2.2 Morphology and Taxonomy Of Palaeozoic Larger Benthic Foraminifera

ORDER PARATHURAMMINIDA MIKHALEVICH 1980

In this order all forms are unilocular, large globular or tubular, and occurred both as free and as attached forms. The wall is thin, calcareous microgranular, simple to bilamellar with an inner hyaline pseudofibrous layer. Apertures are terminal at the top of a hollow neck. Silurian to Permian.

Superfamily PARATHURAMMINOIDEA Bykova, 1955

In this superfamily all forms are unilocular, globular or tubular, and occurred both as free and as attached forms. The wall structure is simple calcareous microgranular or with granulo-fibrous layers (see Fig. 2.2). They range from Early Silurian to Permian.

Family Archaesphaeridae Malakhova, 1966

The representatives of this family mainly have one or more globular to elongate chamber(s), with a test with no apparent aperture. They range from Late Silurian to Early Permian.

• Archaesphaera Suleymanov, 1945 (Type species: Archaesphaera minima Suleymanov, 1945). The test is globular and smooth. Devonian to Carboniferous (Tournaisian), (Fig. 2.2).

• Diplosphaerina Derville, 1952 (Type species: Diplosphaera inaequalis Derville, 1931). The two-chambered test, has a small proloculus enveloped within a larger chamber. The wall is dark, granular, nonperforated and single layered. Middle Devonian to Carboniferous (Tournaisian), (Plate 2.1, figs 15, 16; Plate 2.2, Fig. 10; Fig. 2.3).

Family Parathuramminidae Bykova, 1955

The representatives of the Parathuramminidae include globular, or irregular forms with a simple calcareous agglutinated wall and multiple apertures at the end of a tubular neck. They range from Silurian to Carboniferous (Mississippian) and they are the ancestral group of all the fusulinids.

• Parathurammina Suleymanov, 1945 (Type species: Parathurammina dagmarae Suleymanov, 1945). The test is globular, with apertures at the ends of numerous tubular protuberances. Late Silurian to Carboniferous (Tournaisian) (Plate 2.2, fig.19).

Family Chrysothuramminidae Loeblich and Tappan, 1988

Members of this family have a globular to irregular test in outline, with the aperture at the end of tubular projections. They range from Middle Devonian to Carboniferous (Tournaisian).

• Chrysothurammina Neumann, Pozaryska and Vachard, 1975. (Type species: Chrysothurammina tenuis Neumann, Pozaryska and Vachard, 1975). The test is composed of a single subspherical chamber with an aperture on a neck-like protrusion. Middle Devonian to Carboniferous (Tournaisian) (Plate 2.3, Fig. 1B).

Family Ivanovellidae Chuvashov and Yuferev, 1984

The representatives of this family are spherical with a thick calcareous wall with two layers, a thin dark compact granular inner layer and a thick grey radially fibrous outer layer. This family ranges from Late Silurian to Early Carboniferous (Tournaisian).

• Elenella Pronina, 1969 (Type species: Neoarchaesphaera (Elenella) multispinosa Pronina, 1969). The outer surface has tiny projections. There is no aperture. Late Silurian to Carboniferous (Tournaisian) (Plate 2.3, Fig. 2B).

Family Marginaridae Loeblich and Tappan, 1986

The members of this family are globular in shape, with a wall made of three layers, the inner and outer ones are dark and the median one light grey, with canals running through it. These canals open into the chamber cavity and make conical projections outside the wall. This family ranges from the Middle to Late Devonian.

• Marginara Pertova, 1984 (Type species: Parathurammina tamarae Petrova, 1981). The test is spherical, with some projections from the surface with canals opening through the projections. Middle Devonian.

Family Uralinellidae Chuvashov, Yuferev and Zadorozhnyy, 1984

The Uralinellidae are small, unilocular forms with a three-layered wall, the median layer is thick and clear and the surrounding layers dark. A multiple apertural neck is projected from the test. They range from Middle Devonian (Givetian) to Early Carboniferous (Visean).

• Sogdianina Saltovskaya, 1973 (Type species: Sogdianina angulata Saltovskaya, 1973). Similar to Elenella, but with a thicker wall and a very prominent apertural necks. Carboniferous (Visean) (Plate 2.3, Fig. 3).

• Uralinella Bykova, 1952 (Type species: Uralinella bicamerata Bykova, 1952). The test is subglobular with neck-like tubular projections. Middle Devonian to Carboniferous (Givetian to Tournaisian).

Family Auroriidae Loeblich and Tappan, 1986

The Auroriidae have an ovate test with a two-layered wall, a thin inner finely porous dark layer and a thick outer canaliculated dark layer. They range from Middle to Late Devonian.

• Auroria Poyarkov, 1969 (Type species: Auroria singularis Poyarkov, 1969). The test is irregularly globular. Middle to Late Devonian (Plate 2.3, Fig. 4B).

Family Usloniidae Miklukho-Maklay, 1963

Members of this family have a globular test with no distinct aperture. They range from Middle Devonian to Early Carboniferous.

• Bisphaera Birina, 1948 (Type species: Bisphaera malevkensis Birina, 1948). The test is subglobular, with a constriction. Middle Devonian to Carboniferous (Givetian to Tournaisian).

• Parphia Miklukho-Maklay, 1965 (Type species: Cribrosphaeroides (Parphia) robusta Miklukho-Maklay, 1965). The test is spherical to ovate with a wall that is dark and evenly porous. Late Devonian (Plate 2.3, Fig. 5).

Family Eovolutinidae Loeblich and Tappan, 1986

The representatives of this family have a globular test with a micro-granular wall and a distinct single aperture. They range from Late Silurian to Late Devonian.

• Eovolutina Antropov, 1950 (Type species: Eovolutina elementa Antropov, 1950). The test is tiny, with the proloculus completely surrounded by the second chamber. Late Silurian to Late Devonian (Plate 2.3, fig.6).

Family Tuberitinidae Miklukho-Maklay, 1958

Members of the Tuberitinidae have a simple attached test, made of one or more globular chamber with calcareous microgranular walls (see Fig. 2.2, 2.3). They occur between the Silurian and the Permian.

• Draffania Cummings, 1957 (Type species: Draffania biloba Cummings, 1957). The test is flask-shaped with an aperture at the end of an elongate neck. Carboniferous (Tournaisian to Visean) (Plate 2.1, figs 6, 10, 13, 17, 18; Plate 2.2, Fig. 16; Plate 2.3, Fig. 5B).

• Eotuberitina Miklukho-Maklay, 1958 (Type species: Eotuberitina reitlingerae Miklukho-Maklay, 1958). The test is hemispherical with a basal disc, and with a wall that is microgranular and finely perforate. Middle Devonian to Permian (Lopingian) (Plate 2.2, figs 6, 8, 9; Fig. 2.3).

• Paratuberitina Miklukho-Maklay, 1957 (Type species: Tuberitina collosa Reytlinger, 1950). The test is highly hemispherical with a basal disk, with a microgranular, coarsely perforate wall. Carboniferous (Visean to Moscovian) (Plate 2.2, Fig. 17; Fig. 2.3).

• Tubeporina Pronina, 1960 (Type species: Tubeporina gloriosa Pronina, 1960). The test is hemispherical with a basal disc, and with a coarsely perforate three-layered wall, a hyaline layer between two microgranular layers. Middle Devonian to Carboniferous (Visean) (Plate 2.2, fig.7; Fig. 2.3).

• Tuberitina Galloway and Harlton, 1928 (Type species: Tuberitina bulbacea Galloway and Harlton, 1928). Shows bulbous chambers in an arcuate or straight series. The wall is thick and finely perforate. Late Carboniferous to Late Permian (Bashkirian to Lopingian) (Plate 2.2, Fig. 11; Plate 2.3, Fig. 8).

ORDER MORAVAMMINIDA POKORNY, 1951

Members of this order have an attached tubular test, irregularly septate. They may also have closer affinity to algae than foraminifera (Vachard, 1994; Vachard and Cozar, 2010). Late Silurian to Early Carboniferous (Tournaisian).

Superfamily MORAVAMMINOIDEA Pokorny, 1951

This superfamily (Fig. 2.2) had simple free or attached tests, consisting of a proloculus and a rectilinear second chamber that was subseptate (i.e. with partial septa). They had simple walls and evolved from the Parathuramminoidea in the Silurian, but died out in the Early Carboniferous (Tournaisian).

Family Moravamminidae Pokorny, 1951

In this possible foraminiferal family, the test is attached and irregularly septate with an enrolled rectilinear part. They occurred in the Middle to Late Devonian.

ORDER ARCHAEDISCIDA POJARKOV AND SKVORTSOV 1979

The members of this order have free discoidal, lenticular to conical involute or rarely evolute test. It is composed of a proloculus followed directly by a planispiral trochospirally, or streptospirally tubular second chamber. The wall is calcareous, bilayered formed of an inner dark microgranular layer and a hyaline and radially fibrous outer layer. They range from the Early Carboniferous to the Late Permian (Visean to Lopingian).

Superfamily ARCHAEDISCOIDEA Cushman, 1928

This superfamily is characterized by having a small, free test, consisting of a proloculus followed by an enrolled chamber. The wall is formed of one or more layers. They range from the Early Carboniferous to the Late Carboniferous (Visean to Moscovian), or unilayered and pseudofibrous.

Family Archaediscidae Cushman, 1928

The Archaediscidae (Fig. 2.2) have a discoidal test, composed of a proloculus followed by a streptospirally enrolled second chamber. The wall is formed of a dark inner granular layer and a clear radially fibrous or granulo-fibrous outer layer. They range from the Early Carboniferous (Visean) to the Late Carboniferous (Moscovian).

• Archaediscus Brady, 1873 (Type species: Archaediscus karreri Brady, 1873). The test is free, streptospiral, with an undivided tubular second chamber and thickened wall. Carboniferous (Visean to Moscovian) (Plate 2.3, Fig. 6; Plate 2.4, Fig. 17C; Fig. 2.5; Plate 2.5, Fig. 18B; Plate 2.6, figs 3-16; Plate 2.7, Fig. 6).

• Asteroachaediscus Miklukho-Maklay, 1956 (Type species: Archaediscus baschkiricus Krestovnikov and Teodorovitch, 1936). The test is small, lenticular with sigmoidal coiling, stellate and with occluded chamber cavity. Carboniferous (Visean to Moscovian) (Plate 2.8, Fig. 12A; Plate 2.7, Fig. 3).

• Glomodiscus Malakhova, 1973 (Type species: Glomodiscus biarmicus Malakhova, 1973). The test is discoidal, with a globular proloculus followed by an involutely coiled, undivided tubular second chamber. Carboniferous (Visean).

• Hemiarchaediscus Miklukho-Maklay, 1957 (Type species: Hemiarchaediscus planus Miklukho-Maklay, 1957). The test is small, discoidal with a glomospiral early whorl, later becoming planispiral; chamber lumen are always open. Carboniferous (late Visean to early Serpukhovian) (Plate 2.7, figs 1, 2).

• Permodiscus Dutkevich, 1948 (Type species: Permodiscus vetustus Dutkevich, in Chernysheva, 1948). The test is planispirally enrolled with a tubular undivided second chamber, later whorls have small nodosities partially filling the chamber lumen. Carboniferous (Visean to Bashkirian).

• Planoarchaediscus Miklukho-Maklay, 1957 (Type species: Archaediscus spirillinoides Rauzer-Chernousova, 1948). The test is small, discoidal with a glomospiral early stage, followed by a planispiral adult. Chamber lumen remain open. Carboniferous (Visean to Serpukhovian) (Plate 2.7, Fig. 5).

• Propermodiscus Miklukho-Maklay, 1953 (Type species: Hemigordius ulmeri Mikhaylov, 1939). The test is small, lenticular with glomospiral early whorls and a planispiral, involute adult. Carboniferous (Visean) (Plate 2.7, Fig. 4).

Superfamily LASIODISCOIDEA Reitlinger in Vdovenko et al., 1993

The members of this superfamily have a bilayered wall with finely granular dark inner layer and a radially fibrous outer layer. Tubercles of pseudofibrous fillings or pillars fill the umbilical region. The aperture is simple and terminal, but additional supplementary apertures may occur along the spiral sutures of the successive whorls. The range is between the Early Carboniferous (Visean) and the Permian.

Family Lasiodiscidae Reytlinger, 1956

In this family, the test is discoidal to conical, composed of a proloculus and an undivided enrolled tubular second chamber. The radial fibrous outer layer is mainly concentrated in the umbilical region, where it may form pillars, or a series of tubercles on the surface of one side of the test. The range is between the Early Carboniferous (Visean) and the Permian.

• Eolasiodiscus Reytlinger, 1956 (Type species: Eolasiodiscus donbassicus Reytlinger, 1956). The test is discoidal and concavo-convex, with the umbilical filling on the concave side. Middle to Late Carboniferous.

• Glomotrocholina Nikitina, 1977 (Type species: Glomotrocholina pojarkovi Nikitina, 1977). The test is conical. Second chamber is initially streptospirally enrolled, later forming an irregular trochospiral filled with shell material. Late Permian (Lopingian).

• Howchinia Cushman, 1927 (Type species: Patellina bradyana Howchin, 1888). The test is conical with septal bridges. The umbilical region is filled with fibrous calcite forming pillars (Fig. 2.2). Carboniferous (Visean to Moscovian) (Fig. 2.2; Fig. 2.22; Plate 2.1, Fig. 11A; Plate 2.5, Fig. 1; Plate 2.10, figs 1-7, 13-14; Plate 2.14, figs 1-7, 13-14).

• Lasiodiscus Reichel, 1946 (Type species: Lasiodiscus granifer Reichel, 1946). The test is planispiral with tubular extensions. Middle Carboniferous to Late Permian (Fig. 2.2).

• Lasiotrochus Reichel, 1946 (Type species: Lasiotrochus tatoiensis Reichel, 1946). The test is conical with tubular chamberlets curved towards the proloculus, with distinct pillars filling the centre of the test. Permian (Cisuralian to Lopingian).

• Monotaxinoides Brazhnikova and Yartseva, 1956 (Type species: Monotaxinoides transitorius Brazhnikova and Yartseva, 1956). The test is low conical to nearly planispiral, with a spherical proloculus followed by a semi-cylindrical second chamber. Carboniferous.

ORDER EARLANDIIDA SABIROV IN VDOVENKO ET AL., 1993

EMEND. VACHARD ET AL., 2010

This order had simple free or attached tests, consisting of a proloculus and a rectilinear second chamber. Aperture terminal, simple. Late Silurian to Early Triassic.

Superfamily EARLANDIOIDEA Cummings, 1955

This superfamily is characterised by having a free, non-septate test with a globular first chamber and a straight tubular second chamber. Members range from the Late Silurian to Early Triassic.

Family Earlandiidae Cummings, 1955

The Earlandiidae have a single free chamber (Fig. 2.2) and range from the Late Silurian to Early Triassic.

• Aeolisaccus Elliott, 1958 (Type species: Aeolisaccus dunningtoni Elliott, 1958). Free test, elongate and slightly tapering. Late Permian (Lopingian).

• Earlandia Plummer, 1930 (Type species: Earlandia perparva Plummer, 1930). Free, elongate test, composed of a globular proloculus followed by an undivided straight tubular chamber. The wall is calcareous microgranular. Late Silurian to Early Triassic (Plate 2.1, figs 1-5; Plate 2.3, Fig. 1A; Plate 3.2, fig.3).

• Gigasbia Strank, 1983 (Type species: Gigasbia gigas Strank, 1983). The test is free and elongate, consisting of a globular proloculus and an undivided tubular chamber. Early Carboniferous (Tournaisian).

Family Endotebidae Vachard, Martini, Rettori and Zaninetti, 1994

The test is free, planispiral, in early stages, but later uniserial to biserial. The walls are calcareous, grey, thick, calcareous agglutinated, and the apertures are simple. Late Permian to Triassic.

• Endoteba Vachard and Razgallah, 1988 emend. Vachard et al. 1994 (Type species: Endoteba controversa Vachard et al., 1994). Axial view compressed. Late Permian to Late Triassic (Late Kungurian to Rhaetian) (Fig. 3.5).

Family Pseudoammodiscidae Conil and Lys, 1970

The Pseudoammodiscidae are small, with a simple aperture and globular proloculus followed by a planispiral, trochospiral or streptospirally coiled, undivided second chamber. They occur from the Devonian to the Permian.

• Brunsia Mikhaylov, 1935 (Type species: Spirillina irregularis von Möller, 1879). The proloculus is followed by an early streptospiral coil, and later a planispirally enrolled tubular chamber. Devonian to Carboniferous (Tournaisian) (Plate 2.3, Fig. 9).

• Brunsiella Reytlinger, 1950 (Type species: Glomospira ammodiscoidea Rauzer-Chernousova, 1938). The globular proloculus is followed by an undivided second chamber. Early whorls are streptospiral, later ones planispiral and evolute. Carboniferous (Visean) to Early Permian (Plate 2.2, Fig. 15B; Plate 2.3, figs 9,11,12; Plate 2.11, figs 7-9).

Family Pseudolituotubidae Conil and Longerstaey, 1980

The Pseudolituotubidae have attached tests, consisting of a single enrolled chamber with a simple terminal aperture and a compound calcareous microgranular wall. They range from the Early Carboniferous (Visean) to the Late Carboniferous (Moscovian).

• Pseudolituotuba Vdovenko, 1971 (Type species: Lituotuba? gravata Conil and Lys, 1965). The proloculus is followed by an undivided streptospirally coiled tube. Carboniferous (Visean to Moscovian).

Superfamily CALIGELLOIDEA Reytlinger 1959

This superfamily (Fig. 2.2) had simple free or attached tests, consisting of a proloculus and a rectilinear second chamber that was subseptate (i.e. with partial septa). They had simple walls and evolved from the Parathuramminoidea in the Late Silurian, but died out in the Early Carboniferous (Tournaisian).

Family Caligellidae Reytlinger, 1959

In this Family, the test is attached and subseptate. They ranged from the Late Silurian to the Early Carboniferous (Tournaisian).

• Paracaligella Lipina, 1955 (Type species: Paracaligella antropovi Lipina, 1955). The test is composed of a sub-spherical proloculus followed by an irregular tubular chamber partially divided by incipient septa. Devonian to Carboniferous (Late Tournaisian) (see Fig. 2.2).

Family Paratikhinellidae Loeblich and Tappan, 1984

This family was free with a elongate, sub-septate test and with a simple microgranular wall. They ranged from Middle Devonian to Carboniferous (Tournaisian).

• Paratikhinella Reytlinger, 1954 (Type species: Tikhinella cannula Bykova, 1952). The test is elongate and composed of a subglobular proloculus followed by a cylindrical tubular chamber with incipient septation. Late Devonian to Early Carboniferous (Plate 2.4, Fig. 5).

• Saccaminopsis Sollas, 1921 (Type species: Saccammina carteri Brady, 1871). The test is free, uniserial with ovate chambers, a thin wall and a terminal aperture. Late Devonian to Carboniferous (Tournaisian). (Plate 2.1, Fig. 11B; Plate 2.2, figs 12-15).

Superfamily PTYCHOCLADIOIDEA Elias, 1950

These were attached forms with no distinct apertures. Their walls were microgranular, banded with transverse tubuli. They range from Late Devonian to Late Carboniferous.

Family Ptychocladiidae Elias, 1950

Here the test is uniserial and attached. They ranged from the Late Devonian to Late Carboniferous (Pennsylvanian).

• Ptychocladia Ulrich and Bassler, 1904 (Type species: Ptychocladia agellus Ulrich and Bassler, 1904). The wall is calcareous microgranular, with two layers with no distinct aperture. Carboniferous (Late Pennsylvanian) (see Fig. 2.2).

ORDER PALAEOTEXTULARIIDA HOHENEGGER AND PILLER, 1975

Members of this order have biserial or uniserial tests, with a microgranular calcareous wall, commonly with an inner radial fibrous layer and a finely granular outer layer. The aperture is generally single but may be multiple in later stages. Their range is from Late Devonian to Permian.

Superfamily PALAEOTEXTULARIOIDEA Galloway, 1933

This superfamily has a test that is biserial or uniserial, with a single or two-layered microgranular calcareous wall. The aperture is generally single but may be multiple in later stages. Their range is from Late Devonian to Permian.

Family Semitextulariidae Pokorny, 1956

Here the test is biserial and flattened, but often becoming monoserial with broad chambers, and is fully septate (see Fig. 2.2). They range from the Devonian to the Carboniferous (Pennsylvanian).

• Koskinobigenerina Eickhoff, 1968 (Type species: Koskinobigenerina breviseptata Eickhoff, 1968). The test is elongate, biserial in the early stage, later becoming uniserial. Early Carboniferous to Late Carboniferous (Visean to Gzhelian).

• Koskinotextularia Eickhoff, 1968 (Type species: Koskinotextularia cribriformis Eickhoff, 1968). The test is elongate, biserial throughout with a single layered wall. The aperture is single in the early stage but later becoming cribrate. Early Carboniferous to Late Carboniferous (Visean to Gzhelian) (Plate 2.4, Fig. 6).

Family Palaeotextulariidae Galloway, 1933

This family (see Fig. 2.2 and 2.3) has biserial to uniserial genera that closely resembles the Textulariidae, but they have a dark granular calcareous outer wall and an inner clear to yellowish “fibrous” layer with stacks of granules perpendicular to the surface. Cummings (1956) demonstrated that these forms evolved from simple agglutinated forms in the Devonian (see Fig. 2.4). Early Carboniferous (Tournaisian) to Permian (Lopingian).

• Climacammina Brady, 1873 (Type species: Textularia antiqua Brady, in Young and Armstrong, 1871). The test has a biserial early stage followed by a uniserial stage. The aperture in the early stage is at the base of the last chamber, later becoming areal, multiple and cribrate. Carboniferous to Permian (Visean to Early Lopingian) (Plate 2.4, Fig. 7; Plate 2.5, figs 7, 9-10).

• Cribrogenerina Schubert 1908 (Type species: Bigenerina sumatrana Volz, 1904). The test has a short early biserial stage followed by a uniserial stage. The aperture is a slit in the early stage, later becoming areal, multiple and cribrate in the uniserial stage. Carboniferous to Permian (latest Gzhelian to Lopingian) (see Fig. 2.4).

• Cribrostomum von Möller, 1879 (Type species: Cribrostomum textulariforme von MöIler, 1879). The test is biserial throughout. The aperture in the early part is basal, but in the last two whorls becomes areal, multiple and cribrate. Carboniferous (late Tournaisian to Gzhelian) (Plate 2.4, figs 1, 5, 13, 15; see Fig. 2.2; 2.4).

• Deckerella Cushman and Waters, 1928 (Type species: Deckerella clavata Cushman and Waters, 1928). The test is biserial in the early stage, later becoming uniserial and rectilinear. The aperture is a low slit in the early stage, later becoming terminal with two parallel slits separated by a partition. Carboniferous to Permian (late Visean to early Lopingian) (Plate 2.4, Fig. 8; see Fig. 2.4).

• Deckerellina Reytlinger, 1950 (Type species: Deckerellina istiensis Reytlinger, 1950). The test is biserial throughout. The aperture, in the early stage, is a low slit, but the adult aperture is two parallel slits. Carboniferous (Visean to Moscovian) (Plate 2.4, Fig. 11, 16; see Fig. 2.4).

• Monogenerina Spandel, 1901 (Type species: Type species: Monogenerina atava Spandel, 1901). The test is biserial in the early stage, later becoming uniserial. The aperture extends throughout the test. Permian (Cisuralian to Lopingian) (see Fig. 2.4).

• Palaeobigenerina Galloway, 1933 (Type species: Bigenerina geyeri Schellwien, 1898). The test is biserial in the early stage, but later uniserial, with a slit opening in the biserial stage, followed by a single rounded aperture. Carboniferous to Permian (late Serpukhovian to early Lopingian) (see Fig. 2.4).

• Palaeotextularia Schubert, 1921 (Type species: Palaeotextularia schellwieni Galloway and Ryniker, 1930). The test is biserial throughout. The wall is thick, and the aperture is a low slit opening at the base of the final chamber. Carboniferous to Permian (Tournaisian to Cisuralian) (Fig. 2.4; Plate 2.4, figs 2-4, 9-10, 12, 14, 17-18; Plate 2.5, Fig. 6).

Family Biseriamminidae Chernysheva, 1941

In this family (Fig. 2.2), the test is biserial becoming planar in later stages, with a microgranular wall with one or more layers. They range from the Carboniferous (Tournaisian) to Permian (Lopingian).

• Biseriammina Cherhysheva, 1941 (Type species: Biseriammina uralica Cherhysheva, 1941). The test is planispirally enrolled. Early Carboniferous (Tournaisian).

• Biseriella Mamet, 1974 (Type species: Globivalvulina parva Chernysheva, 1948). The early stage is tightly coiled, while later it has an open helicoid spire. Late Carboniferous (Bashkirian to Moscovian) (Plate 2.3, Fig. 10).

• Dagmarita Reytlinger, 1965 (Type species: Dagmarita chanakchiensis Reyltinger, 1965). The test is flattened with spine-like projections at the outer corners of the angular chambers. Late Permian (Lopingian).

• Globispiroplectammina Vachard, 1977 (Type species: Globispiroplectammina mameti Vachard, 1977). The test is biserial with the early stage enrolled, later becoming uncoiled. Carboniferous (Visean).

• Globivalvulina Schubert, 1921 (Type species: Valvulina bulloides Brady, 1876). The test is planispirally coiled. Middle Carboniferous (Bashkirian) to Late Permian (Wuchiapingian).

• Lipinella Malakhova, 1975 (Type species: Lipinella notata Malakhova, 1975). The test is planispirally enrolled following a globular proloculus. Middle Carboniferous to Late Permian (Bashkirian to Lopingian).

• Louisettita Altiner and Brönnimann, 1980 (Type species: Louisettita elegantissima Altiner and Brönnimann, 1980). The test is trochospirally enrolled in the early stage. In the biserial stage, chambers are subdivided by vertical partitions perpendicular to the septa and have spine-like projections. Late Permian (Wuchiapingian).

• Paraglobivalvulina Reytlinger, 1965 (Type species: Paraglobivalvulina mira Reytlinger, 1965). The test is spherical with the biserial chambers enrolled trochospirally. Late Permian (Wuchiapingian).

• Paradagmarita Lys, 1978 (Type species: Paradagmarita monodi Lys, 1978). The test is small with the early stage completely enrolled, but the later stage is uncoiled. Late Permian (Wuchiapingian).

• Paraglobivalvulinoides Zaninetti and Jenny-Deshusses, 1985 (Type species: Paraglobivalvulina? septulifera Zaninetti and Jenny-Deshusses, 1981). The test is globular with biserially, enrolled, strongly enveloping chambers. The wall is calcareous, microgranular, a single layer. The aperture is rimmed with a well-developed tongue that bends inward. Late Permian (Wuchiapingian).

ORDER TETRATAXIDA MIKHALEVICH 1981

Members of this order have a conical, trochospiral test with an evolute spiral side and involute umbilical side. The wall is microgranular calcareous with one or two distinct layers. Early Carboniferous to Late Permian (Tournaisian to Wuchiapingian).

Superfamily TETRATAXOIDEA Galloway, 1933

The Tetrataxoidea have a conical to spreading test with secondary internal partitions. The walls are microgranular with one or two layers. They range from the Early Carboniferous to Late Permian (Tournaisian to Wuchiapingian).

Family Tetrataxidae Galloway, 1933

The test of the Tetrataxidae is conical leaving an open central umbilicus where the chambers are partially overlapping (Fig. 2.2). The wall is calcareous, microgranular, and in two layers. They occur from the Early Carboniferous to the Late Carboniferous (Tournaisian to Moscovian).

• Globotetrataxis Brazhnikova, 1983 (Type species: Tetrataxis (Globotetrataxis) elegantula Brazhnikova, in Brazhnikova and Vdovenko, 1983). The test enlarges gradually with a last large hemispherical umbilical chamber that forms a convex base. Early Carboniferous (late Visean).

• Polytaxis Cushman and Waters, 1928 (Type species: Polytaxis laheei Cushman and Waters, 1928). The test is low conical. Later chambers form many whorls and result in a spreading test. Late Carboniferous (Moscovian).

• Tetrataxis Ehrenberg, 1854 (Type species: Tetrataxis conica Ehrenberg, 1854). The test is circular in plan with chambers strongly overlapping on the umbilical side. The outer layer is dark and microgranular and the inner layer is light and fibrous. Carboniferous (late Tournaisian to Visean) (Fig. 2.5; Plate 2.2, figs 1, 2, 5; Plate 2.4, Fig. 17B; Plate 2.7, figs 8-18).

Family Pseudotaxidae Mamet, 1974

In this family, the wall is calcareous, microgranular and single layered. They range from Early Carboniferous to Late Carboniferous (Late Tournaisian to Early Bashkirian).

• Pseudotaxis Mamet, 1974 (Type species: Tetrataxis eominima Rauzer-Chernousova, 1948). The test is conical with the proloculus followed by irregular trochospiral coil. Early Carboniferous to Late Carboniferous (late Tournaisian to Early Bashkirian).

• Vissariotaxis Mamet, 1970 (Type species: Monotaxis exilis Vissarionova, 1948). The test is conical with trochospirally coiled chambers and a wide aperture. Carboniferous (early Bashkirian) (Plate 2.10, figs 8-12).

Family Valvulinellidae Loeblich and Tappan, 1984

Here the test is conical with subdivided chambers and a single layer microgranular wall. The range is from the Early to Late Carboniferous (Visean to early Bashkirian).

• Valvulinella Schubert, 1908 (Type species: Valvulina youngi Brady, 1876). Chambers are subdivided by numerous vertical pillars and one or two horizontal ones. Carboniferous (Visean to Early Bashkirian). (Plate 2.2, figs 3-4; Plate 2.9, fig. 4; Fig. 2.22C).

Family Abadehellidae Loeblich and Tappan, 1984

Representatives of this family differ from the Valvulinellidae in having two double-layered walls instead of a single layered wall. They occur in the Late Permian (Wuchiapingian).

• Abadehella Okimura and Ishii, 1975 (Type species: Abadehella tarazi Okimura and Ishi, in Okimura et al., 1975). The test is conical, with numerous whorls. Chambers are subdivided by regularly spaced pillars. Late Permian (Wuchiapingian) (Plate 2.3, Fig. 12).

ORDER TOURNAYELLIDA HOHENEGGER AND PILLER 1973

The representatives of this order have walls that range from homogeneously calcareous microgranular to those that are differentiated into two or more layers in the most advanced forms. Late Devonian to Carboniferous.

Superfamily TOURNAYELLOIDEA Dain, 1953

In this superfamily, the proloculus is followed by a planispiral or streptospiral non-septate to fully septate test. The wall is made of microgranular calcite evolving into forms with a thick outer layer, called a tectum, and an inner granulo-fibrous layer. Late Devonian to Carboniferous (Frasnian to Early Bashkirian).

Family Tournayellidae Glaessner, 1945

The Tournayellidae are small, with early streptospiral or trochospiral coiling, tending to become planispiral in the adult. The aperture is simple and open at the end of a tubular chamber. The Tournayellidae evolved in the Devonian from simple planispiral, non-septate forms (e.g. Eotournayella, see Fig. 2.6) with an initial proloculus and an aperture at the end of an uncoiled tube, to forms with a compound wall and rudimentary septa in the Late Tournaisian (e.g. Tournayella Dain, 1953, see Fig. 2.6; Carbonella Dain, 1953, see Fig. 2.6; Plate 2.9, Fig. 14) to almost complete septa in the late Visean (Mstinia Dain, Plate 2.9, figs 8, 10). The streptospiral tubular forms (Glomospiranella Lipina, 1951, see Fig. 2.6) gradually also developed rudimentary septa in the Late Tournaisian (e.g. Brunsiina Lipina, 1953, see Fig. 2.6). The Tournayellidae range from the Devonian to the Carboniferous.

• Bogushella Conil and Lys, 1977 (Type species: Mistinia ziganensis Grozdilova and Lebedeva, 1960). The test is streptospirally enrolled in the early stage, later becoming rectilinear with no true septa. The aperture is cribrate. Early Carboniferous (Visean).

• Brunsiina Lipina, 1953 (Type species: Brunsiina uralica Lipina, 1953). The test is streptospirally enrolled in early stage, later becoming planispiral, without rudimentary constrictions. Late Devonian to Early Carboniferous (Fammenian to Visean) (Fig. 2.6).

• Carbonella Dain, 1953 (Type species Carbonella spectabilis Dain, 1953). The test is planispiral with septa that are well developed only in the final whorl. Early Carboniferous (Late Tournaisian to Early Visean) (Fig. 2.6; Plate 2.9, Fig. 14).

• Conilites Vdovenko, 1970 (Type species: Ammobaculites? Dinantii Conil and Lys, 1964). The test is planispirally enrolled and undivided in the early undivided part, but the later part is uncoiled rectilinear with distinct septa. The aperture is multiple and cribrate. Late Carboniferous (Late Tournaisian to Early Visean).

• Costayella Conil and Lys, 1977 (Type species: Tournayella costata Lipina, 1955). The test has an early part that is streptospiral, but is later planispiral, with later whorls having slight constrictions opposite basal supplementary deposits. Early Carboniferous (Tournaisian to Visean).

• Chernobaculites Conil and Lys, 1977 (Type species: Ammobaculites sarbaicus Malakhova subsp. beschevensis Brazhnikova, in Brazhnikova et al., 1967). The test is streptospirally enrolled in the early part, later becoming rectilinear with straight and horizontal septa. Late Carboniferous (Bashkirian).

• Chernyshinella Lipina, 1955 (Type species: Endothyra glomiformis Lipina, 1948). The test is streptospirally enrolled throughout with a tubular undivided early portion, and a final whorl with strongly asymmetrical and teardrop-shaped chambers. Late Devonian to Early Carboniferous (Late Famennian to Visean)

• Chernyshinellina Reytlinger, 1959 (Type species: Ammobaculites? pygmaeus Malakhova, 1954). The test is streptospirally enrolled in the early stage and has teardrop-like chambers in the later whorl, which are separated by distinct horizontal septa. Carboniferous (Tournaisian) (Plate 2.3, Fig. 16).

• Condrustella Conil and Longerstaey, 1977 (Type species: “Mstinia” modavensis Conil and Lys, 1967). The test is streptospirally enrolled in the early part, later becoming planispiral with teardrop-shaped chambers. Early Carboniferous (Late Tournaisian to Visean).

• Elbanaia Conil and Marchant, 1977 (Type species: Plectogyra michoti Conil and Lys, 1964). The test is streptospirally enrolled in the early part, but later evolute and septate. Early Carboniferous (Late Tournaisian to Early Visean).

• Endochernella Conil and Lys, 1980 (Type species: Plectogyra (Latiendothyra) quaesita Ganelina, 1966). The test is streptospirally enrolled in the early part, but later with radial septa and numerous chambers in the whorl. Early Carboniferous (Early Tournaisian to Visean).

• Eoforschia Mamet, 1970 (Type species: Tournayella moelleri Malakhova, in Dain and Grozdilova, 1953). The test is planispirally enrolled lacking definite septa. Early Carboniferous (Tournaisian to Visean).

• Eotournayella Lipina and Pronina, 1964 (Type species Tournayella (Eotournayella) jubra Lipina and Pronina, 1964). The test is streptospiral to planispiral in later stages, with slight growth constrictions. Late Devonian (Fig. 2.8).

• Forschia Mikhaylov, 1935 (Type species: Spirillina subangulata von Möller, 1879). The test is evolutely coiled without true septa. The aperture is cribrate. Early Carboniferous (Visean).

• Forschiella Mikhaylov, 1935 (Type species: Forschiella prisca Mikhaylov, 1935). The test is evolutely coiled without true septa in the early stage, later it is uncoiled. The aperture is cribrate. Early Carboniferous (Visean).

• Glomospiranella Lipina, 1951 (Type species: Glomospiranella Asiatica Lipina, 1951). The test is streptospiral in the early stage, later becoming streptospiral, without distinct septation, but with slight constrictions. Late Devonian to Early Carboniferous (Visean) (Fig. 2.6).

• Glomospiroides Reytlinger, 1950 (Type species: Glomospiroides fursenkoi Reytlinger, 1950). The test is streptospirally enrolled, being undivided in the early stage, but becoming rectilinear and irregularly divided with thin pseudosepta. Middle to Late Carboniferous.

• Laxoseptabrunsiina Vachard, 1977 (Type species: Laxoseptabrunsiina valuzierensis Vachard,1977). The test is streptospirally enrolled and undivided, later becoming planispiral with true septa. Early Carboniferous (Visean).

• Lituotubella Rauzer-Chernousova, 1948 (Type species: Lituotubella glomospiroides Rauzer-Chernousova, 1948). The test is streptospirally coiled in the early stage, later becoming rectilinear with no true septa. Early Carboniferous (Visean).

• Mstinia Dain, 1953 (Type species: Mstinia bulloides Dain, in Dain and Grozdilova, 1953). The test is involute, with teardrop-like chambers. The aperture is cribrate. Early Carboniferous (Late Visean).

• Mstiniella Conil and Lys, 1977 (Type species: Mstinia fursenkoi Dain, in Dain and Grozdilova, 1953). The test is streptospirally, undivided in the early part, but later planispiral with septa. The aperture is cribrate. Early Carboniferous (Visean) (Plate 2.9, figs 8, 10).

• Neobrunsiina Lipina, 1965 (Type species: Glomospiranella finitima Grozdilova and Lebedeva, 1954). The test is streptospirally enrolled in the early stage, but later planispiral with constrictions in the wall. Early Carboniferous (Tournaisian to Visean).

• Nevillea Conil and Lys, 1980 (Type species: Georgella dytica Conil and Lys, 1977). The test is streptospirally enrolled in the early stage, but later rectilinear, divided by arched septa. The aperture is multiple with a cribrate aperture occupying the entire terminal chamber surface. Early Carboniferous (Visean).

• Nodochernyshinella Conil and Lys, 1977 (Type species: Chernyshinella tumulosa Lipina, 1955). The test is streptospirally enrolled in the early part, but later planispiral with teardrop-like chambers. Early Carboniferous (Tournaisian).

• Pohlia Conil and Lys, 1977 (type species: Septatournayella henbesti Skipp, in Skipp et al., 1966). The test is planispirally enrolled with distinct septa in the final whorl. Early Carboniferous (Visean).

• Pseudolituotubella Vdovenko, 1967 (Type species: Pseudolituotubella multicamerata Vdovenko, 1967). The test is streptospirally enrolled in the early stage, but later is planispiral becoming rectilinear with true septa. The aperture is cribrate. Early Carboniferous (Late Tournaisian to Early Visean).

• Rectoseptatournayella Brazhnikova and Rostovceva, 1963 (Type species: Rectoseptatournayella stylaensis Brazhnikova and Rostovceva, 1963). The test is planispirally coiled in the early stage, but uncoiled in the final stage with septa and distinct chambers. Early Carboniferous (Tournaisian).

• Rectotournayellina Lipina, 1965 (Type species: Tournayellina (Rectotournayellina) Lipina, 1965). The test is streptospirally enrolled in the early part, but later rectilinear. Late Devonian to Early Carboniferous (Famennian to Tournaisian).

• Septabrunsiina Lipina, 1955 (Type species: Endothyra? Krainica Lipina, 1948). The test is streptospirally enrolled in the early nonseptate stage, but later it is planispiral with distinct septa. Early Carboniferous (Tournaisian) (Fig. 2.6).

• Septaforschia Conil and Lys, 1977 (Type species: Tournayella questita Malakhova, in Dain and Grozdilova, 1953). The test is enrolled, with later stages having true septa. Early Carboniferous (Tournaisian).

• Septaglomospiranella Lipina, 1955 (Type species: Endothyra primaeva Rauzer-Chernousova, 1948) Early Carboniferous (Tournaisian) (Fig. 2.6).

• Septatournayella Lipina, 1955 (Type species: Tournayella segmentata Dain, in Dain and Grozdilova, 1953). The test is a planispirally enrolled tube, with slight constrictions in the early stage, but later with complete septa forming distinct chambers. Late Devonian (Famennian) to Early Carboniferous (Visean).

• Spinobrunsiina Conil and Longerstaey, 1980 (Type species: Septabrunsiina (Spinobrunsiina) ramsbottomi Conil and Longerstaey, 1980). The test is streptospirally enrolled in the early stage, but later becoming planispiral and septate. Early Carboniferous (Tournaisian to Visean).

• Spinolaxina Conil and Naum, 1977 (Type species: Plectogyra pauli Conil and Lys, 1964). The test is streptospirally coiled in the early stage, later becoming nearly planispiral. Early Carboniferous (Visean).

• Spinotournayella Mamet, 1970 (Type species: Plectogyra tumula Zeller, 1957). The test is involute with early parts being streptospiral, but later becoming planispiral with endothyroid chambers. Early Carboniferous (Late Tournaisian).

• Tournayella Dain, 1953 (Type species: Tournayella discoidea Dain, 1953). The test is enrolled without definite septa and supplementary deposits. Early Carboniferous (Tournaisian to Visean) (Fig. 2.5).

• Tournayellina Lipina, 1955 (Type species: Tournayellina vulgaris Lipina, 1955). The test is enrolled with few chambers enlarging rapidly. Late Devonian to Early Carboniferous (Famennian to Tournaisian).

• Uviella Ganelina, 1966 (Type species: Uviella aborigena Ganelina, 1966). The test is streptospirally enrolled in the early undivided stage, but later is planispiral with constrictions in the early part followed in the final whorl by true septa. Early Carboniferous (Tournaisian to Visean).

• Viseina Conil and Lys, 1977 (Type species: Septatournayella? Conspecta Conil and Lys, 1977). The test is enrolled with later chambers divided by true septa. The aperture is cribrate in the final stage. Early Carboniferous (Visean).

Family Palaeospiroplectamminidae Loeblich and Tappan, 1984

The members of this family represent a biserial development from the Tournayellidae, and may be ancestral to the Palaeotextularioidea. They are initially streptospiral with teardrop-like or endothyroid-like chambers, then become planispiral and in later stages the test become biserial. The range is Late Devonian to Early Carboniferous.

• Endospiroplectammina Lippina, 1970 (Type species: Spiroplectammina venusta Vdovenko, 1954). The test is streptospirally enrolled, with distinctly endothyroid chambers, later becoming planispiral, uncoiled with a later biserial stage with a single aperture. Early Carboniferous (Late Tournaisian to Middle Visean).

• Eotextularia Mamet, 1970 (Type species: Palaeotextularia diversa Chernysheva, 1948). The test is irregularly coiled in the early part, later becoming uncoiled with a few pairs of biserial chambers. Early Carboniferous (Late Tournaisian to Middle Visean).

• Halenia Conil, 1980 (Type species: Halenia legrandi Conil, 1980). The test has a biserial elongate stage, followed by a broad uniserial stage with horizontal septa. The aperture is cribrate in the adult. Early Carboniferous (Visean).

• Palaeospiroplectammina Lipina, 1965 (Type species: Spiroplectammina tchernyshinensis Lipina, 1948). The test has a biserial uncoiled stage with curved septa, but may become uniserial with last chambers. Late Devonian to Carboniferous (Early Visean) (Plate 2.3, Fig. 11).

• Rectochernyshinella Lipina, 1960 (Type species: Spiroplectammina mirabilis Lipina, 1948). The test is streptospirally enrolled in a large early stage, the later stage uncoiled with few biserially arranged chambers. Late Devonian to Early Carboniferous (Late Famennian to Early Tournaisian).

ORDER ENDOTHYRIDA FURSENKO 1958

Members of this order have a lenticular test, planispirally coiled. The wall is dark microgranular, but sometimes can be bilayered or multilayered. Aperture simple, basal or cribrate. Late Devonian to Triassic.

Superfamily ENDOTHYROIDEA Brady, 1884 nom. translat. Fursenko, 1958

Members of this superfamily have typical endothyroid coiling with streptospiral to planispiral tests and constant deviation of the axis of coiling, with many chambers followed by a rectilinear stage in some forms (see Fig. 2.9). The walls are microgranular and calcareous, but some forms evolved two to three distinct layers, others may develop an inner perforate or keriothecal layer. Notably, they ranged from the Late Devonian to the Triassic (Famennian to Rhaetian).

Family Endotebidae Vachard, Martini, Rettori and Zaninetti, 1994

Test: free planispiral, in early stages, later uniserial to biserial. Wall: calcareous, grey, thick, calcareous agglutinated. Aperture: simple. Late Permian to Triassic.

• Endoteba Vachard and Razgallah, 1988 emend. Vachard, Martini, Rettori and Zaninetti, 1994 (Type species: Endoteba controversa Vachard and Razgallah, 1988 emend. Vachard, Martini, Rettori and Zaninetti, 1994). The axial view is compressed. Late Permian to Late Triassic (Late Kungurian to Rhaetian) (Fig. 3.4).

Family Endothyridae Brady, 1884

Here the test is small, multilocular, enrolled, with evolute planispiral coiling and well developed septa in early whorls (e.g. Loeblichia Cummings in the Visean), or initially streptospiral to planispiral characterised by the development of secondary deposits of calcite (chomata) on the chamber floor (e.g. Endothyra Phillips, 1846, Early Carboniferous to Permian). They exhibit a low equatorial aperture. Advanced forms may become uniserial (e.g. Haplophragmella Rauzer-Chernousova, 1936, Visean, see Fig. 2.7) with a cribrate terminal aperture. Their range is Late Devonian to Permian.

• Andrejella Malakhova, 1975 (Type species: Andrejella laxiformis Malakhova, 1975). The test is streptospirally coiled in the early part, with later chambers rapidly increasing in height. Early Carboniferous (Visean).

• Banffella Mamet, 1970 (Type species: Endothyra? banffensis McKay and Green, 1963). The test is streptospirally coiled in early whorl, later it is nearly planispiral and evolute, with straight and slightly oblique septa well developed only in early whorls. Early Carboniferous (Visean).

• Corrigotubella Ganelina, 1966 (Type species: Corrigotubella posneri Ganelina, 1966). The test is streptospirally enrolled in the early stage, later it is planispiral and rectilinear, with a few short and broad chambers. Septa in the enrolled stage are short, while those of the rectilinear portion are horizontal. Early Carboniferous (Late Tournaisian to Early Visean)

• Cribranopsis Conil and Longerstaey, 1980 (Type species: Cribranopsis fossa Conil and Longerstaey, in Conil et al., 1980). The test is enrolled, with a streptospiral early stage and a planispiral late coil, expanding rapidly. The septa are short and thick. Early Carboniferous (Visean).

• Cribrospira von Möller, 1878 (Type species: Cribrospira panderi von Möller, 1878). The test is streptospirally enrolled in the early stage, but later coiling is planispiral with numerous chambers per whorl. Septa are nearly radial, short and thick. Early Carboniferous (Visean).

• Elergella Conil, 1984 (Type species: Elergella simakoyi Conil, 1984). The test is streptospirally enrolled in the early stage, later becoming planispiral and evolute with short, slightly oblique septa. Early Carboniferous (Tournaisian)

• Endostaffella Rozovskaya, 1961 (Type species: Endothyra parya von Möller, 1879). The test is streptospirally enrolled in the early stage, later becoming planispiral and evolute. Early Carboniferous (Late Tournaisian to Visean).

• Endothyra Phillips, 1846 (Type species: Endothyra bowmani Phillips, 1846). The text is partially involute, planispiral with an early part streptospirally enrolled. It is formed by few whorls, with strong development of secondary deposits, such as nodes, ridges, or hooks on the chamber floor. The wall is calcareous, microgranular, with two or three layers, a thin dark outer layer or tectum and a thicker, fibrous to alveolar inner layer or diaphanotheca. The aperture is simple, basal. Early Carboniferous to Permian (Plate 2.5, Fig. 20; Plate 2.12, Fig. 8; Plate 2.6, figs 1-2; Plate 2.13, figs 1-3; see Fig. 2.7, 2.8).

• Endothyranella Galloway and Harlton, 1930 (Type species: Ammobaculites powersi Harlton, 1927). Streptospiral test becoming planispiral and then rectilinear. The aperture is simple and rounded. Late Carboniferous (Moscovian to Ladinian) (Plate 2.9, Fig. 1; Plate 3.5, Fig. 10).

• Endothyranopsis Cummings, 1955 (Type species: Involutina crassa Brady, in Moore, 1870). The test is subglobular involute, being almost planispiral with not more than 3 and a half whorls. They have a thick wall with no secondary deposits. Early coiling plectogyroid with a large proloculus. Sutures are perpendicular to the spirotheca. The aperture is a low basal equatorial arch. Carboniferous (Visean) (Plate 2.9, figs 3, 6, 7; Plate 2.12, figs 7, 10, 11; Plate 2.14, figs 1, 4, 5).

• Eoendothyra Miklukho-Maklay, 1960 (Type species: Endothyra communis Rauzer-Chernousova, 1948). The test is involute and streptospirally enrolled in the early stage, later becoming nearly planispiral, with numerous chambers. Late Devonian to Early Carboniferous (Famennian to Visean).

• Eoendothyranopsis Reytlinger and Rostovzeva, 1966 (Type species: Parastaffella pressa Grozdilova, in Lebedeva, 1954). The test is planispirally enrolled, involute, with oblique septa and secondary deposits at the base of the chambers that appear as a forward projecting hook or spine in the final chamber. Early Carboniferous (Early Visean).

• Eoquasiendothyra Durkina, 1963 (Type species: Endothyra bella Chernysheva, 1952). The test is streptospirally enrolled in the early stage, but later becoming planispiral, with inflated chambers and septa slightly oblique to the outer wall. Late Devonian to Early Carboniferous (Tournaisian).

• Euxinita Conil and Dil, 1980 (Type species: Dainella? efremoyi Vdovenko and Rostovtseva, in Brazhnikova et al., 1967). The test is streptospirally enrolled in the early stage with numerous chambers per whorl, later it becomes nearly planispiral. Early Carboniferous (Visean to Bashkirian).

• Globochernella Hance, 1983 (Type species: Globochernella braibanti Hance, 1983). The test is enrolled, with chambers enlarging rapidly as added. The septa are thick at the base and tapering rapidly to a very thin at the inner edge. Early Carboniferous (Visean).

• Globoendothyra Bogush and Yuferev, 1962 (Type species: Globoendothyra pseudoglobulus Bogush and Yuferev, 1962). The test is streptospirally enrolled in the early stage, but later is nearly planispiral with oblique septa. Secondary deposits on the chamber floors. Early Carboniferous (Visean to Moscovian).

• Granuliferella Zeller, 1957 (Type species: Granuliferella granulosa Zeller, 1957). The test is involute, streptospirally enrolled in the early stage, but later nearly planispiral, with relatively few chambers per whorl. The septa are short and slightly oblique. Late Devonian (Famennian) to Early Carboniferous (Visean).

• Granuliferelloides McKay and Green, 1963 (Type species: Granuliferelloides jasperensis McKay and Green, 1963). The test is streptospirally enrolled in the early stage, but later are nearly planispiral, with a few slightly inflated chambers and oblique septa, becoming uncoiled, with short cylindrical chambers and nearly horizontal septa. Early Carboniferous (Late Tournaisian).

• Haplophragmella Rauzer-Chernousova and Reytlinger, 1936 (Type species: Endothyra panderi von Möller,1879). The test is streptospirally enrolled in the early stage, with few chambers per whorl, later chambers become uncoiled and rectilinear. The aperture is simple in the early stage, becoming multiple and cribrate in the later stages. Early Carboniferous (Visean) (Fig. 2.7).

• Haplophragmina Reytlinger, 1950 (Type species: Haplophragmina kashkirica Reytlinger, 1950). The test is involute, planispirally enrolled in the early stage, becoming uncoiled in the late stage. Late Carboniferous (Moscovian).

• Holkeria Strank, 1982 (Type species: Rhodesina avonensis Conil and Longerstaey, in Conil et al., 1980). The test is streptospirally enrolled with the plane of coiling oscillating in the early stage, but later it is nearly planispiral and evolute with short septa short, which follow the curvature of the chambers. Early Carboniferous (Middle Visean).

• Klubonibelia Conil, 1980 (Type species: Klubonibelia immanis Conil, 1980). The test is streptospirally enrolled in the early stage, later becoming planispiral, with numerous sub-quadrate chambers per whorl. The final stage is uncoiled and rectilinear. Early Carboniferous (Middle to Late Visean).

• Latiendothyra Lipina, 1963 (Type species: Endothyra latispiralis Lipina, in Grozdilova and Lebedeva, 1954). The test is inflated, with a broadly rounded periphery and flattened sides. The early stage is streptospiral, later becoming planispiral, with rapidly enlarging whorls, with a moderate number of chambers per whorl (about six to eight in the final whorl). The septa are short, thick, and slightly oblique, projecting toward the aperture. The wall is calcareous, thick, microgranular, dark, single layered, with secondary deposits that result in septal thickening. The aperture is simple at the base of the apertural face. Early Carboniferous (Late Tournaisian to Early Visean).

• Latiendothyranopsis Lipina, 1977 (Type species: Endothyra latispiralis Lipina, in Grozdilova and Lebedeva, 1954). The test is streptospiral and enrolled in the early stage, later becoming planispiral, with rapidly enlarging whorl and short, thick, slightly oblique septa. Early Carboniferous (Late Tournaisian to Early Visean).

• Laxoendothyra Brazhnikova and Vdovenko, 1972 (Type species: Endothyra parakosyensis Lipina, 1955). The test is streptospirally enrolled in the early stage, later is planispiral, with elongate chambers in rapidly enlarging whorls and short septa. Early Carboniferous (Tournaisian to Visean).

• Mediopsis Bogush, 1984 (Type species: Planoendothyra? kharaulakhensis Bogush and Yuferev, 1966). The test is involute, streptospirally enrolled in the first half to one and a half whorls, later being planispiral. Early Carboniferous (Late Tournaisian to Visean).

• Melatolla Strank, 1983 (Type species: Melatolla whitfieldensis Strank, 1983). The test is streptospirally enrolled in the early stage, later becoming planispiral, and finally tending to uncoil, with massive and robust chomatal deposits in the last chambers. The aperture is areal and cribrate in the uncoiled part. Early Carboniferous (Visean).

• Mikhailovella Ganelina, 1956 (Type species: Endothyrina? gracilis Rauzer-Chernousova, 1948). The test is streptospiral to rectilinear with a short rectilinear part. The aperture is a low basal slit in the early enrolled part, but cribrate and in the uncoiled stage. Carboniferous (Middle Visean) (Plate 2.3, Fig. 19).

• Mirifica Shlykova, 1969 (Type species: Endothyra mirifica Rauzer-Chernousova, 1948. The test is involute, streptospirally enrolled in the early stage, later nearly planispiral with numerous chambers per whorl and short, hooked sutures. The aperture is cribrate in the final whorl. Early Carboniferous (Late Visean).

• Omphalotis Shlykova, 1969 (Type species: Endothyra omphalota Rauzer-Chernousova and Reytlinger, in Rauzer-Chernousova and Fursenko, 1937). The test is involute, with early streptospiral coiling, later becoming planispiral. The sutures are radial to slightly oblique. Carboniferous (Middle Visean to Early Bashkirian).

• Paradainella Brazhnikova, 1971 (Type species: Paradainella dainelliformis Brazhnikova and Vdovenko, 1971). The test is streptospirally enrolled in the early stage, but the final whorls are nearly planispiral with secondary deposits forming massive chomata that may cover the inner surface of the whorls. The aperture is simple, basal. Early Carboniferous (Late Tournaisian).

• Paraendothyra Chernysheva, 1940 (Type species: Paraendothyra nalivkini Chernysheva, 1940). The test is enrolled and biumbilicate. The early stage is slightly streptospiral, later becoming nearly completely planispiral. The sutures are radial with a slight serrate appearance, and the final few septa are hook-like in section. The aperture is in a median septal concavity. Early Carboniferous (Middle Tournaisian).

• Paraplectogyra Okimura, 1958 (Type species: Paraplectogyra masanae Okimura, 1958). The test is streptospirally enrolled in the early stage, involute, later becoming planispiral with whorls expanding rapidly with straight, radial septa. Early Carboniferous (Late Tournaisian to Early Visean).

• Planoendothyra Reytlinger, 1959 (Type species: Endothyra aljutovica Reytlinger, 1950). The test is compressed, streptospiral in the early stages, later becoming planispiral and evolute, and slightly asymmetrical with supplementary deposits. Carboniferous (Visean to Bashkirian) (Plate 2.9, Fig. 18; Plate 2.12, Fig. 12).

• Plectogyra Zeller, 1950 (Type species: Plectogyra plectogyra Zeller, 1950). A streptospiral test with a two-layered wall. There are 4 to 12 chambers in the final whorl. Late Devonian/Early Carboniferous to Permian (Plate 2.3, Fig. 2C; Plate 2.12, figs 1, 3-5; Plate 2.13, Fig. 10B; Plate 2.14, figs 3, 6-9, 11, 12).

• Plectogyranopsis Vachard, 1977 (Type species: Endothyra convexa Rauzer-Chernousova, 1948). The test is planispiral, biumbilicate with short, straight, thick septa. Early Carboniferous (Early Visean) to Late Carboniferous (Early Bashkirian).

• Pojarkovella Simonova and Zub, 1975 (Type species: Pojarkovella honesta Simonova and Zub, 1975). The test is streptospirally enrolled in the early stage, biumbilicate, tightly coiled and involute, later becoming planispiral with radiate and straight septa and massive secondary deposits consisting of angular chomata. Early Carboniferous (Visean).

• Priscella Mamet, 1974 (Type species: Endothyra prisca Rauzer-Chernousova and Reytlinger, in Rauzer-Chernousova et al., 1936). The test is streptospirally enrolled in the early stage, later becoming nearly planispiral with long strongly oblique septa and no distinct basal secondary deposits. Early Carboniferous (Tournaisian to Bashkirian).

• Quasiendothyra Rauzer-Chernousova, 1948 (Type species: Endothyra kobeitusana Rauzer-Chernousova, 1948). The test is discoidal with concave sides and slightly streptospiral early whorls, with septa as thick as the outer wall. Secondary deposits are chomata-like, at each side of the median line against the previous whorl. Late Devonian (Famennian) to Early Carboniferous (Tournaisian).

• Rectoendothyra Brazhnikova, 1983 (Type species: Endothyra (Rectoendothyra) donbassica Brazhnikova, 1983). The test is streptospirally coiled in the early stage, becoming planispiral with the final whorl enlarging rapidly and may tend to uncoil. Septa are straight and radial. Chomata-like deposits occur on chamber floor. The aperture is multiple in the final chamber. Early Carboniferous Tournaisian.

• Rhodesinella Conil and Longerstaey, 1980 (Type species: Cribrospira pansa Conil and Lys, 1965). The test is streptospiral enrolled in the early part, later becoming planispiral with a tendency to uncoil in the later stage. Septa moderately short, straight, and inclined toward the aperture. The aperture is multiple a in the final chamber. Early Carboniferous (Tournaisian).

• Semiendothyra Reytlinger, 1980 (Type species: Semiendothyra surenica Reytlinger, 1980). The test is streptospirally enrolled with early whorls, involute, but later slightly evolute, planispiral with slightly arched septa. Secondary deposits are extensive, small chomata-like mounds near the aperture, spine-like in the final chamber. The aperture is simple, basal, and low. Middle Carboniferous (Bashkirian).

• Spinoendothyra Lipina, 1963 (Type species: Endothyra costifera Lipina, in Grozdilova and Lebedeva, 1954). The test is closely coiled in the early stage, with later whorls planispiral. Sutures are slightly curved without any thickenings at the ends. Secondary deposits are spine-like, well developed on chamber floors, anteriorly directed. The aperture is simple and basal. Carboniferous (Late Tournaisian to Early Visean) (Plate 2.9, Fig. 15).

• Spinothyra Mamet, 1976 (Type species: Endothyra pauciseptata Rauzer-Chernousova, 1948). The test is streptospirally enrolled, involute, biumbilicate, with short thick septa, whorls enlarge rapidly and secondary chomata-like deposits occur on the floor of the chambers. Carboniferous (Middle Visean to Early Bashkirian).

• Timanella Reytlinger, 1981 (Type species: Endothyra eostaffelloides Reytlinger, 1950). The test is planispirally enrolled with rapidly enlarging whorls and straight oblique septa, with deposits covering the chamber floor and filling the lateral areas. Late Carboniferous (Moscovian).

• Tuberendothyra Skipp, 1969 (Type species: Endothyra tuberculata Lipina, 1948). The test is streptospirally enrolled in the early stage, later becoming planispiral with long, curved septa, long and secondary spine-like deposits on the floor of all chambers. Early Carboniferous (Late Tournaisian to Early Visean).

• Urbanella Malakhova, 1963 (Type species: Quasiendothyra urbana Malakhova, 1954). The test is streptospirally enrolled in the early whorls, with later whorls increasing in height, becoming planispiral and evolute, with secondary small rounded chomata at the margins of the aperture. Early Carboniferous (Tournaisian to Early Visean).

• Zellerinella Mamet, 1981 (Type species: Endothyra discoidea Girty, 1915. The test is streptospirally enrolled in early stage, later becoming planispiral and evolute with pseudochomata. Early Carboniferous (Late Visean to Early Bashkirian).

Family Bradyinidae Reytlinger, 1950

The Endothyridae evolved into forms such as Bradyina von Möller, 1878 (see Fig. 2.9). These forms have nautiloid planispiral coiling with few chambers. The microgranular calcareous wall is perforate, with distinct radial lamellae with supplementary septal pores opening into the septal aperture. The primary aperture is also cribrate. They occur in the Carboniferous (Visean) to the Permian (Sakmarian).

• Bibradya Strank, 1983 (Type species Bibradya inflata Strank, 1983). The test is streptospirally enrolled with short, thick septa which bifurcate close to the outer wall. Early Carboniferous (Late Visean)

• Bradyina von Möller, 1878 (Type species: Bradyina nautiliformis Möller, 1878). The test is planispiral and involute with supplementary pores and septal partitions. The wall has a microgranular tectum and keriothecal structure. Carboniferous (Tournaisian to Gzhelian) (Fig. 2.7, 2.8; Plate 2.8, Fig. 13; Plate 2.9, Fig. 20; Plate 2.12, Fig. 2; Plate 2.13, Fig. 10A).

• Glyphostomella Cushman and Waters, 1928 (Type species: Ammochilostoma? triloculina Cushman and Waters, 1927). The test is planispirally enrolled, involute, with a complex wall, with plates and partitions and large and branching pores in the wall. Late Carboniferous to Permian (Kasimovian to Changhsingian).

• Janischewskina Mikhaylov, 1935 (Type species: Janischewskina typica Mikhaylov, 1935). The test is planispirally enrolled, and involute with septal chamberlets as in Bradyina, but without the alveolar structure of Bradyina. Early Carboniferous (Late Visean).

• Postendothyra Lin, 1984 (Type species Postendothyra scabra J. X. Lin, 1984). The test is planispirally enrolled and involute, with outer tectum and inner coarsely alveolar keriotheca. Early Permian (Asselian).

Family Loeblichiidae Cummings, 1955

Members of this family have atypical endothyroid planispiral tests with numerous chambers, pseudochomata, a basal aperture, two-layered walls, an inner clear fibrous light layer (called early tectorium) and a dark granular outer layer (called tectum) (Fig. 2.8A). They range from Middle Devonian to Late Carboniferous (Moscovian).

• Dainella Brazhnikova, 1962 (Type species: Endothyra? chomatica Dain, in Brazhnikova, 1962). The test is streptospiral and involute throughout, with secondary deposits in the form of massive chomata. Early Carboniferous (Early Visean)

• Loeblichia Cummings, 1955 (Type species: Endothyra ammonoides Brady, 1873). The test is small, flattened, discoidal and is planispiral throughout, with up to 20 sub-rectangular chambers in the final whorl, but lacks secondary deposits. Early Carboniferous (Visean) (Fig. 2.7, 2.8; Plate 2.7, Fig. 7; Plate 2.8, Fig. 12B; Plate 2.13, Fig. 11B).

• Lysella Bozorgnia, 1973 (Type species: Lysella gadukensis Bozorgnia, 1973). The test is streptospirally enrolled in the early stage, later becoming planispiral, involute with many chambers per whorl that appear sub-quadrate in section. Septa are slightly inclined toward the aperture. The wall has chomata-like structures. Early Carboniferous (Visean).

• Novella Grozdilova and Lebedeva, 1950 (Type species: Novella evoluta Grozdilova and Lebedeva, 1950). The test is planispirally enrolled and evolute with straight septa and well-developed chomata in later whorls. Late Carboniferous (Bashkirian to Early Moscovian).

• Seminovella Rauzer-Chernousova, 1951 (Type species: Eostaffella (Seminovella) elegantula Rauzer-Chernousova, in Rauzer-Chernousova et al., 1951). The test is planispiral, involute becoming later evolute, with weakly developed pseudochomata. Late Carboniferous (Late Bashkirian to Moscovian).

ORDER FUSULINIDA WEDEKIND, 1937

This order includes all larger benthic foraminifera with a homogeneously microgranular primary test made of low-Mg calcite, in which the crystal units have no optical alignment and various foreign particles might be incorporated (see Rauzer-Chernousova, 1948; Loeblich and Tappan 1964; Brazhnikova and Vdovenko 1973; Tappan and Loeblich 1988; Rigaud et al., 2014). Advanced forms have two or more differentiated layers in the wall. They range from the Carboniferous to the Permian.

The Fusulinida have special morphological diagnostic characters, which are unique to this order (see Haynes, 1981; Loeblich and Tappan, 1988; and Fig. 2.9, 2.10). The traces of the septa on the external surface are called “septal furrows”. These furrows extend from pole to pole of the test and marks the early part of the partitions between the chambers, called “septa”. The apertural face of the test is the “antetheca” while the external wall of the test is the “spirotheca”. Both spirotheca and antetheca are finely perforate by numerous small openings “septal pores” and the test lacks primary apertures. Communication between chambers occurs at the base of septa and aided by resorption of a tunnel in the central part of the test in many fusulinides or of several small circular tunnels, foramina throughout the length of the test in others. In some advanced fusulinides, the antetheca is corrugated into uniformly or irregular spaced waved, called fluting. Thin sections of the fusulinides reveal highly complicated internal structures which are essential for identifying and classifying the genera and species. The first chamber is spherical to sub-spherical, called proloculus. Dense calcite, called chomata, were deposited along the margins of the tunnel in many fusulinides, and ridges of dense calcite, parachomata were developed between adjacent foramina in those with multiple foramina). Deposition of dense calcite and simultaneous formations of chomata and parachomata, axial fillings occurred in axial regions of some fusulinides. The spirotheca consists of a thin, dense, primary layer, tectum, which is sometimes covered by layers of tectoria. In more advanced forms, the tectum is augmented by a transparent layer, the diaphanotheca, or a thick layer of honeycomb-like structure, keriotheca. Ridges, called septula spread down from the lower surface of the spirotheca to subdivide the chambers in the neoschwagerinoids.

Since the studies made by Loeblich and Tappan (1964, 1980) and Haynes (1981), many authors have tried to subdivide the fusulinides by considering the highly complicated internal structures, such as the wall structure, the presence or absence of the chomata, and the degree of septal folding in the test. On the basis of these main morphological features, the Fusulinida are divided here into six superfamilies (see Fig. 2.1 and 2.2), the Fusulinoidea, the Ozawainelloidea, the Staffelloidea, the Schubertelloidea, the Fusulinoidea, the Schwagerinoidea, and the Neoschwagerinoidea (= Verbeekinoidea). These superfamilies are considered as orders by Rauzer-Chernousova et al. (1996) and Mikhalevich (2013).

The structure of the spirotheca plays an important role in differentiating the fusulinids. Five types of wall structures (see Fig. 2.10) characterise these superfamilies:

Forms with two layered walls, an inner clear fibrous light layer (called early tectorium) and a dark granular outer layer (called tectum) (Fig. 2.10A)

The first four types of wall characterise the ozawainelloids, staffelloids, and the fusulinoids and the most advanced final type of wall belong to the schwagerinoids and neoschwagerinoids (discussed below).

Over 100 fusulinides genera are recognised. Below are the main taxonomic classifications of the fusulinides, but for an even more detailed taxonomic description at the generic and specific levels see Loeblich and Tappan (1988) and Rauser-Chernousova et al. (1996).

Superfamily OZAWAINELLOIDEA Slovieva, 1978

The test is small, lenticular to rhombic to discoid and involute. The early coiling may be streptospiral. The wall is simple in the early forms, composed of tectum with upper and lower tectoria (Fig. 2.10B), but with diaphanotheca occurring between tectum and lower tectorium (Fig. 2.10C) in advanced forms. The proloculus is small and spherical. The pseudochomata develop into weak to strong chomata. The aperture is simple and basal. This superfamily ranges from the Early Carboniferous to the Late Permian (Visean to Capitanian).

Family Ozawainellidae Thompson and Foster, 1937

Here, the tests are discoidal to elongate in shape. They are involute to slightly evolute with moderate to strong secondary deposits of calcite (chomata) (Fig. 2.11). The wall is of a primitive fusulinid type, with a thin dense, primary layer (tectum) dark with organic matter, surrounded by a thicker, less dense layer of upper and lower tectoria (Fig. 2.10B). Carboniferous to the Late Permian (Late Tournaisian to Capitanian).

This family includes genera such as:

• Chenella Miklukho-Maklay, 1959 (Type species: Orobias kueichihensis S. Chen, 1934). The test is planispirally enrolled, with the final whorl increasing abruptly in height. Late Permian (Capitanian).

• Eoparastaffella Vdovenko, 1954 (Type species. Parastaffella (Eoparastaffella) simplex Vdovenko, 1954). Test is almost planispiral. The wall is of two very thin, and dark microgranular tecta surrounded by thick, pale brown-gray zone. Early Caroboniferous (Late Tournaisian to Early Visean).

• Eostaffella Rauzer-Chernousova, 1948 (Type species: Staffella (Eostaffella) parastruvei Rauzer-Chernousova, 1948). The test is lenticular, involute and planispiral with a sub-acute periphery. Septa are thick, curved, and often with truncated edges. The wall is layered with tectum and upper and lower tectoria. There are discontinuous knobs on either side of the tunnel but no continuous chomata. Carboniferous to Permian (Visean to Capitanian) (Plate 2.3, Fig. 4A; Plate 2.5, Fig. 19; Plate 2.9, Fig. 5; Plate 2.12, figs 4, 9, 13, 14; Plate 2.14, figs 14, 15; Plate 2.15, Fig. 4).

• Eostaffelloides Miklukho-Maklay, 1959 (Type species: Eostaffelloides orientalis Miklukho-Maklay, 1959). The test is lenticular, planispirally enrolled with secondary deposits forming triangular chomata. Late Permian (Capitanian).

• Millerella Thompson, 1942 (Type species: Millerella marblensis Thompson, 1942; see Fig. 2.9). Minute discoidal and partially evolute test. The septa are slightly arched forward. Carboniferous (Serpukhovian to Gzhelian) (Fig. 2.13; Plate 2.3, Fig. 2A; Plate 2.12, Fig. 6; Plate 2.14, Fig. 10; Plate 2.8, Fig. 11).

• Ozawainella Thompson, 1935 (Type species: Fusulinella angulata Colani, 1924). The test is angular with massive chomata. Late Carboniferous to Permian (Serpukhovian to Capitanian).

• Pamirina Leven, 1970 (Type species: Pamirina darvasica Leven, 1970). The test is small and varies in shape from nautiloid to subspherical. The spirotheca is thin, one-layered in inner whorls and two-layered consisting of tectum and protheca in later whorls. The septa are straight and flat. The chomata vary from rudimentary to well developed. Early Permian (Sakmarian to Kungurian).

• Pseudonovella Kireeva, 1949 (Type species: Pseudonovella irregularis Kireeva, 1949). The test is lenticular with early whorls evolute and final whorl increasing rapidly in height and becoming partially evolute. The pseudochomata are weakly developed. Late Carboniferous (Late Bashkirian to Moscovian).

• Quasireichelina Ueno 1992 (Type species: Quasireichelina expansa Ueno, 1992). The test is involute in the early part, but later is uncoiled with numerous unfluted septa. Secondary deposits form small chomata. Middle Permian (Guadalupian).

• Reichelina Erk, 1942 (Type species: Reichelina cribroseptata Erk, 1942). The test is planispirally enrolled in the early stage, but the final whorl uncoils to appear almost peneropliform. Secondary deposits consist of broad chomata, extending and thickening poleward from the tunnel or the small circular foramina. Late Permian (Capitanian).

• Sichotenella Tumanskaya, 1953 (Type species: Sichotenella sutschanica Tumanskaya, 1953). The test is lenticular, resembling Chenella, but the final whorl increases rapidly in height and uncoils. The uncoiled stage is relatively large. The wall is three layered, with diaphanotheca. Late Permian (Capitanian).

• Zarodella Sosnina, 1981 (Type species: Zarodella zhamoidai Sosnina, 1981). The test is small, slightly biumbilicate. The proloculus is large in size compare to the adult test which comprises three to five whorls. The septa and flat and flat and the spirotheca is weakly differentiated. Early Permian (Sakmarian).

Family Pseudostaffellidae Putrya 1956

The Pseudostaffellidae have an involute test which may be a streptospiral in the early stage. Early walls are weakly differentiated, later ones have a tectum, thin diaphanotheca and two tectoria (Fig. 2.10D) with well-developed chomata. Carboniferous to Permian (Serpukhovian to Capitanian).

• Hubeiella Lin, 1977 (Type species: Hubeiella simplex J. X. Lin, 1977). The test is planispirally enrolled and involute in the early stage, later stages may be evolute, and the final whorl is greatly expanded in height and flares. The septa are straight, and the wall shows a tectum and fine keriotheca with small but distinct chomata present only in the inner whorls. Early Permian (Serpukhovian).

• Kangvarella Saurin, 1962 (Type species: Kangvarella irregularis Saurin, 1962). The test is involute in the early part, and later whorls increase rapidly in height. The wall has a tectum, diaphanotheca, and inner and outer tectoria. Irregular and asymmetrical parachomata may be present in the second and third whorls. Late Permian (Capitanian)

• Mediocris Rozovskaya, 1961 (Type species: Eostaffella mediocris Vissarionova, 1948). The test is streptospirally enrolled in the early part, later becoming planispiral and involute. The wall has a dark tectum and a poorly defined tectoria, with strong secondary deposits in the axial region. Carboniferous (Middle Visean to Early Moscovian).

• Neostaffella Miklukho-Maklay, 1959 (Type species: Pseudostaffella sphaeroidea Ehrenberg, in Rauzer-Chernousova et al., 1951). Chomata are very broad, extending from the edge of the narrow tunnel outward to the umbilical region. Late Carboniferous (Moscovian) (Plate 2.16, Fig. 9).

• Ninella Malakhova, 1975 (Type species: Endothyra staffelliformis Chernysheva, 1948). The test is streptospirally enrolled in the early part, later becoming nearly planispiral, with straight septa and widely spaced rounded pseudochomata in all whorls. Early Carboniferous (Visean).

• Pseudostaffella Thompson, 1942 (Type species: Pseudostaffella needhami Thompson, 1942). The test is streptospirally coiled in early whorls, later becoming planispiral with straight, long, unfluted and perpendicular to the outer wall septa, slightly curved in the polar regions. Chomata are well developed. Late Carboniferous (Bashkirian to Moscovian)

• Rauserella Dunbar, 1944 (Type species: Rauserella erratica Dunbar, 1944). The test is planispirally enrolled in the early whorls. The whorls become much wider and the test fusiform with irregular septa in outer whorls. Septal pores are present in the outer whorls. Middle Permian (Kungurian)

• Toriyamaia Kanmera, 1956 (Type species: Toriyamaia latiseptata Kanmera, 1956). The test is fusiform, the first whorl is evolute, later whorls are sub-cylindrical, involute, increasing rapidly in height. Septa are sparse and unfluted. Early Permian (Cisuralian).

Superfamily STAFFELLOIDEA Slovieva, 1978 emend. Vachard et al., 2013

In this order, the tests are small, spherical to discoidal and strong chomata are present. The wall structure of this order has been discussed by many authors, but is still ill-resolved (see Jin-Zhang, 1990). The wall is usually composed of tectum and diaphanotheca (Fig. 2.11) and may have tectoria, but it is mainly found recrystallized and altered by secondary mineralisation, suggesting that they secreted an aragonite or high-Mg calcite wall (Groves et al., 2005). Early Carboniferous to Late Permian (Visean to Wuchiapingian).

Family Staffellidae Miklukho-Maklay, 1949

Staffellidae have a primitive wall structure, consisting of a tectum followed by a transparent layer (diaphanotheca) which is formed of clear calcite (Fig. 2.11). They occur from the Early Carboniferous to Late Permian (Visean to Wuchiapingian).

• Chenia Sheng, 1963 (Type species: Chenia kwangsiensis Sheng, 1963). The test is lenticular with plane, unfluted septa. Chomata are well developed in all whorls with poorly developed and discontinuous parachomata present in the outer four or five whorls. Middle Permian (Kungurian).

• Nankinella Lee, 1934 (Type species: Staffella discoides Lee, 1931). The test is discoidal with plane septa and distinct chomata. Early to Late Permian (Sakmarian to Wuchiapingian).

• Pisolina Lee, 1934 (Type species: Pisolina excessa Lee, 1934). The test is spherical with thick, plane septa and well defined, low, asymmetrical chomata. Early Permian (Asselian to Kungurian).

• Pseudoendothyra Mikhaylov, 1939 (Type species: Fusulinella struvii von Möller, 1879). The test is lenticular, compressed, planispiral and involute, with plain septa and supplementary deposits adjacent to the aperture but not forming continuous chomata. The aperture is low and broad. Early Carboniferous to Early Permian (Visean to Sakmarian). (Plate 2.9, figs 2, 13; Plate 2.12, Fig. 15A; Plate 2.14, figs 2, 13).

• Staffella Ozawa, 1925 (Type species: Fusulina sphaerica Abich, 1859). It is the main representative of this group. It has a small lenticular test with a wall composed of a tectum and diaphanotheca. Permian (Artinskian to Kungurian) (Fig. 2.11).

Family Kahlerinidae Leven, 1963

This family is monotypic; the test is globose with minor chomata and rarely with small parachomata. Middle Permian.

• Kahlerina Kochansky-Devidé and Ramovš, 1955 (Type species: Kahlerina pachytheca Kochansky-Devidé and Ramovš, 1955). Septa are long, planar and unfluted. Chomata are only rudimentarily present in the initial whorls. Middle Permian (Guadalupian).

Superfamily SCHUBERTELLOIDEA Vachard in Vachard et al., 1993

The shell is minute, fusiform or irregular in shape. The wall is composed of tectum with lower tectorium only (Fig. 2.10A), or tectum surrounded by an upper and lower tectoria (Fig. 2.7B), or tectum with diaphanotheca and tectoria (Fig. 2.10C). Late Carboniferous to Late Permian (Bashkirian to Changhsingian).

Family Schubertellidae Skinner, 1931

Schubertellidae have fusiform tests with a streptospiral early coiling. Early septa are flat but evolve into fluted septa in advanced forms. The wall varies, as in some forms the diaphanotheca is present (Fig. 2.10B), while in others only the tectum and early tectorium are present (see Fig. 2.10A; Fig. 2.11). This family includes spirothecal structures of the post-keriotheca phase. Carboniferous to Permian (Bashkirian to Changhsingian)

Significant genera include:

• Biwaella Morikawa and Isomi, 1960 (Type species: Biwaella omiensis Morikawa and Isomi, 1960). The test is large, fusiform to sub-cylindrical, with subglobose and tightly coiled initial whorls followed by ovoid whorls that increase rapidly in size. The wall in first whorls is composed of a thin, dark tectum and a thicker but lighter tectorium. The wall in the final whorl is perforated with coarse mural pores, which do not develop into keriotheca. The septa are nearly straight and widely separated. The chomata are small and prominent in all the test, except for the final whorl. (Davydov, 2007; 2010). Late Carboniferous to Early Permian (Early Gzhelian to Kungurian).

• Boultonia Lee, 1927 (Type species: Boultonia willsi Lee, 1927). The test is fusiform, with strongly fluted septa throughout and asymmetric chomata. Early Permian (Asselian to Sakmarian).

• Codonofusiella Dunbar and Skinner, 1937 (Type species: Codonofusiella paradoxica Dunbar and Skinner, 1937). The test is irregularly coiled, becoming uncoiled in later stages, small, fusiform, with strongly fluted septa. Chomata are present but tunnels are not clearly defined. Late Permian (Lopingian).

• Dunbarula Ciry, 1948 (Type species: Dunbarula mathieui Ciry, 1948). The test is an elongate ellipsoid, irregularly coiled, with septa strongly fluted throughout. Late Permian (Capitanian).

• Dutkevichites Daydov, 1984 (Type species: Dutkevichites darvasica Davydov, 1984). The test is cylindrical. Septa are few, but strongly fluted with an increase in intensity towards the poles. Minute chomata are present in outer whorls. Late Carboniferous to Early Permian (Gzhelian to Asselian) (Fig. 2.14).

• Eoschubertella Thompson, 1937 (Type species: Schubertella lata Lee and Chen, in Lee et al., 1930). The test is elongate with early whorls close coiled and showing a broad tunnel, bordered by low chomata. The wall is unilayered. Late Carboniferous (Moscovian).

• Fusiella Lee and Chen, 1930 (Type species: Fusiella typica Lee and Chen, in Lee et al., 1930). The test is elongate, early whorls are discoidal and endothyroid, later becoming fusiform with axial fillings prominent. Late Carboniferous (Moscovian) (Plate 2.15, Fig. 6).

• Gallowaiina Chen, 1934 (Type species: Gallowaiina meitiensis Chen, 1934). The test is fusiform with very thin septa, closely fluted throughout, with folds extending over most of the chamber height. No chomata are present, fillings only found in the axial region. Late Permian (Changhsingian).

• Grovesella Davydov and Arefifard, 2007 (Type species: Grovesella tabasensis Davydov and Arefifard, 2007). Test: Discoidal, nearly planispirally coiled with a large proloculus and two-layered poorly developed wall lacking chomata. Late Carboniferous to Permian (middle Bashkirian up to Wordian).

• Kwantoella Sakagami and Omata, 1957 (Type species: Kwantoella fujimotoi Sakagami and Omata, 1957). The test is fusiform with numerous, straight and unfluted septa. Chomata are poorly developed, and axial fillings are more extensive in later whorls. Early Permian (Sakmarian).

• Lantschichites Tumanskaya, 1952 (Type species: Codonofusiella (Lantschichites) maslennikovi Tumanskaya, 1953). The test is cylindrical with strongly developed and fluted septa. Late Permian (Lopingian).

• Mesoschubertella Kanuma and Sakagami, 1957 (Type species: Mesoschubertella thompsoni Sakagami, in Kanuma and Sakagami, 1957). The test is fusiform with straight to slightly fluted septa in the polar ends. The wall is thick and composed of a thin, dark tectum and a diaphanotheca between a well-developed upper tectorium and a lower tectorium. Chomata are small, but well developed and asymmetrical. Early Permian (Cisuralian to Guadalupian).

• Minojapanella Fujimoto and Kanuma, 1953 (Type species: Minojapanella elongata Fujimoto and Kanuma, 1953). The test is subcylindrical, with intensely fluted septa, but without developing cuniculi (i.e. multiple tunnels, see Fig. 2.9). Chomata are massive. Late Permian (Capitanian to Changhsingian) (see Fig. 2.14).

• Nanlingella Rui and Sheng, 1981 (Type species: Nanlingella meridionalis Rui and Sheng, 1981). The test has a small proloculus. Septa are strongly fluted except in the median part. The wall is thin and composed of a tectum and diaphanotheca. Chomata are present only in the first two whorls. Late Permian (see Fig. 2.14).

• Neofusulinella Deprat, 1912 (Type species: Neofusulinella praecursor Deprat, 1913). The test is ovoid in the early stage, later becoming fusiform, planispirally coiled throughout with flat, slightly fluted septa. Chomata are large and asymmetrical. Late Carboniferous (Early Moscovian).

• Neoschubertella Saurin, 1962 (Type species: Neoschubertella sisophonensis Saurin, 1962). The test is fusiform, with slightly curved but not fluted septa and continuous pseudochomata. Early Permian to Late Permian (Sakmarian to Capitanian).

• Palaeofusulina Deprat, 1912 (Type species: Palaeofusulina prisca Deprat, 1913). The test has the spirotheca composed of a tectum and an early transparent diaphanotheca. The Palaeofusulina-type fusulinids became abundant after most of the Schwagerina-Verbeekina-Neoschwagerina types had disappeared (Jin-Zhang, 1990). Lattermost Permian (Changhsingian) (see Fig. 2.14).

• Paradoxiella Skinner and Wilde, 1955 (Type species: Paradoxiella pratti Skinner and Wilde, 1955). The test is discoidal, flaring and uncoiling with intensely fluted septa, and with well-developed cuniculi and low chomata. Late Permian (Capitanian).

• Paradunbarula Skinner, 1969 (Type species: Paradunbarula dallyi Skinner, 1969). The test is fusiform with intensely fluted septa and moderately wide, weak chomata. Late Permian (Capitanian)

• Russiella Miklukho-Maklay, 1957 (Type species: Russiella pulchra Mildukho-Maklay, 1957). The test is fusiform, and the first two whorls endothyroid. Septa are strongly fluted, with heavy axial fillings. Chomata are weakly developed except in the last two revolutions. Late Permian (Capitanian to Changhsingian).

• Schubertella Staff and Wedekind, 1910 (Type species: Schubertella transitoria Staff and Wedekind, 1910). The test is small becoming fusiform in later stages, with straight to weakly fluted septa at the polar ends. The wall is differentiated into three layers that are penetrated by relatively coarse pores. Chomata are prominent, low and asymmetrical. Late Carboniferous to Permian (Moscovian to Wordian) (see Fig. 2.11; Plate 2.17, Fig. 1).

• Schubertina Marshall, 1969 (Type species: Schubertina circuli Marshall, 1969). The test is subglobose to ovoid in outline, with a thin wall clearly differentiated into two layers, poorly developed secondary deposits and straight septa. Late Carboniferous to Permian (Bashkirian to Lopingian).

• Sphaeroschwagerina Miklukho-Maklay, 1959 (Type species: Schwagerina sphaerica var. karnica Shcherbovich, in Rauzer-Chernousova and Shcherbovich, 1949). The test is almost spherical with weakly fluted septa at the poles and small chomata. Permian (Plate 2.18, figs 6, 7, 9).

• Yangchienia Lee, 1934 (Type species: Yangchienia iniqua Lee, 1934). The test is fusiform with plane unfluted septa and massive, asymmetrical chomata that extend nearly to the poles. Late Permian (Asselian to Capitanian).

Superfamily FUSULINOIDEA von Möller, 1878

The test of the Fusulinoidea is large and varies from spherical to fusiform. The development of secondary deposits of calcite (chomata) is prominent in most forms, but the numerous chambers are subdivided by folds or septula (Fig. 2.10, 2.12). Late Carboniferous to Permian (Bashkirian to Kungurian).

Family Fusulinidae von Möller, 1878

In the Fusulinidae, the test is spherical or elongate fusiform, and mostly planispiral. The Fusulinidae family belongs to the pre-keriotheca stage of fusulinides development. It includes genera with a wall composed of a tectum surrounded by a late and an early tectorium (Fig. 2.10B) (e.g., the type of wall typical for Profusulinella, see Fig. 2.12). However, it also includes more evolved genera such as the cylindrical Fusulina and the fusiform Beedeina (see Fig. 2.12) with four layered walls, with the late and early tectorium surrounding the tectum and a glossy layer called the diaphanotheca (Fig. 2.10C). Carboniferous to Permian (Bashkirian to Kungurian).

Significant genera include:

• Beedeina Galloway, 1933 (Type species: Fusulinella girtyi Dunbar and Condra, 1928). The test is fusiform with plane to weakly fluted septa in the early whorls, later septa are strongly fluted. Secondary fillings also coat the septa. Chomata are massive, high and broad in the early stage, but lower in the adult. Late Carboniferous (Moscovian) (Fig. 2.12).

• Eofusulina Rauzer-Chernousova, 1951 (Type species: Fusulina triangula Rauzer-Chernousova and Belyaev, in Rauzer-Chernousova et a1.,1936). The test is fusiform with septa strongly fluted throughout, with high and narrow folds. Carboniferous (Early Moscovian).

• Eowedekindellina Ektova, 1977 (Type species: Eowedekindellina fusiformis Ektova, 1977). The test is fusiform with fluted septa, mainly in final whorl. Chomata are extended poleward, forming a basal layer. Late Carboniferous (Bashkirian).

• Fusulina Fisher de Waldheim, 1829 (Type species: Fusulina cylindrica Fischer de Waldheim, 1830). The test is cylindrical with a spirotheca composed of a tectum, diaphanotheca and thick late and early tectoria. The septa are fluted throughout the test but more intensely near the polar ends of the test and the chomata are strong. Late Carboniferous (Moscovian) (Plate 2.8, figs 6, 9; Plate 2.19, Fig. 2).

• Fusulinella von Möller, 1877 (Type species: Fusulinella bocki von Möller, 1878). The test is fusiform with the septa being fluted in the polar region. Chomata are prominent and asymmetrical. Late Carboniferous (Moscovian) (Fig. 2.12, 2.13).

• Hemifusulina von Möller, 1877 (Type species: Hemifusulina bocki von Möller, 1878). The test has its early whorls closely coiled, later they become more loosely coiled. Septa are slightly to moderately fluted. Chomata are rounded and well developed. Middle Carboniferous (Moscovian).

• Obsoletes Kireeva, 1950 (Type species: Fusulina obsoleta Schellwien, 1908). The test is elongate fusiform with tightly coiled early whorls becoming rapidly enlarged in the last whorls. Septa are fluted at the poles. Chomata are well developed and wide. Late Carboniferous (Kasimovian) (Fig. 2.13).

• Paraeofusulina Putrya, 1956 (Type species: Eofusulina (Paraeofusulina) trianguliformis Putrya, 1956). The test is elongate fusiform with strongly fluted septa forming strongly arched loops and poorly differentiated pseudochomata in the early whorls. Late Carboniferous (Moscovian).

• Parafusulinella Stewart, 1970 (Type species: Parafusulinella propria Stewart, 1970). The test is small, fusiform with septa undulating in the central region and slightly fluted towards the apices. Late Carboniferous (Moscovian).

• Profusulinella Rauzer-Chernousova and Belyaev, 1936 (Type species: Profusulinella pararhomboides Rauzer-Chernousova and Belyaev, in Rauzer-Chernousova et al., 1936). The test is fusiform with septa fluted in the polar regions. Chomata are prominent, asymmetrical with secondary thickening extending laterally towards the poles. Late Carboniferous (Middle Bashkirian to Early Moscovian) (Fig. 2.12, 2.13; Plate 2.8, figs 1-4).

• Protriticites Putrya, 1948 (Type species: Protriticites globulus Putrya, 1948). The test is fusiform, and septa are moderately fluted in the axial region. The wall has four-layers, with massive asymmetrical chomata. Late Carboniferous (lattermost Moscovian to Kasimovian) (Fig. 2.13).

• Quasifusulina Chen, 1934 (Type species: Fusulina longissima Moeller, 1878). The test is elongate with poles bluntly rounded. Septa are intensely fluted and may form cuniculi. The wall is very thin, with a tectum and diaphanotheca and a poorly defined tectorium. Chomata are weakly developed, axial filling is heavy. Late Carboniferous (Moscovian) to Early Permian (Sakmarian) (Plate 2.15, Fig. 8; Plate 2.16, figs 3, 5, 8, 10; Plate 2.20, 2-3; Fig. 2.13).

• Skinnerella Coogan, 1960 emend. Skinner, 1971 (Type species: Parafusulina schuberti Dunbar and Skinner, 1937). The test is elongate with strong secondary deposits that may completely fill the chambers of the early whorls and tend to spread away from the axis to outer whorls. Low cuniculi are well developed. Early Permian (Kungurian).

• Wedekindellina Dunbar and Henbest, 1933 (Type species: Fusulinella euthusepta Henbest, 1928). The test is fusiform with straight septa in the central area of the test, but may be slightly fluted toward the extremities. Late Carboniferous (Moscovian).

Superfamily SCHWAGERINOIDEA Vachard, Pille and Gaillot, 2010

The schwagerinoids are large and fusiform to irregularly cylindrical. The test is planispirally enrolled, involute to irregularly coiled. The spirotheca is thick, composed of tectum and alveolar keriotheca (Fig. 2.10E). The septa are fluted in last whorls in primitive genera and completely across the test, forming multiple tunnels or cuniculi in more advanced genera (Fig. 2.14). Late Carboniferous to Permian (Kasimovian to Capitanian).

Family Schwagerinidae Dunbar and Henbest, 1930

The Schwagerinidae have tests that are large, fusiform to sub-cylindrical, planispiral and involute. They have spirothecal structures showing a tectum and a glossy layer, keriotheca (including a finely or coarsely alveolar) phase (Fig. 2.10E). In the most advanced forms, such Polydiexodina Dunbar and Skinner 1931, the folded septa with a reduced single-layer wall have multiple tunnels or cuniculi throughout the test and heavy fillings along the axis (see Fig. 2.14; Plate 2.19; Plate 2.21; Plate 2.22). During the early phase of the keriotheca, the Schwagerinidae possess chomata that range from thin to massive (Fig. 2.14–2.16). The intensity of the folded septa varies from genus to genus. Late Carboniferous to Permian (Kasimovian to Capitanian).

Significant genera include:

• Carbonoschwagerina Ozawa, Watanabe and Kobayashi, 1992 (Type species: Pseudoschwagerina morikawai Igo, 1957). The test is subspherical, septa are closely spaced, and weakly fluted. Chomata are distinct, and massive in inner whorls. This form has a smaller proloculus and much heavier chomata than in Pseudoschwagerina. Late Carboniferous (Gzhelian).

• Chalaroschwagerina Skinner and Wilde, 1965 (Type species: Chalaroschwagerina inflata Skinner and Wilde, 1965). The test is small fusiform, septa are strongly fluted in the early part, but merely wavy in the late part. Weak chomata occur on the proloculus. Early Permian (Artinskian to Kungurian).

• Chusenella Hsu, 1942 (Type species: Chusenella ishanensis Hsu, 1942). The test is fusiform, and later septa are tightly fluted. The wall has a tectum and keriotheca. Chomata may be lacking throughout but axial filling is prominent. Early to Middle Permian (Sakmarian to Capitanian) (Fig. 2.13).

• Cuniculinella Skinner and Wilde, 1965 (Type species: Cuniculinella tumida Skinner and Wilde, 1965). The test is fusiform, and septa are intensely fluted with folds reaching the top of the septa. Those of adjacent septa join to form chamberlets. Chomata are present only on the proloculus. Permian (Artinskian to Kungurian).

• Daixina Rozovskaya, 1949 (Type species: Daixina ruzhencevi Rozovskaya, 1949). The test is fusiform, having septa with strong and irregular fluting. Chomata are rarely present in the early whorls. Late Carboniferous to Early Permian (Gzhelian to Asselian)

• Dunbarinella Thompson, 1942 (Type species: Dunbarinella ervinensis Thompson, 1942). The test is fusiform with a straight axis of coiling. It differs from Schwagerina and Pseudofusulina in the presence of heavy axial fillings and distinct chomata. Late Carboniferous to Early Permian (Gzhelian to Sakmarian) (Plate 2.8, Fig. 5; Plate 2.16, Fig. 11; Plate 2.23, Fig. 8; Plate 2.24, Fig. 4).

• Eoparafusulina Coogan, 1960 (Type species: Fusulina gracilis Meek, 1864). The test is subglobular, tightly coiled and expanding uniformly. Late parts of the septa are planar, while early parts are strongly fluted throughout the test, producing chamberlets, with low tunnel bordered by pseudochomata. Rudimentary chomata are present in in first whorls. Early Permian (Late Asselian to Artinskian and?Kungurian).

• Eopolydiexodina Wilde, 1975 (Type species: Polydiexodina afghanensis Thompson, 1946). Septa are intensely fluted throughout the entire length and in all whorls of the test, with cuniculi well developed. It differs from Polydiexodina in lacking a median tunnel and in having only sporadic supplementary tunnels. Permian (Middle Guadalupian).

• Monodiexodina Sosnina, 1956 (Type species: Schwagerina wanneri var. sutschanica Dutkevich, 1934). The test is subcylindrical, with septal fluting with folds restricted to the early part, forming low cuniculi. Early whorls have chomata. Early to Middle Permian (Cisuralian to Late Guadalupian).

• Montiparus Rosovskaya, 1948 (Type species: Alveolina montipara Ehrenberg, 1854, emend. Möller, 1878). The test is fusiform. Septa are irregularly plicate. Chomata are sub-quadrate. Carboniferous (Kasimovian) (Fig. 2.13; Plate 2.8, Fig. 7).

• Parafusulina Dunbar and Skinner, 1931 (Type species: Parafusulina wordensis Dunbar and Skinner, 1931). The test is very large with intense folding but with no chomata (Fig. 2.9). Middle to Late Permian (Guadalupian to Lopingian) (Fig. 2.14; Plate 2.16, figs 1-2; Fig. 2.19, figs 5, 7).

• Paraschwagerina Dunbar and Skinner, 1936 (Type species: Schwagerina gigantea White, 1932. The test is fusiform to subspherical. The septa are distinctly fluted throughout test, with high, strong, and parallel sided folds. The wall is thin, composed of thin tectum and coarsely alveolar keriotheca. The chomata are small in the first whorls, but absent in the later whorls. Early Permian (Asselian to Sakmarian) (Plate 2.17, figs 4,6; Plate 2.18, Fig. 10).

• Polydiexodina Dunbar and Skinner, 1931 (Type species: Polydiexodina capitanensis Dunbar and Skinner, 1931). Septa are numerous, regularly and strongly fluted, with narrow septal folds, the fluting of adjacent septa producing cuniculi where in contact. Permian (Late Guadalupian) (Fig. 2.14; Plate 2.17, Figs 3, 7).

• Pseudofusulina Dunbar and Skinner, 1931 (Type species: Pseudofusulina huecoensis Dunbar and Skinner, 1931). The test is fusiform with a large proloculus and fluted septa, most strongly towards the poles where it is densely folded to produce chamberlets. Late Carboniferous to Permian (lattermost Gzhelian to Kungurian) (Fig. 2.13; Plate 2.18, Fig. 13).

• Pseudoschwagerina Dunbar and Skinner, 1936 (Type species: Schwagerina uddeni Beede and Kniker, 1924). The test has inflated outer whorls and strongly folded septa throughout (Fig. 2.9). Chomata are well marked at juvenile stage only. They are rudimentary or lacking in the final stage. Early to Middle Permian (Asselian to Kungurian) (Fig. 2.14; Plate 2.19, Fig. 5B; Plate 2.18, figs 3, 4, 8, 11; Pate 2.22, figs 3, 5).

• Schwagerina von Mölle, 1877 (Type species: Borelis princeps Ehrenberg, 1842). Septa are regularly and intensely fluted, and folds of septa divide the early part of the chambers into small chamberlets. Chomata are weak or absent, with narrow tunnels. Early Permian (Sakmarian to Artinskian) (Plate 2.15, Fig. 5; Plate 2.16, figs 4,6; Plate 2.17, Fig. 5A; Plate 2.20, figs 4, 5; Plate 2.23, figs 3-4, 6; Plate 2.24, Fig. 6; Plate 2.25, figs 1, 3-4).

• Skinnerina Ross, 1964 (Type species: Skinnerina typicalis Ross, 1964). Septa are numerous and intensely fluted from pole to pole, resulting in high and well developed cuniculi. Permian (Early Guadalupian).

• Taiyuanella Zhuang, 1989 (Type species: Taiyuanella subsphaerica Zhuang, 1989). The test is fusiform, with spirotheca composed of a tectum and stalactotheca. Septa are strongly fluted throughout the length of the test. Chomata are weakly developed. Middle Permian (Wordian) (Fig. 2.13).

• Triticites Girty, 1904 (Type species: Miliolites secalicus Say, in James, 1823). Septa are weakly folded over the equator, with massive chomata (Fig. 9). Late Carboniferous to Early Permian (Kasimovian to Sakmarian) (Fig. 2.14–2.16; Plate 2.8, Fig. 10; Plate 2.15, figs 1-3, 7; Plate 2.20, Fig. 6; Plate 2.25, figs 5-6; Plate 2.11, figs 1-2).

• Zellia Kahler and Kahler, 1937 (Type species: Pseudoschwagerina (Zellia) heritschi Kahler and Kahler, 1937). The test is globular with a thick wall composed of a tectum and coarsely alveolar keriotheca, with low, widely spread chomata. Permian (Sakmarian) (Plate 2.25, Fig. 2).

Superfamily NEOSCHWAGERINOIDEA Solovieva, 1978

Found in the Late Permian showing the last stage of the keriotheca phase, these Neoschwagerinoids are different from the fusulinoids in having a wall made of a tectum and a glossy layer with alveoles running through it (the keriotheca) (Fig. 2.10E), and they differ from the schwagerinoids in having secondary layers and multiple chomata or parachomata, with multiple tunnels. Permian (Asselian to Wordian).

Family Verbeekinidae Staff and Wedekind, 1910

The test is globose with discontinuous parachomata. Permian (Asselian to Early Lopingian).

• Verbeekina Staff, 1909 (Type species: Fusulina verbeeki Geinitz, in Geinitz and von der Marck, 1876). Spherical with a tiny proloculus, parachomata are absent or rudimentary (Fig. 2.9). Permian (Roadian to Capitanian) (Fig. 2.9, 2.14; Plate 2.15, Fig. 9).

• Brevaxina Schenck and Thompson, 1940 (Type species: Doliolina compressa Deprat, 1915). The test is subspherical with a small proloculus, followed by an endothyroid early stage. The wall is thick, with tectum, high and broad parachomata and numerous foramina. Permian (Kungurian to early Wordian).

Family Misellinidae Miklukho-Maklay, 1958

The test is fusiform with well-developed parachomata. Permian (Sakmarian to Early Lopingian).

• Misellina Schenck and Thompson, 1940 (Type species: Doliolina ovalis Deprat, 1915). The test is ellipsoidal with numerous chambers, and numerous plane, unfluted septa. The keriotheca is thick and the parachomata are “saddle-shaped”. Permian (Kungurian to early Wordian).

Family Pseudodoliolinidae Leven, 1963

Ellipsoidal test with planar septa and narrow, high parachomata. The parachomata supplement the primary pair of ridges in the equatorial zone of the shell laterally and polewards, regularly intercalated between supplementary tunnels until the polar end of the chamber Permian (Artinskian to Capitanian).

• Pseudodoliolina Yabe and Hanzawa, 1932 (Type species: Pseudodoliolina ozawai Yabe and Hanzawa, 1932). The test is bluntly circular with thin wall and reduced keriotheca. Septa are planar and unfluted. The parachomata are high, continuous and may reach the top of the chambers adjacent to the septa, appearing as septula in axial section. Permian (Artinskian to Capitanian).

Family Neoschwagerinidae Dunbara and Condra, 1927

In the Neoschwagerinidae (see Fig. 2.11) the finely alveolar keriotheca join the parachomata to form transverse septula which form complete partitions in the chamber. They occur in the Permian (Asselian to Early Lopingian).

• Cancellina Hayden, 1909 (Type species: Fusulina (Neoschwagerina) primigena Hayden, 1909). The test is fusiform and broadly inflated in the centre. The septa are arcuate and broad transverse septula are formed by the extension of the lower part of the keriotheca. Broad parachomata are present. Early to Late Permian (Roadian to Capitanian).

• Neoschwagerina Yabe, 1903 (Type species: Schwagerina craticulifera Schwager, 1883). The test is subspherical. Chambers are divided into chamberlets by transverse septula of first order which connect with the poorly developed parachomata. The wall consists of a tectum and a very thick keriotheca that may be differentiated into an upper finely alveolar layer and a lower layer with fewer and coarser alveoli and with upper and lower tectoria. Early to Middle Permian (Wordian to Capitanian) (Plate 2.20, Fig. 1; Plate 2.23, figs 1-2; Plate 2.26, figs 5-7).

• Sumatrina Volz, 1904 (Type species: Sumatrina annae Volz, 1904). The test is elongate fusiform with a large proloculus and long, thin septa. Up to four short secondary septula develop between primary septula. The wall is very thin with a thin keriotheca. Massive parachomata and numerous foramina occur throughout the length of the test. Middle Permian (Capitanian).

• Yabeina Deprat, 1914 (Type species: Neoschwagerina (Yabeina) inouyei Deprat, 1914). The test is large and fusiform with well-developed parachomata connected to the early ends of the septula. Up to three secondary spiral septula in outer whorls and nine axial septula per chamber. Middle Permian (Guadalupian) (Fig. 2.14; Plate 2.24, figs 1-2).

ORDER LAGENIDA DELAGE AND HÉROUARD, 1896

As an order the Lagenida are characterised by having walls made of orientated calcite crystals. It appears that this characteristically evolved independently, in parallel several times in the Palaeozoic, and that the superfamilies Robuloidoidea and Nodosinelloidea have different phylogenetic roots (see Fig. 2.1).

Members of this order (Late Silurian to Holocene) were referred to the Nodosariida Calkins 1926 by Vachard et al. (2010) and Mikhalevich (2013). They have alveoles. Primitive taxa are without secondary lamination, more advanced forms have a secondary lamination. The wall structures in selected lagenides have been restudied recently (Groves et al., 2004). “Monolamellar” refers to the primary single-layered septal wall. In many lagenides, secondary lamellar develop, where extensions of the primary wall of a given chamber overlap some of the previous chambers. Reiss (1963) proposed an extensive reclassification of lamellar forms, and was among the first to note that radial walls are secreted on an organic substrate. Grønland and Hansen (1976) by examining Holocene lagenides developed a new terminology for lamellarity:

• “Atelo-monolamellar” refers to forms that lack secondary lamellarity and they represent the earliest lagenides, e.g. the simple Nodosaria (Plate 2.5, Fig. 2.11);

• “plesio-monolamellar” refers to forms in which secondary lamellarity envelops some but not all previous chambers;

• “ortho-monolamellar” refers to forms in which the primary wall of each chamber secondarily envelops all previous chambers; and

• “poly-monolamellar” refers to forms in which yet another lamella secondarily covers the entire test.

The plesio-monolamellar forms are thought to have originated in Early Jurassic, followed by ortho-monolamellar and poly-monolamellar types in the Cretaceous and Cenozoic (Groves et al., 2004).

Superfamily ROBULOIDOIDEA Reiss, 1963

The tests of members of this superfamily are those of a typical Lagenida, but without secondarily lamellar or with slight lamination in younger taxa. The aperture is primitive and cylindrical. Late Silurian to Early Cretaceous.

Family Syzraniidae Vachard, 1981

The test has two chambers and is elongate, composed of a proloculus and an undivided or subseptate tubular chamber partitioned to varying degrees by internal thickenings of the wall, which is calcareous and may have two layers. Late Silurian to Late Permian (Pridolian to Lochkovian; Tournaisian to Changhsingian).

• Amphoratheka Mamet and Pinard, 1992 (Type species: Amphoratheka iniqua Mamet and Pinard, 1992). The septa are slightly more pronounced than in Tezaquina. Late Carboniferous to Early Permian (Gzhelian to Sakmarian).

• Rectostipulina Jenny-Deshusses, 1985 (Type species: Rectostipulina quadrata Jenny-Deshusses, 1985). The test consists of a long narrow tapering tube with a polygonal transverse section. Septa are short and nearly perpendicular to the test wall. The wall is atelo-monolamellar. The aperture is a simple opening in the center of the apertural face. Late Permian (Wuchiapingian).

• Syzrania Reytlinger, 1950 (Type species: Syzrania bella Reytlinger, 1950). The test has a spherical proloculus followed by a non-septate tubular chamber. The wall has a “fibrous” layer and an inner microgranular layer. The aperture is a simple terminal opening. This genus may have evolved from another simple robuloidoid or most likely from the earlandiides Earlandia in Middle Pennsylvanian. Carboniferous to Late Triassic (Moscovian to Rhaetian) (Fig. 3.3).

• Tezaquina Vachard, 1980 (Type species: Tezaquina clivuli Vachard, 1980). Groves in Groves and Boardman (1999) emended Tezaquina to include nonseptate syzraniid forms whose tubular second chamber is subdivided into pseudo-chambers by weak internal thickenings of the wall, which is thick hyaline fibrous and finely perforate. The test consists of a spherical proloculus followed by a second chamber that is partitioned into elongate, subcylindrical pseudo-chambers by weak internal thickenings of the wall. The aperture is a simple terminal opening, and the wall is atelo-monolamellar. Carboniferous to Late Triassic (late Moscovian to Rhaetian) (Fig. 3.3).

• Tuborecta Pronina, 1980 (Type species: Tuborecta vagranica Pronina, in Petrova and Pronina, 1980). The test is small, forming a long narrow tube. The wall is calcareous, of two layers, a thin inner finely granular, dark layer and a thick radial, light outer layer. The aperture is terminal, at the end of the tube. Late Silurian to Early Devonian (Pridolian to Lochkovian).

Family Protonodosariidae Mamet and Pinard 1992

This family includes the uniserial Lagenida with atelo-monolamellar or plesio-monolamellar wall structures consisting of radial-fibrous forms. Late Carboniferous to Late Permian (Kasimovian to Changhsingian).

• Protonodosaria Gerke, 1959 (Type species: Nodosaria proceraformis Gerke, 1952). The test is elongate, circular in thin section. Sutures are horizontal and straight. The aperture is round. Late Carboniferous to Late Permian to Late Triassic (Kasimovian to Rhaetian).

Family Ichtyolariidae Loeblich and Tappan, 1986

The test is elongate uniserial with a single layered wall, and may show some secondary lamination, with a simple terminal aperture. Early Permian to Early Cretaceous (Artinskian to Albian).

• Pseudotristix Miklukho-Maklay, 1960 (Type species: Tristix (Pseudotrislix) tcher-dvnzevi Miklukho-Maklay, 1960). The test consists of a spherical proloculus followed by a uniserial arrangement of triangular chambers. The sutures are nearly horizontal, the aperture is a simple opening, and the wall is atelo-monolamellar. Late Permian (Lopingian).

Family Robuloididae Reiss, 1963

The test is uniserial, enrolled with an atelo-monolamellar wall and a terminal aperture. Middle Permian to Late Jurassic (Guadalupian to Kimmeridgian).

• Calvezina Sellier de Civrieux and Dessauvagie, 1965 (Type species: Calvezina ottoman Sellier de Civrieux and Dessauvagie, 1965). The test consists of a proloculus followed by five or more uniserial chambers that are weakly trochospirally enrolled in the early stage, later becoming rectilinear. The chambers are slightly compressed laterally and their size increases rapidly from early hemispherical ones to later lobate or irregular ones. The aperture is a simple terminal opening accompanied by a slight thickening of the wall. Late Permian (Lopingian).

• Eocristellaria Miklukho-Maklay, 1954 (Type species: Eocristellaria permica Miklukho-Maklay, 1954). The test is planispirally enrolled with chambers increasing rapidly in breadth, but slowly in height, so the apertural face extend back towards the proloculus. The calcareous wall is composed of an outer hyaline radiate layer and an inner dark microgrannular layer. Late Permian (Lopingian).

• Robuloides Reichel, 1946 (Type species: Robuloides lens Reichel, 1946). The test is planispiral with oblique septa, arching back towards the periphery. Permian to Middle Triassic (Artinskian to Ladinian).

Superfamily NODOSINELLOIDEA Rhumbler, 1895

The Nodosinelloidea evolved from the Earlandiida which in turn evolved from the Parathuramminoidea (see Fig. 2.2). Their tests were free, with more than one distinct chamber, partially septate or fully septate, with a simple microgranular wall or evolving into two layers with an extra fibrous inner layer. The Nodosinelloidea range from the Late Silurian to the Permian.

Family Earlandinitidae Loeblich and Tappan, 1988

The Earlandinitidae (Fig. 2.2) evolved from an earlandioid, Caligellidae, by becoming elongate, uniserial, semi-septate, with a simple microgranular wall, and range from Late Devonian to Late Carboniferous (Bashkirian).

• Darjella Malakhova, 1964 (Type species: Darjella monilis Malakhova, 1964). The test is uniserial, with globular chambers and strongly constricted sutures. Early Carboniferous (Early Visean).

• Earlandinita Cummings, 1955 (Type species: Nodosinella perelegans Plummer, 1930). The test is subseptate with a terminal round aperture. Carboniferous (See Fig. 2.2).

• Lugtonia Cummings, 1955 (Type species: Nodosinella concinna Brady, 1876. The test is rectilinear with depressed chambers overhanging the sutures. Carboniferous.

Family Nodosinellidae Rhumbler, 1985

This family (Fig. 2.2) evolved from the Earlandioidea, and is characterised by having a uniserial test which develops in the Late Carboniferous to having a double layered wall, with an inner fibrous layer and an outer microgranular layer. They range from the Late Silurian to Permian.

• Biparietata Zolotova, 1980 (Type species: Biparietata ampula Zolotova, 1980). The test is rectilinear with elongate chambers and sutures not apparent externally. Middle Permian (Guadalupian).

• Nodosinella Brady, 1876 (Type species: Nodosinella digitata Brady, 1876). The test is free and fully septate. Permian (see Fig. 2.2, 2.4).

Superfamily COLANIELLOIDEA Fursenko, 1959

In this superfamily, the tests are uniserial and fully septate, and chambers are strongly overlapping with partitions. The walls occur with an outer vitreous layer and an inner granular layer. They range over the Late Devonian to Late Permian.

Family Colaniellidae Fursenko, 1959

The family evolved from the Nodosinellidae by developing strong internal partitions or septa (Fig. 2.2). They range from the Late Devonian to Late Permian.

• Colaniella Likharev, 1939 (Type species: Pyramis parva Colani, 1924). Elongate and uniserial, having a calcareous wall with a fibrous structure with strongly overlapping sutures. The aperture is terminal radiate. Late Permian (see Fig. 2.2).

• Cylindrocolaniella Loeblich and Tappan, 1985 (Type species: Wanganella ussuriensis Sosnina, in Kiparisova et al., 1956). The test is slightly arcuate, narrow and cylindrical in form with numerous short septa. Late Permian (Lopingian).

• Pseudowanganella Sosnina, 1983 (Type species: Pseudowanganella tenuitheca Sosnina, 1983). The test is narrow, flattened and elliptical in cross section. Chambers are curved centrally and overlapping near the periphery. Septa are more thickened near the aperture. Late Permian (Lopingian).

Superfamily GEINITZINOIDEA Bozorgnia, 1973

The tests of this superfamily are uniserial and appear similar to the Nodosinelloidea, but the microgranular layer is the inner dark layer while the radially fibrous layer is an outer layer. Advanced forms have secondary lamellarity. Late Devonian to Middle Triassic.

Family Geinitzinidae Bozorgnia, 1973

The tests are free and uniserial. They evolve from species that are rounded in cross-section to species that are compressed with a terminal aperture. The range is Late Devonian to Late Permian.

• Geinitzina Spandel, 1901 (Type species: Geinitzella (Lunucammina) permiana Spandel, 1898). The test is uniserial elongate, laterally compressed, exhibiting in transverse view two planes of bilateral symmetry and an oval aperture in the centre of the apertural face. Permian (Cisuralian to Lopingian) (Fig. 3.3 and 3.4).

• Howchinella Palmieri, 1985 (Type species: Frondicularia woodwardi How-chin, 1895). The test is laterally compressed, with arched or chevron sutures. The wall is calcareous granular with a dark thin inner organic layer and an outer hyaline layer of optically radial calcite. The aperture is radiate with a tooth. Early Permian (Sakmarian).

• Nodosinelloides Mamet and Pinard, 1992 (Type species: Nodosinelloides potievskayae Mamet and Pinard, 1996 (for Nodosaria gracilis Potievskaya, 1962 preoccupied)). The test consists of a spherical proloculus followed by as many as 10 or more uniserial circular chambers with pronounced septal inflection and a simple aperture. The wall lacks secondary lamellarity (atelo-monolamellar) and the aperture is a simple terminal opening. Carboniferous to Triassic (Kasimovian to Rhaetian) (Fig. 3.3).

Family Pachyphloiidae Loeblich and Tappan, 1984

This family has a free uniserial compressed test with a simple calcareous microgranular wall, but with secondary thickenings on both sides of the test. The range is from Early to Late Permian.

• Pachyphloia Lange, 1925 (Type species: Pachyphloia ovata Lange, 1925). The test consists of a globular proloculus followed by a uniserial succession of laterally compressed, strongly overlapping chambers. The wall is plesio- or ortho-monolamellar with thin dark inner layer and thicker hyaline outer layer. The aperture is terminal, oval, radiate (toothed). Early Permian to Late Permian (Sakmarian to Changhsingian) (Plate 2.5, figs 2-4; Fig. 3.3).

ORDER MILIOLIDA DELAGE AND HÉROUARD, 1896

The miliolides have tests that are porcellaneous and imperforate, which are made of high Mg-calcite with fine randomly oriented crystals. They range from the Carboniferous to the Holocene.

Superfamily CORNUSPIROIDEA Schultze, 1854

The test is free or attached, and composed of a globular proloculus followed by a tubular enrolled chamber. The coiling is planispiral or trochospiral, evolute or involute and may become irregular. There is a simple aperture at end if the tube. Early Carboniferous to Holocene.

Family Cornuspiridae Schultze, 1854

The tests are free or attached, composed of a proloculus followed by an undivided planispiral to streptospiral, involute or evolute second tubular chamber. Early Carboniferous (Visean) to Holocene.

• Cornuspira Schultze, 1854 (Type species: Orbis foliaceus Philippi, 1844). The test is discoidal with a globular proloculus and undivided planispirally enrolled and tubular second chamber. Carboniferous (Moscovian) to Holocene.

• Rectocornuspira Warthin, 1930 (Type species: Rectocornuspira lituiformis Warthin, 1930). Test elongate, with a globular proloculus and planispirally enrolled to later uncoiling and rectilinear undivided tubular second chamber. Late Carboniferous to Early Triassic (Moscovian to Induan).

Family Hemigordiidae Reitlinger, 1993

Characterized by a test with at least an early trochospiral stage, but that may later become streptospiral. Carboniferous.

• Hemigordius Schubert, 1908 (Type species: Cornuspira schlumbergeri Howchin, 1895). The test has early whorls that are streptospiral, later becoming planispiral and evolute. Early Carboniferous to Late Carboniferous (Late Visean to Gzhelian) (Fig. 3.5).

• Neohemigordius Wang, 1973 (Type species: Neohemigordius maopingensis Wang and Sun, 1973). The test is lenticular, with a lamellar thickening over the umbilical area. Early Permian (Asselian to Sakmarian) (Fig. 3.5).

• Pseudovidalina Sosnina, 1978 (Type species: Pseudovidalina Sosnina, 1978). The test is discoid with a globular proloculus followed by planispirally, evolute, enrolled tubular undivided second chamber with secondary thickenings in the umbilical region. Middle Permian (Kungurian).

Family Hemigordiopsidae Nikitina, 1969

Forms in this family have a test with at least an early streptospiral stage, that later may be planispiral. Early Carboniferous (Visean) to Holocene.

• Hemigordiopsis Reichel, 1945 (Type species: Hemigordiopsis renzi Reichel, 1945). The test is globular with a proloculus followed by an undivided tube, which is strongly involute. Early to Middle Permian (Guadalupian) (Fig. 3.3).

• Shanita Brönnimann, Whittaker and Zaninetti, 1978 (Type species: Shanita amosi Brönnimann, Whittaker and Zaninetti, 1978). The proloculus followed by an undivided enrolled second chamber, with a streptospiral in early coiling, becoming planispiral and involute. Later whorls with alternating vertical pillars. Late Permian (Wuchiapingian to Changhsingian) (Fig. 3.6).