New insight into the systematics and evolution of the foraminifera

New insight into the systematics and evolution of the foraminifera Valeria I. Mikhalevich Zoological Institute, Russian Academy of Sciences, Universitetskaya naberezhnaya, 1, St. Petersburg, Russia, 199134 email: mikha07@mail.ru; vm38600@gmail.com ABSTRACT: A new suprageneric classification of the Foraminifera is here presented based on fundamentally new concepts of their evolution and classification. The predominant significance is given to the shell morphology as the most conservative feature, while the wall composition and shell wall ultrastructure are considered as having important but nevertheless subordinate meaning. Foraminifera are regarded as a phylum that includes five classes: Astrorhizata Saidova 1981, Spirillinata Mikhalevich 1992, Miliolata Saidova 1981, Nodosariata Mikhalevich 1992 and Rotaliata Mikhalevich 1980. Each of the five classes unite forms that can be characterized by a complex of common features of their shell morphology reflecting the building plan of the organism (number of chambers, their form, the predominant mode of coiling, position and character of the aperture and its inner structures, the presence or absence of additional apertures, the presence or absence of integrative systems and their peculiarities, and some other features – all of them having evolutionary significance). Each of these classes represents independent and well-outlined phyletic lines. Some characters of the cell structure and nuclear apparatus are also used as taxonomic features of some higher-ranking taxa where the accumulated data permit. Isomorphic agglutinated forms differing from their calacareous analogues in their shell wall composition are separated as subclasses within the appropriate classes: the subclasses Ammodiscana Mikhalevich 1980, Miliamminana Mikhalevich 1980, Hormosinana Mikhalevich 1992, Textulariana Mikhalevich 1980 within the Spirillinata, Miliolata, Nodosariata and Rotaliata correspondingly (the latter also includes two calcareous subclasses – the Rotaliana Mikhalevich 1980 and Globigerinana Mikhalevich 1980). Within the class Astrorhizata, subclasses with organic (Lagynana Mikhalevich 1980) and agglutinated (Astrorhizana Saidova 1981) shell walls are included. In total, the phylum Foraminifera embraces 73 orders, 27 suborders, 98 superfamilies, 499 families and 368 subfamilies among which 2 orders (Cymbaloporida, Cassigerinellida), one suborder Duostominina, two families (Cymbaloporettidae, Haynesinidae), and two subfamilies (Cushmanellinae and Tristixinae) are described as new. The composition of the classes and subclasses is also partially revised. The largest changes were made within the classes Spirillinata, Miliolata and Nodosariata: thus the Fusulinids were included into the Miliolata, the Chapmaninids - into the Spirillinata, the Stilostomellids, Pleurostomellids and Paleozoic Nodosariids – into the Nodosariata. The former suborder Textulariina (= Textulariacea Ehrenberg 1838 sensu lata Loeblich and Tappan 1987) was shown to be heterogenous and its representatives are split out into several subclasses of the different classes according to their shell morphology. The composition of the subclasses is here given up to the family level; most of the subclasses need further revision at the family and generic level. Under the new approach the morphologically similar agglutinated and calcareous shells within each class could be regarded as closely related rather than convergent forms. The rise and development of the classes took place independently and in parallel in each of the phylogenetic lines examined. Key words: Foraminifera, taxonomy, shell morphology, wall ultrastructure, nuclear apparatus, evolution. INTRODUCTION Since the end of the 1970s and beginning of 1980s, two strong scientific schools – the American school with the leadership of Levine and the Russian school in the Laboratory of Protozoology of the Zoological Institute, Russian Academy of Sciences (the latter uniting the group of scientists including M. Krylov, Ja. Starobogatov, L. Seravin, I. Raikov, S.Podlipaev, S. Shulman and some others) began the revision of the previous phylum Protista and split it into 7 or 9 phyla. The previous Sarcodina were also split into several groups (Krylov et al. 1980). Thus the heterogeneity of the former Sarcodina was already stated, and later confirmed by the molecular data (Nikolaev et al. 2004, Pawlowski and Burki 2009, Burki et al. 2010, Adl et al. 2012). The first attempt of the revision of the foraminiferal classification within the framework of the Russian school was performed by the author of this article (Mikhalevich 1980a) and the task of reclassification was later continued with subsequent profound changes (Mikhalevich 1981, 1992, 1998, 1999, 2000, 2004, 2009). In the process of revision it became obvious that the strict adherence to the principle of the predominant significance of the shell wall composition which I initially tried to follow gives an absurd result. Consequently, the first step toward uniting agglutinated and calcareous forms of similar morphology was made, resulting in the separation of the class Rotaliata Mikhalevich 1980. Following the author’s initial report in Baku (Mikhalevich 1980b) the only specialist who accepted the new approach was Kh. Saidova, who made the next step in this direction, uniting agglutinated and calcareous isomorphs in the class Miliolata and unilocular forms in the class Astrorhizata (Saidova 1981). Later the new concept was actively supported (in chronological order) by G.P. Nestell, J.-P. Debenay, M. McGann, and M.A. Kaminski. Micropaleontology, vol. 59, no. 6, plate 1, text-figs. 1–10, table 1, pp. 493–527, 2013 493 V. I. Mikhalevich: New insight into the systematics and evolution of the foraminifera During the next decade, the comparative morphological analysis of the multitude of ancient and Recent foraminiferal forms allowed me to outline a new macrotaxonomy of the phylum Foraminifera. The phylum now consists of 5 classes (Astrorhizata Saidova 1981, Spirillinata Mikhalevich 1992, Miliolata Saidova 1981, Nodosariata Mikhalevich 1992, and Rotaliata Mikhalevich 1980) each representing an independent and well-outlined phyletic line (Plate 1). The majority of the lower-ranking groups within these classes still awaits further investigations and revisions. DISCUSSION In the widely adopted previous classification systems, in which predominant taxonomic significance was given to the character of the shell wall, the multitude of the agglutinated representatives uniting all the possible types of the foraminiferal shell structure were placed within the superfamily Textulariacea including the astrorhizid forms as well (Loeblich and Tappan 1987). This group was later elevated to the order rank – Textulariina (Loeblich and Tappan 1992). However, in spite of some changes made in the later publications, and the elevation of rank of some taxa, the same principal approach that stressed the significance of the shell wall composition was preserved. In the Loeblich and Tappan (1992) classification, the astrorhizids were separated into a separate suborder, those with an organic wall were grouped into the suborder Allogromiina, and all the calcareous forms were separated into 9 different suborders. All the enumerated suborders were given the equal taxonomic rank (without their subordinate ranking). In this scheme, all the shell types included into Textulariina were morphologically isomorphic to representatives of the nine calcareous suborders. In the new classification scheme offered here the composition of the former Textulariacea is outlined with a dashed line (Plate 1). Thus the groups with a mirror-like appearance are evident (see below). Their morphological similarity was regarded previously as a result of evolutionary convergence. At this point it would be appropriate to stress that without the outstanding labour of the classics of foraminiferal taxonomy (Loeblich and Tappan 1987) who thoroughly demonstrated all the multitude of foraminiferal shells in a single volume (where the images of genera could be seen page by page without breaks) and provided the clearest picture of their diversity and changes through time, it would be hardly possible to make the next crucial step and elaborate the new macrotaxonomy of this group. The results of the comparative analysis of shell morphology of different ancient and recent foraminiferal groups as well as their shell wall ultrastructure and of some of their cytological features are discussed below. COMPARATIVE ANALYSIS OF DIFFERENT FORAMINIFERAL GROUPS Similarity of the shells in the subclasses Lagynana (=Allogromiina) and Astrorhizana Shells belonging to these subclasses have similar morphology: sphaerical (Text-fig. 1, figures 2– with reticulopodia, 9), hemisphaerical (Text-fig. 1, figures 1, 8), oval (text-fig. 1, figures 4, 5, 11, 12), with an elongated neck (Text-fig. 1, figures 3; 10), elongated and tubular (Text-fig. 1, figures 7 – with reticulopodia, 6; 13, 14). Each of such forms may lack an aperture (text-fig. 1, figures 1, 8), may have two apertures (text-fig. 494 5, figures 5 - with reticulopodia, 12) or multiple apertures (Text-fig. 1, figures 2 – with reticulopodia, 9), the aperture may be at the end of an elongated neck (text-fig. 1, figures 3, 10). This profound similarity is especially demonstrative in the shells having an entosolenian tube (text-fig. 1, figures 4, 11). The organic (tectinous) shell wall of the lagynids (=allogromiins) was considered to be their only distinctive diagnostic feature. Nevertheless, their shell wall may also be partially covered by sand grains (text-fig. 1, figure 1). And, vice-versa, noncovered areas of the organic wall may be present in some astrorhizids (Vanhoeffenellinae Saidova 1981). Such a profound morphological similarity provides the basis to unite them into a single class: the Astrorhizata (Mikhalevich 1999, 2000). Their close genetic relationship was also confirmed by molecular data (Pawlowski 2000, 2005). Similarity of shells with agglutinated and calcareous walls in the classes Nodosariata, Spirillinata and Rotaliata In the other foraminiferal classes there are many examples of morphological isometry. In the Nodosariata - unilocular (text-fig. 2, figures 1, 3-2, 4), pseudomultichambered (text-fig. 2, figures 5–6), multichambered uniserial throughout (text-fig. 2, figures 7–8) or in the final part only (text-fig. 2, figures 9–10) (in the latter case with V-shaped chambers, characteristic for this class only). The polymorphinoid shell type (text-fig. 2, figures 11, 12–13, 14, 15) is also present in this class only, in both of its subclasses. The isomorphic forms of both subclasses of the class Spirillinata are seen in text-fig. 2, figures 16–17 and 18–19 : 16–17 – conical forms with a long tubular second chamber, 18–19 – multichambered forms with chambers forming a high spire with one (or a little more than one) chamber per whorl. The parallel rows of isomorphic shells of both of the Rotaliata subclasses can be seen in text-fig. 2, figures 20, 22, 24–21, 23, 25: trochoid (20–21), planispiral (22–23), and triserial (24–25). In contrast to the shells of the two previous classes (as well as of the Miliolata) the aperture in the rotaliata is positioned at the base of the septal face of the last chamber (at least initially) rather than having a terminal position. Similarity of shell form in representatives with agglutinated and calcareous shell wall in the class Miliolata In the class Miliolata such similarity is apparent even more visually. Two rows of forms (agglutinated and porcelaneous) repeat each other in a nearly mirror-like way with respect to both chamber-form and their number per whorl (two tubular chambers per whorl in figures 1–3; 7–9 of Text-fig. 3, three widely-tubular chambers in the final whorls (text-fig. 3, figures 4; 10) (in spite of the distinctions of their early stages which are quinqueloculine in Pseudoflintina Saidova 1981 and triloculine in Flinitina Cushman 1921), multiple wide and slightly elongated in Text-fig. 3, figure 5; 11, multiple and very wide in Text-fig. 3, figures 6; 12). The similarity is also apparent in their mode of coiling: in more than 5 planes (text-fig. 3, figures 1, 7), in 5 planes (quinqueloculine type – text-fig. 3, figures 2, 8), sigmoiline coiling (text-fig. 3, figures 3, 9), fully planispiral (text-fig. 3, figures 6, 12) and combined glomerate and planispiral types (text-fig. 3, figures 4, 10 and 5, 11). Examples with triloculine (Trilocularena Loeblich and Tappan 1955, Falsagglutinella Loeblich and Tappan 1994 – Triloculina d’Orbigny 1826) or spiroloculine types could also be added. Micropaleontology, vol. 59, no. 6, 2013 PLATE 1 Scheme of the composition of the phylum Foraminifera d’Orbigny 1826 (the right dotted line marks the border between subclasses within the same class, the curved dotted line encircles the taxa which were united in the former Textulariacea, not including Lagynana). 495 V. I. Mikhalevich: New insight into the systematics and evolution of the foraminifera only for the two subclasses (Miliolana and Miliamminana) of the class Miliolata. Position of the aperture in the different foraminiferal classes In the complex of diagnostic features not only the number of chambers, their form and arrangement are important but especially their aperture, its position and additional structures (especially the inner apertural structures), the presence or absence of additional apertures, of the inner integrative systems (apertural integrative systems, stolons, canals – Mikhalevich 1981, 1992; Mikhalevich and Debenay 2001) because of the great functional importance of these structures providing better communication between the chambers and communication of the whole cell with the environment. The position of the aperture is connected with the building plan structure of the whole shell. In multichambered shells of the classes Miliolata (text-fig. 4, figures 1, 2, 3, 7), Spirillinata (text-fig. 4, figure 6), and Nodosariata (text-fig. 4, figure 4) the aperture is terminal. In the Rotaliata the building plan of the shell underwent profound changes and their aperture has initially a basal position (Text-fig. 4, figures 5, 9), at the base of the final chamber, though in some genera it may become terminal in the later stages of development. Just the position of the aperture (along with some other features) permits the easy distinction between the Nodosariata and Rotaliata multichambered forms with a planispiral coiling (Text-fig. 4, figure 6 – likewise figures 5 and 8). The plan of the shell structure and position of the aperture are to a great degree interrelated, thus its position in the umbilical area of multichambered trochoid forms provides a better means of communication with the environment to all the chambers. Hence the trochoid shell type could be considered as the most progressive one. It is characteristic and abundant in two classes only – the Rotaliata and Spirillinata (especially in the former), and can be used as the diagnostic feature of these classes having some special peculiarities in each of them (Mikhalevich 1981, 2000, Mikhalevich and Debenay 2001). Similarity in position of the aperture and in the inner and outer apertural structures in agglutinated and calcareous shells of different classes TEXT-FIGURE 1 Similarity of shells in the subclasses Lagynana (=Allogromiina) (A) and Astrorhizana (B). Thus the types of coiling and the change of the type of coiling during ontogenesis within the species or the genus are also similar in agglutinated and calcareous miliolata genera. In all the enumerated cases the aperture is terminal (even in text-fig. 3, figures 6; 12 it is above the base of apertural face). This feature is especially demonstrative distinguishing miliolata shells from the Rotaliata shells with similar planispiral or trochospiral (text-fig. 4, figures 7 and 8) coiling. Such shell types are not met in the other Foraminiferal classes and are characteristic 496 Apertural types with an entosolenian tube (text-fig. 5, figures 1, 2) and an asymmetric fissured aperture (text-fig. 5, figures 3, 4) characteristic for the Nodosariata only is present both in their agglutinated (text-fig. 5, figures 1, 3) and calcareous forms (text-fig. 5, figures 2, 4), Similarly, in both types of shell wall in the Miliolata, similarly constructed and disposed inner apertural teeth are observed (text-fig. 5, figures 5–7). Not only the structure but also the tendencies of their ontogenic and phylogenic development from singular to the multiple apertures and then to the formation of a trematophore (text-fig. 5, figures 8, 9) are similar. In the Miliolata with a fusulinoid shell type the aperture consists of a row of openings elevated above the base of the last chamber – i.e., also terminal. The profound similarity in the structure of the inner apertural tooth-plates is revealed in the Rotaliata (text-fig. 5, figures 12–19). The similarity of such complex apertural plates as in some buliminids (text-fig. 5, figures 13, 15, 17) and valvulinids (text-fig. 5, figures 12, 14, 16) is striking. Well-developed additional apertures in the Rotaliata may be disposed similarly in both their subclasses as for instance on the spiral side of some Micropaleontology, vol. 59, no. 6, 2013 TEXT-FIGURE 2 Similarity of shells with agglutinated (À) and calcareous (Â) walls in the classes Nodosariata, Spirillinata and Rotaliata (scheme). 497 V. I. Mikhalevich: New insight into the systematics and evolution of the foraminifera nated form has its own mirror reflection in forms with a calcareous shell wall as if the two mirror systems exist in parallel. The nuclear apparatus in the different foraminiferal classes The nuclear apparatus has been studied in only a few foraminiferal species. Even these data permit to elucidate its peculiarities in several foraminiferal classes. Representatives of the Astrorhizata are uninucleate or multinucleate but homocariotic (their nuclei are similar in their form, size and function). Spirillinata are multinucleate and homocariotic. In the multinucleate Miliolata and Rotaliata nuclear dimorphism arose when nuclei are differentiated in their size and function into somatic ones (of larger size and more usually singular) and multiple and small generative nuclei (text-fig. 6). Heterokariocity arose in the most advanced phylogenetic lines and among the other unicellular eukariotes it is known only in one other phylum – in the Infusoria (though having in it some peculiar features – Mikhalevich 1980; Starobogatov and Mikhalevich 1985). Therefore, such important cytological features of the structure of the living cell such as the peculiarities of their nuclear apparatus may also be used as diagnostic features of class rank in four of the five classes separated here (data on the Nodosariata are still lacking, maybe just because of the wrong understanding of their taxonomic position their study lacked special attention). Development of the cytological structures also evolved in parallel in several foraminiferal classes, thus heterocariosity was achieved in two classes – the Miliolata and Rotaliata. Special organelles (microvilli and cryptosomes) arose only in the planktonic subclass Globigerinana and may be considered as having taxonomic significance. Scheme of the type of ultrastructure of the calcareous wall The subclasses with a calcareous shell wall of the different phylogenetic lines (classes) differ from each other in the ultrastructure of their calcareous wall, which displays different patterns of the packing of their crystals (text-fig. 7, 1–4). Thus the taxonomic weight of this feature has rather important but nevertheless subordinate significance, and is characteristic only for some subclasses, not classes (Mikhalevich 1992, 2002). Transitional character of shell wall structure TEXT-FIGURE 3 Similarity of shell form in representatives with an agglutinated (À) and calcareous (Â) shell wall in the class Miliolata. trochamminids (Textulariana) and oridorsaliids (Rotaliana) (text-fig. 5, figures 18, 19). In all the classes represented here the agglutinated and calcareous shells have nearly mirror-like similarity. Although convergent similarity is widely known in different plant and animal groups it hardly would be possible that practically each aggluti498 Sometimes in a given species different types of the shell wall may be combined showing the transitional character of some of them, as for instance in Dentostomina agglutinans d’Orbigny 1839 (text-fig. 8), where in transverse section the thick agglutinated layer can be seen underlined from the inside by a thin white porcellaneous layer (text-fig. 8, a). The ultrustructure of the porcellaneous layer is characteristic for the typical porcellaneous miliolid wall, and can be clearly seen under higher magnification (text-fig. 8, b) (Mikhalevich et al. 1986). Skeletal features are more conservative and stable during the evolutionary process than the wall structure. Scheme of the development of different foraminiferal classes in geologic history The most ancient shells of classes Astrorhizata and Spirillinata are known from the Cambrian, Spirillinata and Miliolata – from the Ordovician, Rotaliata – from the Mesozoic (though some of their more primitive forms – from the Carboniferous). The scheme given in the text-fig. 9 is based on the geological data. It is thought that representatives of the class Astrorhizata with a Micropaleontology, vol. 59, no. 6, 2013 TEXT-FIGURE 4 Position of the aperture in the different foraminiferal classes. soft shell may have evolved much earlier than the Cambrian (Mikhalevich 1980; Starobogatov and Mikhalevich 1985; Mikhalevich and Debenay 1998, 2001, Pawlowski et al. 1999), The origin of the Miliolata and of some other foraminiferal groups was also thought to be earlier than their geological record suggests. Plotnikova (1991) discovered some foraminiferal remnants in the Vendian (Ediacarian). Data of molecular analysis of different animal groups usually show an earlier time of origin than what the fossil record suggests (Fedonkin 2009). At present with the use of the Bayesian method, the age of the appearance of foraminifera is dated between 650 and 920 millions years (Groussin et al. 2011), which is in accordance with the data of other authors on the early Eukariotic origins in the Neoproterozoic (Fedonkin 2009). Obviously, in the group of soft walled unilocular representatives the roots of all the other classes can be found (text-fig. 10) and hence this group was believed to be heterogeneous (Mikhalevich 1992). The lack of morphological features in this group makes it the most urgent object of the molecular studies. According to Groussin et al. (2011), the results of the Bayesian method are mostly in agreement with the fossil records except the Miliolida (= class Miliolata) whose remains are known from the Carboniferous (~ 350 Ma) but the estimation of their age by molecular analysis gives a time of their origin of 490 Ma; Devonian and Silurian). With the inclusion of fusulinoids into the composition of the class Miliolata (Mikhalevich, 2004, 2009) this conflict seems to be resolved. This example shows the necessity and importance of collaboration of the classical taxonomists and specialists in the molecular methods. The main evolutionary processes took place in each of the foraminiferal phyletic lines (classes) independently and in parallel, for in stance nearly in each line the rise of multichamberedness, of inner apertural structures, of additional apertures, of integrative systems took place. At the same time form of the chambers, position of additional apertures, character of inner apertural structures and integrative systems have a lot of specific features in each of them. The classification composed according to the new understanding of the foraminiferal evolutionary development is given below. Taxa with doubtful position are marked with a question mark, those with strikingly reassigned position with mark *. The new genera published after Loeblich and Tappan (1987) were added when the publications were available, especially 499 V. I. Mikhalevich: New insight into the systematics and evolution of the foraminifera those of the taxa with a calcareous secreted wall, as the new agglutinated genera are monitored in a series of papers by Kaminski (2000, 2004, 2008, 2011) and are also considered by Mikhalevich (2004a,c). – the references for the authors of the taxa mentioned in the “Ñlassification“ are given only for those published after Loeblich and Tappan (1987), as all the previous references can be found in their two-volume book. In the cases of the change of taxonomic rank the authorship is preserved within the same category (e.g. – subfamily, family, superfamily), and is changed with the transfer to a higher category (e.g. – from the family to the order rank). CLASSIFICATION Supergroup RHIZARIA Cavalier-Smith 2002 Phylum FORAMINIFERA d’Orbigny 1826 nomen translat. Mikhalevich, 1980 Class ASTRORHIZATA Saidova 1981 (= class Monothalamids Pawlowski et al. 2003, part) Shells unilocular, sometimes pseudocolonial or pseudochambered, of subsphaerical, elongated, tubular, stellate, branching or other irregular form; wall tectinous, agglutinated or microgranular, sometimes the pseudopores are visible, may be simple or partly subdivided by the partitions, may be thickened from the inside (spongy, labyrinthic); aperture may be entirely absent, may be primarily multiple (openings often irregular in form and position or situated at the end of tubes), or single in more advanced forms (usually of definite form and position, may be slightly elevated on a neck, may have a small lip); free-living or (very often) attached; agamonts uninuclear, nuclear dualism unknown. Cambrian – Holocene. Remarks. The superfamily Moravamminacea Pokorný 1951 is excluded here from the ranks of the phylum Foraminifera after Vachard (1994) and Vachard and Cozar (2010) demonstrated the affinity of moravamminids and some other closely related genera to algae. Subclass *LAGYNANA Mikhalevich 1980 Order LAGYNIDA Mikhalevich 1980 Families: Lagynidae Schultze 1854 (with the subfamily Belariinae Mikhalevich 1999 icluding representatives whose shells have an inner septa, genera Belaria and Cystophrys), Maylisoriidae E.V. Bykova 1961, Blysmasphaeridae Brönnimann 1988. Order ALLOGROMIIDA Fursenko 1958 Remarks. A number of new genera of allogromiids were described during the last few years (Niveus Altin et al. 2009, Edaphoallogromia Pawlowski and Holzmann 2002, Xiphophaga Goldstein et al. 2010 and some others) but a lot of them still remain unidentified and often could only be distinguished by molecular methods as they lack distinctive morphological features. Families: Allogromiidae Rhumbler 1904, Shepheardellidae Loeblich and Tappan 1948, Argillotubidae Avnimelech 1952, Labyrinthochitinidae Mikhalevich 1999 (wall labyrinthic, may be with inner subpartitions, genera Labyrinthochitinia, Chitinolagena). 500 Order AMMOSCALARIIDA Mikhalevich 1980 Remarks. A heterogenous group including forms of different shell structure (uniserial – Hospitellidae, trochoid – Phthanotrochus, milioloid – Periptigma, planispiral – Ammoscalariidae). Such distinctions are regarded in the higher advanced Foraminifera as characteristic for the orders. Because the evolution of multichamberedness is here at its earliest stage (e.g., in Periptigma the chambers are not stable, often fall off, the trochoid spire in Phthanotrochus may erect itself, may become of irregular form), and owing to the poor knowledge of all these forms they are for the present time kept in this taxon. If it is the case that the Periptigma is the ancestral form of Miliolata, and Phthanotrochus – of the Rotaliata, their systematic position would need to be changed. Families: Phthanotrochidae Arnold 1978 (including Periptigma), Ammoscalariidae Mikhalevich 1982 (with Starobogatovella Mikhalevich 1994), Hospitellidae Loeblich and Tappan 1984 (including free-living uniserial forms such as Nodellum Rhumbler 1913 in a special subfamily). Subclass ASTRORHIZANA Saidova 1981 This subclass includes many transitional forms whose features permit to include them in Astrorhizana or with some reservation (provisionally) into other subclasses: thus the Vanhoeffenellidae, Causiinae and Amphifenestrellinae are transitional from the subclass Lagynana; the Ammovoluminidae – to the subclass Ammodiscana, many of the Hippocrepinida (Earlandiidae, Hormosinellidae, Caligellidae, Paratikhinallidae) – to the subclasses Hormosinana or Nodosariana. Obviously the whole order Hippocrepinida ought to be placed in the subclass Hormosinana but it would be preferable previously to make a thorough revision of all its taxa before taking such a step. Some of the microgranular forms included previously in the lower Fusulinacea are transferred to Astrorhizana as the distinction between the agglutinated and microgranular wall is often transitional. Order ASTRORHIZIDA Haeckel 1894 Superfamily ASTRORHIZOIDEA Brady 1881 Families: Astrorhizidae Brady 1881 [(including Globodendrina Plewes, Palmer and Haynes 1993), the genus Clados Schroeder, Medioli and Scott 1989 assigned by its authors to the Astrorhizinae belongs more probably in the Komokiida), Vanhoeffenellidae Saidova 1981, Rhabdamminidae Brady 1884 (with Rhabdammininae Brady 1884, Bathysiphoninae Avnimelech 1952), Hippocrepinellidae Loeblich and Tappan 1984 (including the genus Lakites Nestell and Tolmacheva 2004, the family differs from the closely related family Rhabdamminidae in the shorter shell, tapered apertural end, and structure of the aperture). Order DENDROPHRYIDA Haeckel 1894 Superfamily DENDROPHRYOIDEA Haeckel 1894 Families: Dendrophryidae Haeckel 1894 (with the subfamilies Dendrophryinae Haeckel 1894, Spiculidendroninae Mikhalevich 2004),*Notodendrodidae De Laca, Lipps and Hessler 1980 (included earlier in Hippocrepinacea), Dryorhizopsidae Loeblich and Tappan 1984, Arboramminidae Shires, Gooday Micropaleontology, vol. 59, no. 6, 2013 TEXT-FIGURE 5 Similarity in position of the aperture and in the inner and outer apertural structure in the agglutinated (À) and calcareous (B) shells of different classes (scheme). 501 V. I. Mikhalevich: New insight into the systematics and evolution of the foraminifera TEXT-FIGURE 6 The nuclear apparatus in the different foraminiferal classes. and Jones 1994 (including Luffammina Kamenskaya, Bagirov and Simdianov 2002), *Komokiidae Tendal and Hessler 1977 (including Rhizammina according Cartwright, Gooday and Jones 1989 and genera Calos, Cactos, Cerebrum, Crambis, Reticulum, Tuber Schroeder, Medioli and Scott 1989 and Skeletonia Gooday, Kamenskaya and Cedhagen 2007), *Baculellidae Tendal and Hessler 1977 (including Neocatena Özdikmen, 2009 pro Catena Schroder, Medioli and Scott 1989 (non Richter 1975), *Normaninidae Mikhalevich 1994. Superfamily SCHIZAMMINOIDEA Norvang 1961 Families Schizamminidae Nørvang 1961, Halyphysematidae Loeblich and Tappan 1984, nom. corrected, Diffusilinidae Loeblich and Tappan 1961. Order PSAMMOSPHAERIDA Haeckel 1894 Suborder STEGNAMMININA Mikhalevich, G.P.Pronina, and M. Nestell 2000 Superfamily STEGNAMMINOIDEA Moreman 1930 Families: Stegnamminidae Moreman 1930 (with subfamilies Stegnammininae Moreman 1930, Amphifenestrellinae Mikhalevich 1995, Hemisphaerammininae Loeblich and Tappan 1961), Colonamminidae Rauser-Chernousova and Reytlinger 1993 in Vdovenko et al. 1993 (stat. elevated, with subfamilies Causiinae Mikhalevich 1995, Colonammininae Rauser-Chernousova and Reytlinger 1993 in Vdovenko et al. 1993, Tholosininae Mikhalevich 1995). Superfamily CRITHIONINOIDEA Goes 1894 Families: Crithionidae Goes 1894 (differing from the Oryctodermatidae in the presence of inner septa subdividing the test lumen and in the absence of a labyrintic wall) (with the subfamilies Daitroninae Mikhalevich 1995, Crithioninae Goes 1894), Oryctodermatidae Saidova 1981 [= Subfam. Masonelinae Saidova 1981 (as “Marsonellinae”), part]. 502 Suborder PSAMMOSPHAERINA Haeckel 1894 Families: Psammosphaeridae Haeckel 1894 (with subfamilies Psammosphaerinae Haeckel 1894, Telammininae Loeblich and Tappan 1985), Polysaccamminidae Loeblich and Tappan 1984 (with subfamilies Polysaccammininae Loeblich and Tappan 1984, Saccamminidinae Mikhalevich 1995, Amphicervicinae Mikhalevich 1995); Lacustrinellidae Mikhalevich 1995. Îrder *PARATHURAMMINIDA Mikhalevich 1980 Superfamily PARATHURAMMINOIDEA E.V. Bykova 1955. Families: Archaesphaeridae Malakhova 1956 [with the subfamilies Archaesphaerinae Malakhova 1956, Neoarchaesphaerinae Sabirov 1987, Insolentithecinae Loeblich and Tappan 1986, Usloniinae A.D. Miklukho-Maklay 1963, Eoammosphaeroidinae Mikhalevich 1995, Atjussellinae Zadorozhny 1987 (subfamily reinstated by Vdovenko et al. 1993)], Parathuramminidae E.V. Bykova 1955 (with the subfamilies Parathurammininae E.V. Bykova 1955, Irregularininae Zadorozhnyy and Juferev 1984, Chrysothurammininae Loeblich and Tappan 1986). Superfamily MARGINARIOIDEA Loeblich and Tappan 1986 (= Ivanovelloidea Chuvashov and Yuferev 1984). Families Marginaridae Loeblich and Tappan 1986 (with the subfamilies Marginarinae Loeblich and Tappan 1986, and Uralinellinae Chuvashev and Yuferev 1984), Tuberitinidae A.D. Miklukho-Maklay 1958 (with the subfamilies Tuberitininae A.D. Miklukho-Maklay 1958, Hemithurammininae Mikhalevich 1995), Auroriidae Loeblich and Tappan 1986, Eovolutinidae Loeblich and Tappan 1986, Rauserinidae Sabirov 1987, Cribrosphaeroididae Sabirov 1987. Order HIPPOCREPINIDA Saidova 1981 Superfamily HIPPOCREPINOIDEA Rhumbler 1895 Families: Hippocrepinidae Rhumbler 1895 ((with the subfamilies Hippocrepininae Rhumbler 1895 = Hyperamminoid- Micropaleontology, vol. 59, no. 6, 2013 TEXT-FIGURE 7 Scheme of the type of ultrastructure of the calcareous wall (iol – inner organic lining, mol – middle organic lining (POM)). idae Loeblich and Tappan 1984 (part) and Jaculellinae Mikhalevich 1995 = Hyperamminoididae Loeblich and Tappan 1984 (part)), Hyperamminidae Eimer and Fickert 1899 ((with the subfamilies Hyperammininae Eimer and Fickert 1899 = Hyperamminoididae Loeblich and Tappan 1984 (part), Saccorhizinae Eimer and Fickert 1899)), Ammovoluminidae Chernych 1967. Superfamily BOTELLINOIDEA Chapman and Parr 1936 (= Caligelloidea Reytlinger 1959) Family: Botellinidae Chapman and Parr 1936. Class SPIRILLINATA Mikhalevich 1992 (=Tubothalamea Pawlowski et al. 2012, part, in Adl et al. 2012) Shells pseudotwochambered, with a long tubular second chamber coiled around the proloculus, in higher taxa with the tendency of transition to multichamberedness, as a rule in the last coils, often with a very limited chamber number (2–3 per coil), at the same time an undivided tubular second part around the proloculus may be preserved during 1–3 volutions; types of coiling - glomerate, planispiral, trochospiral, the predominant type of shells - subsphaerical, lenticular, flat, patellinoid; high conical; in higher more advanced forms a weakly developed canal system is rarely present; wall monofontinal (monolamellar), insufficiently studied, agglutinated in the lower less advanced forms (Ammodiscana), usually with a ferrugenous cement, or microgranular, fully secreted in higher ones (Spirillinana), in the ancient forms often two layers are marked – microgranular and radial, in the involutinids – usually radial, in recent Spirillinids – radial crystal units act as a single crystal, shell wall may be pitted by regular pseudopores, may have inner and outer additional skeletal sculpture, pillars, typical layers of growth are not formed but secondary layers of thickening of a special character, often multilamellar are often formed, especially in the umbilical area; aperture usually at the end of the tubular chamber, may be multiple in advanced agglutinated and calcareous forms, simple, in Patellina Williamson 1858 with a special T- shaped plate, additional apertures rarely present; agamonts homokariotic, their proloculus larger than that of the gamonts (the reverse correlation), gametes amoeboid; benthic, mostly free living forms. Cambrian – Holocene. Subclass AMMODISCANA Mikhalevich 1980 Order AMMODISCIDA Mikhalevich 1980 Families: Ammodiscidae Reuss 1862 (with Ammodiscinae Reuss 1862 (without Agatamminoides) and Paulbronnimanniinae Rettori and Zaninetti 1993 (nom. corr. pro Paul- bronnimanninae Rettori and Zaninetti 1993)), *Pseudoammodiscidae Conil and Lys 1970 (from the Fusulinacea, but without Warnantella), Turritellellidae Saidova 1981 (fam. reinstated and given stat. nov), ? Tolypamminidae Cushman 1928 (without Ammolagena transferred by Kaminski et al. 2009 to the subclass Hormosinana as having a second aperture that opens out of the proloculus opposite the tubular chamber and including later described Ammodiscellites of Resig and Glenn 1997), Turriglominidae Zaninetti, in Limongi et al. 1987. Order AMMOVERTELLINIDA Mikhalevich 1999 Families: Usbekistaniidae Vyalov 1968 (without Turritellella and Repmanina), Ammovertellinidae Saidova 1981 [with Ammovertellininae - having an uncoiled part, Glomospirellinae Ciarapica and Zaninetti 1985 (without an uncoiled part, subfamily reinstated from Ammovertellininae of Loeblich and Tappan 1987), and Pilammininae Urosevic 1992], *Pseudolituotubidae Conil and Longerstaey 1980 from the Fusulinacea. Order PLAGIORAPHIDA Mikhalevich 2003. Family: Plagioraphidae Mikhalevich 1995 (Plagioraphe). Order *?ATAXOPHRAGMIIDA Fursenko 1958 Superfamily ATAXOPHRAGMOIDEA Schwager 1877 Families: Duotaxidae Mikhalevich 2003 (differs from representatives of the Verneuilinidae where it was previously placed by the character of the shell and chamber shape) (type genus Duotaxis Kristan 1957), Ataxophragmiidae Schwager 1877 (with Ataxophragmiinae Schwager 1877 - only genera having Ataxophragmium-like largely involute spiral coiling with early whorls not visible and with high chambers turned to the axis of coiling are retained here) (Ataxophragmium, Arenobulimina, ?Ataxoorbignyna, but without Hagenowella and Sabulina (both transferred to the Globotextulariidae) and without Pityusina and Praechrysalidina), Dobrogelinidae Mikhalevich 1992 (Dobrogelina). Superfamily TEXTULARIELLOIDEA Groenhagen and Luterbacher 1966 Families: Pernerinidae Loeblich and Tappan 1984 [with Pernerininae Loeblich and Tappan 1984 (only genera having Ataxophragmium-like largely involute spiral coiling with not visible early whorls and with high chambers and intercameral sutures inclined to the axis of coiling and shells lacking a uniserial part are retained here) (Pernerina, Orbignyna, Voloshinoides, ?Opertum, ?Anatoliella, without Agglutisolena, 503 V. I. Mikhalevich: New insight into the systematics and evolution of the foraminifera TEXT-FIGURE 8 Transitional character of shell wall structure: Dentostomina agglutinans d’Orbigny 1839, à – transverse section of the thick agglutinated wall (×3000); b - ultrastructure of the thin porcellaneous layer underlying the wall from the inside (×10 000). Coprolithina and Kaeveria transferred to the subclass Hormosinana and without Crenaverneuilina, Hagenowina and Voloshinovella) and Voloshinovellinae Mikhalevich 2003 (differing from the Pernerininae in its small coiled part, with most of the shell formed by the uniserial rectilinear widening part giving it a subconical appearance, and from the Coskinolininae in its endoskelandal features: more simple and situated in the marginal part of the chambers (type genus Voloshinovella Loeblich and Tappan 1964, Trochamijiella Athersuch, Banner and Simmons 1992 (originally put into the Biokovinidae but differing in its early trochoid coil)], Textulariellidae Groenhagen and Luterbacher 1966 [Textulariella, Cuneolinella, Guppyella) (without Alveovalvulina, Hagenowinoides transferred to the Crenaverneuilinidae)], ?Syrianidae Kaminski 2004 (Syriana Fourcade and Mouti 1995 - if its initial part is really trochospiral (unclear, broken), final part uniserial and compressed), Cuneolinidae Saidova 1981 [(with Cuneolininae Saidova 1981 (Cuneolina - in the understanding of Neagu 2000), Palaeolituonella, Pseudotextulariella, ?Vercorsella), Scythiolininae Neagu 2004 in Kaminski 2004 (Histerolina Neagu 2000, Scythiolina Neagu 2000 – these two last genera with an initial scarce planispiral part according to Neagu 2000) and Montsaleviinae Zaninetti, Salvini-Bonnard, Charollais and Decrouez 1987 (type genus Montsalevia described by Zaninetti, Salvini-Bonnard, Charollais and Decrouez 1987)], Dicyclinidae Loeblich and Tappan 1964 (Dicyclina and Nakkadyia described by Ismail, Hussein-Kamel, Boukhary and 504 Ghadour 2009), Pfenderinidae Smout and Sugden 1962 [(with Pfenderininae Smout and Sugden 1962 (Pfenderina, Drevennia, Pfenderinella, without Accordiella transferred to the Chrysalidininae), Pseudopfenderininae Septfontaine 1988 (without tunnels, subfamily reinstated here from the synonymy of Pfenderinidae in Loeblich and Tappan 1987) (Pseudopfenderina, Pseudoeggerella), Paleopfenderininae Septfontaine 1988 (Paleopfenderina Septfontaine in Kaminski 2000, Conicopfenderina, Chablasia (transferred from the Biokovinidae), Satorina, Sanderella, ?Steinekella (chambers subdivided), and Kurnubiinae Redmond 1964 (Kurnubia, Gyroconulina, Praekurnubia, ?Conicokurnubia)]; Coskinolinidae Moullade 1965 [(Coskinolina, Coleiconus, Coskinon, Lituonelloides, Pseudolituonella); Hauraniidae Septfontaine 1988 (Haurania, ?Cymbriaella Fugagnoli 1999, Gutnicella, Meyendorfina, ?Plathyhaurania Bassoullet and Boutakiout 1996, Bostia Bassoullet 1998, ?Socotraina Banner, Whittaker, Boudagher-Fadel and Samuel 1997, Spiraloconulus (from Spirocyclinidae), (without Amijiellinae)]; Parurgoninidae Septfontaine 1988 (Parurgonina), Orbitolinidae Martin 1980 [(with Orbitolininae Martin 1980 (Orbitolina, Alpillina, Conicorbitolina, Eopalorbitolina, Eygalierina, Mesorbitolina, , Montseciella Cherchi and Schroeder 1999, Naupliella, Neoiraqia, Neoorbitolinopsis, Palorbitolina, Palorbitolinoides, Praeorbitolina (the genus Praeorbitolinoides Matsumaru 2005 is regarded by Cherchi and Schroeder (2009) as a synonym of Praeorbitolina based on the similarity of their embryonic appa- Micropaleontology, vol. 59, no. 6, 2013 TEXT-FIGURE 9 Scheme of the development of different foraminiferal classes in geological history. ratus), Rectodictyoconus, Valserina), Dictyoconinae Moullade 1965 (Dictyoconus, Abrardia, Calveziconus, Campanellula, Carinoconus, Coskinolinoides, Cribellopsis, Cushmania, Daviesiconus, Dictioconella, Fallotella, Falsurgonina, Handerocoskinolina, Iraqia, Karsella Sirel 1997, Orbitolinella, Orbitolinopsis, Paleodictyoconus, Pseudorbitolina, Simplorbitolina, Urgonina, Valdanchella, Verseyella and Barattolites Vecchio and Hottinger 2007), Praedictyorbitolininae Schroeder 1990 (including Praedictyorbitolina Schroeder 1990 in Schroeder, Clavel and Charollais 1990, Paracoskinolina) and Dictyorbitolininae Schroeder 1990 in Schroeder et al. 1990 (Dictyorbitolina Cherchi and Schroeder 1976]. Subclass SPIRILLINANA Mikhalevich 1992 Superorder ARCHAEDISCOIDA Pojarkov and Skvortsov 1979. Order ARCHAEDISCIDA Pojarkov and Skvortsov 1979 Superfamily ARCHAEDISCIDOIDEA Cushman 1928 Families: Archaediscidae Cushman 1928 [with Archaediscinae Cushman 1928, Eosigmoilininae Brazhnikova and Vdovenko 1977 (reinstated from the synonymy of Archaediscinae (Loeblich and Tappan 1987) according to Ponomareva 2012), including two genera – Eosigmoilina Ganelin 1956 and Brenckleina Zaninetti and Altiner 1979 previously entering Asteroarchaediscidae), Ammarchaediscinae Conil and Pirlet 1974 in Pirlet and Conil 1974 including Donodiscus Vdovenko 1988), Uralodiscinae Grozdilova 1993 in Vdovenko et al. 1993, Kasachstanodiscinae Marfenkova 1983 – the three latter subfamilies reinstated in Vdovenko et al. 1993], Asteroarchaediscidae A.D. Miklukho-Maklay 1957, Planoarchaediscidae Gubenko 1989, Tubispirodiscidae Mikhalevich 1996 (fully planispirally and evolutely enrolled). Superfamily GLOMODISCIDOIDEA Mikhalevich 1998 Family: Glomodiscidae Mikhalevich 1998 (Glomodiscus, Uralodiscus, Tournarchaediscus, Permodiscus). Order *TETRATAXIDA Mikhalevich 1981 Families: Tetrataxidae Galloway 1933, Pseudotaxidae Mamet 1974, ?Valvulinellidae Loeblich and Tappan 1984 [(with Valvulinellinae Loeblich and Tappan 1984, Abadehellinae Loeblich and Tappan 1984 (chambers subdivided)], Akcayidae Özdikmen 2009 pro Sabaudiidae Brönnimann, Decrouez and Zaninetti 1983 (transferred from the Textularielloidea as having a hyaline radial layer of the wall in the embrional stage), Endotaxidae Bogush and Brazhnikova 1996 in RauserChernousova et al. 1996. Order LASIODISCIDA Mikhalevich 1993 Families: Lasiodiscidae Reytlinger 1956 (with Lasiodiscinae Reytlinger 1956, Vissariotaxinae Reytlinger in Vdovenko et al. 1993), Monotaxinoididae Mikhalevich 1999 (slightly trochoid 505 V. I. Mikhalevich: New insight into the systematics and evolution of the foraminifera shell lacking the additional apertures and bridges), Howchiniidae R. Martini and Zaninetti 1988, Pseudovidalinidae Altiner 1988. Superorder INVOLUTINOIDA Hohenegger and Piller 1977 Order INVOLUTINIDA Hohenegger and Piller 1977 Superfamily INVOLUTINOIDEA Butschli 1880 Families: Involutinidae Butschli 1880 (including the genus Bancilina of Neagu 1995), Planispirillinidae Piller 1978, Triadodiscidae Zaninetti 1984 [with Triadodiscinae Zaninetti 1984, Lamelliconinae Zaninetti, Ciarapica, Decrouez 1987 (the latter subfamily regarded by Loeblich and Tappan 1987 as a synonym of Triadodiscinae)], Aulotortidae Zaninetti 1984 [with Aulotortinae Zaninetti 1984 (including the genus of Neagu 1994 – Andersenolina), Auloconinae Zaninetti, Ciarapica, Decrouez 1987 (the latter subfamily placed by Loeblich and Tappan 1987 into the synonymy of the Aulotortinae) and Parvalamellinae Rigaud, Martini and Rettori 2012, with ball-like coiling and reduced lateral laminar extensions (Rigaud et al. 2012)], Hirsutospirellidae Zaninetti, Ciarapica, Cirilli and Cadet 1985, ?Ventrolaminidae Weynschenck 1950. Superfamily TRIASINOIDEA Loeblich and Tappan 1986 Family: Triasinidae Loeblich and Tappan 1986. Order HOTTINGERELLIDA Mikhalevich 1993 Family: Hottingerellidae Mikhalevich 1993. Superorder SPIRILLINOIDA Hohenegger and Piller 1975 Order SPIRILLINIDA Hohenegger and Piller 1975 Suborder SPIRILLININA Hohenegger and Piller 1975 Families: Spirillinidae Reuss and Fritsch 1861, Conicospirillinidae Mikhalevich 1995 (with a trochospiral shell and the umbilical area filled with secondary shell matter). Suborder SPIROTROCHOLININA Mikhalevich 1993 Family: Spirotrocholinidae Mikhalevich 1993. Order PATELLINIDA Mikhalevich 1992 Superfamily HERGOTTELLOIDEA Loeblich and Tappan 1984 Families: Hergottellidae Loeblich and Tappan 1984, Ungulatellidae Seiglie 1964 (the shell of Ungulatella – the type genus of the family is close in its structure to that of the agglutinated genus Plagioraphe), Placentulinidae G.K. Kasimova, Poroshina, Geodakchan 1980 [with Placentulininae G.K. Kasimova, Poroshina, Geodakchan 1980, (obviously, including the genus Mandapatellina from the Ungulatellidae), Ashbrookininae Loeblich and Tappan 1984], Pannellainidae Loeblich and Tappan 1984 (possibly the genus Heronallenita close in its structure to this family also needs to be included here but the structure of its aperture is unclear). Superfamily PATELLINOIDEA Rhumbler 1906 Families: Patellinidae Rhumbler 1906 (including the new genus of Neagu and Cîrnaru 2001 – Rumanolina), *Annulopatellinidae Loeblich and Tappan 1964, Paleopatellinidae 506 Mikhalevich 1999 - shell similar in its structure to the patellinid shell, even the inner columella may be present, but the tubular undivided chamber enrolled around the proloculus is absent (Paleopatellina, Pseudopatellina, Patellinella, Subpatellinella, ? Paalzowella from Ashbrookininae, ? Pseudopatellinoides), Patellinellidae Bugrova, 1990, *Chapmaninidae Thalmann, 1938, Dictyoconoidessidae Bugrova, 1990 (placed by the author of the family in the rotaliids). Order *?CYMBALOPORIDA Mikhaelevich ord. nov. Shell initially trochospiral, with a few or multiple small chambers (rarely two chambers per whorl) visible on the spiral side and only the chambers of the last whorl visible on umbilical side, later chambers in cyclic series in a single flat to conical layer (usually those of the successive cycles alternating in position) or in flaring tiers (Fabianiidae); umbilical area with symmetrically radial rather wide and incised sutures, either widely opened or may be obscured by the plate-like chamber extensions forming a flange-like plate or a complex series of perforated plates (Cymbaloporettidae) or may be filled with lamellae and pillars (Halkyardiidae); wall perforated on the spiral side and imperforate on the umbilical side, often forming lamellae obscuring the spiral side (Cymbaloporidae) or filling the umbilicus (Halkyardiidae); apertures in the trochoid stage at the open umbilical ends of chambers, may also be on both sides of the umbilical chambers in sutures as one or a series of openings, which sometimes may be bordered by rim-like lips, lips of the apertures at the central ends of the umbilical chambers may form tubes, funnels or flange-like plates (Cymbaloporettidae), in tiered chambers – it consists of pore-like openings on the terminal face; in gamont generations in some genera the float and balloon chambers of different degree of complexity formed beneath the umbilical side are known, balloon chambers may have large rounded openings bordered by lips or may have branching channels (Millettiana Banner, Pereira and Desai 1985). Upper Cretaceous; Paleocene – Holocene Remarks. The representatives of the new order are transferred here from the former Rotaliina on the basis of their shell structure. Their unique pelagic mode of life forming rather complex balloon and float chambers is not encountered in any other planktonic foraminiferal group and could be regarded as a special parallel way to partial occupying the pelagic niche compared with the Globigerinana. Such peculiarities of their shell structure as right symmetrical rather wide and incised radial sutures with all the chambers of umbilical side having the equal dimensions, the hollow centres of the trochoid shells, the tendency towards the lamellar covering of the umbilical area characteristic for many ancient representatives of the Sprillinana (archaediscids, involutinids), special character of flange-like plates and funnels in the umbilical area of Cymbaloporetta Cushman 1928, characteristic pore-like apertures with elevated lips and some other minor features of their appearance make it possible to put them closer to the other Spirillinana. The true billamellar character of their shell wall was not observed. The new order differs from the order Patellinida also having a trochoid shell in the structure of the umbilical side and aperture, and also in its unique adaptation to a pelagic mode of life in one of the generations. Further study is necessary to elucidate the resemblance of Cymbaloporids with Tretomphalus Moebius 1880 – are their so similar floating chambers formed as convergent structures or are the both groups in close relationship? The order Micropaleontology, vol. 59, no. 6, 2013 is tentatively placed in this subclass - perhaps this transfer will inspire special future attention and reinvestigation of the representatives of this group. Such a study ought to help to decide the problem whether radially-symmetrical structure of the umbilical side of representatives of the order is a feature characteristic for many Spirillinata taxa or a feature acquired by rotaliate form in its adaptation to the pelagic mode of life. Superfamily CYMBALOPOROIDEA Cushman, 1927 stat. elevated Families: Cymbaloporidae Cushman 1927, Cymbaloporettidae fam. nov. (shell similar to Cymbalopora and Millandtiana, but lips of the apertures at the central ends of the umbilical chambers form tubes, funnels, flange-like plates or a complex series of such perforated plates over the center of umbilicus, type genus Cymbaloporetta Cushman, 1928), Halkyardiidae Kudo 1931, stat. elevated. Superfamily FABIANIOIDEA Deloffre and Hamaoui 1973, stat. elevated Familiy: Fabianiidae Deloffre and Hamaoui 1973 stat. elevated. Order ?SEABROOKIIDA Mikhalevich 1980 Family: Seabrookiidae Cushman 1927. Class MILIOLATA Saidova 1981 (= class Miliolicea Saidova 1981) TEXT-FIGURE 10 Scheme of the supposed origin of different foraminiferal classes. gametes were previously described as biflagellate (and having an axostyle?) but in the recent investigations only biflagellate gamets were observed; benthic forms, mostly free living, more rarely attached. Ordovician – Silurian – Permian - Carboniferous - Holocene. Subclass MILIAMMINANA Mikhalevich 1980 (= class Tubothalamea Pawlowski et al. 2012, part, in Adl et al. 2012) (= Rzehakinicae Saidova, 1981, = Schlumbergerinana Mikhalevich 1992) Shells unilocular (very seldom), pseudotwo-chambered, pseudomultichambered in the more primitive forms, in the advanced forms multichambered and supermultichambered, the predominant type of coiling - irregular- and regular glomerate, planispiral more often combined with one of the glomerate ones, (trochospiral only as an exception, rarely biserial); shells of the higher most advanced representatives could be cyclical and fusulinoid in form; the piece of the tubular chamber (flexostyle) is typical of the majority of the groups; chambers of the later forms (Miliolana) predominantly tubular, usually two per volution (in Soritida and rarely in some Milioloida - broad and more numerous per whorl), in Paleozoic forms multiple mostly broad chambers per whorl; chamber lumen may be subdivided by septa (sometimes by the primary and the secondary ones), by pillars, advanced forms may have integrative systems (stolons, tunnels), inner apertural systems not developed, canal system absent; shell wall - monofontinal, agglutinated or agglutinated and microgranular in the lower forms (subclass Miliamminana), in the higher ones (subclass Miliolana) porcellaneous, with irregularly scattered not put into shape pseudopores between irregular randomly oriented needles of crystal units, of irregular character, not formed externally; chamber growth occurs by the gradual addition of the wall part by part, organic and calcareous secreted elements are lain down simultaneously; aperture terminal in position, simple in the most primitive forms, in more complex ones - with an outer (flap) or inner tooth of a special structure (the same apertural structures called in Paleozoic forms in a different way), sometimes aperture secondarily multiple, additional apertures not developed (only as an exception); nuclear apparatus heterokariotic in higher calcareously secreted forms, sometimes with polymerization of somatic nuclei (Sorites); in Triloculina Order LITUOTUBIDA Mikhalevich 1992 (emend) Shell with the second tubular chamber with irregular often alternating (glomerate, planispiral or combined) plan of coiling and undivided chamber lumen, with the following subdivision of the tubular chamber lumen into rather irregular pseudochambers, sometimes forming true chambers in the last volutions, usually of somewhat irregular form and dimensions, bearing some tubular features, the final part may be erected and irregularly rectilinear; wall agglutinated; aperture at the open end of the tubular chamber. Cretaceous – Holocene. Remarks. The representatives of this order are similar in their shell structure to some tournayelids but have an agglutinated rather than microgranular shell wall, though microgranular tournayelids also often have agglutinated particles in their wall. Families: Lituotubidae Loeblich and Tappan 1984, Trochamminoididae Haynes and Nwabufo-Ene 1998 (emend., stat. elevated.) - without an uncoiled part. Order SCHLUMBERGERINIDA Mikhalevich 1980 (= ord. Rzehakinida Saidova 1981) Families: Rzehakinidae Cushman 1933 ((without Rothina which is considered to be a synonym of Hormosinella, representing its broken chamber (Bubik 1997, Kaminski 2004b, Kaminski and al 1996), and without Ammoflintina, Spirolocammina, Spirosigmoilinella, Silicosigmoilina, Trilocularena, Silicomassilina placed into other families according to their mode of coiling and shell structure, and Miliammina, 507 V. I. Mikhalevich: New insight into the systematics and evolution of the foraminifera Birsteiniolla transferred into Miliammininae) [with Rzehakininae Cushman 1933, Psamminopeltinae Mikhalevich and Kaminski 2008, (with an evolute shell), including Spiroglutininae Mikhalevich 1983 as synonym because the type species of the genus Spiroglutina – Spiroloculina asperula (Karrer 1868) ought to be assigned to the genus Psamminopelta], Miliamminidae Saidova 1981, stat. nov, (with Birsteiniollinae Mikhalevich and Kaminski 2008, Miliammininae Saidova 1981 ((=Siphonapertinae Saidova 1975 (part), =Agglutinellinae El-Nakhal 1985) and with Rudoloculina Guibault and Patterson 1998 as a synonym of Siphonaperta Vella 1957 – see Mikhalevich and Kaminski 2008], Trilocularenidae Mikhalevich and Kaminski 2008, Ammomassilinidae Mikhalevich and Kaminski 2008 (with Silicomassilininae Mikhalevich and Kaminski 2008, Ammomassilininae Mikhalevich and Kaminski 2008), Sigmoilopsidae Vella 1957 (with Spirolocammininae Mikhalevich and Kaminski 2008, Sigmoilopsinae Vella 1957), Schlumbergerinidae Mikhalevich 1980, Ammoflintinidae Mikhalevich and Kaminski 2008 (with Ammoflintininae Mikhalevich and Kaminski 2008, Pseudoflintininae Mikhalevich and Kaminski 2008). Order HAPLOPHRAGMIIDA Loeblich and Tappan 1989 Superfamily RECURVOIOIDEA Alekseychik-Mitskevich 1973 Families: Cystamminidae Mikhalevich 2003 (differs from all other families in the Recurvoioidea by its strongly inflated shell and few chambers), Recurvoididae Alekseychik-Mitskevich 1973, Ammobaculinidae Saidova 1981 (with Ammobaculininae Saidova 1981, Telatynellinae Gawor-Biedowa 1987), Acupeinidae Brönnimann and Zaninetti 1984 (Acupeina, Navarella – transferred from the Ammobaculinidae as having multiple apertures), Plectorecurvoididae Loeblich and Tappan 1964. Superfamily HAPLOPHRAGMIOIDEA Eimer and Fichert 1899 Families: Haplophragmiidae Eimer and Fickert 1899, *Mesoendothyridae Voloshinova 1958 (transferred from the former Loftusiacea). Superfamily LABYRINTHIDOMATIDOIDEA Loeblich and Tappan 1987 Family: Labyrinthidomatidae Loeblich and Tappan 1987. Order SPHAERAMMINIDA Mikhalevich and Kaminski 2003 in Mikhalevich 2003 Family: Sphaeramminidae Cushman 1933 (with Sphaerammininae having a tooth (Sphaerammina, Ammosphaerulina, Canepaia) and Praesphaerammininae Kaminski and Mikhalevich 2003 in Mikhalevich 2003 – similar to Sphaerammininae but without a tooth). Order LITUOLIDA Lankester 1885 Superfamily LITUOLOIDEA de Blainville 1827 (Nom. corrected pro Lituolinoidea) Families: Lituolidae de Blainville 1827 (with Ammobaculitinae Alekseychik-Mitskevich, in Subbotina et al. 1981 and Lituolinae de Blainville, 1827 (as Ammobaculitinae, but with 508 multiple apertures), Buzasinidae Mikhalevich 2003 [(differs from Haplophragmoididae where it was originally placed by the terminal character of the aperture (areal rather than basal), from the other families of this superfamily by the special character and position of its areal aperture, from the family Lituolidae in the absence of the uncoiled part of the shell)], Mayncinidae Loeblich and Tappan 1985 (it is necessary in some genera to study the aperture at the initial stage, without Deuterospira with basal aperture at the initial stage) (with Mayncininae Loeblich and Tappan 1985 and Comaliamminae Mikhalevich 2004b), Placopsilinidae Rhumbler 1913 (attached) (with Placopsilininae Rhumbler 1913, Acruliammininae Mikhalevich 2004b, Flatschkofeliinae Kaminski 2004 and Adhaerentiinae Loeblich and Tappan 1986). Superfamily BARKERINOIDEA Smout 1956 Families: Barkerinidae Smout 1956, Biplanatidae Mikhalevich 1992 (transferred from Nezzazatidae where it was originally separated as a subfamily). Superfamily BIOKOVINOIDEA Gusic 1977 Families: Charentiidae LoeblichandTappan 1985, Lituoliporidae Gusic and Velic 1978, Biokovinidae Gusic 1977 (without Chablaisia and Bosniella), Labyrinthinidae Septfontaine 1988 ((with Labyrinthininae Septfontaine 1988, Planiseptinae Septfontaine 1988, nom. nudum (Planisepta Septfontaine in Kaminski 2000) and ?Levantinellinae Fourcade, Mouty and Teherani 1997 (the structure of the type genus is somewhat unclear)), Buccicrenatidae Loeblich and Tappan 1985 (including Evertycyclammina), Choffatellidae Maync 1958 (Banner and Whittaker 1991 showed the planispiral character of the genus Alveosepta and its similarity with Choffatella), Hottingeritidae Loeblich and Tappan 1985 ((with Hottingeritinae Loeblich and Tappan 1985 (its planispiral character was also shown by Banner and Whittaker 1991) and Popoviinae Mikhalevich 2004b)), Pseudochoffatellidae Loeblich and Tappan 1946, Spirocyclinidae Munier-Chalmas 1887. ? Superfamily COSCINOPHRAGMATOIDEA Thalmann 1951 Families: Haddoniidae Saidova 1981 (Haddonia, Stylolina reinstated by Popescu et al. 1998 from the synonymy of Lituola), Coscinophragmatidae Thalmann 1951. Order CYCLOLINIDA Mikhalevich 1992 Superfamily CYCLOLINOIDEA Loeblich and Tappan 1964 Family: Cyclolinidae Loeblich and Tappan 1964. Superfamily CYCLOPSINELLOIDEA Loeblich and Tappan 1984 Families: Cyclopsinellidae Loeblich and Tappan 1984 (with Cyclopsinellinae Loeblich and Tappan, 1984 and Levatinellinae Fourcade, Mouty and Teherani 1997), Ilerdorbidae Hottinger and Caus 1982, Orbitopsellidae Hottinger and Caus 1982. Superfamily VANIINOIDEA Mikhalevich 2003 Family: Vaniidae Mikhalevich 2003 [with subfamilies Vaniinae Mikhalevich 2003 (without pillars) and Thomasellinae Mikhalevich 2003 (with pillars)]. Micropaleontology, vol. 59, no. 6, 2013 Order LOFTUSIIDA Kaminski and Mikhalevich 2003 in Mikhalevich 2003 (= Loftusiida Kaminski and Mikhalevich, 2004a) Family: Loftusiidae Brady 1884 (Loftusia, Praereticulinella, Reticulinella). Subclass MILIOLANA Saidova 1981 *Group of superorders FUSULINOIDS Fursenko 1958 Superorder ENDOTHYROIDA Fursenko 1958 Order TOURNAYELLIDA Hohenegger and Piller 1973 (= order Tournayellida Dain 1953 in Vdovenko et al. 1996) Superfamily TOURNAYELLOIDEA Dain 1953 Families: Tournayellidae Dain 1953 (with Tournayellinae Dain 1953, Forschiinae Dain 1953), Lituotubellidae A.D. MiklukhoMaklay 1963 [with Lituotubellinae A.D. Miklukho- Maklay 1963(including Alticonilites Hance, Hou and Vachard 2011), Septabrunsiininae Conil and Lys 1977, Septaglomospiranellinae Reytlinger 1996 in Rauser-Chernousova et al. 1996], Laxoendothyridae Hance, Hou and Vachard 2011(including Crassiseptella Brenckle and Hance 2005 and Endolaxina Hance, Hou and Vachard 2011). Superfamily CHERNYSHINELLOIDEA Reytlinger 1958 Families: Chernyshinellidae Reytlinger 1958 (with Chernyshinellinae Reytlinger 1958, Tournayellininae Reytlinger 1996 in Rauser-Chernousova et al. 1996), Mstiniidae Lipina 1989 (with Mstiniinae Lipina, 1989, Eotextulariinae Hance, Hou and Vachard 2011, Darjellinae Hance, Hou and Vachard 2011), Palaeospiroplectamminidae Loeblich and Tappan 1984 (with Palaeospiroplectammininae Loeblich and Tappan 1984). Superfamily BRADYINOIDEA Reytlinger 1996 in RauserChernousova et al. 1996 Families: Bradyinidae Reytlinger 1950 (with Bradyininae Reytlinger 1950, Glyphostomellinae A.D. Miklukho-Maklay, 1963), Janischewskinidae Reytlinger 1996 in RauserChernousova et al. 1996. Superorder OZAWAINELLOIDA Solovieva 1980 (nom. corrected pro Ozawainelloidea) Order OZAWAINELLIDA Solovieva 1980 Families: Eostaffellidae Mamet 1970 (including Bozorgniella Cozar and Vachard 2001), Ozawainellidae Thompson and Foster 1937, Reichelinidae A.D. Miklukho-Maklay 1959 stat. elevated (including Quasireichelina Ueno 1992 originally placed by the author of the genus into Staffellidae), and Incertae family with the genera Eostaffelloides, Kangvarella and Hubeiella). Superorder FUSULINOIDA Fursenko 1958 Order STAFFELLIDA A.D. Miklukho-Maklay 1949 Families: Pseudoendothyridae Mamet 1970, Nankinellidae A.D. Miklukho-Maklay 1963, Cheniidae F. Kahler and G. Kahler 1966, Staffellidae A.D. Miklukho-Maklay 1949, Pisolinidae Rauser 1985, Kahlerinidae Leven 1963, Pseudostaffellidae Putrja 1956. Order SCHUBERTELLIDA Skinner 1931 Families: Schubertellidae Skinner 1931, Boultoniidae Skinner and Wilde 1954, Palaeofusulinidae A.D. Miklukho-Maklay 1963, Yangchieniidae Leven 1987. Order FUSULINIDA Fursenko 1958 Order ENDOTHYRIDA Fursenko 1958 Superfamily ENDOTHYROIDEA Brady 1884 Families: Endothyridae Brady 1884 (with Endothyrinae Brady 1884, Omphalotinae Vdovenko 1996, Endospiroplectammininae Loeblich and Tappan 1986), Haplophragmellidae Reytlinger 1959 (Haplophragmellinae Reytlinger 1959, Paraendothyrinae Lipina 1996 in Rauser-Chernousova et al. 1996), Endothyranopsidae Reytlinger 1958 (with Endothyranopsinae Reytlinger 1958, Eoendothyranopsinae Reytlinger 1996 in Rauser-Chernousova et al. 1996, Neoendothyrinae Reytlinger, 1996 in Rauser-Chernousova et al. 1996), Globoendothyridae Reytlinger in Voloshinova and Reytlinger 1959 [family reinstated by Hance, Hou and Vachard 2011, with Globoendothyrinae Reytlinger in Voloshinova and Reytlinger 1959 (includig genera Carbotarima described by Brenckle 2004 and Planogloboendothyra described by Hance, Hou and Vachard 2011 , Eblanainae Hance, Hou and Vachard 2011], Dainellidae Cozar and Vachard 2001 (including Paralysella Cozar and Vachard 2001), Loeblichiidae Cummings 1955 [with Loeblichiinae Cummings 1955, Urbanellinae Hance, Hou and Vachard 2011 (type genus Urbanella Malakhova in Anonyme 1963, emend. Brenckle 1997), Mediocrinae Hance, Hou and Vachard 2011, Quasiendothyrinae Reytlinger 1961], Endostaffellidae Loeblich and Tappan 1984 (including Praeeostaffellina Cozar et al. 2008 and Praeplectostaffella Cozar et al. 2008), Endotebidae Vachard, Martini, Rettori and Zaninetti 1994, Endotriadidae Vachard, Martini, Rettori and Zaninetti 1994. Families: Profusulinellidae Solovieva 1996 in RauserChernousova et al. 1996, Aljutovellidae Solovieva 1996 in Rauser-Chernousova et al. 1996, Fusulinidae Moeller 1878 (with Fusulininae Moeller 1878, Beedeininae Solovieva 1996, Eofusulininae Rauser and Rosovskaya 1959, Quasifusulininae Putrja 1956), Hemifusulinidae Putrja 1956, Fusulinellidae Staff and Wedekind 1910 (with Fusulinellinae Staff and Wedekind 1910, Pulchrellinae Solovieva 1983), Wedekindellinidae F. Kahler and G. Kahler 1966. Order SCHWAGERINIDA Solovieva 1985 Families: Triticitidae Davydov 1986, Rugosofusulinidae Davydov 1980, Schwagerinidae Dunbar and Henbest, 1930 [with Schwagerininae Dunbar and Henbest 1930 (= Biwaellinae Davydov 1984), Pseudoschwagerininae Chang 1963, Paraschwagerininae Bensh 1996], Pseudofusulinidae Dutkevich 1934 (with Pseudofusulininae Dutkevich 1934, Chusenellinae F. Kahler and G. Kahler 1966, Monodiexodininae Kanmera, Ishii and Toriyama 1976), Polydiexodinidae A.D. MiklukhoMaklay 1953 (with Parafusulininae Bensh 1996 in RauserChernousova et al. 1996, Polydiexodininae A.D. MiklukhoMaklay 1953). Order NEOSCHWAGERINIDA Minato and Honjo 1966 Families: Misellinidae A.D. Miklukho-Maklay 1958, Verbeekinidae Staff and Wedekind 1910, Neoschwagerinidae Dunbar and Condra 1927 (with Neoschwagerininae Dunbar and Condra 509 V. I. Mikhalevich: New insight into the systematics and evolution of the foraminifera 1927, Lepidolininae A.D. Miklukho-Maklay 1958, Gifuellinae Kobayashi, Ross C.F. and Ross J.R.P. 2010 based on their new genus Gifuelloides), Sumatrinidae Silvestri 1933, Pseudodoliolinidae Leven 1963. Families: Nubeculariidae Jones 1875, Ophthalmidiidae Wiesner 1920, Meandroloculinidae Bogdanovich 1981, Wiesnerellidae Saidova 1981, Nodobaculariellidae Bogdanovich 1981, Spiriamphorellidae Senowbary - Daryan and Zaninetti 1986. Group of superorders MILIOLOIDS Delage and Hérouard 1896 Superorder SQUAMULINOIDA Mikhalevich 1988 Superfamily DISCOSPIRINOIDEA Wiesner 1931 Order SQUAMULINIDA Mikhalevich 1988 Family: Squamulinidae Reuss and Fritsch 1861 (with Brasiliellinae Loeblich and Tappan 1986 and Squamulininae Reuss and Fritsch 1861). Superorder CORNUSPIROIDA Jirovec 1953 (= Order Cornuspirida Mikhalevich 1980 (part), suborder Cyclogyrina Saidova 1981) Order CORNUSPIRIDA Jirovec 1953 (=Order Cornuspirida Mikhalevich 1980 (part), suborder Cyclogyrina Saidova 1981) Suborder CORNUSPIRINA Jirovec 1953 Family: Cornuspiridae Cushman 1919 [with Cornuspirinae Schultze 1854 (including Barbuspira Neagu 1999), Cornuspiroidinae Saidova 1981, Calcivertellinae Loeblich and Tappan 1964, Pseudocornuspirinae Reytlinger 1993 in Vdovenko et al. 1993, Glomospiroidinae Reytlinger 1993 in Vdovenko et al. 1993, Falsotubinellinae Bogdanovich and Mikhalevich 1983], Hoyenellidae Rettori 1994. Suborder HEMIGORDIOPSINA Mikhalevich 1986 Superfamily HEMIGORDIOPSIDEA Nikitina 1969 Family: Hemigordiopsidae Nikitina 1969 (with Hemigordiopsinae Nikitina 1969, Meandrospirinae Saidova 1981, Turriglomininae Zaninetti in Limongi et al. 1987, Orthovertellinae Mikhalevich 1988, Kamuraninae Trifonova 1984), Hemigordiidae Reytlinger 1993 in Vdovenko et al. 1993. Superfamily SHANITOIDEA Loeblich and Tappan 1986 (nom corrected pro Shanitinidea Mikhalevich 1988) Families: Shanitidae Loeblich and Tappan 1986 (= Shanitidae Mikhalevich 1986), Baisalinidae Loeblich and Tappan 1986 (with Baisalininae Loeblich and Tappan 1986, Septagathammininae Mikhalevich 1988), Neodiscidae Lin 1984 – nomen translat. Gaillot and Vachard 2007, pro subfamily, (after Vachard et al. 2008), with the new genera Uralogordius (in Gaillot and Vachard 2007, and Glomomidiella (in Vachard et al. 2008). Superorder COSTIFEROIDA Mikhalevich 1988 Order COSTIFERIDA Mikhalevich 1988 Family: Costiferidae Senowbary-Daryan and Zaninetti 1986. Superorder MILIOLOIDA Delage and Hérouard 1896 Order NUBECULARIIDA Jones 1875 (=order Cornuspirida Mikhalevich 1980 (part), suborder Nubeculariina Saidova 1981) Superfamily NUBECULARIIDEA Jones 1875 510 Family: Discospirinidae Wiesner 1931 Order MILIOLIDA Delage and Hérouard 1896 Suborder MILIOLINA Delage and Hérouard 1896 Superfamily QUINQUELOCULINOIDEA Cushman 1917 Families: Hauerinidae Schwager 1876 [with Hauerininae Schwager 1876 (including Dervieuxina Popescu and Crihan 2002, firstly placed by the authors of the new genus into Miliolinellinae), Lorettaoidinae Mikhalevich 1988, Flintininae Saidova 1981, Nummoloculininae Saidova 1981, Polysegmentininae Mikhalevich 1986, Danubiellinae Mikhalevich 2005, Hechtininae Mikhalevich 2005, Flintinellinae Mikhalevich 2005, Kayseriellinae Mikhalevich 2008], Tubinellidae Rhumbler 1906 (with Tubinellinae Rhumbler 1906, Articulariinae Mikhalevich 1987, Poroarticulininae Saidova 1981), Pavoninoididae Saidova 1981), Quinqueloculinidae Cushman 1917 (with Quinqueloculininae Cushman 1917, Cribrolinoidinae Haynes 1981, Scutulorinae Mikhalevich 1987, Pseudoschlumbergerininae Mikhalevich 2008, Labalininae Mikhalevich 1988), Spiroloculinidae Wiesner 1920 (with Spiroloculininae Wiesner 1920, Kaloshinae Mikhalevich 1988, Cribrospiroloculininae Mikhalevich 1988, Stellarticulininae Mikhalevich 1988, Parahauerinoidinae Mikhalevich 2005, Elasigellinae Mikhalevich 2008), Planispiroidinidae Saidova 1981, Massilinidae Thalmann 1941 (with Pseudomassilininae Mikhalevich 1988, Rectomassilininae Mikhalevich 1988, Massilininae Thalmann 1941, Miliolinellinae Vella 1957, Flintininae Saidova 1981, Tortonellinae Mikhalevich 1988), Sigmoilinitidae £uczkowska 1974 (with Sigmoilinitinae £uczkowska 1974, Mesosigmoillininae Mikhalevich 1988, Sigmoinellinae Mikhalevich 1987, Longiapertininae Mikhalevich 1987), Glomulinidae Saidova 1981 ((with Glomulininae Saidova 1981, Orthellinae Mikhalevich 2005 (transferred from the Hemigordiopsinae as having true chambers according to Azbel in Subbotina et al. (1981), though its chambers are rather primitive)), Triloculinidae Bogdanovich 1981 (with Triloculininae Bogdanovich 1981, Triloculinoidinae Mikhalevich 1988, Triloculinellinae Mikhalevich 1988, Involvohauerininae Mikhalevich 1986), Pyrgoidae Mikhalevich 1983 (with Pyrgoinae Mikhalevich 1983, Biloculinellinae Mikhalevich 1986, Cribropyrgoinae Mikhalevich 1986, Idalininae Mikhalevich 1988), Miliolechinidae Zaninetti, Ciarapica and Cirilli 1985. Superfamily MILIOLOIDEA Ehrenberg 1839 Families: Miliolidae Ehrenberg 1839, Neaguitinidae Andersen 1984. Superfamily RIVEROINOIDEA Saidova 1981 [(= Reveroininea Saidova 1981, p. 32 (nom. incorr.)] Family: Riveroinidae Saidova 1981. Superfamily AUSTROTRILLINOIDEA Loeblich and Tappan October 9, 1986 (=Mikhalevich October 27, 1986) Micropaleontology, vol. 59, no. 6, 2013 Families: Peneroplidae Schultze 1854 [with Peneroplinae Schultze 1854 (Hottinger (2007) also placed here his new genus Penarchaias], Vandenbroeckiinae Mikhalevich 1988, Renulininae Mikhalevich 1978, Dendritininae Saidova 1981), *Fischerinidae Millandt 1899, *Fischerinellidae Saidova 1981, *Zoyaellidae Saidova 1981. etal plates and integrative systems (such as stolons, tunnels, canal systems) not developed (except in one genus – Delosina having a special canal system and integrated entosolenian tubes in Pleurostomellids); wall monofontinal (= monolamellar), agglutinated in the lower representatives (Hormosinana), may be with calcareous cement and microgranular, in higher representatives (Nodosariana) - hyaline-radial, with regular pseudopores having cylindrical pseudopore canaliculi but without additional seive-plates (basal seive-plate present), growth interrupted, process of forming new chambers unstudied; aperture terminal even in the coiled forms, simple (circular or slit-like or of other configuration) or radial, may be with an entosolenian tube (non homologous to the apertural structures of Rotaliata) or with ampula (= apertural chamberlet), slit-like aperture may be asymmetrical, hooked, the last four apertural types (radial, with entosolenian tube, with ampula, hooked aperture) are unique and are found only in this class, additional apertures and integrated apertural systems absent (with one exception– in Delosina and integrated entosolenian tubes in Pleurostomellids); data on nuclear apparatus and gamete structure absent; mostly free living, benthic forms, rarely parasitic. Ordovician – Holocene. Superfamily MEANDROPSINOIDEA Henson 1848 (nom corr. pro Meandropsinidea Saidova 1981 and Mikhalevich 1988). Subclass HORMOSINANA Mikhalevich 1992 Order *SACCAMMINIDA Lankester 1885 Families: Meandropsinidae Henson 1848, *Rhapydionidae Keijer 1945 (with Rhapydioniae Keijer 1945, Crateritinae Saidova 1981), Fusarchaiasidae Saidova 1981, Hottingerinidae Mikhalevich 1988, ? Orduellidae Sirel 1998 (nom. corr. pro Orduellinidae Sirel 1998). Families: Saccamminidae Brady 1884 [with Saccammininae Brady 1884 (including genus Lavella Nestell and Tolmacheva 2004), Ovammininae Mikhalevich 2003, Caudammininae Mikhalevich 2003, Pilulininae Brady 1884], Thuramminidae A.D. Miklukho-Maklay 1963]. Superfamily SORITIDOIDEA Ehrenberg 1839 Order HORMOSINIDA Mikhalevich 1980 Superfamily HORMOSINELLOIDEA Rauser-Chernousova and Reytlinger 1986, stat. elevated Families: Austrotrillinidae Loeblich and Tappan 1986, Brebinidae Mikhalevich 1986, Pseudohauerinidae Mikhalevich 1986 , stat. elevated, (with Pseudohauerininae Mikhalevich 1986 and *Malatyninae Hottinger 2007). Suborder ALVEOLININA Mikhalevich 1980 (= Alveolinidae Loeblich, Tappan 1964 (including Helenalveolina described by Hottinger et al. 1989); Alveolinellinina Saidova 1981) Families: Fabulariidae Ehrenberg 1839, Alveolinidae Ehrenberg 1839, *Keramosphaeridae Brady 1884 (from Soritida). Îrder SORITIDA Schultze 1854 (= Orbitolinida Ehrenberg 1839, Orbitolitida Wedekind 1937) Superfamily PENEROPLIDEA Schultze 1854 Families: Soritidae Ehrenberg 1839 [with Soritinae Ehrenberg 1839 (with inclusion by Hottinger (2007) of his two new genera Neorhipidionina and ?Neotaberina?, the taxonomic position of the latter obviously needs to be changed), Opertorbitolitinae Loeblich and Tappan 1986], Archaiasidae Cushman 1927, Praerhapydionidae Hamaoui and Fourcade 1973, Cycledomiidae Mikhalevich 1988. INCERTAE SEDIS Family: Milioliporidae Bronnimann et Zaninetti 1971. and a group of families which obviously is heterogenous and belongs to the subclass Miliolana only partially: Pyreninidae Mikhalevich 1999 (Pyrenina ) and Nezzazatidae Hamaoui and Saint-Marc 1970 with Nezzazatinae Hamaoui et Saint-Marc 1970 (Nezzazata, Trochospira, Nezzazatinella) and Coxitinae Hamaoui et Saint-Marc 1970 (Coxites, ?Antalyna, Rabanitina). All these taxa needs furter study. Class NODOSARIATA Mikhalevich 1992 Shells unilocular and multichambered (pseudomultichambered as an exception), predominant shell types - uniserial, hightrochospiral (in a special polymorphinoid spire, absent in any other taxa), planispiral (nearly exclusively involute), typical trochospiral type is fully absent; chambers often subcircular, triangular, V-shaped, palmate, inner additional skeletal chamber septa only in one fossil taxa (Colaniellidae), outer shell sculpture well developed in secreted forms, but outer additional skel- Family: Hormosinellidae Rauser-Chernousova and Reytlinger 1986. Superfamily HORMOSINOIDEA Haeckel 1894 Families: Hormosinidae Haeckel 1894 [(with Hormosininae Haeckel 1894 (including Reophanus – as having true chambers), Nodosininae Saidova 1981 ( without Cribratinoides lacking inner ribs), Polychasmininae Kaminski, 2004)], Ginesinidae Mikhalevich 2003, (stat. elevated), Aschemocellidae Vyalov 1966, Reophacidae Cushman 1910 ((with Reophacinae Cushman 1910 (without Adelungia), Bireophacinae Mikhalevich 2003)), *Oxinoxisidae Vyalov 1968, *Glaucoamminidae Saidova 1981 (transferred from the former Textulariacea by Mikhalevich ( 1999)), Dusenburyinidae Loeblich and Tappan 1984, Cuneatidae Loeblich and Tappan 1984, *Earlandinitidae Loeblich and Tappan 1984, Kunklerinidae Rauser and Reytlinger 1986 (including Leptohalysis from the Reophacinae), Ammolagenidae Kaminski et al. 2009. Superfamily CRIBRATINOIDEA Loeblich and Tappan 1984 Families: Cribratinidae Loeblich and Tappan 1984 (Pseudotriplasia is transferred from Pavonitininae as having a uniserial shell), Thomasinellidae Loeblich and Tappan 1984. 511 V. I. Mikhalevich: New insight into the systematics and evolution of the foraminifera Order AMMOMARGINULINIDA Mikhalevich 2002 Superfamily *AMMOMARGINULINOIDEAPodobina 1978 Subclass NODOSARIANA Mikhalevich 1992 Order LAGENIDA Delage and Hérouard 1896 Families: Ammomarginulinidae Podobina 1978 ((with Ammomarginulininae Podobina 1978 (?Adelungia (transferred from the Reophacidae looking as having arisen from a coiled forms), ?Haymanellinae Mikhalevich 2003, Flabellammininae Podobina 1978 and ?Ammoastutinae Loeblich and Tappan 1984)), Bykoviellidae Loeblich and Tappan 1984 ((reinstated from Vialovinae in which it was placed in synonymy (Loeblich and Tappan 1987) and given stat. nov.) (with Bykoviellinae Loeblich and Tappan 1984, Ponceammininae Seiglie 1991 in Seiglie et al. 1991 and Agardhellinae Mikhalevich 2003)), Orectostominidae Mikhalevich 2003 (type genus Orectostomina Seiglie 1965 is transferred from the Spiroplectamminidae), Lacroixinidae Saidova 1981 [with Lacroixininae Saidova 1981 (reinstated here from the sinonymy of Pseudobolivinidae in Loeblich and Tappan 1987) and Monotaleinae Brönnimann, Whittaker and Zaninetti 1992]. Families: Lagenidae Reuss 1862 (with Lageninae Reuss 1862, Cribrolageninae Mikhalevich 1993), Ellipsolagenidae A.Silvestri 1923 (with Ellipsolageninae A. Silvestri 1923, Oolininae Loeblich and Tappan 1961, Parafissurininae R.W. Jones 1984), Sipholagenidae Patterson and Richardson 1987, Asipholagenidae Reddy and Rajshekhar 2007 (according to the original diagnosis having as in Sipholagenidae a double wall but in distinction of the latter lacking an entosolenian tube). Superfamily *FLABELLAMMINOPSIDOIDEA Mikhalevich 2003 Families: Flabellamminopsidae Mikhalevich 2002 (with Flabellamminopsinae Mikhalevich 2003 and ?Amijiellinae Septfontaine 1988), *Spiropsammiidae Loeblich and Tappan 1961 (from the Pavonitinacea). Order *NOURIIDA Mikhalevich 1980 Superfamily NOURIOIDEA Chapman and Parr 1936 (nom. corr. pro Nouriidoidea Mikhalevich 2004) Families: *Nouriidae Chapman and Parr 1936), Reophacellidae Mikhalevich and Kaminski 2003 in Mikhalevich 2003 [with Reophacellinae Mikhalevich and Kaminski 2003 in Mikhalevich 2003 (the character of the inner structure of Falsogaudryinella and Uvigerinammina Kaminski et al. 1995 entering this subfamily was shown by Kaminski et al. 1995), Pseudoreophaxinae Mikhalevich and Kaminski 2003 in Mikhalevich 2003 and ?Cylindroclavulininae Mikhalevich 2003 (character of the inner “tooth” needs further study)], Barbourinellidae Saidova, 1981 (with ?Spiroplectinatinae Cushman 1928 and Barbourinellinae Saidova 1981), Conotrochamminidae Saidova 1981 Superfamily *LIEBUSELLOIDEA Saidova 1981, stat. n. Families: Liebusellidae Saidova 1981, Marieitidae Loeblich and Tappan 1986. Order *PSEUDOPALMULIDA Mikhalevich 1992 Families: Pseudopalmulidae E.V. Bykova 1959, ? Semitextulariidae Pokorný 1956. Incertae Sedis: family Agglutisolenidae Mikhalevich 2003 (type genus Agglutisolena Senowbari-Daryan 1984, transferred from the Pernerinidae, unclear genus with quite unusual structure). 512 Order NODOSARIIDA Calkins 1926 Suborder SYZRANIINA Mikhalevich 1992 Superfamily CALIGELLOIDEA Reytlinger 1959 Family: Caligellidae Reytlinger 1959 (including Ademassa Vachard in Vachard et al. 1993). Superfamily SYZRANIOIDEA Vachard 1981 Family: Syzraniidae Vachard 1981 (including the recent genus Grigelis Mikhalevich 1981), Earlandiidae Cummings 1955 (including Lobatiquinella Vachard 1994 and Reitlingerlandia Vachard 1994), Eogeinitzinidae Vachard 1994, Paratikhinellidae Loeblich and Tappan 1984 (with Paratikhinellinae Loeblich and Tappan 1984, Saccorhininae Mikhalevich 1995, Juferevellinae Vdovenko et al. 1993). Superfamily EONODOSARIOIDEA Rauser 1993 in Vdovenko et al. 1993 Shell may have secondary radially going septae. Family: Eonodosariidae Rauser 1993 in Vdovenko et al. 1993. Suborder NODOSARIINA Calkins 1926 Superfamily PTYCHOCLADIOIDEA Elias 1950 Family: Ptychocladiidae Elias 1950. Superfamily NODOSARIOIDEA Ehrenberg 1838 Families: Nodosariidae Ehrenberg 1838 (including Rectoglandulina Loeblich and Tappan 1955 reinstated by Karavaeva and G. Nestell 2007 who followed K. Miklukho-Maklay 1969 – cited after Karavaeva and G. Nestell 2007 and including Fingerina Hayward 2012 in Hayward et al. 2012 ), ?Dentalinidae Schwager, 1877, Rimulinidae R.W. Jones, 1984 (stat. elevated, reinstated from the synonymy of Lingulininae), Frondiculariidae Reuss 1860 [belonging of the genus Acostinella of Sarkar and Gupta, 2008 to Reussellidae Cushman 1933 where it was initially placed by the authors of the new genus is doubtful, its radial aperture looks more like nodosariat radial aperture, the fully uniserial hyaline shell with a small pores is also characteristic for the Nodosariana representatives, the genus is closer in its structure to nodosariid Tristix and together with the genus Tribrachia and the genus Acostina (also uniserial entirely and earlier assigned to Reussellidae) probably ought to be united in a separate subfamily Tristixinae subfam. nov. differing from the Frondiculariinae in its shell form – triangular (or quadrangular) rather than strongly flattened, with chambers triangular or quadrangular and with sutures only slightly arched at the center (rather than chambers Micropaleontology, vol. 59, no. 6, 2013 being extremely narrow and of prominent V-shaped character characteristic for Frondiculariinae), aperture may be simple (Tribrachia), radiate (Tristix) or crybrate, deriving from the radiate one (Acostina). Acostinella probably is synonymic with Tristix but the comparison of the material is necessary for the final conclusion], *Nodosinellidae Rhumber 1895, Geinitzinidae Bozorgnia 1973 (including the genus Omoloniella Karavaeva and Pronina 2007), Pachyphloiidae Loeblich and Tappan, 1984 Ichthyolariidae Loeblich and Tappan 1986, Daucinoididae Mikhalevich 1993, Protonodosariidae Mamet and Pinard 1992, Glandulonodosariidae Silvestri 1901 (with Neugeborina Popescu in Cicha et al. 1998), Chrysalogoniidae Mikhalevich 1993 (with 4 new genera in Hayward et al. 2012: Anastomosa Hayward 2012, Cribroconica Hayward and Kawagata 2012, Lotostomoides Hayward and Kawagata 2012, Scallopostoma Hayward and Kawagata 2012). Fuchs 1967], Glandulinidae Reuss 1860 ((with Glandulininae Reuss 1860 (including genus Bombulina Mikhalevich 1983 published in Russian literature only slightly available in the West), Entolingulininae Saidova 1981)), Plectofrondiculariidae Cushman 1927 (with Plectolingulina Hayward 2012 in Hayward et al. 2012), Partisaniidae Loeblich and Tappan 1984, ?Ramulinidae Brady 1884. Suborder *PLEUROSTOMELLINA Mikhalevich 1993 Family: Stilostomellidae Finlay 1947 (with the new genera Carchariostomoides Hayward and Kawagata 2012, Caveastomella Hayward and Kawagata 2012 Toddostomella Hayward 2012, Toddostomella Hayward 2012 – all in Hayward et al. 2012). Families: Pleurostomellidae Reuss 1860 [with Pleurostomellinae Reuss 1860 ( including Czarkowyella Gawor-Biedowa 1987, Delphinoidella of Popescu 1992, Neopleurostomella Srinivasan and Rai 1992, Obesopleurostomella Hayward 2012, Ossaggittia Hayward and Kawagata 2012 (the two latter in Hayward et al. 2012)), Wheelerellinae Petters 1954, Cribropleurostomellinae Owen 1971, Pazdroellinae Gawor-Biedowa 1987 ((with Pazdroella, Triaperturina and Quadryaperturina of Gawor-Biedowa, 1987, the latter genus with unclear structure of the initial part)]], Ellipsoidinidae Silvestry 1923, [a family placed by Loeblich and Tappan (Loeblich and Tappan, 1987) into the synonymy of Pleurostomellidae, reinstated by Hayward et al. 2012 who included the new genus Lateroihiatus (Hayward, 2012)]. Superfamily *COLANIELLOIDEA Fursenko 1959 Order DELOSINIDA Revets 1989, stat. elevated Family: Colaniellidae Fursenko 1959 (with Colaniellinae Fursenko 1959, Multiseptidinae Rauser 1993 in Vdovenko et al. 1993). Family: Delosinidae Parr 1950 Order *PALAEOTEXTULARIIDA Hohenegger and Piller 1975 Superfamily PALAEOTEXTULARIOIDEA Galloway 1933 Families: Vaginulinidae Reuss 1860, Lenticulinidae Chapman, Parr, Collins 1934, Marginulinidae Wedekind 1937, Robuloididae Reiss 1963, Palmulidae Saidova 1981, Lingulinidae Loeblich and Tappan 1961, stat. elevated (with Spirolingulininae Loeblich and Tappan 1986 as its junior synonym), Nanicellidae Fursenko 1959. Superfamily STILOSTOMELLOIDEA Finlay 1947 Families: Palaeotextulariidae Galloway 1933 (including Consorbinella Mamet and Pinard 1992). Superfamily LOUISETITIOIDEA Loeblich and Tappan 1984 Family: Louisettitidae Loeblich and Tappan 1984. Order *BISERIAMMINIDA Mikhalevich 1981 Families Biseriamminidae N. Chernysheva 1941 (including the new genus Parabiseriella published by Cozar and Somerville 2012, Globivalvulinidae Reitlinger 1950 [with Globivalvulininae Reitlinger 1950 (including the four new genera of Marfenkova (1991): Admiranda, Dzhamansorina, Koktjubina, Ulanbela, and the genera Charliella Altiner and Ozcan-Altiner 2001, Labioglobivalvulina Gaillot and Vachard 2007, Retroseptellina Gaillot and Vachard 2007, Septoglobivalvulina Lin, Li and Sun 1990, Verispira Palmieri 1988), Paraglobivalvulininae Gaillot and Vachard 2007, Dagmaritinae Bozorgnia 1973 including Sengoerina Altiner 1999, Paradagmaritinae Gaillot and Vachard 2007 (all cited after Hance et al. 2011)]. Order POLYMORPHINIDA Mikhalevich 1980 Suborder POLYMORPHININA Mikhalevich 1980 Families: Polymorphinidae d’Orbigny 1839 [with Polymorphininae d’Orbigny 1839 (including the new genus of Neagu and Cîrnaru 2001 – Scythimorphina), Falsoguttulininae Loeblich and Tappan 1986, Webbinellinae Rhumbler 1904, Edithaellinae Order VAGINULINIDA Mikhalevich 1993 Class ROTALIATA Mikhalevich 1980 ( = class Globothalamea Pawlowski et al. 2012, in Adl et al. 2012) Shell multichambered only, all regular types of chamber arrangement well presented (except for the glomerate type which is practically absent), mostly trochospiral, planispiral and derived from both of them biserial and triserial ones (rarely secondarily uniserial), in advanced forms often cyclical; inner complications of chamber lumen are strongly developed: from thickened layers of the wall of different structure (labyrintic, spongy, alveolar - in the subclass Textulariana), pillars, septa and septula (of the first and second order) in higher representatives of the Textulariana and Rotaliana, and also of curved plates, valves, tongues, grooves mostly in higher Rotaliata; in higher representatives of both subclasses, chamber number could increase to some hundreds and thousands, and in this case they are often differentiated in their structure (e.g. – embrionic chambers, arcuate median chambers, lateral chambers of varied form in Orbitoides, embriionic chambers, chambers without chamberlets, then chambers with chamberlets and secondary septa in Heterostegina) (additional skeletal plates above the sutures and umbilicus are often, especially in higher Rotaliata; integrative systems developed strongly (stolons in Textulariana, stolons and very complex canal systems in Rotaliana); wall in Textulariana monofontinal (monolamellar), agglutinated and microgranular, with pseudopores (irregular or as canaliculi), with various complication of the inner wall, which could be 513 V. I. Mikhalevich: New insight into the systematics and evolution of the foraminifera cancellate, alveolar, spongy, in Rotaliana – bifontinal (= bilamellar), with true regular pores, having cylindrical pore canaliculi with some transverse seave-plates besides of the basal one, growth interrupted, the contours of every new chamber forming entirely at once, organic and calcareous secretion are divided in the time, alternating; aperture usually at the base of the apertural face both in coiled and in some biserial and triserial forms (in the last ones could became secondarily terminal), may also be areal and siave-like, rotaliate aperture may have inner tooth (of another structure and origin than in Miliolata), higher Rotaliata may have various inner tooth-plates and open subcylindrical or closed cylindrical plates, which could be interconnected and could form a single system, additional apertures well developed in both subclasses (sutural, umbilical, peripheral); some groups of Rotaliana gave rise to planctonic forms with morphological adaptations to the pelagic mode of life (subclass Globigerinana); agamonts of the higher Rotaliata multinucleate, heterokariotic, gametes biflagellate (Glabratella has several somatic nuclei and three-flagellate gametes). Mostly free living, benthic forms (Globigerinana planctonic), rarely attached, some parasitic species are known. Carboniferous - Holocene. Subclass TEXTULARIANA Mikhalevich 1980 Order NAUTILOCULINIDA Mikhalevich 2003 Superfamily NAUTILOCULINOIDEA Loeblich and Tappan 1985 Families: Nautiloculinidae Loeblich and Tappan 1985, Haplophragmoididae Maync 1952, [(with Haplophragmoidinae Maync 1952, (lacking supplementary apertures) (without Ammosiphonina, Buzasina), Debarininae Kaminski 2004 and Trematophragmoidinae Mikhalevich 2003)]. Superfamily ALVEOLOPHRAGMIOIDEA Stschedrina 1936 Families: Alveolophragmiidae Saidova 1981 [with Alveolophragmiinae Saidova 1981 (without Popovia and Sabellovoluta having a terminal aperture from the very beginning), Hemicyclammininae Banner 1966 (without Flabellamminopsis)], Discamminidae Mikhalevich 1980 (without Ammoscalariidae Mikhalevich 1982 with partially tectinous wall and without Glaphyrammina), Cyclamminidae Marie 1941. Order SPIROPLECTAMMINIDA Mikhalevich 1992 Superfamily SPIROPLECTAMMINOIDEA Cushman 1927 Family: Spiroplectamminidae Cushman 1927 [(with Spiroplectammininae Cushman 1927 (without Orectostomina), Vulvulininae Saidova 1981, ?Morulaplectinae Saidova 1981 (coiling unclear – streptospiral?)]. Superfamily SPIROTEXTULARIOIDEA Saidova 1975 Order TEXTULARIIDA Delage and Hérouard 1896 Superfamily TEXTULARIOIDEA Ehrenberg 1938 Families: Pseudobolivinidae Wiesner 1931 (without Lacroixina and Parvigenerina with a terminal aperture from the initial stages), Textulariopsidae Loeblich and Tappan 1982 (without Plectinella), Kaminskiidae Neagu 1999, Textulariidae Ehrenberg 1938 [with Textulariinae Ehrenberg, 1938, Norvanganinae Mikhalevich 2003 – shell like in Textularia, but with supplementary peripheral apertures at the ends of peripheral outgrowths (type genus Norvanganina Mikhalevich 1981, genus reinstated from the synonymy of the Textularia in Loeblich and Tappan 1987 by Mikhalevich 2012), ?Siphotextulariinae Loeblich and Tappan 1985 and Planctostomatinae Loeblich and Tappan 1984], Bigenerinidae Saidova 1981 [with Bigenerininae Saidova 1981, subfam. reinstated from the synonymy of Textulariinae in Loeblich and Tappan 1987 and Cribrobigenerininae Mikhalevich 2003 (type genus Cribrobigenerina Andersen 1961, from Planctostomatinae), Siphobigenerininae Loeblich and Tappan 1986 (transferred from the Valvulinidae)], Textularioididae Loeblich and Tappan 1984 (attached forms). Superfamily SEPTOTEXTULARIOIDEA Loeblich and Tappan 1985 Families: Septotextulariidae Saidova 1975, Tawitawiidae Loeblich and Tappan 1961. Order VERNEUILINIDA Mikhalevich and Kaminski 2003 in Mikhalevich 2003 Superfamily VERNEUILINOIDEA Cushman 1911 Families: Verneuilinidae Cushman 1911 [(without Spiroplectinatinae and Barbourinellinae), with Verneuilinoidinae Suleymanov 1973 (Belorussiella is transferred from the Spiroplectinatinae which are characterized by the terminal aperture), without Duotaxis transferred to the Duotaxidae Mikhalevich 2003, without Eggerellina, Vialovella – transferred to the Globotextulariidae, and without Falsogaudryinella, Paleogaudryina, Paragaudryina, Pseudoreophax, Reophacella, Talimuella, Uvigerinammina transferred to the subclass Hormosinana) and Verneuilininae Cushman 1911 (including Heterostomella transferred from Barbourinellinae), (without Latentoverneuilina)], Prolixoplectidae Loeblich and Tappan, 1985 [with Prolixoplectinae Loeblich and Tappan 1985 (without Karrerulina), Gerochellinae Kaminski 2004 and Caroniinae Brönnimann, Whittaker and Zaninetti 1992], Piallinidae Rettori and Zaninetti 1993, Globotextulariidae Cushman 1927 (with Globotextularia, Cribroturretoides, Gravellina, Rhumblerella, Tetrataxiella, Verneuilinulla and Eggerellina and Vialovella from Verneuilinidae and Hagenowella and Sabulina from Ataxophragmiinae), Tritaxidae Plotnikova 1979, Plectoverneuilinellidae Cetean and Kaminski 2011. Superfamily CRENAVERNEUILINOIDEA Mikhalevich 2003 Families: Spirotextulariidae Saidova 1975 [with Spirotextulariinae Saidova 1975, Duquepsammininae Seiglie and Baker 1987 (Duquepsammina Seiglie and Baker, 1987 is very close to Septigerina)], Novalesiidae Loeblich and Tappan 1984, ?Ecougellidae Loeblich and Tappan 1985, ?Pavonitinidae, Loeblich and Tappan 1961 (without Pseudotriplasia). 514 Family: Crenaverneuilinidae Mikhalevich 2003 Superfamily VALVULINOIDEA Berthelin 1880 Families: Valvulinidae Berthelin 1880 (with Valvulininae Berthelin 1880, Cribrobulimininae Mikhalevich 2003 and Micropaleontology, vol. 59, no. 6, 2013 Vacuovalvulininae Mikhalevich 2003), Eggerellidae Cushman 1937 [with Dorothiinae Balakhmatova 1972, Minouxiinae Loeblich and Tappan 1986, Eggerellinae and Colominellinae Popescu 1998 (in Cicha et al. 1998, p. 71), with the genus Cubanina transferred from the ataxophragmiids by Kaminski and Cetean (2011) on the basis of its canaliculated shell wall], Pseudogaudryinidae Loeblich and Tappan 1985 (with Pseudogaudryininae Loeblich and Tappan 1985 and Siphoniferoidinae Loeblich and Tappan 1985), Clavulinidae Balakhmatova 1973 (with Clavulininae Balakhmatova 1973, Goesellinae Mikhalevich 2003). Middle Triassic – Jurassic. ?Superfamily CHRYSALIDININOIDEA d’Orbigny 1839 Suborder ASTERIGERINIDA d’Orbigny 1839 Superfamily ASTERIGERINOIDEA d’Orbigny 1839 Families: Chrysalidinidae Neagu 1968 [with Chrysalidininae Neagu 1968, ?Pseudodictyopsellinae Septfontaine and De Matos 1998, Tritaxilininae Loeblich and Tappan 1986, Paravalvulininae Septfontaine and De Matos 1998 (without Pseudomarssonella and Riyadhoides], Olgiidae Mikhalevich 2011. Incertae Sedis (to the whole order): Siphovalvulina Septfontaine 1988. Order TROCHAMMINIDA Saidova 1981 Superfamily TROCHAMMINOIDEA Schwager 1877 Remarks. Jadammina Bartenstein and Brand 1938 is a junior synonym of Entzia Daday 1883 as shown by studies at the morphological and molecular level (Filipescu and Kaminski 2011; Holzmann et al. 2012) but according to the rules of Zoological Record the family name ought not be changed. Families: Trochamminidae Schwager 1877 [with the subfamilies Trochammininae Schwager 1877, Trochamminellinae Brönnimann, Zaninetti and Whittaker 1983, Toretammininae Brönnimann, 1986, Zavadovskininae Brönnimann and Whittaker 1988, Arenoparrellinae Saidova 1981, Jadammininae Saidova 1981, Polystomammininae Brönnimann and Beurlen 1977, Ammogloborotaloidinae Kaminski and Contreras 2011 (with the type genus Ammogloborotaloides Kaminski and Contreras 2011 in Kaminski and Contreras 2011)], Vialoviidae Suleymanov 1983, Rotaliamminidae Saidova 1981, Adercotrymidae Brönnimann and Whittaker 1988, Ammosphaeroidinidae Cushman 1927, Valvulamminidae Loeblich and Tappan 1986. Superfamily REMANEICIOIDEA Loeblich and Tappan 1964 Families: Remaneicidae Loeblich and Tappan 1964, Carterinidae (with Carterininae Loeblich and Tappan 1955 and Zaninettinae Brönnimann and Whittaker 1983), Asterotrochamminidae Brönnimann, Zaninetti and Whittaker 1983, Dictyopsellidae Brönnimann, Zaninetti and Whittaker 1983. Subclass ROTALIANA Mikhalevich 1980 Superorder ?ROBERTINOIDA Mikhalevich 1980 Order ROBERTINIDA Mikhalevich 1980 (= Robertinida Haynes 1981) Families: Duostominidae Brotzen 1963, Asymmetrinidae Brotzen 1963, Oberhauserellidae Fuchs 1970. Suborder CERATOBULIMININA Mikhalevich 1980 Families: Ceratobuliminidae Cushman 1927, Epistominidae Wedekind 1937 (with Epistomininae Wedekind 1937, Garantellinae Grigyalis 1977), Reinholdellidae Seiglie and Bermúdez 1965, Epistominoididae Saidova 1981. Families: Asterigerinidae d’Orbigny 1839, Epistomariidae Hofker 1954 (with Epistomariinae Hofker 1954, Eponidellinae Sieglie and Bermúdez 1965, Nuttallidinae Saidova 1981, Palmerinellinae Loeblich and Tappan 1984), Asterigerinatidae Reiss 1963, Amphisteginidae Reiss 1963, Alfredinidae S.N.Singh and Kalia 1972, Actinosiphonidae Adams 1987. Superfamily CLYPEORBIOIDEA Sigal 1952 Families: *Clypeorbidae Sigal 1952, Boreloididae Reiss 1963. Suborder CONORBOIDINA Mikhalevich 1992 Shell low to high trochospiral (at least initially or in one of the generations), at the later stages with a reduced number of chambers per whorl up to two and one; aperture in trochospiral forms at the umbilical side, extending along the base of the septal margin, in elongated forms – terminal, a prominent tooth plate of hemicylindrycal or pillar-like form projects inward from the aperture as a columella-like structure changing its orientation from chamber to chamber even in uniserial representatives thus reflecting the ancestral coiled state. Lower Cretaceous – Upper Cretaceous. Family: Conorboididae Thalmann 1952. Suborder ROBERTININA Mikhalevich 1980 Families: Robertinidae Reuss 1850, Alliatinidae McGowran 1966 ((with Alliatininae McGowran 1966 and Cushmanellinae subfam. nov., type genus Cushmanella Palmer and Bermúdez 1936, (including the genus Sidebottomina Seiglie 1964) – shell nearly planispiral, oval in outline or elongated, biserial (presumably secondarily biserial as the initial part of the shell is slightly asymmetrically curved), with umbilical or peripheral additional chamberlets, main aperture areal or an arch at the base of the terminal face, additional sutural apertures present in both the genera of the subfamily distinguishing the new subfamily from the other representatives of the family). Superorder DISCORBOIDA Ehrenberg 1838 Order ROSALINIDA Delage and Hérouard 1896 Superfamily ROSALINOIDEA Reiss 1963 Suborder DUOSTOMININA Mikhaelevich subord. nov. Shell planispiral to trochospiral and high trochospiral; wall nonlamellar and may be with some foreign particles; aperture single or double, at the base of apertural face. Families: Bagginidae Cushman 1927 (with Baggininae Cushman 1927, Serovaininae Sliter 1968), Cancrisidae Chapman, Parr and Collins 1934 (with Gyroidinoidinae Saidova 1981 and Valvulineriidae Brotzen 1942 placed into synonymy 515 V. I. Mikhalevich: New insight into the systematics and evolution of the foraminifera of Cancrisidae by Haynes 1981 and Revets 1996), Eponididae Hofker 1951 (with Eponidinae Hofker 1951, Sestronophorinae Saidova 1981, Rectoeponidinae Saidova 1981), Neoeponididae Hauser and Keller Grunig 1993, ?Heleninidae Loeblich and Tappan 1987, Mississippinidae Saidova 1981 (with Mississippininae Saidova 1981, Stomatorbininae Saidova, 1981), Rosalinidae Reiss 1963 [with Conorbinidae Reiss 1963 as synonym according to Revets 1996, Conorbina Brotzen 1936, including new genus Albertinopsis of Revets 2002a, which he assigned to this family, though tentatively, without Gavelinopsis Hofker, 1951 (according Revets 2002c) and without Tretomphalus (which either should not receive systematic recognition being only a life stage of Rosalina (according Hansen and Revets 1992) or may be a junior synonym of Cymbaloporetta or Tretomphaloides, both transferred here though tentatively to Cymbaloporidae], Bronnimanniidae Loeblich and Tappan 1984. Superfamily DISCORBINELLOIDEA Sigal 1952 Families: Discorbinellidae Sigal 1952, Pseudoparrellidae Voloshinova 1952 (with Pseudoparrellinae Voloshinova 1952, Concavellinae Saidova 1981, ?Stetsoniinae Saidova 1981), Planulinoididae Saidova 1981, Svratkinidae Bugrova 1989. Superfamily GAVELINELLOIDEA Hofker 1956, stat. elevated. Families: Gavelinellidae Hofker 1956 [(with Gavelinellinae Hofker, 1956 (without Gyroidina according to Revets, 1996), Brotzenellinae Bugrova 1990, Pseudogavelinellinae Bugrova 1990, without Gyroidinoidinae Saidova 1981 placed by Haynes (1981) and Revets (1996) into the Cancrisidae), ?Anomalinidae Cushman 1927 [(with Parrelloididae Hofker 1956, Heterolepidae Gonzales-Donoso 1969 and Oridorsalidae Loeblich and Tappan 1984 as synonyms of Anomalinidae (after Revets (1996), at that he obviously means only part of the Oridorsalidae as the genus Oridorsalis was included by Revets (1996) into the Alabaminidae)], Lingulogavelinellidae Scheibnerova 1972 [(fam. reinstated by Bugrova 1990) (with Lingulogavelinellinae Scheibnerova 1972, Orithostellinae Bugrova 1990)], ?Karreriidae Saidova 1981, Trichohyalidae Saidova 1981. Families: Globorotalitidae Loeblich and Tappan 1984 (became extinct in Cretaceous), Osangulariidae Loeblich and Tappan 1964 (Revets, 1996 placed this family into synonymy of Alabaminidae), Alabaminidae Hofker 1951 (with Oridorsalidae Loeblich and Tappan 1984 placed by Revets 1996 as synonym of this family (with the type genus Oridorsalis) and partially of the family Anomalinidae – see above), Coleitidae Loeblich and Tappan 1984, ?Quadrimorphinidae Saidova 1981. Order GLABRATELLIDA Mikhalevich 1994 Families: Glabratellidae Loeblich and Tappan 1964, Rotaliellidae Loeblich and Tappan 1964 (family transferred to the glabratellids by Pawlowski and Zaninetti from the Discorbacea in Loeblich and Tappan 1987, with the new genus Rossyatella of Pawlowski and Zaninetti 1993), ?Buliminoididae Seiglie 1970 (without Elongobula Finlay 1939 according to Revets 1993), ?Heronallenidae Loeblich and Tappan 1986. Order PLANORBULINIDA Mikhalevich 1992 Suborder PLANORBULININA Mikhalevich 1992 Families: Planorbulinidae Schwager 1877 (with Planorbulininae Schwager 1877, Caribeanellinae Saidova 1981), Cibicididae Cushman 1927, Victoriellidae Chapman and Crespin 1930 (with Victoriellinae Chapman and Crespin 1930, Carpenteriinae Saidova 1981, Rupertininae Loeblich and Tappan 1961), Planulinidaå Bermúdez 1952, Stichocibicididae Saidova 1981, Annulocibicididae Saidova 1981, Stensioeinidae Bugrova 1990 (nom. correct. pro Stensioinidae Bugrova 1990). Suborder ACERVULININA Schultze 1854 Families: Acervulinidae Schultze 1854, Menoellidae Cherchi and Schroeder 2005 (obviously Menaellidae nom corr. pro Menoellidae Cherchi and Schroeder 2005 as type genus is Menaella Cherchi and Schroeder 2005), Homotrematidae Cushman 1927. Order ROTALIIDA Lankester 1885 (nom. corr. Calkins 1909 (after Loeblich and Tappan 1992) Superfamily SPHAEROIDINOIDEA Cushman 1927 Families: Sphaeroidinidae Cushman 1927, ?Pegidiidae HeronAllen and Earland 1928, Tremachoridae Lipps and Lipps 1967, Bueningiidae Saidova 1981. Superfamily ? SIPHONINIDOIDEA Cushman 1927 Family: Siphoninidae Cushman 1927 (with Siphonininae Cushman 1927, Siphonidinae Saidova 1981, Siphoninoidinae Loeblich and Tappan 1984). Order DISCORBIDA Ehrenberg 1838 Superfamily DISCORBINOIDEA Ehrenberg 1838 Families: Discorbidae Ehrenberg 1838 (including the genus Acarotrochus described by Kawagata, Yamasaki and Jordan in 2005), Torresinidae Loeblich and Tappan 1984. Superfamily ALABAMINOIDEA Hofker 1951 516 Families: Rotaliidae Ehrenberg 1839 [with Rotaliinae Ehrenberg 1839 (including the new genera Rotaliconus Hottinger, 2007 and Urnummulites of Boukhary and Scheibner 2009), Sakesariinae Bugrova 1990], Calcarinidae Schwager 1876, Miogypsinidae Vaughan 1928, Pseudorbitoididae M.G. Rutten 1935 (with Pseudorbitoidinae M.G. Rutten 1935, Vaughanininae MacGillavry 1963, Pseudorbitellinae Hanzawa 1962, Pararotaliinae Reiss 1963), Faujasinidae Bermúdez 1952, Notorotaliidae Hornibrook 1961, Cuvillierinidae Loeblich and Tappan 1964, Ammoniidae Saidova 1981 (with Gavelinopsis Hofker 1951 according to Revets 2002 c), Dictioconoidessidae Bugrova 1990. Order ? CHILOSTOMELLIDA Haeckel 1894 Families: Chilostomellidae Brady 1881, Pallaimorphinidae Loeblich and Tappan 1987, stat. nov., Chilostominidae Finger and Gaponoff 1986, Lublinidae Gawor-Biedowa 1987. Micropaleontology, vol. 59, no. 6, 2013 Scheffen 1932 (with Lepidocyclininae Scheffen 1932, Helicolepidininae Tan 1936), Linderinidae Loeblich and Tappan 1984, Droogerinellidae Popescu and Brotea 1995 (type genus Droogerinella Popescu and Brotea 1995), Orbitosiphonidae Matsumaru and Jauhri 2003. Superorder NONIONOIDA Saidova 1981 Order NONIONIDA Saidova 1981 Suborder NONIONINA Saidova 1981 Superfamily NONIONOIDEA Schultze 1854 Families: Nonionidae Schultze 1854 (with Nonioninae Schultze 1854, Pulleniinae Schwager 1877, Melonisinae Voloshinova 1958), Nonionellidae Voloshinova 1958, ?Spirotectinidae Saidova 1981. Superfamily ALMAENOIDEA Mjatliuk 1959 Family: Almaenidae Mjatliuk 1959 (with Almaeninae Mjatliuk 1959, Anomalinellinae Cushman 1927). Order NUMMULITIDA Carpenter, Parker and Jones 1862 Families: Nummulitidae de Blainville 1827, Heterosteginidae Galloway 1933, Discocyclinidae Galloway 1928, Pellatispiridae Hanzawa 1937, Asterocyclinidae Brönnimann 1951. Superorder BULIMINOIDA Saidova 1981 Order BOLIVINITIDA Saidova 1981 Suborder SPIROBOLIVINA Mikhalevich 1998 Suborder ASTRONONIONINA Saidova 1981 Families:Astrononionidae Saidova Loeblich and Tappan 1987. 1981, *Bisacciidae Order ELPHIDIIDA Saidova 1981 Families: Elphidiidae Galloway 1933 (including the genus Bangiana Drobne, Ogorelec and Riccamboni 2007); Haynesinidae fam. nov., Cribroelphidiidae Voloshinova 1958. The new family Haynesinidae with the type genus Haynesina Banner and Culver 1978 (transferred from the Nonioninae) is separated here: shell planispiral, at least at the adult, involute or partially evolute (Protelphidium), moderately compressed, with broadly rounded periphery, biumbilicate, may have asymmetrical sides, chambers gradually increasing in size, not numerous (usually 8 to 10), sutures gently curved, deeply incised towards the umbilicus especially in the later chambers, umbilicus slightly depressed, pustules and prominent tubercules which may become pillarlike strongly developed in the umbilical area along the sutures, may also be present on the apertural face; aperture at the base of apertural face, a low narrow arc, may be obscured by pustules, additional integrative apertural systems developed to a different degree (intercameral lacunae along the septa formed by the bend of the chamber walls going inward posteriorly and fusing to the preceding septal face to form intercameral spaces (in Haynesina), intercommunicating umbilical cavities between septal flaps and supporting pillars in Protelphidium). Paleocene - Holocene. Such inner structures, represented by additional inner spaces or umbilical cavities represent the formations of the secondary level of organization of the apertural system and are not mentioned in the diagnosis of the Nonionidae, being absent in the rest of the nonionid genera. These features (differing in each of the aforementioned genera) make Haynesina and Protelphidium closer to elphidiids though their secondary integrative apertural structures are simpler than the elphidiid canal system. The absence of septal or spiral canals, retral processes or fossettes clearly distinguishes the new family from the family Elphidiidae, as well as absence of any canals – from the Cribroelphidiidae. Order ORBITOIDIDA Copeland 1956 Families: Orbitoididae Schwager 1876 (with Orbitoidinae Schwager 1876, Omphalocyclinae Vaughan 1928), Lepidorbitoididae Vaughan 1933 1933 (with Lepidorbitoidinae Vaughan 1933 and Neumannitinae Rahaghi 1992), Lepidocyclinidae Families: Lacosteinidae Sigal 1952, Spirobolivinidae Saidova 1981 (with Spirobolivininae Saidova 1981 and Sigmavirgulininae Saidova 1981). Suborder BOLIVINITINA Saidova 1981 Superfamily LOXOSTOMATOIDEA Loeblich and Tappan 1962 Families: Loxostomatidae Loeblich and Tappan 1962, Bolivinellidae Hayward 1980, Tortoplectellidae Loeblich and Tappan 1985. Superfamily BOLIVINITOIDEA Cushman 1927 Families: Bolivinitidae Cushman 1927 (with subfamilies Bolivinitinae Cushman 1927, Sagrininae Revets 1996 including the genus Alectinella Revets 1996), Bolivinidae Glaessner 1937, Bolivinoididae Loeblich and Tappan 1984. Superfamily PARABRIZALININOIDEA Revets 1996, stat. elevated. Family: Parabrizalinidae Revets 1996, stat. elevated. Superfamily EOUVIGERINOIDEA Cushman 1927 Families: Eouvigerinidae Cushman 1927, Siphogenerinoididae Saidova 1981. Superfamily PAVONININOIDEA Eimer and Fickert 1899, nom. correct. pro Pavoninoidea Mikhalevich 2000 Family: Pavoninidae Eimer and Fickert 1899. Superfamily MILLETTIOIDEA Saidova 1981 Family: Millettiidae Saidova 1981. Order CASSIDULINIDA d’Orbigny 1839 Families: Cassidulinidae d’Orbigny 1839, Ehrenberginidae Cushman 1927, Cassidulinitidae Saidova 1981, Orthoplectidae Loeblich and Tappan 1984 ((with Elongobula Finlay 1939 according to Revets 1993), the rank of the subfamily elevated by Revets and Whittaker 1991), Cassidinidae Boukhary and Scheibner 2011. 517 V. I. Mikhalevich: New insight into the systematics and evolution of the foraminifera TABLE 1 Summary of the classification presented in this publication. Order BULIMINIDA Saidova 1981 Suborder TURRILININA Saidova 1981 Families: Turrilinidae Cushman 1927 (with Turrilininae Cushman 1927 and Sporobulimininae Mikhalevich 1998 – with multiple aperture, including genera Sporobulimina and Sporobuliminella), Tosaiidae Saidova 1981, Virgulinellidae Loeblich and Tappan 1984, ?Praeplanctoniidae Georgescu 2009 (nom corr. pro Praeplanctonidae) (type genus Praeplanctonia Georgescu 2009 with type species Praeplanctonia globifera Georgescu 2009). Suborder BULIMININA Saidova 1981 518 Families: Buliminidae Jones 1875, Uvigerinidae Haeckel 1894 (with Uvigerininae Haeckel 1894, Angulogerininae Galloway 1933), Reussellidae Cushman 1933 (without Acostina and Acostinella of Sarkar and Gupta 2008 initially placed by the authors of the new genus into this family), Buliminellidae Hofker 1951, ?Biedafranciszkinidae Gawor-Biedowa 1992, Fursenkoinidae Loeblich and Tappan 1961 (without Coryphostoma and Cassidella), Stainforthiidae Reiss 1963, Trimosinidae Saidova 1981, Pappinidae Haunold 1990. Suborder TUBULOGENERININA Mikhalevich 1998 Family: Tubulogenerinidae Saidova 1981 (obviously including the genus Rectobulimina from the Turrilinidae). Micropaleontology, vol. 59, no. 6, 2013 TABLE 1 continued. Suborder CAUCASININA Mikhalevich 1998 Family: Ñaucasinidae N.K. Bykova 1959 (without the genus Tremachora)(with Ñaucasiniae N.K. Bykova 1959, Baggatellinae N.K. Bykova 1959). Subclass GLOBIGERINANA Mikhalevich 1980 Order HETEROHELICIDA Mikhalevich 1992 Superfamily HETEROHELICIDOIDEA Cushman 1927 Families: Heterohelicidae Cushman 1927 (with Heterohelicinae Cushman 1927, Gublerininae Alyulla 1977, Pseudo- gumbelininae Alyulla 1977), Guembelitriidae Montanaro Gallitelli 1957, Chiloguembelinidae Reiss 1963. Order GLOBOROTALIIDA Mikhalevich 1980 Superfamily ROTALIPOROIDEA Sigal 1958 Families: Rotaliporidae Sigal 1958 (with Rotaliporinae Sigal 1958, Rotundininae Bellier and Salaj 1977, Helvetoglobotruncaninae Lamolda 1976, Ticinellinae Longoria 1974), Hedbergellidae Loeblich and Tappan 1961, Praehedbergellidae Banner and Desai 1988. Superfamily GLOBOTRUNCANOIDEA Brotzen 1942 519 V. I. Mikhalevich: New insight into the systematics and evolution of the foraminifera TABLE 1 continued. Families: Globotruncanidae Brotzen 1942 (with Globotruncaninae Brotzen 1942, Globotruncanellinae Maslakova 1964, Abathomphalinae Pessagno 1967, Archaeoglobigerininae Salaj 1987), Rugoglobigerinidae Subbotina 1959. Superfamily GLOBOROTALIOIDEA Cushman 1927 Families: Globorotaliidae Cushman 1927, Pulleniatinidae Brönnimann and Bermúdez 1953, Truncorotaloididae Loeblich and Tappan 1961, Candeinidae Cushman 1927 (with Candeininae Cushman 1927, Tenuitellinae Banner 1982), Catapsydracidae Bolli, Loeblich and Tappan 1957 [with Catapsydracinae Bolli, Loeblich and Tappan 1957 (probably without Protentella), Soldaniellinae Brönnimann, Whittaker and Parisi 1987 (transferred from Globigerinidae by Loeblich and Tappan 1992, as nonspinose)]. 520 Order HANTKENINIDA Mikhalevich 1980 Superfamily HANTKENININOIDEA Cushman 1927 Families: Hantkeninidae Cushman 1927, Globanomalinidae Loeblich and Tappan 1984. Superfamily PLANOMALINOIDEA Bolli, Loeblich and Tappan 1957 Families: Planomalinidae Bolli, Loeblich and Tappan 1957, Globigerinelloididae Longoria 1974 (with Globigerinelloidinae Longoria 1974,? Eohastigerinellinae Loeblich and Tappan 1984), Schackoinidae Pokorný 1958. Order CASSIGERINELLIDA, Mikhaelevich ord. nov. Micropaleontology, vol. 59, no. 6, 2013 TABLE 1 continued. Shell biserial and planispirally enrolled, flattened or subglobular, with chambers from slightly to strongly inflated becoming nearly subsphaerical; umbilicus closed; aperture at the base of the apertural face, as an arched opening or a loop-shaped elongated opening extending up from the base of the apertural surface in parallel to the peripheral margin. Upper Eocene – Lower Miocene. Remarks. Biserially enrolled chambers clearly distinguish this order from all the other orders of the subclass Globigerinana. Superfamily CASSIGERINELLIDOIDEA Bolli, Loeblich and Tappan 1957 Family: Cassigerinellidae Bolli, Loeblich and Tappan 1957. Order GLOBIGERINIDA Carpenter, Parker and Jones 1862 Superfamily FAVUSELLOIDEA Longoria 1974 Families: Favusellidae Longoria, 1974, Globuligerinidae Loeblich and Tappan 1984 Superfamily GLOBIGERINOIDEA Carpenter, Parker and Jones 1862 Families: Globigerinidae Carpenter, Parker and Jones 1862 (with Globigerininae Carpenter, Parker and Jones 1862, Porticulasphaerinae Banner 1982), Marginotruncanidae Pessagno 1967 (with Pessagnoites AlShuaibi 2011), Orbulinidae Schultze 1854 (without the genus Globigerinatella, which is transferred here to the cymbaloporids), Hastigerinidae Bolli, Loeblich and Tappan 1957, Globigerinitidae Bermúdez 1961 (with Globigerinitinae Bermudez 1961 and *Tenuitellinae Banner 1982). For the better understanding of the new classification, a summary down to superfamily level is given in Table 1. CONCLUSIONS Foraminifera are regarded here as a phylum embracing totally 5 classes, 11 subclasses, 73 orders, 27 suborders, 98 superfamilies, 499 families and 368 subfamilies among which two orders (Cymbaloporida, Cassigerinellida), suborder Duostominina, two families (Cymbaloporettidae, Haynesinidae), and two subfamilies (Cushmanellinae and Tristixinae) are new. The macrosystem is built hierarchically with the attempt to reflect the phylogenetic relations of the larger groups in their different ranks. Special attention was paid to the evolutionary significant features and to the degree of their development, reflecting the level of differentiation and integration of the skeletal and cyto521 V. I. Mikhalevich: New insight into the systematics and evolution of the foraminifera plasmic structures. These evolutionary important characters were used to describe the diagnoses of the high rank taxa with a tendency to use the categories of the features of a single level in the description of the taxa of the same taxonomic level. REFERENCES The former Textulariacea and Fusulinacea are considered to be heterogenous groups and their components were distributed among the several classes correspondingly according to their shell morphology and the character of their shell wall ultrastructure. Some taxa with hyaline calcitic shell walls are transferred from the Rotaliana to the Spirillinana (patellinids, chapmaninids) and the Nodosariana (stilostomellids, pleurostomellids). The former inclusion of these groups (nodosarians and spirillinans) into the Rotaliana (the previous Rotaliacea, then - Rotaliina) resulted in the polyphyletic nature of the former rotalians. Some Rotaliana orders and families are possibly still heterogenous (though their heterogeneity more often may be of lower rank than class level). Nevertheless the classification of Rotaliana of Loeblich and Tappan (1987) (with some addition of later data) is to a great degree followed here as being up to the present the most complete and reliable among the others. The new approach to foraminiferal systematics together with a new understanding of their evolution requires a profound revision of many of the taxa within the separated classes and subclasses at a lower (order and family) taxonomic level (especially of agglutinated groups, of the class Spirillinata, of some major groups within the Rotaliana) and to some extent provides a roadmap for such revision. ALTIN, D. Z., HABURA, A. and GOLDSTEIN, S. T., 2009. A new allogromiid foraminifer Niveus flexilis nov. gen., nov. sp., from Coastal Georgia, USA: fine structure et gametogenesis. Journal of Foraminiferal Research, 39: 73–86. The analysis of all the multitude of all the main foraminiferal morphotypes has shown that the morphological regularities are the same for the agglutinated and isomorphic calcareous forms within the separate classes and the difference is more significant between the agglutinated forms of different classes than between the agglutinated and calcareous analogues of the same class. Under such an approach the morphologically similar agglutinated and calcareous shells within each class can be regarded as closely related rather than convergent forms. The origin and development of the classes (including subclasses with agglutinated and calcareous shell wall) took place independently and in parallel in each of the phylogenetic lines examined above. 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