; Groves 2000; Vachard & Krainer 2001b; Filimonova 2010). Cisuralian citations (e.g. Groves & Wahlman 1997) correspond most probably to oblique sections of Geinitzina. Pachyphloia has its acme in the Guadalupian–Lopingian (especially in Tunisia, Turkey and Iran); its LAD is most probably latest Permian (e.g. Lin et al. 1990; Pronina-Nestell & Nestell 2001; Shang et al. 2003; Wang & Ueno 2003; Groves et al. 2007). Robustopachyphloia is late Capitanian, Wuchiapingian and Changhsingian. Aulacophloia is rather early Changhsingian in age, but was mentioned in the late Changhsingian (Wang et al. 2010). Biogeography. Pachyphloia is cosmopolitan. Robustopachyphloia is known from South China, central Japan, New Mexico, southern Turkey (Hazro), Zagros and Oman (Hauser et al. 2000; Gaillot & Vachard 2007), as well as the central Pamirs (Pronina 1996), Texas, USA (Nestell et al. 2006) and Tunisia (Ghazzay et al. 2015). Aulacophloia exists in Turkey, Saudi Arabia and southern Tibet. Cryptoseptida was described in the ‘Calcaires à Bellerophon’ of Turkey: that is, a composite unit probably Guadalupian and Lopingian in age, but Cryptoseptida was never really re-found – for example, its type species C. anatoliensis re-illustrated by Groves et al. (2005, fig. 23.26) is more likely to be Langella ocarina Sellier de Civrieux & Dessauvagie, 1965. Family Ichthyolariidae Loeblich & Tappan, 1986b Permian genus. Ichthyolaria sensu Sellier de Civrieux & Dessauvagie, 1965 (part.) non Wedekind, 1937. Remarks. The presence of true Ichthyolaria (i.e. defined by the type species Frondicularia bicostata d’Orbigny, 1850), or simply of ichthyolariids, in the Permian seems to be very doubtful. Furthermore: (1) the Permian taxon is often misspelled Ichtyolaria or even Icthyolaria; and (2) the Permian Ichthyolaria are generally confused with the genus Ichthyofrondina, which differs in its wall microstructure. Biostratigraphy. The genus Ichthyolaria and its probable homeomorphs are cited from the Permian to the Jurassic. Biogeography. Permian homeomorphs of Ichthyolaria were mentioned in the Early Permian of 231 Australia (Palmieri 1984; Palmieri et al. 1994); Middle –Late Permian of Turkey (Sellier de Civrieux & Dessauvagie 1965); Middle Permian of Cyprus (Nestell & Pronina 1997); Middle Permian of Russia, Volga–Kama region (Pronina 1998, 1999a), Taymir (Pronina 1999a) and NE Siberia (Karavaeva & Nestell 2007); late Guadalupian of New Mexico (Nestell & Nestell 2006) and South China (Gu et al. 2007; Zhang & Gu 2015). Group incertae sedis misinterpreted as Nodosariata It would take too long to discuss the history and the assignment of the genus Sphairionia Nguyen Duc Tien, 1989b non 1987 ¼ Pseudosphairionia Pronina, 1996 (see e.g. Pronina 1996; Vachard et al. 2001a; Sone 2008). The genus is interesting because it is a biomarker of the early Midian (Pronina 1996). This regional substage is the equivalent of the early Capitanian (e.g. Henderson et al. 2012; Davydov & Arefifard 2013; Ghazzay et al. 2015). Moreover, Sphairionia is a good biogeographical marker, being probably only peri-Gondwanan and/or Cimmerian (see the Discussion later). Pronina (1996) attributed this genus to the nodosariates: however, its morphology totally differs (especially when the sections of the Pseudosphairionia type are integrated into the reconstruction of this microfossil, as in Vachard et al. 2001a). Furthermore, the microstructure of each layer of the bilayered wall differs from that of the nodosariates, whereas, as indicated by several authors (Vachard 1980; Mertmann 2000; Vachard et al. 2001a), this microstructure is more similar to that of the umbellacean microproblematica. The umbellacean, generally assigned to the Charophyta (e.g. Vachard 2000 and references therein), are known from the late Givetian to the Tournaisian (?early Visean), but similar forms have been mentioned up to the Miocene (Vachard et al. 1982). A Lazarus effect during the Permian is not excluded. Class Textulariata Mikhalevich, 1980 The taxa originally considered to have an agglutinated test, like Ammodiscus, Glomospira, Tolypammina, Hyperammina, Reophax or Ammobaculites (e.g. Sosipatrova 1970; Ukharskaya 1970; Woszczynska 1987; Pronina 1998), should be assigned to Fusulinata and/or Miliolata genera (e.g. Vachard et al. 2010; Hance et al. 2011). Notice that when an agglutinated genus name existed in the literature, the earliest authors often gave the name of a Textulariata: for example, Rectoseptournayella chappelensis was initially called Ammobaculites chappelensis (see Vachard et al. 2010). Today, the trend is to give a Palaeozoic name (e.g. Pseudoammodiscus, Pseudoglomospira, Earlandia and so on) to these taxa: that is, to suggest that they Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 232 D. VACHARD possess a calcitic dark-migrogranular wall, not agglutinated. Recently, the pendulum has swung back, and the names Ammodiscus and Hyperammina have, again, been proposed (Nestell et al. 2015). This problem is therefore very complex. As indicated by Rigaud et al. (2014), large benthic agglutinated and microgranular foraminifers show significant morphological and microstructural similarities. The two groups have yet to be regarded as being part of the distinct lineages on the basis of a gap exceeding 50 myr in the record of the first large alveolar agglutinated Mesozoic forms (Textulariida) and their last known Palaeozoic microgranular homologues (Fusulinana). In fact, both Fusulinana and the Textulariida may possess a microgranular test, potentially containing some optically discernible agglutinated grains and a keriothecal/alveolar texture. Three major hypotheses can explain these striking wall similarities: (1) the Textulariida and the Fusulinana have experienced a convergent evolution; (2) the Endotebidae belong to the Textulariida and have been erroneously included in the Fusulinana; and (3) all Fusulinana have agglutinated microparticles, the microgranular appearance of their wall being related to the nature and size of the grains that they incorporate. In the latter case, the Fusulinana would have survived the Permo-Triassic and the Triassic–Jurassic major extinction events and still have representatives among some lineages of Textulariata. In consequence, there would be no major extinct groups of foraminifers, and molecular phylogeny, combined with fossil data, should allow the foraminiferal evolution to be retraced to a very advanced level. As a result, the Textulariata reappear, or seem to reappear, in the late Middle Permian, with the first Ataxophragmiidae (Gaillot 2006). Undoubtedly, the family is represented by well-identified loose specimens in the Kazanian (¼Middle Permian) of Russia (Ukharskaya 1970). Another correlation is possible: an inner organic lining exists among the Ataxophragmiidae. The first unquestionable inner organic linings of foraminifers, found in Palaeozoic sediments, are Late Permian in age (Stancliffe 1989; Groves et al. 2004). Owing to the monophyletism of the Rotaliata (e.g. Flakowski et al. 2005), the lineage admitted between Fusulinata and Rotaliata via the Nodosariata (Gaillot & Vachard 2007), and the connection between Fusulinata and Textulariata suggested herein, it seems difficult to integrate the Textuladiida in the Rotaliata as proposed by Mikhalevich (2003). Order Textulariida Lankester, 1885 Trying to accurately differentiate the Fusulinata and the Textulariata, Vachard et al. (2010) have indicated that the siliceous agglutinate was exclusive to Textulariata, whereas the agglutinates of the Fusulinata, when they existed, were always calcareous. This statement is not entirely true because some Palaeotextularioidea, Endoteboidea and even Endothyroidea (e.g. Endothyranella kamaica Baryshnikov in Baryshnikov et al., 1982) can exhibit a siliceous agglutinate wall (Fig. 9). Among the Palaeotextularioidea with siliceous agglutinates, the genus Palaeobigenerina Galloway, 1933 (type species: Bigenerina geyeri Schellwien, 1898) is probably the most typical. This genus is rarely used, probably because the original description of its apertures is erroneous. It was described with an agglutinating wall, but, in this case, the original description is probably exact. This agglutinated wall is obvious in the Carnic Alps (e.g. Vachard & Krainer 2001b, pl. 4, figs 10 & 14) and northern Afghanistan (Vachard unpublished data). The most developed siliceous agglutinate was probably illustrated in ‘Deckerella sp. 1’ by Nestell et al. (2006, pl. 5, fig. 20). The name Textularia has been Fig. 9. Regularly arranged siliceous agglutinate in an oblique subtransverse section of a palaeotextulariid (the inner pseudo-fibrous hyaline layer is also visible). It should be noted that for Miklukho-Maklay (1956) only the pseudofibrous layer was interpreted as secreted, while the siliceous grains and the calcitic grains of the dark-microgranular layer were considered as agglutinated. Scale bar ¼ 0.100 mm. Late Artinskian of Zweikofel (Carnic Alps, Austria). Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 PERMIAN SMALLER FORAMINIFERS recently used for naming Middle Permian (Kazanian) species (Ukharskaya 1970; Pronina 1998). As proposed by Altıner & Özkan-Altıner (2010), it is possible that Labiodagmarita should be assigned to Palaeotextulariidae and not to Dagmaritidae. In contrast, the similar nomenclatural change, proposed by the same authors, for Bidagmarita is not acceptable because this latter genus possesses an unquestionable wall of dagmaritins. Order Verneuilinida Mikhalevich & Kaminski in Mikhalevich, 2003 First Verneuilinida seem appear in the Kubergandy Formation (late Kungurian–early Roadian) of Tajikistan (Angiolini et al. 2015). Then, Verneuilinida are illustrated in the Capitanian of Oman (Vachard et al. 2002). Many species of Verneuilinoides and Digitina were described in the Middle Permian (Kazanian) of Russia (Ukharskaya 1970; Pronina 1998) and in the Permian of Australia (e.g. Crespin 1958). Sections of Verneuilinidae also correspond to ‘Reophax sp.’ of Flügel et al. (1984, pl. 32, fig. 6) and Climacammina sp. (Flügel et al. 1991, pl. 43, fig. 8), both from the Middle Permian in Croatia and Sicily. See also Ataxophragmiidae indet. in Vachard et al. (2002, pl. 1, figs 1 & 2) from the Capitanian of Oman. Monodorina Lin et al., 1990, described as coming from the Early Permian of South China, is most probably a Triassic species. Order Lituolida Lankester, 1885 Vachardella, attributed here to the Endoteboidea, might alternatively be the first lituolid in the geological story. Discussion: biostratigraphic and palaeobiogeographical implications Biostratigraphy of the Permian smaller foraminifers Smaller foraminifer biozones were introduced in the Early Permian formations of the Pamirs (Tajikistan) by Filimonova (2010), but it was impossible to identify these biozones either in our North American material (Fig. 10) or in our Tethyan material, probably because: (1) the evolution for all the groups has been very slow for Kasimovian–Artinskian times; and (2) many species are poorly defined. During the Middle and Late Permian, the Miliolata and Nodosariata display the best biostratigraphic value of the smaller foraminifers owing to their quite rapid evolution and their strong diversification. Some small Fusulinata, such as Globivalvulinoidea, remain biostratigraphically interesting (see Altıner & Brönnimann 1980; Altıner 1999; Altıner & Özkan-Altıner 2001, 2010; Gaillot & Vachard 233 2007; Gaillot et al. 2009), as well as the Spireitlinoidea genus Endoteba (see Vachard et al. 2013) and Pseudovidalinidae. As recorded for the fusulinids, after recovery in the Wuchiapingian after the end-Guadalupian crisis, the Changhsingian was a period of increasing diversity of the genera of the smaller foraminifers. A biostratigraphic scheme, proposed in this paper, is given in Figure 11. Cisuralian. Many survivals from the Late Pennsylvanian are present and relatively numerous up to the Artinskian: Endothyra, Bradyina, Tetrataxis, calcivertellids and Nodosinelloides. The Asselian is characterized by oncolites built by Ellesmerella (formerly ‘Girvanella’), occasionally associated with Claracrusta Vachard, 1980 (incertae sedis algae) and cyanobacteria (rarely, true Girvanella). No genera of smaller foraminifers are characteristic for the Asselian; only species of Bradyina/Bradyinelloides, Endothyra, Pseudovidalina, Hemidiscus and Tetrataxis can be used in association when characterizing the Asselian. Amongst the Nodosariata, characteristic species belong to the genera Nodosinelloides, Polarisella, Protonodosaria and Geinitzina (the FAD of which occurs in the Carboniferous– Permian boundary interval). Sakmarian. This period has been characterized by Blazejowski (2009) with species of Calcitornella and Midiella, and by Filimonova (2010) with species of Geinitzina and Nodosinelloides. Species of Pseudovidalina and Bradyina could be also useful. Artinskian. The Artinskian sees the LAD of Bradyina and the possible replacement of Protonodosaria (or Nodosinelloides) by Nodosaria: that is, the appearance of the stellate aperture in this group. Pseudovermiporella appeared in the Sakmarian – Artinskian boundary interval and became rapidly common. Kungurian. The main bioevents are the LAD of Mesolasiodiscus and Xingshandiscus. Various species of Globivalvulina are noticeable, such as G. praegraeca Vachard et al., 2015, which heralds the relative gigantism of this genus in the Capitanian –Lopingian. First Septoglobivalvulina appear, or at least the transitional forms between Globivalvulina and Septoglobivalvulina. In association, giant Geinitzina and first atypical Rectoglandulina and Pachyphloia are found, and the FAD of Nestellorella occurs. Roadian. Smaller foraminifers are poorly known during this period. New species of Nodosaria, first typical species of Pachyphloia and Rectoglandulina appear in many areas of western Palaeotethys, and probably all around the world. The gigantism of the Miliolata and Nodosariata could begin during Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 234 ZONATION D. VACHARD SAN BIOMARKERS (and representative samples) Olgaorlovella davydovi Tubiphytes epimonellaeformis Geinitzina indepressa ANDRES Fm Hemigordiellina spp. FORMATIONS Zone 15 Zone 14 REGIONAL STAGES INTERNATIONAL STAGES late KUNGURIAN middle KUNGURIAN Zone 13 GLORIETA Fm CATHEDRALIAN Frondicularia aff. turae Zone 12 Zone 11 barren of foraminifers YESO Gr Ellesmerella rara Nestellorella ? sp. Globivalvulina novamexicana Orthovertellopsis protaeformis Glomomidiella infrapermica Zone 10 Zone 9 Zone 8 ROBLEDO Fm Zone 7 Pseudoreichelina sp. Geinitzina sp. 2 [? multicamerata ] Praeneodiscus sp. Globivalvulina novamexicana Globivalvulina praegraeca Pseudovermiporella spp. Zone 6 Pachyphloia? sp. Geinitzina postcarbonica Nodosinelloides longissima Zone 5 Community Pit Fm Zone 4 L E O N A R D I A N Nodosinelloides pinardae Globivalvulina parapiciformis Geinitzina postcarbonica early KUNGURIAN late ARTINSKIAN middle ARTINSKIAN HESSIAN early ARTINSKIAN W O L F C A M P I A N L E N O X I A N LENOXIAN Hedraites sp. Zone 3 latest SAKMARIAN late SAKMARIAN early SAKMARIAN late ASSELIAN NEALIAN Geinitzina postcarbonica Nodosinelloides netjachewi Zone 2 Zone 1 Shalem Colony Fm Leptotriticites sp. Nodosinelloides longissima Tubiphytes sp., Geinitzina sp. Pseudovidalina sp., Pseudoschwagerina sp. middle ASSELIAN BURSUMIAN early ASSELIANlatest GZHELIAN Fig. 10. Cisuralian biozonation in New Mexico based on smaller foraminifers (according to Lucas et al. 2015; Vachard et al. 2015). Abbreviations: DAA, DAB and DAC designate samples of the sections A, B and C in the Doña Ana Mountains (DA) in New Mexico. this period with the first true Neodiscus, Graecodiscus, Agathammina and Langella, amongst others. Wordian. Wordian smaller foraminifers are also poorly known and often confused with the Capitanian taxa (partly because the two stages were previously linked under the name Murgabian or Murghabian). Giant Nodosariata diversify with Langella, Calvezina and large Nodosaria. The FAD of Postendothyra and Hemigordiopsis, if confirmed, might become characteristic of the Wordian. Capitanian. Numerous FAD of smaller foraminifer genera occur during this period, and especially during the late Capitanian (El Capitan stratotype in the USA, Tebaga, Turkey, Thailand and South China). It is also the acme of Hemigordiopsis and Endoteba, especially in Tebaga (Tunisia). The first colaniellids and the first Altineria appear near the end of the late Capitanian. Middle–Late Permian interval. A possible significant time to distinguish in the Permian history Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 PERMIAN SMALLER FORAMINIFERS 235 Fig. 11. Proposed biomarkers and bioevents among the smaller foraminifers during the Permian. is the Capitanian–Wuchiapingian boundary interval, in areas such as El Capitan (USA), Turkey, Japan, Iran (Abadeh), Tunisia (Tebaga), Armenia and Azerbaijan (e.g. see, Gaillot & Vachard 2007; Ghazzay et al. 2015 and references therein). This period could be characterized by the following fusulinid events: the FAD of true Codonofusiella, the FAD of true Reichelina and the LAD of Chusenella – see, for example, the Khachik Formation in Armenia (Leven 1998), the Sphaerulina beds of Abadeh in Iran (Iranian– Japanese Research Group 1981; Kobayashi & Ishii 2003; Davydov & Arefifard 2013), the ‘barren interval’ of Isozaki et al. (2007) and the Neoendothyra permica Zone in Japan (Kobayashi 2012b). Some smaller foraminifers are also characteristic for this time interval: Shanita, Lysites and Altineria alpinotaurica (Altıner, 1988), and a possible transitional taxon between Syzrania and Rectostipulina (this latter genus having a Wuchiapingian FAD, as well as Paradagmarita: see Gaillot & Vachard 2007). Wuchiapingian and early Changhsingian. Both of these periods in the Lopingian superstage share the same smaller foraminifers, which had already appeared in the late Guadalupian. Langella, Pachyphloia, Pseudolangella, primitive Colaniella, Frondina and Ichthyofrondina dominate among the Nodosariata; Glomomidiellopsis, Pseudovermiporella and Multidiscus among the Miliolata; and Charliella among the globivalvulinids. Late Changhsingian. This substage is dominated by the FAD, and the short duration (because they disappear at the PTB), of Paraglobivalvulinoides, Urushtenella, advanced Colaniella, Paradagmarita monodi and Louisettita (Fig. 9, with the first and last occurrences in Hazro, SW Turkey). Palaeogeography of the Middle – Late Permian smaller foraminifers Many palaeobiogeographical reconstructions have been proposed for the Guadalupian and Wuchiapingian, especially based on the distributions of brachiopods (see Angiolini et al. 2013 and references therein), fusulinids (Kobayashi 1997a, b, 1999; Kobayashi & Ishii 2003; Ueno 2003; Colpaert et al. 2015) and smaller foraminifers (e.g. Hemigordiopsis, Shanita, Abahedella, Paradagmarita and Colaniella: see Şengör et al. 1988; Nestell & Pronina 1997; Altıner et al. 2000; Gaillot & Vachard 2007; Huang et al. 2007; Vachard 2014). The two borders of the Palaeotethys and Neotethys are difficult to discriminate palaeobiogeographically at this time because both are located in the subtropical realm, and because both Tethyan oceanic branches Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 236 D. VACHARD lead to a cul-de-sac and close westwards (Ghazzay et al. 2015). Consequently, all faunal and floral mixtures are possible. Palaeobiogeographical data on Panthalassa are very rare, except in Japan; however, this country itself is composed of numerous terranes that complicate the interpretations and reconstructions. In contrast, Cimmerian assemblages are generally characterized more accurately (Gaetani et al. 2009; Angiolini et al. 2015). Since the remarkable pioneering work of Şengör et al. (1988), Shanita has always been confirmed as an important palaeobiogeographical marker (Sheng & He 1983; Nestell & Pronina 1997; Jin & Yang 2004; Gaillot & Vachard 2007; Huang et al. 2007; Ebrahim Nejad et al. 2015). It is sometimes assigned to Cimmerian terranes (Nestell & Pronina 1997; Ueno 2003), but we propose herein that Shanita is somewhat characteristic of these periGondwanan terranes, initially peri-Gondwanan, which migrated to the Cimmeria and Sibumasu microcontinents after the opening of the Neotethys. In addition, Vachard (2014) indicated how Colaniella, by its evolution and migration, in turn confirms this palaeogeography with: (1) the Salt Range (Pakistan) as the radiation centre; (2) a first migration along the peri-Gondwanan border with primitive species of Colaniella; and (3) a second migration to the peri-Laurentian northern boundary of the Palaeotethys, and the shelves bordering the eastern Tethys and western Panthalassa. During the late Changhsingian, owing to the rapid opening of the Neotethys between the peri-Gondwanan margin and the Cimmerian microcontinent, and the connection with the peri-Gondwanan areas and, consequently, with the Salt Range in Pakistan, the centre of speciation during the Wuchiapingian was interrupted (Fig. 12). Currently, work is being carried out with a team of colleagues (Vachard, Rettori, Altıner and Gennari) on the palaeobiogeography of Altineria. At first, the palaeogeography of all the outcrops where Altineria has been recorded can appear complicated, as a result of the current data in the literature. The source of Altineria is the Taurus Mountains in southern Turkey; Altineria is also known, under the name Angelina, in Armenia (Pronina & Gubenko 1990; Pronina 1996) and Tunisia (Ghazzay et al. 2015; Ghazzay-Souli et al. Fig. 12. Palaeobiogeographical distribution of the genus Colaniella during the Late Permian. The numbers indicated on the map, from 1 to 26, correspond to the localities containing species of Colaniella: (1) Sicily; (2) Tunisia; (3) Albania; (4) Montenegro; (5) Greece; (6) Crimea; (7) NW Turkey; (8) NW Caucasus; (9) central Turkey; (10) Cyprus; (11) Taurus (southern Turkey); (12) Saudi Arabia; (13) Zagros (southern Iran) and Oman; (14) Alborz and central Iran; (15) Central Mountains of Afghanistan; (16) Salt Range; (17) Kashmir; (18) Lamayuru (Ladakh, Himalaya); (19) western Tibet; (20) southern Tibet; (21) South China; (22) Thailand; (23) Malaysia; (24) Indochina (Vietnam); (25) Primorye; (26) Japan terranes (according to Vachard 2014; text-fig. 2). In the late Changhsingian, owing to the rapid opening of the Neotethys between the peri-Gondwanan margin (13 & 16) and the Cimmerian subcontinent (14–20), the connection with the peri-Gondwanan areas and, consequently, with the Salt Range in Pakistan (16), the centre of speciation during the Wuchiapingian, was interrupted. Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 PERMIAN SMALLER FORAMINIFERS 2015). Our team has also found Altineria in limestones of regions that were alternatively assigned to peri-Gondwanan or Cimmerian areas, as for the NW part of Chios Island (Greece), NW Iran and the Abadeh area (central Iran). Since, at all of these localities, Altineria is confined to shallow carbonate platforms in the vicinity of bioconstructions with microbialites, Tubiphytes, inozoan calcisponges and richthofeniid brachiopods, it seems difficult that under these conditions Altineria may have migrated through the Neotethys up to the Cimmerian Block. Therefore, we suggest that Taurus, Armenia, NW Iran (see Ebrahim Nejad et al. 2015) and Chios (and the related Karaburun area), as well as Tunisia and the Abadeh area, could have been located on the same extended carbonate platform. Owing to the demonstrated location of Taurus and Tunisia on the peri-Gondwanan border, this carbonate platform therefore corresponds to the whole of the peri-Gondwanan margin (i.e. southern) of the Tethyan Ocean (eventually, in the yet to open Neotethys). This peri-Gondwanan margin supposedly also exists in Afghanistan (but more probably in the Kabul Block than in the Central Mountains, often abusively called the Helmend Block: see the discussion in Colpaert et al. 2015), in Karakoram, SE Pamirs, Tibet, Baoshan Block, western Myanmar and western Thailand. The presence of Altineria in these regions is possible, although not demonstrated. However, the presence of Altineria in Hubei (China) would be puzzling because this region is generally emplaced far from the Tethyan borders on the traditional palaeomaps. However, it this is easier to explain on the palaeomap of Gaillot & Vachard (2007), where the South China and Iran terranes are located closely together. During the entire Permian, the southern part of the North American Craton (i.e. the modern states of Sonora, New Mexico and Texas) remained relatively isolated from the Laurentian and Gondwanan continents. The endemism of the smaller foraminifers was important during the late Cisuralian (¼Wolfcampian) (Lucas et al. 2015; Vachard et al. 2015), where several Tethyan groups are apparently lacking and probably endemic taxa exist among the globivalvulinid and miliolate foraminifers and the gymnocodiacean algae (Vachard et al. 2015). In contrast, several studies (Vachard et al. 1992; Nestell & Nestell 2006; Nestell et al. 2006) have demonstrated that the endemism of the smaller foraminifers is lesser than for the large fusulinids, during the Middle Permian and, especially, the Capitanian. However, if some genera were found (e.g. Partisania, Abadehella and Charliella), other ones remain unknown (e.g. giant paraglobivalvulinins, baisalinids, hemigordiopsids and langellins). Finally, no Late Permian foraminifers are known in the Americas. 237 Conclusions † Four classes of smaller foraminifers were present during the Permian: Fusulinata, Miliolata, Nodosariata and Textulariata † Biostratigraphically, the most interesting groups are the Lasiodiscoidea, Bradyinoidea, Globivalvulinoidea among the Fusulinata; the Cornuspirida among the Miliolata; and the entire class of the Nodosariata. The class Textulariata is too poorly known during the Permian to permit the characterization of biostratigraphical markers; nevertheless, the appearance of the Verneuilida order is probably very important. † The Lasiodiscoidea are poorly differentiated during the Cisuralian. Pseudovidalina gives rise to two evolute forms: Xingshandiscus during the Kungurian; and Altineria at the Capitanian– Wuchiapingian boundary interval. Lasiodiscus and Lasiotrochus display several species during the Late Guadalupian –Lopingian. † The Bradyinoidea are separated in two lineages during the Permian: one of Bradyina and one of Postendothyra. † The phylogeny of the globivalvulinids, from the Early Pennsylvanian to the late Cisuralian, was only that of the lineages of Globivalvulina. During the Guadalupian –Lopingian, they diversify with three new subfamilies: Paraglobivalvulininae, Dagmaritinae and Paradagmaritinae, the evolutionary trends of which are relatively well known in Turkey, Iran and peri-Gondwanan areas. † The Miliolata remain poorly known, especially in the Cisuralian. It is difficult to understand if the genera recently described in the USA, Praeneodiscus and Olgaorlovella, have also a biostratigraphic importance in the Tethys. The evolution of tubiphytids and ellesmerellids is yet to be discussed, because these taxa are generally assigned to cyanobacteria and/or algae. The Middle-Late Permian is well characterized by giant genera: Neodiscus, Neohemigordius, Baisalina, Glomomidiellopsis, Hemigordiopsis and Shanita. † The Permian families and genera of the Nodosariata, as for those of the Mesozoic and Cenozoic, are still poorly understood. The Cisuralian genera most interesting with respect to biostratigraphy are probably Tezaquina, Nodosinelloides, Geinitzina and Polarisella, and Vervilleina, although they present many problems of nomenclature. The Middle Permian is a period of generic diversification of Pachyphloia, Calvezina, Cryptoseptida, Rectoglandulina, true Nodosaria, Langella and Pseudolangella. The Late Permian is marked by evolved colaniellids and frondinids, whereas at the base of the Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 238 D. VACHARD Triassic several very primitive genera, which appeared in the Cisuralian, Tezaquina, Nodosinelloides and Polarisella, reappear. † Palaeogeographically, the most interesting markers have already been characterized: Shanita and Colaniella, but Altineria should be restudied (work in progress). All of these genera indicate the opening of the Neotethys and its consequence for foraminiferal evolution. † Shanita is somewhat characteristic of these peri-Gondwanan terranes, which migrated to the Cimmeria and Sibumasu microcontinents after the opening of the Neotethys. Our hypothesis is that Altineria, more discrete and minute than Shanita, confirms such a palaeogeographical and geodynamic evolution. Colaniella has the Salt Range (Pakistan) as its radiation centre; a first migration along the peri-Gondwanan border with the primitive species of Colaniella; and a second migration to the peri-Laurentian northern boundary of the Palaeotethys and the shelves bordering the eastern Tethys and western Panthalassa. † The southern part of the North American Craton (i.e. the modern states of Sonora, New Mexico and Texas) remained relatively isolated from the Laurentian and Gondwanan continents. The endemism of the smaller foraminifers, important during the late Cisuralian (¼Wolfcampian), decreased during the late Guadalupian (Capitanian). I wish to thank Galina and Merlynd Nestell and Demir Altıner, who improved the manuscript with their constructive criticisms and English corrections. I appreciated the friendly assistance of Elisabeth Locatelli, Sébastien Clausen (Villeneuve d’Ascq, France) and Karl Krainer (Innsbruck, Austria). Thanks to Drs Lucas and Shen for the invitation to write a paper and to Jessica Pollitt at GSL for the editing. References Adl, S., Simpson, A. et al. 2012. The revised classification of eukaryotes. Journal of Eukaryotic Microbiology, 59, 429– 493. Alipour, Z., Hosseini-Nezhad, S.M., Vachard, D. & Rashidi, K. 2013. Biostratigraphy and description of smaller foraminifers of Dorud Group (uppermost Gzhelian-lower Sakmarian) in Tang-e Shamshirbor section Eastern Alborz, Iran. Geological Journal, 48, 385– 402. Altıner, D. 1981. Recherches stratigraphiques et micropaléontologiques dans le Taurus Oriental au NW de Pinarbasi (Turquie). PhD thesis, University of Geneva. Altıner, D. 1984. Upper Permian foraminiferal biostratigraphy in some localities of the Taurus Belt. In: Tekeli, O. & Göncüoğlu, M.C. (eds) Geology of the Taurus Belt: Proceedings of the International Symposium 26– 29 September 1983, Ankara, Turkey. MTA, Ankara, Turkey, 255–268. Altıner, D. 1988. Pseudovidalinidae n. fam. and Angelina n. gen. from the Upper Permian of south and southeast Turkey. Revue de Paléobiologie, Special Volume, 2, 25–36. Altıner, D. 1999. Sengoerina argandi, n. gen., n. sp., and its position in the evolution of late Permian biseriamminid foraminifers. Micropaleontology, 45, 215–220. Altıner, D. & Brönnimann, P. 1980. Louisettita elegantissima, n. gen. n. sp., un nouveau foraminifère du Permien supérieur du Taurus oriental (Turquie). Notes du Laboratoire de Paléontologie de l’Université de Genève, 6, 39–43. Altıner, D. & Özkan-Altıner, S. 1998. Baudiella stampflii n. gen., n. sp., and its position in the evolution of late Permian ozawainellid. Revue de Paléobiologie, 17, 163– 175. Altıner, D. & Özkan-Altıner, S. 2001. Charliella rossae n. gen., n. sp., from the Tethyan realm: remarks on the evolution of late Permian biseriamminids. Journal of Foraminiferal Research, 31, 309–314. Altıner, D. & Özkan-Altıner, S. 2010. Danielita gailloti n. gen., n. sp., within the evolutionary framework of Middle– Late Permian dagmaritins. Turkish Journal of Earth Sciences, 19, 497–512, https://doi.org/10. 3906/yer-0905-4 Altıner, D. & Savini, R. 1997. New species of Syzrania from the Amazonas and Solimöes basins (North Brazil): remarks on the generic and suprageneric position of syzraniid foraminifers. Revue de Paléobiologie, 16, 7 –20. Altıner, D. & Zaninetti, L. 1977. Kamurana broennimanni, n. gen., n. sp., un nouveau foraminifère porcelané perforé du Permien supérieur du Taurus oriental, Turquie. Notes du Laboratoire de Paléontologie de l’Université de Genève, 1, 1 –6. Altıner, D., Özkan-Altıner, S. & Koçyigit, A. 2000. Late Permian foraminiferal biofacies belts in Turkey: palaeogeographic and tectonic implications. In: Bozkurt, E., Winchester, J.A. & Piper, J.D.A. (eds) Tectonics and Magmatism in Turkey and the Surrounding Area. Geological Society London, Special Publications, 173, 83– 96, https://doi.org/10.1144/GSL.SP. 2000.173.01.04 Angiolini, L. & Rettori, R. 1995. Chitralina undulata gen. n. sp. n. (Foraminiferida) from the Late Permian of Karakorum (Pakistan). Rivista Italiana di Paleontologia e Stratigrafia, 100, 477 –491. Angiolini, L., Crasquin-Soleau, S., Platel, J.P., Roger, J., Vachard, D., Vaslet, D. & Al-Husseini, M. 2004. Saiwan, Gharif and Khuff formations, Haushi-Huqf Uplift, Oman. In: Al-Husseini, M. (ed.) Carboniferous, Permian and Early Triassic Arabian Stratigraphy. GeoArabia, Special Publications, 3, 149–183. Angiolini, L., Crippa, G., Muttoni, G. & Pignatti, J. 2013. Guadalupian (Middle Permian) paleobiogeography of the Neotethys Ocean. Gondwana Research, 24, 173–184. Angiolini, L., Zanchi, A. et al. 2015. From rift to drift in South Pamir (Tajikistan): Permian evolution of a Cimmerian terrane. Journal of Asian Earth Sciences, 102, 146 –169. Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 PERMIAN SMALLER FORAMINIFERS Angiolini, L., Campagna, M. et al. 2016. Brachiopods from the Cisuralian–Guadalupian of Darvaz, Tajikistan and implications for Permian stratigraphic correlations. Palaeoworld, 25, 539– 568, https://doi.org/10. 1016/j.palwor.2016.05.006. Argyriadis, I., de Graciansky, P.C. & Lys, M. 1976. Datation de niveaux rouges dans le Permien marin péri-égéen. Bulletin de la Société géologique de France, Serie 7, 18, 513–519. Arnaud-Vanneau, A. 1980. Micropaléontologie, paléoécologie et sédimentologie d’une plate-forme carbonatée de la marge passive de la Téthys: l’Urgonien du Vercors septentrional et de la Chartreuse (Alpes occidentales). Géologie Alpine, Mémoire, 11, 269–874. Aw, P.C., Ishii, K.-I. & Okimura, Y. 1977. On Palaeofusulina and Colaniella fauna from the upper Permian of Kelantan, Malaysia. Transactions of the Proceedings of the Palaeontological Society of Japan, 104, 407–417. Baghbani, D. 1993. The Permian sequence in the Abadeh region, Central Iran. In: Nairn, A.E.M. & Koroteev, A.V. (eds) Contributions to Eurasian Geology. Occasional Publication Earth Sciences Research Institute,University of South Carolina, New Series, 9A– B, (part II), 7 –22. Baryshnikov, V.V., Zolotova, V.P. & Kosheva, V.F. 1982. New Species of Artinskian Foraminifers of the Pre-Urals. Akademiya Nauk SSSR, Uralskii Nauchnyi Tsentr, Institut Geologii i Geokhimii, preprint [in Russian]. Baud, A., Jenny, C., Papanikolaou, D., Sideris, C. & Stampfli, G. 1991. New observations on Permian stratigraphy in Greece and geodynamic interpretation. Bulletin of the Geological Society of Greece, 25, 187–206. Bensaid, M., Termier, H., Termier, G. & Vachard, D. 1979. Le Carbonifère (Viséen supérieur-Bachkirien) entre Bou Chber et Ich ou Mellal (Maroc Central). Annales de la Société géologique du Nord, 98, 189–204. Berczi-Makk, A. 1992. Midian (Upper Permian) Foraminifera from the large Mihalovits quarry at Nagyvisnyó (North-Hungary). Acta Geologica Hungarica, 35, 27– 38. Berczi-Makk, A. 1996. Foraminifera of the Triassic formations of Alsó Hill (northern Hungary), part 3: Foraminifer assemblage of the basinal facies. Acta Geologica Hungarica, 39, 413– 459. Blazejowski, B. 2009. Foraminifers from the Treskelodden Formation (Carboniferous– Permian) of south Spitsbergen. Polish Polar Research, 30, 193 –230. Bogush, O.I. 1985. Early Carboniferous foraminifers and stratigraphy from the western Siberian Plain. Akademiya Nauk SSSR, Sibirskoye Otdelenie, Trudy Instituta Geologii i Geofiziki, 619, 49– 68 [in Russian]. Boomgaart, L. 1949. Smaller Foraminifera from Bodjonegoro (Java). Smit & Dontje Publishers, Sappemeer, The Netherlands. Bowser, S.S., Habura, A. & Pawlowski, J. 2006. Molecular evolution of Foraminifera. In: Katz, L. & Bhattacharya, D. (eds) Genomics and Evolution of Microbial Eukaryotes. Oxford University Press, Oxford, 78–93. 239 Bozorgnia, F. 1973. Paleozoic Foraminiferal Biostratigraphy of Central and East Alborz Mountains, Iran. National Iranian Oil Company, Geological Laboratories Publications, 4. Brady, H.B. 1876. A monograph of Carboniferous and Permian foraminifera (the genus Fusulina excepted). Palaeontographical Society of London, 30, 1– 166. Brady, H.B. 1878. On the Reticularian and Radiolarian Rhizopoda (Foraminifera and Polycystina) on the North Polar Expedition of 1875–76. Annals and Magazine of Natural History, Series 5, 1, 425– 440. Brady, H.B. 1884. Report of the foraminifera dredged by H.M.S. Challenger during the years 1873– 76. Reports on the Scientific Results of the Exploratory Voyage of the H.M.S. Challenger, Zoology, 9, 1–814. Brenckle, P.L. & Wahlman, G.P. 1996. Raphconilia, a new name for the Upper Paleozoic pseudovidalinid foraminifer Conilia Brenckle and Wahlman, 1994. Journal of Foraminiferal Research, 26, 184. Brönnimann, P., Whittaker, J.E. & Zaninetti, L. 1978. Shanita a new pillared miliolacean foraminifer from the Late Permian of Burma and Thailand. Rivista Italiana di Paleontologia, 84, 63–92. Brunner, P. 1979. Microfacies y microfosiles permotriásicos en el area El Antimonio, Sonora, Mexico. Revista del Instituto Mexicano del Petróleo, México, 11, 6 –41. Bykova, E.V. & Polenova, E.N. 1955. Devonian Foraminifers, Radiolarians and Ostracods of the Volga– Urals Region. Trudy VNIGRI, 87 [in Russian; French translation, C.E.D.P. No. 1603. BRGM, Paris]. Calkins, G.N. 1926. The Biology of the Protozoa. Lea & Febiger, Philadelphia, PA. Canuti, P., Marcucci, M. & Pirini Radrizzani, C. 1970. Microfacies e microfaune nelle formazioni paleozoiche dell’anticlinale di Hazro (Anatolia sudorientale, Turchia). Bolletino Societá Geologica Italiana, 89, 21– 40. Caridroit, M., Fontaine, H., Jongkanjanasoontorn, Y., Suteethorn, V. & Vachard, D. 1990. First results of a paleontological study of Northwest Thailand. In: Fontaine, H. (ed.) Ten years of CCOP Research on the Pre-Tertiary of East Asia. CCOP Technical Publications, 20, 337– 350. Çatal, E. & Dager, Z. 1974. Toros bölgesi permiyenine ait yeni bir Colaniella türünün tanımı (Description of a new Colaniella species from the Permian of Taurus region). Türkije Jeoloji Kurumu Bülteni (Bulletin of the Geological Society of Turkey), 17, 187– 191. Cavalier-Smith, T. 2002. The phagotrophic origin of eukaryotes and phylogenetic classification of Protozoa. International Journal of Systematic and Evolutionary Microbiology, 52, 297–354. Cavalier-Smith, T. 2003. Protists phylogeny and the high-level classification of Protozoa. European Journal of Protistology, 39, 338–348. Cherdyntsev, W. 1914. On a foraminiferal Permian fauna from western European Russia. Trudy Obshchestva Estestvoispytateley pri Imperatorskomy Kazanskomy Universitety, 46, 3 –88 [in Russian]. Chuvashov, B.I., Shuysky, V.P. & Ivanova, R.M. 1993. Stratigraphical and facies complexes of the Paleozoic calcareous algae of the Urals. In: Barattolo, F., De Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 240 D. VACHARD Castro, P. & Parente, M. (eds) Studies on Fossil Benthic Algae. Bolletino della Societá Paleontologica Italiana, Modena, Special Volume, 1, 93– 119. Ciarapica, G., Cirilli, S., Martini, R. & Zaninetti, L. 1986. Une microfaune à petits foraminifères d’âge permien remaniée dans le Trias moyen de l’Apennin méridional (Formation du Monte Facito, Lucanie occidentale); description de Crescentia vertebralis n. gen. n. sp. Revue de Paléobiologie, 5, 207 –215. Ciarapica, G., Cirilli, S., Passeri, L., Trincati, E. & Zaninetti, L. 1987. ‘Anidriti di Búrano’ et formation du Monte Cetona (nouvelle formation), biostratigraphie des deux séries-types du Trias supérieur dans l’Apennin septentrional. Revue de Paléobiologie, 6, 341– 409. Colpaert, C., Monnet, C. & Vachard, D. 2015. Eopolydiexodina (Middle Permian giant fusulinids of Afghanistan): biometry, morphometry, palaeobiogeography and end-Guadalupian events. Journal of Asian Earth Sciences, 102, 127 –145, https://doi. org/10.1016/j.jseaes.2014.10.028 Conil, R. & Pirlet, H. 1970. Le calcaire carbonifère du synclinorium de Dinant et le sommet du Famennien. Congrès et Colloques de l’Université de Liège, 55, 47–63. Conil, R., Groessens, E. & Lys, M. 1977. Etude micropaléontologique de la tranchée d’Yves-Gomezée (Tn 3c-V 1-V 2, Belgique). Bulletin de la Société belge de Géologie, 82, 201– 239. Conil, R., Longerstaey, P.J. & Ramsbottom, W.H.C. 1980. Matériaux pour l’étude micropaléontologique du Dinantien de Grande-Bretagne. Mémoires de l’Institut Géologique de l’Université de Louvain, 30, 1 –115. Cózar, P. & Somerville, I.D. 2012. Mississippian Biseriamminaceae and their evolutionary development. Journal of Foraminiferal Research, 42, 216– 233. Cózar, P., Medina-Varea, P., Somerville, I.D., Vachard, D., Rodrı́guez, S. & Said, I. 2014. Foraminifers and conodonts from the late Viséan to early Bashkirian succession in the Saharan Tindouf Basin (southern Morocco): biostratigraphic refinements and implications for correlations in the western Palaeotethys. Geological Journal, 49, 271– 302. Cózar, P., Sanz-López, J. & Blanco-Ferrera, S. 2015. Lasiodiscid foraminifers during the late Viséan to Serpukhovian in Vegas de Sotres section (Cantabrian Mountains, NW Spain). Geobios, 48, 213 –238. Crescenti, U. 1969. Biostratigrafia delle facies mesozoiche dell’Appennino centrale. Geologica Romana, 8, 15– 40. Crespin, I. 1958. Permian Foraminifera of Australia. Commonwealth of Australia, Department of National Development, Bureau of Mineral Resources, Geology and Geophysics, 48. Crespin, I. & Belford, D.J. 1957. New genera and species of foraminifera from the Lower Permian of Western Australia. Contributions of the Cushman Foundation for Foraminiferal Research, 8, 73– 76. Cummings, R.H. 1955. Nodosinella Brady, 1876 and associated upper Paleozoic genera. Micropaleontology, 1, 221– 238. Cummings, R.H. 1956. Revision of the upper Palaeozoic textulariid foraminifera. Micropaleontology, 2, 201– 242. Cushman, J.A. 1928. Additional genera of the Foraminifera. Contributions from the Cushman Laboratory for Foraminiferal Research, 4, 1 –8. Cushman, J.A. 1929. A Late Tertiary fauna of Venezuela and other related regions. Contributions from the Cushman Laboratory for Foraminiferal Research, 5, 77–101. Cushman, J.A. & Ozawa, Y. 1930. A monograph of the foraminiferal family Polymorphinidae, Reecnt and fossil. Proceedings of the United States National Museum, 77, 1 –195. Cushman, J.A. & Waters, J.A. 1928a. The development of Climacammina and its allies in the Pennsylvanian of Texas. Journal of Paleontology, 2, 119– 130. Cushman, J.A. & Waters, J.A. 1928b. Some Foraminifera from the Pennsylvanian and Permian of Texas. Contributions from the Cushman Laboratory for Foraminiferal Research, 4, 31–55. Cushman, J.A. & Waters, J.A. 1928c. Upper Paleozoic foraminifera from Sutton County,Texas. Journal of Paleontology, 2, 358–371. Cushman, J.A. & Waters, J.A. 1930. Foraminifera of the Cisco Group. Bulletin University of Texas, 3019, 22–81. Dain, L.G. & Grozdilova, L.P. 1953. Fossil foraminifers of the USSR: tournayellids and archaediscids. Trudy VNIGRI, 74, 7 –63 [in Russian; French translation, M. Jayet, BRGM, 1816, 1– 124]. Davydov, V.I. 1988. Archaediscidae in the Upper Carboniferous and Lower Permian. Revue Paléobiologie, Special Volume, 2, 39– 46. Davydov, V.I. & Arefifard, S. 2013. Middle Permian (Guadalupian) fusulinid taxonomy and biostratigraphy of the mid-latitude Dalan Basin, Zagros, Iran and their applications in paleoclimate dynamics and paleogeography. GeoArabia, 18, 17–62. Davydov, V.I., Nilson, I. & Stemmerik, L. 2001. Fusulinid zonation of the Upper Carboniferous Kap Jungersen and Foldedal Formations, southern Amdrup Land, eastern North Greenland. Bulletin of the Geological Society of Denmark, 48, 31– 77. Davydov, V.I., Krainer, K. & Chernykh, V. 2013. Fusulinid biostratigraphy of the Lower Permian Zweikofel Formation (Rattendorf Group; Carnic Alps, Austria) and Lower Permian Tethyan chronostratigraphy. Geological Journal, 48, 57– 100. Dawson, O., Racey, A. & Whittaker, J.E. 1993. The palaeoecological and palaeobiogeographical significance of Shanita (foraminifer) and associated foraminifera/algae from the Permian of peninsular Thailand. In: International Symposium on Biostratigraphy on Mainland Southeast Asia: Facies & Paleontology, 31 January– 5 February 1993, Chiang Mai, Thailand, Volume 2. Department of Geological Sciences, University of Chiang Mai, Chiang Mai, Thailand, 283–298. Deleau, P. & Marie, P. 1961. Les Fusulinidés du Westphalien C du bassin d’Abadla et quelques autres foraminifères du Carbonifère algérien. Publications Service Carte Géologique Algérie, 25, 45–160. Derville, H. 1952. A propos de Calcisphères (rectification). Comptes Rendus Sommaires de la Société Géologique de France, 6, 236–237. Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 PERMIAN SMALLER FORAMINIFERS d’Orbigny, A. 1826. Tableau méthodique de la classe des Céphalopodes. Annales des Sciences Naturelles, 7, 245–314. d’Orbigny, A. 1850. Prodrome de paléontologie stratigraphique universelle des animaux mollusques et rayonnés, Volume 1. Masson Publisher, Paris. Dronov, V.I. 2004. Carboniferous and Permian stratigraphy of the Rushan-Pshart Pamir (southern Rushan– Pshart Zone, southern Pshart Tectonic Block). Doklady Akademii Nauk, 395, 791– 793 [in Russian]. Ebrahim Nejad, E., Vachard, D., Siabeghodsy, A. & Abbasi, S. 2015. Middle– Late Permian (Murgabian– Djulfian) foraminifers of the northern Maku area (western Azerbaijan, Iran). Palaeontologia Electronica, 18.1.19A, 1 –63. Ehrenberg, C.G. 1838. Über dem blossen Auge unsichtbare Kalkthierchen und Kieselthierchen als Hauptbestandtheile der Kreidegebirge. Bericht über die zu Bekanntmachung geeigneten Verhandlungen der königlichen preussischen Akademie der Wissenschaften zu Berlin, 3, 192– 200. Ehrenberg, C.G. 1854. Zur Mikrogeologie. von Leopold Voss, Leipzig. Elliott, G.F. 1958. Fossil microproblematica from the Middle East. Micropaleontology, 4, 419– 428. Etheridge, R., JR. 1873. Notes on certain genera and species mentioned in the foregoing lists. Scotland Geological Survey, Memoir Explanation Sheet, 23, appendix 2, 93– 107. Fahrni, J.F., Pawlowski, J., Richardson, S., Debenay, J.P. & Zaninetti, L. 1997. Actin suggests Miliammina fusca (Brady) is related to porcellaneous rather than to agglutinated foraminifera. Micropaleontology, 43, 211–214. Filimonova, T.V. 2008. Smaller foraminifers from type sections of the Bolorian Stage, the Lower Permian of Darvaz. Stratigraphy and Geological Correlation, 16, 22– 41. Filimonova, T.V. 2010. Smaller foraminifers of the Lower Permian from Western Tethys. Stratigraphy and Geological Correlation, 18, 687– 811. Filimonova, T.V. 2013. Smaller foraminifers from the Permian of Central Iran. Stratigraphy and Geological Correlation, 21, 18–35. Flakowski, J., Bolivar, I., Fahrni, J. & Pawlowski, J. 2005. Actin phylogeny in foraminifera. Journal of Foraminiferal Research, 35, 93–102. Flügel, E. & Reinhardt, J. 1989. Uppermost Permian reefs in Skyros (Greece) and Sichuan (China): implications for the Late Permian extinction event. Palaios, 4, 502–518. Flügel, E., Kochansky-Devidé, V. & Ramovs, A. 1984. A Middle Permian calcisponge/algal/cement reef: Straza near Bled, Slovenia. Facies, 10, 179– 256. Flügel, E., Di Stefano, P. & Senowbari-Daryan, B. 1991. Microfacies and depositional structure of allochthonous carbonate base-of-slope deposits. The Late Permian Pietra di Salomone megablock, Sosio Valley (western Sicily). Facies, 25, 147– 186. Fontaine, H., NGUYEN DUC TIEN & Vachard, D. 1986. Discovery of Permian limestone South of Tara Island in the Calamian Islands, Philippines. United Nations ESCAP, CCOP Technical Bulletin, 18, 161–167. 241 Fontaine, H., Khoo, H.P. & Vachard, D. 1988a. Discovery of Triassic fossils at Bukit Chuping, in Gunung Sinyum area, and at Kota Jin, Peninsular Malaysia. Journal of Southeast Asian Earth Sciences, 2, 145– 162. Fontaine, H., Lys, M. & NGUYEN DUC TIEN 1988b. Some Permian corals from east Peninsular Malaysia, associated microfossils, paleogeographic significance. Journal of Southeast Asian Earth Sciences, 2, 65– 78. Fontaine, H., Suteethorn, V. et al. 1988c. Late Palaeozoic and Mesozoic fossils of West Thailand and their environments. CCOP Technical Bulletin, 20, 1 –107. Fontaine, H., Suteethorn, V. & Vachard, D. 1993. Carboniferous and Permian limestones in Sop Pong area: unexpected lithology and fossils. International Symposium on Biostratigraphy of Mainland Southeast Asia: Facies and Paleontology, 2, 319 –336. Fontaine, H., Bin Amnan, I., Khoo, H.P., Tien, N.D. & Vachard, D. 1994. The Neoschwagerina and Yabeina–Lepidolina Zones in Peninsular Malaysia, and Dzhulfian and Dorashamian in Peninsular Malaysia, the Transition to the Triassic. Geological Survey of Malaysia, Geological Papers, 4. Fontaine, H., Tansuwan, V. & Vachard, D. 1997. Age of limestones associated with gypsum deposits in Northeast and Central Thailand. CCOP Newsletter, 21–22, 6– 7. Forke, H.C., Kahler, F. & Krainer, K. 1998. Sedimentology, microfacies and stratigraphic distribution of foraminifers of the Lower ‘Pseudoschwagerina’ Limestone (Rattendorf Group, Late Carboniferous), Carnic Alps (Austria/Italy). Senckenbergiana lethaea, 78, 1– 39. Foster, C.B., Palmieri, V. & Fleming, P.J.G. 1985. Plant microfossils, Foraminiferida, and Ostracoda, from the Fossil Cliff Formation (Early Permian, Sakmarian), Perth Basin, Western Australia. Special Publications South Australia Department Mines and Energy, 5, 61–105. Fursenko, A.V. 1958. Main stages of development of foraminiferal fauna in the geological past. Akademiya Nauk Beloruskoi SSR, Trudy Instituta Geologicheskikh Nauk, 1, 10–29 [in Russian]. Gaetani, M., Angiolini, L. et al. 2009. Pennsylvanian– Early Triassic stratigraphy in the Alborz Mountains (Iran). In: Brunet, M.-F., Wilmsen, M. & Granath, J.W. (eds) South Caspian to Central Iran Basins. Geological Society of London, Special Publications, 312, 79–128, https://doi.org/10.1144/SP312.5 Gaillot, J. 2006. The Khuff Formation (Late Permian, Middle East): high-resolution biostratigraphy by means of foraminifers and algae, sequence stratigraphy, depositional environments, and new approach of the Permian/Triassic crisis. PhD thesis, University of Lille. Gaillot, J. & Vachard, D. 2007. The Khuff Formation (Middle East) and time-equivalents in Turkey and South China: biostratigraphy from Capitanian to Changhsingian times (Permian), new foraminiferal taxa and palaeogeographical implications. Coloquios de Paleontologı́a, 57, 37–223. Gaillot, J., Vachard, D., Galfetti, T. & Martini, R. 2009. New latest Permian foraminifers from Laren Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 242 D. VACHARD (Guangxi Province, South China). Palaeobiogeographic implications. Geobios, 42, 141–168. Galloway, J.J. 1933. A Manual of Foraminifera. James Furman Kemp Memorial Series, 1. Principia Press, Bloomington, IN. Galloway, J.J. & Harlton, B.H. 1928. Some Pennsylvanian Foraminifera of Oklahoma with special reference to the genus Orobias. Journal of Paleontology, 2, 338– 357. Galloway, J.J. & Ryniker, C. 1930. Foraminifera from the Atoka Formation of Oklahoma. Oklahoma Geological Survey Circular, 21, 1– 36. Gargouri, S. & Vachard, D. 1988. Sur Hemigordiopsis et d’autres Foraminifères porcelanés du Murghabien du Tebaga (Permien supérieur de Tunisie). Revue de Paléobiologie, Special Volume, 1, 57–68. Gerke, A.A. 1952. Permian microfauna of the Nordvik region and stratigraphic significance. Trudy NauchnoIssledovatel’skii Institut Geologii Arktiki (NIIGA), 28, 1– 209 [in Russian]. Gerke, A.A. 1959. On new Permian nodosarioid foraminifers and precisions on the diagnosis of Nodosaria. Sbornik Statey po Paleontologii i Biostratigrafii, Nauchno-issledovatelskiy Institut Geologii Artiki (NIIGA), 17, 41–59 [in Russian]. Gerke, A.A. 1961. Permian and Lower Mesozoic rectoglandulines from northcentral Siberia. Sbornik statey po Paleontologii i Biostratigrafii, 23, 5– 34 [in Russian]. Ghazzay, W., Vachard, D. & Razgallah, S. 2015. Revised fusulinid biostratigraphy of the Middle–Late Permian of Jebel Tebaga (Tunisia). Revue de Micropaléontologie, 58, 57–83. Ghazzay-Souli, W., Vachard, D. & Razgallah, S. 2015. Carboniferous and Permian biostratigraphy by foraminifers and calcareous algae of Bir Mastoura (BMT-1) and related boreholes of southern Tunisia. Revue de Micropaléontologie, 58, 239– 265. Glaessner, M.F. 1945. Principles of Micropaleontology. Melbourne University Press, Melbourne. Golubic, S., Radoicic, R. & Seon-Joo, L. 2006. Decastronema kotori gen. nov., comb nov.; a mat-forming cyanobacterium on Cretaceous carbonate platforms and its modern counterparts. Carnets de Géologie/ Notebooks on Geology, Article 2006/02. Grant, R.E., Nestell, M.K., Baud, A. & Jenny, C. 1991. Permian stratigraphy of Hydra Island, Greece. Palaios, 6, 479–497. Griffith, J.W. & Henfrey, A. 1875. The Micrographic Dictionary, Volume 1. 3rd edn. Van Voorst, London. Groves, J.R. 1992. Stratigraphic distribution of nonfusulinacean foraminifers in the Marble Falls Limestone (Lower– Middle Pennsylvanian), western Llano region, central Texas. In: Sutherland, P.K. & Manger, W.L. (eds) Recent Advances in MiddleCarboniferous Biostratigraphy. A Symposium. Okhahoma Geological Survey, Circular, 94, 145– 161. Groves, J.R. 1997. Repetitive patterns of evolution in late Paleozoic foraminifers. In: Ross, C.A., Ross, J.R.P. & Brenckle, P.L. (eds) Late Paleozoic Foraminifera, Their Biostratigraphy, Evolution, and Paleoecology, and the Mid-Carboniferous Boundary. Cushman Foundation for Foraminiferal Research, Special Publications, 36, 51– 54. Groves, J.R. 2000. Suborder Lagenina and other smaller foraminifers from uppermost Pennsylvanian-lower Permian rocks of Kansas and Oklahoma. Micropaleontology, 46, 285–326. Groves, J.R. 2002. Evolutionary origin of the genus Geinitzina (Foraminiferida) and its significance for international correlation near the Carboniferous– Permian boundary. In: Hills, L.V., Henderson, C.M. & Bamber, E.W. (eds) Carboniferous and Permian of the World. Canadian Society of Petroleum Geologists, Memoirs, 19, 437– 447. Groves, J.R. & Altıner, D. 2005. Survival and recovery of calcareous foraminifera pursuant to the end-Permian mass extinction. Comptes Rendus Paleovol, 4, 487–500. Groves, J.R. & Boardman, H. 1999. Calcareous smaller foraminifers from the lower Permian Council Grove Group near Hooser, Kansas. Journal of Foraminiferal Research, 29, 243– 262. Groves, J.R. & Wahlman, G.P. 1997. Biostratigraphy and evolution of Late Carboniferous and Early Permian smaller foraminifers from the Barents Sea (offshore Arctic Norway). Journal of Paleontology, 71, 758– 779. Groves, J.R., Altıner, D. & Rettori, R. 2003. Origin and early evolutionary radiation of the order Lagenida (Foraminifera). Journal of Paleontology, 77, 831– 843. Groves, J.R., Rettori, R. & Altıner, D. 2004. Wall structures in selected Paleozoic Lagenide Foraminifera. Journal of Paleontology, 78, 245–256. Groves, J.R., Altıner, D. & Rettori, R. 2005. Extinction, Survival, and Recovery of Lagenide Foraminifers in the Permian– Triassic Boundary Interval, Central Taurides, Turkey. Paleontological Society Memoir, 62. Groves, J.R., Rettori, R., Payne, J.L., Boyce, M.D. & Altıner, D. 2007. End-Permian mass extinction lagenide foraminifers in the Southern Alps (northern Italy). Journal of Paleontology, 81, 415– 434. Gu, S., Feng, Q. & He, W. 2007. The last Permian deepwater fauna: latest Changhsingian small foraminifers from southwestern Guangxi, South China. Micropaleontology, 53, 311–330. Habeeb, K.H. 1979. SEM study of foraminiferal wall structure in the Lower Carboniferous Limestone. In: Abstracts of Papers, Ninth International Congress of Carboniferous Stratigraphy and Geology, 19–26 May 1979. University of Illinois, Urbana-Champaign, IL, 82–83. Hance, L., Hou, H.F. & Vachard, D. 2011. Upper Famennian to Visean Foraminifers and Some Carbonate Microproblematica from South China, Hunan, Guangxi and Guizhou. Geological Science Press, Beijing. Hauser, M., Vachard, D., Martini, R., Matter, A., Peters, T. & Zaninetti, L. 2000. The Permian sequence reconstructed from reworked carbonate clasts in the Batain Plain (Northeastern Oman). Comptes Rendus de l’Académie des Sciences, 320, 273– 279. Haynes, J.R. 1981. Foraminifera. Wiley, New York. Henbest, L.G. 1963. Biology, mineralogy, and diagenesis of some typical late Paleozoic sedimentary foraminifera and algal-foraminiferal colonies. Cushman Foundation for Foraminiferal Research, Special Publications, 6, 1 –44. Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 PERMIAN SMALLER FORAMINIFERS Henderson, C.M., Pinard, S. & Beauchamp, B. 1995. Biostratigraphic and sequence stratigraphic relationships of Upper Carboniferous conodont and foraminifer distribution, Canadian Arctic Archipelago. Bulletin of Canadian Petroleum Geology, 43, 226–246. Henderson, C.M., Davydov, V.I. & Wardlaw, B.R. 2012. The Permian Period. In: Gradstein, F.M., Ogg, J.G., Schmitz, M.D. & Ogg, G.M. (eds) The Geologic Time Scale 2012. Elsevier, Amsterdam, 653– 679. Ho, J. 1959. Triassic Foraminifera from the Chialingchiang Limestone of South Szechuan. Acta Paleontologica Sinica, 7, 387– 418 [in Chinese]. Hohenegger, J. & Piller, W. 1975. Wandstrukturen und Grossgliederung der Foraminiferen. Sitzungsberichte der österreichischen Akademie der Wissenschaften, Mathematisch-naturwissenschaftliche Klasse, Abteilung I, 184, 67–96. Huang, H., Yang, X. & Jin, X. 2007. The Shanita fauna (Permian foraminifera) from Baoshan area, western Yunnan province, China. Frontiers of Biology in China, 2, 114–124 [English translation from Acta Palaeontologica Sinica, 44, 544–555]. Hughes, G.W. 2005. Saudi Arabian Permo-Triassic biostratigraphy, micropaleontology and palaeoenvironment. In: Powell, A.J. & Riding, J.B. (eds) Recent Developments in Applied Biostratigraphy. Micropalaeontological Society, Special Publications, 93, 91– 108. IRANIAN – JAPANESE RESEARCH GROUP 1981. The Permian and the Lower Triassic systems in Abadeh region, Central Iran. Memoirs Faculty Sciences, Kyoto University, Series Geology & Mineralogy, 47, 61–133. Ishii, K.-I., Okimura, Y. & Nakazawa, K. 1975. On the genus Colaniella and its biostratigraphic significance. Journal Geosciences, Osaka City University, 19, 107–129. Isozaki, Y., Kawahata, H. & Minoshima, K. 2007. The Capitanian (Permian) Kamura cooling event: the beginning of the Paleozoic-Mesozoic transition. Palaeoworld, 16, 16–30. Jenny, J. & Jenny-Deshusses, C. 1978. Dorudia dorudensis n. gen., n. sp., et les Tuberitininae du Permien de l’Elbourz oriental en Iran. Note du Laboratoire de Paléontologie de l’Université de Genève, 2, 7–15. Jenny-Deshusses, C. 1983. Le Permien de l’Elbourz Central et Oriental (Iran): stratigraphie et micropaléontologie (foraminifères et algues). PhD thesis, University of Geneva. Jenny-Deshusses, C. 1985. Rectostipulina n. gen. (¼Stipulina Lys, 1978), a Upper Permian, incertae sedis organism of the central-eastern Tethys: morphological description and stratigraphic remarks. Revue de Paléobiologie, 4, 153–158. Jenny-Deshusses, C., Martini, R. & Zaninetti, L. 2000. Discovery of the foraminifer Colaniella Likharev in the Upper Permian of the Sosio valley (Sicily). Comptes Rendus de l’Académie des Sciences de Paris, 330, 799–804. Jin, X. & Yang, X. 2004. Paleogeographic implications of the Shanita-Hemigordius fauna (Permian foraminifer) in the reconstruction of Permian Tethys. Episodes, 27, 273–278. 243 Jin, Y., Wardlaw, B.R., Glenister, B.F. & Kotlyar, G.V. 1997. Permian chronostratigraphic subdivisions. Episodes, 20, 10– 15. Jin, Y.G., Wang, Y., Wang, W., Shang, Q.H., Cao, C.Q. & Erwin, D.H. 2000. Pattern of marine mass extinction near the Permian– Triassic boundary in South China. Science, 289, 432– 436. Jirovec, O. 1953. Protozoologie. Nakladatelstvi Ceskoslovenské Akademie Ved, Prague. Jones, T.R. 1895. A Monograph of the Foraminifera of the Crag, Part 2. Monograph of the Palaeontographical Society, London, 49, (230), 73–210. Karavaeva, N.I. & Nestell, G.P. 2007. Permian foraminifers of the Omolon Massif, northeastern Siberia, Russia. Micropaleontology, 53, 161– 211. Kalia, P., Kumar Pande, P. & Singh, T. 2000. Early Permian foraminifers from eastern Himalaya and their palaeobiogeographic significance. Alcheringa, 24, 207 –227. Kiparisova, L.D., Markovsky, B.P. & Radchenko, G.P. 1956. Materials for palaeontology, new families and genera. Vsesoyuznogo Nauchno-issledovatelskogo Geologicheskii Institut (VSEGEI) novya seriya, Paleontologiya, 12, 1– 354 [in Russian]. Kobayashi, F. 1986. Middle Permian foraminifers of the Gozenyama Formation, southern Kwanto Mountains, Japan. Bulletin of the National Science Museum, Series C (Geology and Paleontology), 12, 131– 163. Kobayashi, F. 1988. Middle Permian foraminifers of the Omi Limestone, Central Japan. Bulletin of the National Science Museum, Series C (Geology and Paleontology), 14, 1– 35. Kobayashi, F. 1996. Middle Triassic (Anisian) foraminifers from the Kaizawa Formation, southern Kanto Mountains, Japan. Transactions and Proceedings of the Paleontological Society of Japan, New Series, 183, 528–539. Kobayashi, F. 1997a. Middle Permian biogeography based on fusulinacean faunas. In: Ross, C.A., Ross, J.R.P. & Brenckle, P.L. (eds) Late Paleozoic Foraminifera: Their Biogeography, Evolution, and Paleoecology; and Mid-Carboniferous Boundary. Cushman Foundation for Foraminiferal Research, Special Publications, 36, 73–76. Kobayashi, F. 1997b. Middle Permian fusulinacean faunas and paleogeography of exotic terranes in the circumPacific. In: Ross, C.A., Ross, J.R.P. & Brenckle, P.L. (eds) Late Paleozoic Foraminifera: Their Biogeography, Evolution, and Paleoecology; and MidCarboniferous Boundary. Cushman Foundation for Foraminiferal Research, Special Publications, 36, 77–80. Kobayashi, F. 1997c. Upper Permian foraminifers from the Iwai–Kanyo area, West Tokyo, Japan. Journal of Foraminiferal Research, 27, 186–195. Kobayashi, F. 1999. Tethyan uppermost Permian (Dzhulfian and Dorashamian) foraminiferal faunas and their paleogeographic and tectonics implications. Palaeogeography, Palaeoclimatology, Palaeoecology, 150, 279– 307. Kobayashi, F. 2001. Faunal analysis of Permian foraminifers of the Kuma Formation in the Kurosegawa Belt of west Kyushu, Southwest Japan. News of Osaka Micropaleontologists, Special Volume, 12, 61–77. Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 244 D. VACHARD Kobayashi, F. 2002. Lithology and foraminiferal fauna of the allochthonous limestone (Changhsingian) in the upper part of the Toyoma Formation in the South Kitakami Belt, northeast Japan. Paleontological Research, 6, 331– 342. Kobayashi, F. 2004. Late Permian foraminifers from the limestone block in the Southern Chichibu Terrane of west Shikoku, SW Japan. Journal of Paleontology, 78, 62– 70. Kobayashi, F. 2006a. Middle Permian foraminifers of Kaize, southern part of the Saku Basin, Nagano prefecture, central Japan. Paleontological Research, 10, 179– 194. Kobayashi, F. 2006b. Latest Permian (Changhsingian) foraminifers in the Mikata area, Hyogo – Late Paleozoic and Early Mesozoic foraminifers of Hyogo, Japan, Part 3. Nature and Human Activities, 10, 15– 24. Kobayashi, F. 2006c. Early Late Permian (Wuchiapingian) foraminifers in the Tatsuno area, Hyogo – Late Paleozoic and Early Mesozoic foraminifers of Hyogo, Japan, Part 4. Nature and Human Activities, 10, 25– 33. Kobayashi, F. 2012a. Permian non-fusuline foraminifers of the Akasaka Limestone (Japan). Revue de Paléobiologie, Genève, 31, 313– 335. Kobayashi, F. 2012b. Middle and Late Permian foraminifers from the Chichibu Belt, Takachiho area, Kyushu, Japan: implications for faunal events. Journal of Paleontology, 86, 669– 687. Kobayashi, F. 2013. Late Permian (Lopingian) foraminifers from the Tsukumi Limestone, southern Chichibu Terrane of eastern Kyushu, Japan. Journal of Foraminiferal Research, 43, 154–169. Kobayashi, F. & Ishii, K. 2003. Permian fusulinaceans of the Surmaq Formation in the Abadeh region, central Iran. Rivista Italiana di Paleontologia e Stratigrafia, 109, 307– 337. Kochansky-Devidé, V. 1973. Ramovsia limes n. g., n. sp. (Problematica), ein Leitfossil der Grenzlandbänke (unteres Perm). Neues Jahrbuch für Geologie, Paläontologie Monathefte, 8, 462 –468. Koehrer, B., Aigner, T., Forke, H. & Pöppelreiter, M. 2012. Middle to Upper Khuff (Sequences KS1 to KS4) outcrop-equivalents in the Oman Mountains: grainstone architecture on a some regional scale. GeoArabia, 17, 59– 104. Kolodka, C., Vennin, E., Vachard, D., Trocme, V. & Goodarzi, M.H. 2012. Timing and progression of the end-Guadalupian crisis in the Fars province (Dalan Formation; Kuh-e Gakhum, Iran) constrained by foraminifers and other carbonate microfossils. Facies, 58, 131–153. Kotlyar, G.V., Zakharov, YU.D. et al. 1984. Late Permian Etaps of the Evolution of the Organic World, Dzhulfian and Dorashamian Stages in the USSR. Akademiya Nauk SSSR, Dalnevostochnyi Nauchnyi Tsentr, Biologo-Pochvennyi Institut, Leningrad [in Russian]. Kotlyar, G.V., Zakharov, YU.D., Kropacheva, G.S., Pronina, G.P., Chediya, I.O. & Burago, V.I. 1989. Late Permian etaps of the evolution of the organic world, Midian Stage in the USSR. Leningrad ‘Nauka’, Leningradskoe Otdelenie, 1, 1– 177 [in Russian]. Kotlyar, G.V., Pronina, G.P., Zakharov, YU.D. & Nestell, M.K. 1999. Changhsingian of the northwestern Caucasus, southern Primorye and southeastern Pamirs. Permophiles, 35, 18–22. Köylüoglu, M. & Altıner, D. 1989. Micropaléontologie (foraminifères) et biostratigraphie du Permien supérieur de la région d’Hakkari (SE Turquie). Revue de Paléobiologie, 8, 467– 503. Krainer, K. & Vachard, D. 2011. The Lower Triassic Werfen Formation of the Karawanken Mountains (southern Austria) and its disaster survivor microfossils, with emphasis on Postcladella n. gen. (Foraminifera, Miliolata, Cornuspirida). Revue de Micropaléontologie, 54, 59–85. Krainer, K., Lucas, S.G., Vachard, D. & Barrick, J.E. 2015. The Pennsylvanian–Permian section at Robledo Mountain, Doña Ana County, New Mexico, USA. In: Lucas, S.G. & DiMichele, W.A. (eds) Carboniferous–Permian Transition in the Robledo Mountains, Southern New Mexico. New Mexico Museum of Natural History and Science Bulletin, 65, 9 –41. Kulagina, E.I., Rumyantseva, Z.C., Pazukhin, V.N. & Kochetova, N.N. 1992. Granitsa nizhnego-srednego Karbona na Yuzhnom Urale i Srednem Tyan-Shane. Rossiiskaya Akademiya Nauk, Uralskoe Otdelenie, Bashkirskii Nauchnyi Tsentr, Institut Geologii, Moskva, Nauka, Moscow [in Russian]. Lamarck, J.B. DE. 1812. Extrait du cours de zoologie du Muséum d’Histoire Naturelle sur les animaux sans vertèbres. D’Hautel, Paris. Lamarck, J.B. DE. 1816. Tableau encyclopédique et méthodique des trois règnes de la nature. Partie 23 – Mollusques et Polypes divers. Mme veuve Agasse, Paris. Lange, E. 1925. Eine mittelpermische Fauna von Guguk Bulat (Padanger Oberland, Sumatra). Verhandelingen van het geologisch mijnbouwkunding Genootschap voor Nederland en Koloniën, geologisch serie, 7, 213–295. Langer, M.R., Lipps, J.H. & Piller, W.E. 1993. Molecular paleobiology of prtists: amplification and direct sequencing of foraminiferal DNA. Micropaleontology, 39, 63–68. Lankester, E.R. 1885. Protozoa. In: Encyclopedia Britannica, Volume 19. 9th edn. Encyclopædia Britannica, London. Leven, E.YA. 1967. Stratigraphy and fusulinids of the Permian strata of Pamir. Trudy Geologicheskogo Instituta Akademiya Nauk SSSR, Moscow, 167, 1– 224 [in Russian]. Leven, E.YA. 1998. Permian fusulinids assemblages and stratigraphy of Transcaucasia. Rivista Italiana di Paleontologia e Stratigrafia, 104, 299– 328. Leven, E.YA. 2003. Permian stratigraphy and fusulinids of the Tethys. Rivista Italiana di Paleontologia i Stratiigrafia, 109, 267–280. Leven, E.YA. & Gorgij, M.N. 2011. Fusulinids and stratigraphy of the Carboniferous and Permian in Iran. Stratigraphy and Geological Correlation, 19, 687–776. Leven, E.YA. & Okay, A.I. 1996. Foraminifera from the exotic Permo-Carboniferous limestone blocks in the Karakaya complex, northwestern Turkey. Rivista Italiana di Paleontologia e Stratigrafia, 102, 139–174. Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 PERMIAN SMALLER FORAMINIFERS Likharev, B.K. 1939. Atlas of the Guide Species Among the Fossil Fauna of the USSR. Tsentralnyi Nauchnoissledovatelskii Geologo-razvedochnyi Institut, Leningrad, 6 (Permian System), 1– 268 [in Russian]. Lin, J.X. 1978. Carboniferous and Permian Foraminiferida. In: HUBEI INSTITUTE OF GEOLOGICAL SCIENCE (eds) Paleontological Atlas of Central South China (Micropaleontological Volume). Geological Publishing House, Beijing, 10–43 [in Chinese]. Lin, J.X. 1980. On the age and stratigraphical significance of the genus Gallowayinella. Bulletin of the Chinese Academy Geological Sciences, Series 8, 1, 37– 46 [in Chinese]. Lin, J.X. 1984. Protozoa. In: YICHAN INSTITUTE OF GEOLOGY & MINERAL RESOURCES (eds) Biostratigraphy of the Yangtze Gorge Area Chiefly, 3) Late Paleozoic Era. Geological Publishing House, Beijing, 110 –177 [in Chinese], 323– 364 [in English]. Lin, J.X. 1985. Late Early Permian foraminifera and its paleoecology in Jiahe, Hunan. Bulletin of the Yichang Institute of Geology and Mineral Resources, Chinese Academy of Geological Sciences, 9, 43– 52 [in Chinese with English abstract]. Lin, J.X., Li, L.X. & Sun, Q.Y. 1990. Late Paleozoic Foraminifers in South China, Volume 1. Science Publication House, Beijing [in Chinese]. Liu, C., Jarochowska, E., Du, Y., Vachard, D. & Munnecke, A. In press. Stratigraphical and d13C records of Permo-Carboniferous platform carbonates: responses to deep icehouse and icehouse– greenhouse transition. Palaeogeography, Palaeoclimatology, Palaeogeography. Loeblich, A.R. & Tappan, H. 1955. A Revision of Some Glanduline Nodosariidae (Foraminifera). Smithsonian Miscellaneous Collections, 126, (3). Loeblich, A.R. & Tappan, H. 1961. Suprageneric classification of the Rhizopodea. Journal of Paleontology, 35, 245–330. Loeblich, A.R. & Tappan, H. 1964. Sarcodina, chiefly ‘Thecamoebians’ and Foraminiferida. In: Moore, R.C. (ed.) Treatise of Invertebrate Paleontology, Part C, Protista 2, Volume 2. Geological Society of America and the University of Kansas Press, Lawrence, KS, C1–C900. Loeblich, A.R. & Tappan, H. 1981. Suprageneric revision of some calcareous Foraminiferida. Journal of Foraminiferal Research, 11, 159– 164. Loeblich, A.R. & Tappan, H. 1982. Classification of the Foraminiferida. In: Broadhead, T.W. (ed.) Foraminifera, Notes for a Short Course Organized by M.A. Buzas and B.K. Sen Gupta. University of Tennessee, Department of Geological Sciences, Studies in Geology, 6, 22–36. Loeblich, A.R. & Tappan, H. 1984. Suprageneric classification of the Foraminiferida (Protozoa). Micropaleontology, 30, 1 –70. Loeblich, A.R. & Tappan, H. 1985. Cylindrocolaniella a new name for the foraminiferal genus Wanganella Sosnina, 1956, non Wanganella Laserow, 1954. Journal of Foraminiferal Research, 15, 218. Loeblich, A.R. & Tappan, H. 1986a. Some new and redefined genera and families of Textulariina, Fusulinina, Involutinina and Miliolina (Foraminiferida). Journal of Foraminiferal Research, 16, 334–346. 245 Loeblich, A.R. & Tappan, H. 1986b. Some new and revised genera and families of hyaline calcareous Foraminiferida (Protozoa). Transactions of the American Microscopical Society, 105, 239 –265. Loeblich, A.R. & Tappan, H. 1987. Foraminiferal Genera and Their Classification. Van Nostrand Reinhold, New York. Loeblich, A.R. & Tappan, H. 1992. Present status of foraminiferal classification. In: Takayanagi, Y. & Saito, T. (eds) Studies in Benthic Foraminifera, Benthos’ 90. Tokai University Press, Tokyo, 93–102. Lucas, S.G., Krainer, K. & Vachard, D. 2015. The Lower Permian Hueco Group, Robledo Mountains, New Mexico (U.S.A.). In: Lucas, S.G. & DiMichele, W.A. (eds) Carboniferous –Permian Transition in the Robledo Mountains, Southern New Mexico. New Mexico Museum of Natural History and Science Bulletin, 65, 43– 95. Lucas, S.G., Krainer, K. & Vachard, D. 2016a. The Pennsylvanian system in the Mud Springs Mountains, Sierra County, New Mexico. New Mexico Museum of Natural History and Science Bulletin, 69, 1–58. Lucas, S.G., Krainer, K. & Vachard, D. 2016b. The Pennsylvanian section at Priest Canyon, southern Manzano Mountains, New Mexico. In: New Mexico Geological Society, 67th Field Conference, Geology of the Belen Area. New Mexico Geological Society, Socorro, NM, 69– 95. Lys, M. & Marcoux, J. 1978. Les niveaux du Permien supérieur des Nappes d’Antalya (Taurides occidentales, Turquie). Comptes Rendus Académie Sciences Paris, 286, série D, 1417–1420. Lys, M., Colchen, M., Bassoullet, J.P., Marcoux, J. & Mascle, G. 1980. La biozone à Colaniella parva du Permien supérieur et sa microfaune dans le bloc calcaire exotique de Lamayuru, Himalaya du Ladakh. Revue de Micropaléontologie, 23, 76–108. MacFadyen, W.A. 1941. Foraminifera from the Green Ammonite Beds, lower Lias, of Dorset. Philosophical Transactions of the Royal Society, London, Series B, 231, 1– 73. Maslakova, N.I. 1990. Criteria for establishing higher taxa in foraminifers. In: Menner, V.V. (ed.) Systematics and Phylogeny of Invertebrates: The Criteria for Establishing Higher Taxa. Izdateltsvo Nauka, Moscow, 22–27 [in Russian]. Maslakova, N.I., Gorbachik, T.N., Alekseev, A.S., Barskov, I.S., Golubev, S.N., Nazarov, B.B. & Petruchevskaya, M.G. 1995. Micropalaeontology (Anonymous: Class Foraminifera, foraminifers, 13-111). Izdatelsvo Moskovskogo Universita, Moscow [in Russian]. Mamet, B. 1996. Late Paleozoic small foraminifers (endothyrids) from South America (Ecuador and Bolivia). Canadian Journal of Earth Sciences, 33, 452–459. Mamet, B. & Pinard, S. 1992. Note sur la taxonomie des petits foraminifères du Paléozoı̈que supérieur. Bulletin de la Société belge de Géologie, 99, 373 –398. Mamet, B.L., Roux, A. & Nassichuk, W.W. 1987. Algues carbonifères et permiennes de l’Arctique canadien. Geological Survey of Canada Bulletin, 342, 1– 143. Maslov, V.P. 1956. Fossil calcareous algae from the USSR. Trudy Instituta Geologichesnikh Nauk, Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 246 D. VACHARD Akademiya Nauk SSSR, 160, 1 –301 [in Russian; French translation BRGM No. 3517]. Mertmann, D. 2000. Foraminiferal assemblages in Permian carbonates of the Zaluch Group (Salt Range and Trans Indus Ranges, Pakistan). Neues Jahrbuch Paläontologie Monathefte, 2000, 129 –146. Mikhalevich, V.I. 1980. Systematics and evolution of the foraminifers in the light of some new data on their cytology and ultrastructure. Trudy Zoologicheskogo Instituta, Akademiya Nauk SSSR, 94, 42– 61 [in Russian]. Mikhalevich, V.I. 1988. System of the subclass Miliolata (Foraminifera). Akademiya Nauk SSSR, Trudy Zoologicheskogo Instituta, 184, 77– 110 [in Russian]. Mikhalevich, V.I. 1993. New higher taxa of the subclass Nodosariata (Foraminifera). Zoosystematica Rossica, 2, 5 –8. Mikhalevich, V.I. 2003. System of the four foraminiferal subclasses with the agglutinated shell wall (Ammodiscana, Miliamminana, Hormosiniana, Textulariana) (Foraminifera). Rossiiskaya Akademiya Nauk, Izvestiya Zoologicheskogo Instituta, 7, 5 –50. Mikhalevich, V.I. 2013. New insight into the systematics and evolution of the foraminifera. Micropalaeontology, 59, 493– 527. Miklukho-Maklay, A.D. 1953. On the systematics of the family Archaediscidae. Ezhegodnik Vsesoyuznogo Paleontologischeskogo Obshchestva (1948– 53), 14, 127– 131 [in Russian]. Miklukho-Maklay, A.D. 1956. On the systematics of the Palaeozoic foraminifers. Vestnik Leningradskogo Universiteta, 6, 58– 66 [in Russian]. Miklukho-Maklay, A.D. 1958. A new foraminiferal family, Tuberitinidae M.Maclay fam. nov. Voprosy Mikropaleontologii, 2, 130 –135 [in Russian]. Miklukho-Maklay, K.V. 1954. Permian foraminifers from North Caucasus. Trudy VSEGEI, Gosgeoltekhizdat, 1, 1 –163 [in Russian; French translation BRGM No. 2683, 1 –216]. Miklukho-Maklay, K.V. 1960a. New Kazanian lagenids from the Russian Platform. In: Markovskyi, B.P. (ed.) New Species of Fossil Plants and Invertebrates from the USSR, pt. 1. Trudy VSEGEI, Gosgeoltekhizdat, 153– 161 [in Russian]. Miklukho-Maklay, K.V. 1960b. On the genus Lunucammina Spandel, 1898 (foraminifers). Ezhegodnik Vsesoyuznogo Paleontologicheskogo Obshchestva, 32, 5 –12 [in Russian]. Mohtat-Aghai, P. & Vachard, D. 2003. Dagmarita shahrezahensis n. sp. globivalvulinid foraminifer (Wuchiapingian, Late Permian, Central Iran). Rivista Italiana di Paleontologia i Stratigrafia, 109, 37–44. Mohtat-Aghai, P. & Vachard, D. 2005. Late Permian foraminiferal assemblages from the Hambast region (Central Iran) and their extinctions. Revista Española de Micropaleontologı́a, 37, 205 –227. Mohtat-Aghai, P., Vachard, D. & Krainer, K. 2009. Transported foraminifera in Palaeozoic deep red nodular limestones exemplified by latest Permian Neoendothyra in the Zal section (Julfa area, NW Iran). Revista Española de Micropaleontologı́a, 41, 197–213. Moix, P., Vachard, D., Allibon, J., Martini, R., Wernli, R., Kozur, H.W. & Stampfli, G.M. 2013. Palaeotethyan, Neotethyan and Hu Lu-Pindos series in the Lycian Nappes (SW Turkey): geodynamical implications. In: Tanner, L.H., Spielmann, J.A. & Lucas, S.G. (eds) The Triassic System. New Mexico Museum of Natural History and Science, 61, 401– 444. Montenat, C., de Lapparent, A.F., Lys, M., Termier, H., Termier, G. & Vachard, D. 1977. La transgression permienne et son substrat éocambrien dans le Jebel Akhdar, montagnes d’Oman, Péninsule Arabique. Annales de la Société géologique du Nord, 96, 239–258. von Möller, V. 1878. Die spiral-gewundenen Foraminiferen des russischen Kohlenkalks. Mémoires de l’Académie Impériale des Sciences de St Pétersbourg, 7th Series, 25, 1 –147. Nestell, G.P. & Nestell, M.K. 2006. Middle Permian (Late Guadalupian) foraminifers from Dark Canyon, Guadalupe Mountains, New Mexico. Micropaleontology, 52, 1 –50. Nestell, G.P., Mestre, A. & Heredia, S. 2009. First Ordovician Foraminifera from South America: a Darriwilian (Middle Ordovician) fauna from the San Juan Formation, Argentina. Micropaleontology, 55, 329–344. Nestell, G.P., Nestell, M.K. et al. 2015. High influx of carbon in walls of agglutinated foram inifers during the Permian– Triassic transition in global oceans. International Geology Review, 57, 411– 427, https://doi.org/ 10.1080/00206814.2015.1010610 Nestell, M.K. & Pronina, G.P. 1997. The distribution and age of the genus Hemigordiopsis. In: Ross, C.A., Ross, J.R.P. & Brenckle, P.L. (eds) Late Paleozoic Foraminifera; Their Biostratigraphy, Evolution, and Paleoecology; and the Mid-Carboniferous Boundary. Cushman Foundation for foraminiferal Research, Special Publications, 36, 105–110. Nestell, M.K., Nestell, G.P., Wardlaw, B.R. & Sweatt, M.J. 2006. Integrated biostratigraphy of foraminifers, radiolarians and conodonts in shallow and deep water Middle Permian (Capitanian) deposits of the ‘Rader slide’, Guadalupe Mountains, West Texas. Stratigraphy, 3, 161–194. Nestler, H. 1973. The types of Tetrataxis conica Ehrenberg, 1854 and Tetrataxis palaeotrochus (Ehrenberg, 1854). Micropaleontology, 19, 366– 369. Neumayr, M. 1887. Die natürlichen Verwandtschaftverhältnisse der schalentragenden Foraminiferen. Sitzungsberichte der kaiserliche Akademie der Wissenschaften in Wien, Mathematisch-Naturwissenschaftliche Klasse, 95, 156–186. NGUYEN DUC TIEN 1979. Etude micropaléontologique (foraminifères) de matériaux du Permien du Cambodge. PhD thesis, Université Paris Sud, Orsay. NGUYEN DUC TIEN 1986a. Foraminifera and algae from the Permian of Kampuchea. In: Fontaine, H., Suteethorn, V. et al. (eds) The Permian of Southeast Asia. CCOP Technical Bulletin, 18, 116– 137. NGUYEN DUC TIEN 1986b. Foraminifera and algae from the Permian of Guguk Bulat and Silungkang, Sumatra. In: Fontaine, H., Suteethorn, V. et al. (eds) The Permian of Southeast Asia. CCOP Technical Bulletin, 18, 138– 147. Nguyen Duc TIEN 1987. Sphairionia sikuoides gen. nov. sp. nov. a Permian incertae sedis organism. 11e Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 PERMIAN SMALLER FORAMINIFERS Congrès International Stratigraphie et Géologie Carbonifère, 1–90, Beijing. NGUYEN DUC TIEN 1989a. Lower Permian Foraminifera of Sumatra. In: Fontaine, H. & Gafoer, S. (ed.) The Pre-Tertiary Fossils of Sumatra and Their Environments. CCOP Technical Publications, 19, 71– 93. NGUYEN DUC TIEN 1989b. Sphairionia sikuoides nov. gen. nov. sp., a Permian incertae sedis organism. 11e Congrès International de Géologie du Carbonifère, Beijing 1987, Compte Rendu, 3, 73– 78. Nikitina, A.P. 1969. Genus Hemigordiopsis (Foraminifera) in the Late Permian of Primorye. Paleontologicheskii Zhurnal, 3, 63–69 [in Russian]. Noé, S.U. 1987. Facies and paleogeography of the marine Upper Permian and of the Permian-Triassic boundary in the southern Alps (Bellerophon Formation, Tesero Horizon). Facies, 16, 89–142. Norvang, A. 1957. The foraminifera of the Lias series in Jutland, Denmark. Meddeleser fra Dansk Geologisk Forening, 13, 1 –135. Okimura, Y. 1972. Permo-Carboniferous endothyraceans from Japan; Part 1: Biseriamminidae. Transactions of the Proceedings of the Paleontological Society Japan, 87, 414–428. Okimura, Y. 1988. Primitive colaniellid foraminiferal assemblage from the Upper Permian Wargal Formation of Salt Range, Pakistan. Journal of Paleontology, 62, 715–723. Okimura, Y. & Ishii, K.-i. 1981. Smaller foraminifera from the Abadeh Formation, Abadehian stratotype, central Iran. Reports of the Geological Survey of Iran, 49, 7 –27. Okimura, Y., Ishii, K.-i. & Nakazawa, K. 1975. Abadehella, a new genus of tetrataxid Foraminifera from the Late Permian. Memoirs of the Faculty of Science Kyoto University, Series Geology and Mineralogy, 41, 45– 48. Okuyucu, C. 2007. New Middle Permian foraminifers (Chitralinidae) from the Karakaya Complex, in northwestern Turkey. Comptes Rendus Palevol, 6, 311–319. Okuyucu, C., Vachard, D. & Göncüoğlu, M.C. 2013. Refinements in biostratigraphy of the foraminiferal zone MFZ11 (middle Visean, Mississippian) in the Cebeciköy Limestone (Istanbul Terrane, NW Turkey), palaeogeographic implications. Bulletin of Geosciences, 88, 621– 645. Okuyucu, C., Ivanova, D., Bedi, Y. & Ergen, A. 2014. Discovery of an earliest Triassic, post-extinction foraminiferal assemblage above the Permian-Triassic boundary, Strandzha nappes, north-west Turkey. Geological Quarterly, 58, 1– 8, https://doi.org/10.7306/ gq.1145 Orlov-Labkovsky, O. 2004. Permian fusulinids (Foraminifera) of the subsurface of Israel: taxonomy and biostratigraphy. Revista Española de Micropaleontologı́a, 36, 389–406. Özdikmen, H. 2009. Substitute names for some unicellular animal taxa (Protozoa). Munis Entomology & Zoology, 4, 233–256. Palmieri, V. 1984. Permian foraminifera in the Bowen Basin, Queensland. Queensland Geology, 6, 1 –126. Palmieri, V. 2004. Flectospiroides fusiformis gen. et sp. nov. (Foraminiferida) from the Late Permian of the 247 Bowen Basin, central Queensland. Memoir of the Association of Australasian Palaeontologists, 29, 259– 264. Palmieri, V., Foster, C.B. & Bondareva, E.V. 1994. First record of shared species of Late Permian small foraminiferids in Australia and Russia: time correlations and plate reconstructions. AGSO Journal of Australian Geology and Geophysics, 15, 359– 365. Pantic, S. 1965. Pilammina densa n. gen. n. sp. and other Ammodiscidae from the Middle Triassic in the Cr. Mnica (Montenegro). Geoloski Vjesnik, 18, 189–192. Pantic, S. 1970. Lithostratigraphy and micropaleontology of the Middle and Upper Permian of Western Serbia. Bulletin Institute Geology Geophysics Research (Geology), A, 27, 239–272. Panzanelli-Fratoni, R., Limongi, P., Ciarapica, G., Cirilli, S., Martini, R., Salvini-Bonnard, G. & Zaninetti, L. 1987. Les Foraminifères du Permien supérieur remaniés dans le ‘Complexe Terrigène’ de la formation triasique du Monte Facito, Apennin méridional. Revue de Paléobiologie, 6, 293–319. Pawlowski, J., Holzmann, M. & Tyszka, J. 2013. New supraordinal classification of Foraminifera: molecules meet morphology. Marine Micropaleontology, 100, 1– 10. Phillips, J. 1846. On the remains of microscopic animals in the rocks of Yorkshire. Proceedings of the Geological and Polytechnic Society of the West Riding of Yorkshire (1844– 45), 2, 274– 285. Pia, J.von. 1937. Die wichtigsten Kalkalgen des Jungpaläozoikums und ihre geologische Bedeutung. Compte Rendu du 2e Congrès Avancement Etudes de Stratigraphie du Carbonifère, Heerlen 1935, 2, 765– 856. Pille, L. 2008. Foraminifères et algues calcaires du Mississippien supérieur (Viséen supérieur-Serpukhovien): rôles biostratigraphique, paléoécologique et paléogéographique aux échelles locale, régionale et mondiale. PhD thesis, Université de Lille. Piller, W.E. 1983. Remarks on the suborder Involutinina Hohenegger and Piller, 1977. Journal of Foraminiferal Research, 13, 191–201. Pinard, S. & Mamet, B. 1998. Taxonomie des petits foraminifères du Carbonifère supérieur-Permien inférieur du basin de Sverdrup, Arctique canadien. Palaeontographica Canadiana, 15, 1 –253. Plummer, H.J. 1930. Calcareous foraminifera in the Brownwood shale near Bridgeport, Texas. Bulletin of the University of Texas, 3019, 5– 21. Plummer, H.J. 1945. Smaller foraminifera in the Marble Falls, Smithwick, and lower Strawn strata around the Llano uplift in Texas. Bulletin University of Texas, 4401, 209– 271. Pokorny, V. 1958. Grundzüge der zoologischen Mikropaläontologie, Bd 1. VEB Deutscher Verlag der Wissenschaften, Berlin. Poty, E., Devuyst, F.X. & Hance, L. 2006. Upper Devonian and Mississippian foraminiferal and rugose coral zonations of Belgium and Northern France, a tool for Eurasian correlations. Geological Magazine, 143, 829– 857. Poyarkov, B.V. & Skvortsov, V.P. 1979. On methodological segregation of local epiboles and local biozones (exemplified by the Early Carboniferous of Tian Shan). Akademiya Nauk SSR, Dalvnevostochnyi Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 248 D. VACHARD Nauchnyi Tsentr, Dalvnevostochnyi Geologicheskii Institut, Vladivostok, 5 –27 [in Russian]. Pronina, G.P. 1988a. The Late Permian smaller foraminifers of Transcaucasus. Revue de Paléobiologie, Special Volume, 1, 89–96. Pronina, G.P. 1988b. Late Permian miliolates of Transcaucasia. Trudy Zoologicheskogo Instituta, Akademiya Nauk SSSR, 184, 49–63 [in Russian]. Pronina, G.P. 1994. Systematics and phylogeny of the order Hemigordiopsida (foraminifers). Paleontologicheskii Zhurnal, l994, 13–24 [in Russian]. Pronina, G.P. 1996. Genus Sphairionia and its stratigraphic significance. In: Reports of Shallow Tethys 4, International Symposium, Albrechtsberg (Austria) 8 – 11 September 1994. Supplement agli Annali dei Musei Civici di Rovereto, Sezione Archeologia, Storia e Scienze Naturali, 11, 105 –118. Pronina, G.P. 1998. Foraminifers. In: Esaulova, N.K., Lozovsky, V.R. & Rozanov, A.Yu. (eds) Stratotypes and Reference Sections of the Upper Permian in the Regions of the Volga and Kama Rivers. International Symposium Upper Permian Stratotypes of the Volga Region. GEOS, Moscow, 163– 169 [in Russian]. Pronina, G.P. 1999a. Correlations of Late Permian deposits of Boreal regions based on smaller foraminifers. In: International Symposium Upper Permian Stratotypes of the Volga Region. GEOS, Moscow, 182 –188 [in Russian]. Pronina, G.P. 1999b. New Late Permian species of smaller foraminifers from Transcaucasia. Voprosy Paleontologii, 11, 4–14 [in Russian]. Pronina, G.P. & Gubenko, T.A. 1990. Upper Permian planoarchaediscids from Transcaucasia. Paleontologicheskii Zhurnal, 2, 119–123 [in Russian; English translation: Paleontological Journal, 1990, 2, 119–123]. Pronina, G.P. & Nestell, M.K. 1997. Middle and Late Permian Foraminifera from exotic limestone blocks of the Alma river basin, Crimea. In: Ross, C.A., Ross, J.R.P. & Brenckle, P.L. (eds) Late Paleozoic Foraminifera; Their Biostratigraphy, Evolution, and Paleoecology; and the Mid-Carboniferous Boundary. Cushman Foundation for Foraminiferal Research, Special Publications, 36, 111 –114. Pronina-Nestell, G.P. & Nestell, M.K. 2001. Late Changhsingian foraminifers of the northwestern Caucasus. Micropaleontology, 47, 205– 234. Rauzer-Chernousova, D.M. & Chermnykh, B.A. 1990. Mesolasiodiscus gen. nov. – new data on the evolution of the Late Palaeozoic. Paleontologicheskii Zhurnal, 1990, 1, 121–125 [in Russian]. Rauzer-Chernousova, D.M. & Fursenko, A.V. 1959. Fundamentals of Paleontology, General Part, Protozoans. Akademiya Nauk SSSR, Moscow [in Russian]. Rauzer-Chernousova, D.M., Bensh, F.R. et al. 1996. Reference-Book on the Systematics of Paleozoic Foraminifers; Endothyrida and Fusulinoida. Rossiiskaya Akademiya Nauk, Geologicheskii Institut, Moscow [in Russian]. Reichel, M. 1945. Sur un Miliolidé nouveau du Permien de l’ı̂le de Chypre. Verhandlungen der Naturforschenden Gesellschaft in Basel, 56, 521 –530. Reichel, M. 1946. Sur quelques foraminifères nouveaux du Permien méditerranéen. Eclogae Geologicae Helvetiae, 38, 524–560. Reiss, Z. 1963. Reclassification of perforate Foraminifera. Bulletin of the Geological Survey of Israel, 35, 1– 111. Reitlinger, E.A. 1950. Middle Carboniferous foraminifers of the central part of Russian Platform (Fusulinidae excepted). Trudy Instituta Geologicheskikh Nauk, geologichevskaya seriya 6, 47, 1– 126 [in Russian; French translation BRGM No. 1456]. Reitlinger, E.A. 1956. New family Lasiodiscidae. Voprosy Mikropaleontologii, 1, 69–78 [in Russian; French translation by Sigal, S. & Sigal, J., BRGM No. 1753, 1– 16]. Reitlinger, E.A. 1958. On the questions of systematics and phylogeny of the superfamily Endothyroidea. Voprosy Mikropaleontologii, 2, 53–73 [in Russian; French translation by Sigal, S. & Sigal, J. 1960, Editions Technip, Paris, 59–81]. Reitlinger, E.A. 1965. On the development of the foraminifera of the Late Permian and Early Triassic in Transcaucasia. Voprosy Mikropaleontologii, 9, 45–70 [in Russian]. Reuss, A.E. 1862. Entwurf einer systematischen Zusammenstellung der Foraminiferen. Sitzungsberichte der kaiserlichen Akademie der Wissenschaften in Wien, Mathematich-Naturwissenschaftlische Klasse (1862), 46, 5 –100. Rettori, R. 1994. Replacement name Hoyenella, gen. n. (Triassic Foraminiferida, Miliolina) for Glomospira sinensis Ho, 1959. Bolletino della Società Paleontologica Italiana, 33, 341–343. Rettori, R. 1995. Foraminiferi del Trias inferiore e medio della Tetide: revisione tassonomica, stratigrafia ed interpretazione filogenetica. Université de Genève, Publications du Département de Géologie et Paléontologie, 18. Rhumbler, L. 1895. Entwurf eines natürlichen Systems der Thalamophoren. Nachrichten von der Gesellschaft der Wissenschaften zu Göttingen, Mathematisch– Physikalische Klasse, 1895, 51– 98. Rigaud, S., Vachard, D. & Martini, R. 2014. Unsuspected phylogenetic links between Paleozoic Fusulinana and Recent Textulariana. In: Marchant, M. & Hromic, T. (eds) International Symposium on Foraminifera Forams 2014, 19– 24 January 2014, Chile, Abstract Volume. Grzybowski Foundation Special Publication, 20, 4. Rigaud, S., Vachard, D., Martini, R. & Rettori, R. 2015a. Agglutinated versus microgranular: end of a paradigm? Journal of Systematic Palaeontology, 13, 75–95, https://doi.org/10.1080/14772019.2013. 863232 Rigaud, S., Vachard, D., Schlagintweit, F. & Martini, R. 2015b. New lineage of Triassic aragonitic Foraminifera and reassessment of the class Nodosariata. Journal of Systematic Palaeontology, 14, 1 –19, https://doi.org/10.1080/14772019.2015. 1112846 Şahin, N., Altıner, D. & Bülent Ercengiz, M. 2014. Discovery of Middle Permian volcanism in the Antalya Nappes, southern Turkey: tectonic significance and global meaning. Geodinamica Acta, 25, 286– 304, https://doi.org/10.1080/09853111.2013.858949 Saidova, KH.M. 1981. On an Up-to-Date System of Supraspecific Taxonomy of Cenozoic Benthic Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 PERMIAN SMALLER FORAMINIFERS Foraminifers. Institut Okeanologii P.P. Shirshova, Akademiya Nauk SSSR, Moscow [in Russian]. Salaj, J., Borza, K. & Samuel, O. 1983. Triassic Foraminifers of the West Carpathians. Geologicky Ustav Dionyza Stura, Bratislava, Slovakia. Sanders, D. & Krainer, K. 2005. Taphonomy of Early Permian benthic assemblages (Carnic Alps, Austria): carbonate dissolution v. biogenic carbonate precipitation. Facies, 51, 522–540, https://doi.org/10. 1007/s10347-005-0065-6 Schellwien, E. 1898. Die Fauna des Karnischen Fusulinenkalks. Theil 2: Foraminifera. Palaeontographica, 44, 237–282. Schubert, R.J. 1908. Beiträge zu einer natürlichen Systematik der Foraminiferen. Neues Jahrbuch für Mineralogie, Geologie und Paläontologie, 25, 232–260. Schubert, R.J. 1921. Paläontologische Daten zur Stammesgeschichte der Protozoen. Paläontologische Zeitschrift, 3, 129 –188. Schultze, M.S. 1854. Über den Organismus der Polythalamien (Foraminiferen) nebst Bemerkungen über die Rhizopoden im allgemeinen. Wielhem Engelmann, Leipzig. Sellier de Civrieux, J.M. & Dessauvagie, T.F.J. 1965. Reclassification de quelques Nodosariidae, particulièrement du Permien au Lias. Maden Tetkik ve Arama Enstitu sü Yayinlarindan (M.T.A.), 124, 1–178. Şengör, A.M.C., Altıner, D., Clin, A., Ustaösmer, T. & Hsü, K.J. 1988. Origin and assembly of the Tethyside orogenic collage at the expense of Gondwana Land. In: Audley-Charles, M.G. & Hallam, A. (eds) Gondwana and Tethys. Geological Society, London, Special Publications, 37, 119– 181, https://doi. org/10.1144/GSL.SP.1988.037.01.09 Senowbari-Daryan, B., Bucur, I., Schlagintweit, F., Sasaran, E. & Matyszkiewicz, J. 2008. Crescentiella, a new name for ‘Tubiphytes’ morronensis Crescenti, 1969: an enigmatic Jurassic–Cretaceous microfossil. Geologica Croatica, 61, 185–214. Shang, Q.H., Vachard, D. & Caridroit, M. 2003. Smaller foraminifera from the late Changhsingian (latest Permian) of southern Guangxi and discussion on the Permian-Triassic boundary. Acta Micropalaeontologica Sinica, 20, 377– 388. Sheng, J.Z. & He, Y. 1983. Permian Shanita-Hemigordius (Hemigordiopsis) (Foraminifera) fauna in western Yunnan, China. Acta Palaeontologica Sinica, 22, 55– 59 [in Chinese with English abstract]. Silvestri, A. 1903. Linguloglanduline e Lingulonodosarie. Atti della Pontifica Accademia Romana dei Nuovi Lincei, Roma (1902–03), 56, 45–50. Sone, M. 2008. Correction of the publication date of Sphaerionia Tiên, a possible foraminifer or incertae sedis. Journal of Foraminiferal Research, 38, 271. Song, H.J., Tong, J.N., Zhang, K.X., Wang, Q.X. & Chen, Z.Q. 2007. Foraminiferal survivors from the Permian–Triassic mass extinction in the Meishan section, South China. Palaeoworld, 16, 105–119. Song, H.J., Tong, J.N., Chen, Z.Q., Yang, H. & Wang, Y.B. 2009. End-Permian mass extinction of foraminifers in the Nanpanjiang Basin, South China. Journal of Paleontology, 83, 718– 738. 249 Song, H., Tong, J. & Chen, Z.Q. 2011. Evolutionary dynamics of the Permian-Triassic foraminifer size: evidence for Lilliput effect in the end-Permian mass extinction and its aftermath. Palaeogeography, Palaeoclimatology, Palaeoecology, 308, 98– 110. Song, H.J., Wignall, P.B., Tong, J.N. & Yin, H.F. 2013. Two pulses of extinction during the Permian–Triassic crisis. Nature Geoscience, 6, 52–56. Sosipatrova, G.P. 1969. Foraminifers of the Starostin Suite of Spitsbergen. Trudy NIIGA, Sbornik Statey po Paleontologii i Biostratigrafii, 30, 35–72 [in Russian]. Sosipatrova, G.P. 1970. Foraminifers. In: Menner, V.V., Sarycheva, T.G. & Chernyak, G.E. (eds) Stratigraphy of the Carboniferous and Permian deposits of the Northern Verkhoyansk. Trudy NIIGA, Vsesoyuznyi Aèrogeologicheskii Trest, 154, 56–70 [in Russian]. Sosnina, M.I. 1969. New name Eomarginulinella for the genus Marginulinella Sosnina, 1967. Paleontologicheskiy Zhurnal, 1969, 101 [in Russian]. Sosnina, M.I. 1977. Late Permian nodosariids of southern Primorye. Ezhegodnik Vsesoyuznogo Paleontologicheskogo Obshchestva, 20, 10–31 [in Russian]. Sosnina, M.I. 1978. On the foraminifers of the Late Permian Chandalaz horizon of Primorye. Akademiya Nauk SSSR, Dalnevostochnyi Nauchniyi Tsentr, Institut Tektoniki i Geofiziki, Moscow, 24–42 [in Russian]. Sosnina, M.I. 1983. Some new Late Permian miliolids and nodosariids (foraminifers) from the southern Primorye. Ezhegodnik Vsesoyuznogo Paleontologischeskogo Obshchestva, 26, 29–47 [in Russian]. Sosnina, M.I. & Nikitina, A.P. 1977. Late Permian smaller foraminifers of Primoriye. Akademiya Nauk SSSR, Dalnevostochnyi Nauchniyi Tsentr, Dalnevostochnyi Geologicheskii Institut, Moscow, 27–52 [in Russian]. Spandel, E. 1898. Die Foraminiferen des deutschen Zechsteins (vorläufige Mitteilung) und ein zweifelhaftes mikrokopisches Fossil ebendaher. Verlag des Verlags-Instituts ‘General-Anzeiger’, Nürnberg. Spandel, E. 1901. Die Foraminiferen des Permo-Karbons von Hooser, Kansas, Nord Amerika. Festschrift der Naturhistorischen Gesellschaft, Nürnberg, 175–194. Stancliffe, R.P.W. 1989. Microforaminiferal linings: their classification, biostratigraphy and paleoecology, with special reference to specimens from British Oxfordian sediments. Micropaleontology, 35, 337–352. Termier, G., Termier, H. & Vachard, D. 1977. Etude comparative de quelques Ischyrosponges. Géologie Méditerranéenne, 4, 139– 180. Théry, J.M., Vachard, D. & Dransart, E. 2007. Late Permian limestones and Permian-Triassic Boundary: new biostratigraphic, palaeobiogeographic and geochemical data in Caucasus and Eastern Europe. In: Alvaro, J.J., Aretz, M., Boulvain, F., Munnecke, A., Vachard, D. & Vennin, E. (eds) Palaeozoic Reeefs and Bioaccumulations: Climatic and Evolutionary Controls. Geological Society, London, Special Publications, 275, 255–274, https://doi.org/10. 1144/GSL.SP.2007.275.01.16 Ukharskaya (¼ Ucharskaja), L.B. 1970. New Kazanian (Permian) arenaceous foraminifers of the Russian Platform. Paleontologicheskii Zhurnal, 4, 468–476. Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 250 D. VACHARD Ueno, K. 2003. The Permian fusulinoidean faunas of the Sibumasu and Baoshan blocks: their implications for the paleogeographic and paleoclimatologic reconstruction of the Cimmerian Continent. Palaeogeography, Palaeoclimatology, Palaeoecology, 193, 1 –24. Ueno, K. & Sakagami, S. 1993. Middle Permian foraminifers fauna from Ban Nam Suai Tha Sa-at, Changwat Loei, Northeast Thailand. Transactions and Proceedings of the Paleontological Society of Japan, 172, 277– 291. Ueno, K. & Tsutsumi, S. 2009. Lopingian (Late Permian) foraminiferal faunal succession of a Paleo-Tethyan mid-oceanic carbonate buildup: Shifodong Formation in the Changning-Menglian Belt, West Ynnan, Southwest China. Island Arc, 18, 69–93. Ueno, K., Miyahigashi, A. & Chaoentitirat, T. 2010. The Lopingian (Late Permian) of mid-oceanic carbonates in the Eastern Palaeotethys: stratigraphical outline and foraminiferal faunal [sic] succession. Geological Journal, 45, 285 –307. Ünal, E., Altıner, D., Ömer Yilmaz, I. & ÖzkanAltıner, S. 2003. Cyclic sedimentation across the Permian-Triassic boundary (Central Taurides, Turkey). Rivista Italiana di Paleontologia i Stratigrafia, 109, 359– 376. Vachard, D. 1980. Téthys et Gondwana au Paléozoı̈que supérieur; les données afghanes: biostratigraphie, micropaléontologie, paléogéographie. Documents et Travaux IGAL, Institut Géologique Albert de Lapparent, 2, 1– 463. Vachard, D. 1990. Fusulinoids, smaller foraminifera and pseudo-algae from southeastern Kelantan (Malaysia) and their biostratigraphic and paleogeographic value. In: Fontaine, H. (ed.) Ten years of CCOP Research on the Pre-Tertiary of East Asia. CCOP Technical Publications, 20, 43–167. Vachard, D. 1994. Foraminifères et moravamminides du Givétien et du Frasnien du domaine Ligérien (Massif Armoricain, France). Palaeontographica, A, 231, 1 –92. Vachard, D. 2000. On some umbellinids (carbonate microproblematica) from the Frasnian (Late Devonian) of Chah Riseh area (central Iran). Annales de la Société Géologique du Nord, 2nd series, 8, 75– 80. Vachard, D. 2014. Colaniella, wrongly named, welldistributed Late Permian nodosariate foraminifers. Permophiles, 60, 16– 24. Vachard, D. & Beckary, S. 1991. Algues et foraminifères bachkiriens des coal balls de la Mine Rosario (Truebano, Léon, Espagne). Revue de Paléobiologie, 10, 315–357. Vachard, D. & Ferrière, J. 1991. Une association à Yabeina (foraminifère fusulinoı̈de) dans le Midien (Permien supérieur) de la région de Whangaroa (Baie d’Orua, Nouvelle-Zélande). Revue de Micropaléontologie, 34, 201– 230. Vachard, D. & Krainer, K. 2001a. Smaller foraminifers of the Upper Carboniferous Auernig Group, Carnic Alps (Austria/Italy). Rivista Italiana di Paleontologia e Stratigrafia, 107, 147– 168. Vachard, D. & Krainer, K. 2001b. Smaller foraminifers, characteristic algae and pseudo-algae of the latest Carboniferous/Early Permian Rattendorf Group, Carnic Alps (Austria/Italy). Rivista Italiana de Paleontologia e Stratigrafia, 107, 169–195. Vachard, D. & Miconnet, P. 1990. Une association à Fusulinoı̈des du Murghabien supérieur au Monte Facito (Appennin méridional, Italie). Revue de Micropaléontologie, 32, 297 –318. Vachard, D. & Moix, P. 2013. Kubergandian (Roadian, Middle Permian) of the Lycian and Aladag Nappes (Southern Turkey). Geobios, 46, 335– 356, https:// doi.org/10.1016/j.geobios.2013.02.002 Vachard, D. & Montenat, C. 1981. Biostratigraphie, micropaléontologie et paléogéographie du Permien de la région de Tezak (Montagnes Centrales d’Afghanistan). Palaeontographica, B, 178, 1 –88. Vachard, D. & Razgallah, S. 1988. Importance phylogénétique d’un nouveau foraminifère endothyroı̈de Endoteba controversa n. gen. n. sp. (Permien du Jebel Tebaga, Tunisie). Geobios, 21, 805– 811. Vachard, D., Ott d’Estevou, P. & Briend, M. 1982. Un micro-organisme problématique du Miocène d’Espagne voisin des Umbellaceae paléozoı̈ques: Mitrigalia n. gen. Revue de Micropaléontologie, 25, 57–68. Vachard, D., Oviedo, A., Flores de Dios, A., Malpica, R., Brunner, P., Guerrero, M. & Buitrón, B.E. 1992. Barranca d’Olinalá: une coupe de référence pour le Permien du Mexique central; étude préliminaire. Annales de la Société Géologique du Nord, 2ème série, 2, 153–160. Vachard, D., Clift, P. & Decrouez, D. 1993a. Une association à Pseudodunbarula (Fusulinoı̈de) du Permien supérieur (Djoulfien) remaniée dans le Jurassique d’Argolide (Grèce). Revue de Paléobiologie, 12, 217–242. Vachard, D., Martini, R., Zaninetti, L. & Zambetakis-Lekkas, A. 1993b. Révision micropaléontologique (Foraminifères, Algues) du Permien inférieur (Sakmarien) et supérieur (Dorashamien) du Mont Beletsi (Attique, Grèce). Bolletino della Societá Paleontologica Italiana, 32, 89–112. Vachard, D., Martini, R., Rettori, R. & Zaninetti, L. 1994. Nouvelle classification des Foraminifères endothyroı̈des du Trias. Geobios, 27, 543–557. Vachard, D., Hauser, M., Martini, R., Zaninetti, L., Peters, T. & Matter, A. 2001a. New algae and microproblematica with algal affinity from the Permian of the Aseelah Unit in the Batain Plain, East Oman. (Nouvelles algues et formes affines microproblématiques du Permien de l’Unité d’Asselah) (Batain Plain, Est de l’Oman). Geobios, 34, 375–404. Vachard, D., Martini, R. & Zaninetti, L. 2001b. Earliest Artinskian (Early Permian) fusulinid reworked in the Triassic Lercara Formation (NW Sicily). Journal of Foraminiferal Research, 31, 33–47. Vachard, D., Hauser, M., Martini, R., Zaninetti, L., Matter, A. & Peters, T. 2002. Middle Permian (Midian/Capitanian) fusulinid assemblages from the Aseelah Unit (Batain Group) in the Batain Plain, East-Oman: their significance to Neotethys paleogeography. Journal of Foraminiferal Research, 32, 155–172. Vachard, D., Zambetakis-Lekkas, A., Skourtsos, E., Martini, R. & Zaninetti, L. 2003. Foraminifera, Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 PERMIAN SMALLER FORAMINIFERS algae and carbonate microproblematica from the late Wuchiapingian/Dzhulfian (Late Permian) of Peloponnesus (Greece). Rivista Italiana di Paleontologia i Stratigrafia, 109, 339– 358. Vachard, D., Gaillot, J., Vaslet, D. & Le Nindre, Y.M. 2005. Foraminifers and algae from the Khuff Formation (late Middle Permian–Early Triassic) of central Saudi Arabia. GeoArabia, 10, 137– 186. Vachard, D., Rettori, R., Angiolini, L. & Checconi, A. 2008. Glomomidiella gen. n. (Foraminifera, Miliolata, Neodiscidae): a new genus from the late Guadalupian-Lopingian of Hydra Island (Greece). Rivista Italiana di Paleontologia e Stratigrafia, 114, 349–361. Vachard, D., Pille, L. & Gaillot, J. 2010. Palaeozoic Foraminifera: systematics, palaeoecology and responses to the global changes. Revue de Micropaléontologie, 53, 209 –254. Vachard, D., Krainer, K. & Lucas, S. 2012. Pennsylvanian (Late Carboniferous) calcareous microfossils from Cedro Peak (New Mexico, USA); Part 1: Algae and Microproblematica. Annales de Paléontologie, 98, 225–252. Vachard, D., Krainer, K. & Lucas, S. 2013. Pennsylvanian (Late Carboniferous) calcareous microfossils from Cedro Peak (New Mexico, USA); Part 2: smaller foraminifers and fusulinids. Annales de Paléontologie, 99, 1– 42. Vachard, D., Krainer, K. & Lucas, S. 2015. Late Early Permian (late Leonardian; Kungurian) algae, microproblematica, and smaller foraminifers from the Yeso Group and San Andres Formation (New Mexico; USA). Palaeontologia Electronica, 18.1.21A, 1– 77, palaeo-electronica.org/content/2015/1160-kungurianof-new-mexico Vachard, D., Rettori, R., Altıner, D. & Gennari, V. In press. The Permian foraminiferal family Pseudovidalinidae and the genus Altineria emend. herein. Journal of Foraminiferal Research. Vdovenko, M.V., Rauzer-Chernousova, D.M., Reitlinger, E.A. & Sabirov, A.A. 1993. Guide for the Systematics of Paleozoic Smaller Foraminifers. Rossiiskaya Akademiya Nauk, Komissiya po Mikropaleontologii Nauka, Moscow [in Russian]. Vologdin, A.G. 1932. Archaeocyaths of Siberia. Issue 2: Fauna of the Cambrian Limestones of Altaya. State Scientific and Technical Geology Exploratory Publishing House, Moscow [in Russian]. Vuks, G.P. 1984. First discovery of Geinitzinita (foraminifer) in the Permian of Transcaucasia. Paleontologicheskii Zhurnal, 3, 131– 133 [in Russian]. Vuks, G.P. & Chediya, I.O. 1986. Foraminifers from the Lyudyansk Suite (southern Primorye). In: Correlations of Permian–Triassic Deposits of Eastern USSR. Academy of Sciences USSR, Far-Eastern Scientific Centre, Institute of Biology and Pedology, Vladivostok (Project IGCP No. 203, 82–88) [in Russian]. Wang, G.-L. 1966. On Colaniella and its two allied new genera. Acta Palaeontologica Sinica, 14, 206– 221 [in Chinese]. Wang, K.L. & Sun, X.F. 1973. Carboniferous and Permian Foraminifera of the Chinling Range and its geologic significance. Acta Geologica Sinica, 2, 137–178 [in Chinese with English abstract]. 251 Wang, Y. & Ueno, K. 2003. Latest Permian fusulinoideans and smaller foraminifers in the Meili section, Guangxi, S. China. Permophiles, 42, 19–21. Wang, Y., Ueno, K., Zhang, Y.C. & Cao, C.Q. 2010. The Changhsingian foraminiferal fauna of a Neotethyan seamount: the Gyanyima Limestone along the Yarlung–Zangbo Suture in southern Tibet, China. Geological Journal, 45, 308– 318. Wang, Y.J. 1974. A Handbook of the Stratigraphy and Paleontology in Southwest China. Science Press, Beijing [in Chinese]. Warthin, A.S., JR. 1930. Micropaleontology of the Wetumka, Wewoka, and Holdenville formations. Bulletin of the Oklahoma Geological Survey, 53, 1– 95. Wedekind, P.R. 1937. Einführung in die Grundlagen der historischen Geologie. Band II. Mikrobiostratigraphie die Korallen und Foraminiferenzeit. Ferdinand Enke, Stuttgart. Wignall, P.B. & Hallam, A. 1996. Facies change and the end-Permian mass extinction in S.E. Sichuan, China. Palaios, 11, 587–596. Wignall, P.B., Védrine, S. et al. 2012. Capitanian (Middle Permian) mass extinction and recovery in western Tethys: a fossil, facies, and d13C study from Hungary and Hydra Island (Greece). Palaios, 27, 78–89. Wilde, G.L. 2006. Pennsylvanian-Permian fusulinaceans of the Big Hatchet Mountains, New Mexico. New Mexico Museum Natural History and Science Bulletin, 38, 1 –311. Wood, G.D., Groves, J.R., Wahlman, G.P., Brenckle, P.L. & Alemán, A.M. 2002. The paleogeographic and biostratigraphic significance of fusulinacean and smaller foraminifers, and palynomorphs from the Copacabana Formation (Pennsylvanian-Permian), Madre de Dı́os basin, Peru. In: Hills, L.V., Henderson, C.M. & Bamber, E.W. (eds) Carboniferous and Permian of the World: XIV ICCP Proceedings. Canadian Society of Petroleum Geologists, Memoirs, 19, 630– 664. Woszczynska, S. 1987. Foaraminifera and ostracods from the carbonate sediments of the Polish Zechstein. Acta Palaeontologica Polonica, 32, 115–205. Yanagida, J., Sakagami, S. et al. 1988. Biostratigraphic study of Paleozoic and Mesozoic groups in central and northern Thailand, an interim report. Geological Survey Division, Department of Mineral Resources, Bangkok, 1 –47. Zaninetti, L. & Jenny-Deshusses, C. 1985. Les Paraglobivalvulines (foraminifères) dans le Permien supérieur téthysien; répartition géographique et description de Paraglobivalvulinoides n. gen. Revue de Paléobiologie, 4, 343–346. Zaninetti, L., Altıner, D. & Çatal, E. 1981. Foraminifères et biostratigraphie dans le Permien supérieur du Taurus oriental, Turquie. Notes du Laboratoire de Paléontologie de l’Université de Genève, 7, 1– 38. Zhang, M. & Gu, S. 2015. Latest Permian deep-water foraminifers from Daxiakou, Hubei, South China. Journal of Paleontology, 89, 448–464, https://doi.org/10. 1017/jpa.2015.19 Zhang, Z., Wang, Y. & Zheng, Q.-F. 2015. Middle Permian smaller foraminifers from the Maokou Downloaded from http://sp.lyellcollection.org/ by guest on March 10, 2018 252 D. VACHARD Formation at theTieqiao section, Guangxi, South China. Palaeoworld, 24, 263– 276. Zhang, Z.H. & Hong, Z.Y. 2004. Smaller foraminiferal fauna from the Changhsing Formation of Datian, Fujian. Acta Micropaleontologica Sinica, 21, 64– 84 [in Chinese with English abstract]. Zhao, J.-K., Sheng, J.-Z., Yao, Z.-Q., Liang, X.-L., Chen, C.-Z., Lin, R. & Liao, Z.-T. 1981. The Changhsingian and Permian–Triassic boundary of SouthChina. Bulletin Nanjing Institute Geology and Paleontology, Academia Sinica, 2, 58– 69 [in Chinese with English abstract]. Zheng, H. 1986. The smaller foraminifers faunas in Qixia stage (Early Permian) of Daxiakou, Xingshan County, Hubei Province. Earth Science, Journal of Wuhan College of Geology, 11, 489– 497 [in Chinese]. Zolotova, V.P. & Baryshnikov, V.V. 1980. Foraminifers of the Kungurian Stage in the stratotype localities. In: Rauzer-Chernousova, D.M. & Chuvashov, B.I. (eds) Biostratigraphy of the Artinskian and Kungurian Stages of the Urals. Akademiya Nauk SSSR, Uralskii Nauchnyi Tsentr, Sverdlovsk, 72–109 [in Russian].

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Permian smaller foraminifers: taxonomy, biostratigraphy and biogeography

Permian smaller foraminifers: taxonomy, biostratigraphy and biogeography

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