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Mikhalevich, V.I. , 2004. On the heterogeneity of the former Textulariina (Foraminifera). In: Bubik, M. & Kaminski, M.A. (eds), 2004. Proceedings of the Sixth International Workshop on Agglutinated Foraminifera. Grzybowski Foundation Special Publication, 8: 317 – 349.



ABSTRACT _______________________________________________________________ A new suprageneric system of the agglutinated foraminifera is offered consisting of five subclasses (Astrorhizana, Ammodiscana, Miliamminana (= Schlumbergerinana), Hormosinana, and Textulariana) belonging to five different classes, 27 orders, 156 families, 147 subfamilies, and 849 genera. The whole foraminiferal group is regarded as a phylum. The taxonomical significance of the different taxonomic features is discussed. In contrast to the majority of classification schemes adopted, the basic morphological characteristics of the test such as number of chambers (one, two or many), mode of coiling, test and chamber form, structure and position of the main and additional apertures, and the presence or absence of integrative systems are considered as having preferential significance compared with the character of the shell wall. The latter having important but subordinate value. The forms with different kinds of agglutinated wall are considered as ancestral stages in the evolutionary development of various phyletic lineages having different types of calcareous shell microstructure in their most advanced forms. Thus, the former Textulariacea are split into five groups according to their morphology. The taxonomic composition of each group elaborated up to the generic level is given as a preliminary scheme. Five new orders (Plagioraphida, Sphaeramminida Mikhalevich & Kaminski, Loftusiida Kaminski & Mikhalevich, Nautiloculinida, Verneuilinida Mikhalevich & Kaminski), five new families (Cystamminidae, Vaniidae, Reophacellidae Mikhalevich & Kaminski, Agglutisolenidae, Duotaxidae, Crenaverneuilinidae) and 15 new subfamilies (Praesphaerammininae Kaminski & Mikhalevich, this work, Ovammininae, Caudammininae, Agardhellinae, Reophacellinae Mikhalevich & Kaminski, Pseudoreophaxinae Mikhalevich & Kaminski (with the type species Pseudoreophax cisovnicensis Geroch, 1961, non Pseudoreophacinae Suleymanov, 1963 – invalid according to Loeblich & Tappan, 1987 (with type sp. Pseudoreophax marginulinae Suleymanov, 1963 (= Adelungia Suleymanov, 1966)), Vaniinae, Cylindroclavulininae, Trematophragmoidinae, Norvanganinae, Cribrobigenerininae, Voloshinovellinae, Cribrobulimininae, Goesellinae, Vacuovalvulininae) are described.
Mikhalevich, V.I. , 2004. On the heterogeneity of the former Textulariina
(Foraminifera). In: Bubik, M. & Kaminski, M.A. (eds), 2004. Proceedings of the
Sixth International Workshop on Agglutinated Foraminifera. Grzybowski
Foundation Special Publication, 8: 317 – 349.
On the heterogeneyty of the former Textulariina (Foraminifera)
Zoological Institute Russian Academy of Sciences, Universitetskaja nab., 1,
199034, S-Petersburg, Russia,
The new system of the agglutinated Foraminifera is offered consisting of five
subclasses (Astrorhizana, Ammodiscana, Rzehakinana (=Schlumbergerinana),
Hormosinana and Textulariana) belonging to five different classes, 27 orders, 155
families, 149 subfamilies, 842 genera. The whole Foraminiferal group is regarded as a
phylum. The taxonomical significance of the different taxonomic features is
discussed. Opposite to the majority of the classifications adopted, the basic
morphological character of the shell such as number of chambers (one, two or many),
mode of coiling, shell and chamber form, structure and position of the main and
additional apertures, presence or absence of the integrative systems are considered as
having the preferential significance compared with the character of the shell wall.
The latter having important but subordinate value. The forms with the different kinds
of agglutinated wall are considered as ancestral stages (the preceding steps) in the
evolutionary development of (the different) various phyletic lineages having different
types of calcareous shell microstructure in their most advanced forms. Thus, the
former Textulariacea are split into five groups according to their morphology, the
taxonomic composition of each of them elaborated up to the generic level is given as
a preliminary scheme. Eight (7) new orders (Plagioraphida, Haplophragmiida,
Sphaeramminida Mikhalevich et Kaminski, Cyclolinida, Loftusiida,
Ammomarginulinida, Nautiloculinida, Verneuilinida ), eight new families
(Cystamminidae, Vaniidae, Orectostominidae, Lacroixinidae, Flabellamminopsidae,
Reophacellidae Mikhalevich et Kaminski, Agglutisoleniidae, Duotaxidae,
Crenaverneuilinidae, )and 18 new subfamilies (Streptocyclammininae,
Praesphaerammininae Kaminski in Mikhalevich, this work, Biplanatinae,
Ovammininae, Caudammininae, Ginesininae, Bireophacinae, Haymanellinae,
Agardhellinae, Lacroixininae, Reophacellinae Mikhalevich et Kaminski,
Pseudoreophaxinae Mikhalevich et Kaminski (with type species Pseudoreophax
cisovnicensis Geroch, 1961, non Pseudoreophacinae Suleymanov, 1963 invalid
according to Loeblich & Tappan, 1987 (with type.sp. Pseudoreophax marginulinae
Suleymanov, 1963 (= Adelungia Suleymanov, 1966)), Cylindroclavulininae,
Trematophragmoidinae, Norvanganinae, Cribrobigenerininae, Voloshinovellinae,
Cribrobulimininae, Goesellinae, Vacuovalvulininae) are described.
In Foraminifera the agglutinated shell wall is more primitive comparing with the
fully secreted one and it originated earlier than the last one. Hence the foraminiferal
species with the agglutinated wall represent the lower step in the development of the
whole group. To elucidate the position of the agglutinated forms within (among)
Foraminifera and their relations to the other Foraminiferal taxa we have firstly to see
this whole group, to understand how it is organised. We also have to weight (= to
evaluate) the taxonomic significance of the different morphological features of
the shell and of the different kinds of the Foraminiferal shell wall using the method of
the comparative morphology.
Schultze (1854) was the first who distinguishing the major foraminiferal groups relies
himself mostly on the composition and structure of their shell wall evaluating it
then the shell structure. This approach was followed by Reuss (1862), Schwager
(1877), Brady (1884) and till most of the recent classifications including the
remarkable labours of Loeblich and Tappan (1964, 1988). (We don’t regard here the
classifications of Haynes, 1981, Lee, 1990 and some others as though bringing some
valuable changes they follow the main principals of the Loeblich and Tappan
approach). The classifications of the Russian school headed by A. Fursenko and D.
Rauser-Chernousova (1959) though also following this prevailing tendency
nevertheless more emphasized the significance of the shell morphology. In the more
generally accepted recent classifications Foraminifera are regarded as an order or
class including 12 taxa (suborders or orders correspondingly) of equal high rank.
These taxa are separated out according not only to their shell morphology but strongly
to the ultrastructure and composition of their shell wall: ( Allogromiina, Textulariina,
Involutinina, Spirillinina, Miliolina, Silicoloculinina, Lagenina, Fusulinina,
Carterinina, Robertinina, Rotaliina, Globigerinina). Later Loeblich and Tappan
(1992) had added 4 new orders (Astrorhizida, Lituolida, Trochamminida and
Buliminida) and took off 2 of the previous ones (Involutinina and Silicoloculinina).
As a result the whole group included 14 orders but the approaches of the authors
stayed in principal the same. In the further taxonomic work M. Kaminski (2001,
2003 this volume) reclassified the agglutinated forams and recognized the
agglutinated representatives as entering three orders (Astrorhizida, Lituolida,
Textulariida) of one subclass Textulariana eliminating Trochamminida and
Carterinida as a separate orders. He regarded the Trochamminid forms as superfamily
Trochamminacea within the order Lituolida and the Carterinid forms entering the
family Trochamminidae as a separate subfamily. His colossal and necessary work
(Kaminski, 2000, 2003, Kaminski et al.,1995, Geroch et Kaminski, 1995 and others)
permitting him to give the right taxonomic position of all the new agglutinated
genera and families described since 1988 year is hard to overestimate.
In parallel with this studies the new conception of the foraminiferal macrosystem was
offered and developed by Mikhalevich (1980, 1981, 1985, 1988, 1992, 1995, 2000),
Saidova (1981) and Maslakova and Gorbachic (1995). According to this new
conception the predominant significance is given to the morphology of the shell, to
the plan of its structure which is connected with the functions of the whole organism
and in Foraminifera is reflected and preserved in their skeleton. The skeleton is the
most conservative structure of the organism and its form changes even less than its
chemical and structural composition. According to this conception the Foraminifera
are composed of the less number of large groups of equal rank of the classes
Astrorhizata, Spirillinata, Miliolata, Nodosariata and Rotaliata (including subclass
Globigerinana which I leave apart in this article devoted to the agglutinated forms).
All of them have in their composition the subclasses whose representatives have the
agglutinated shell wall (subclasses Astrorhizana, Ammodiscana, Schlumbergerinana,
Hormosinana and Textulariana correspondingly) (pl. 1). The main morphological
features of the forms belonging to the different subclasses within the one class are
There is also the disparity between the different authors in understanding of the
rank of the whole group. It ranges from the order and class and up to the phylum
level. I regard them as a phylum as they sharply differ from all the other protistan
phyla and from the groups of the former Sarcodina (Mikhalevich,1980, Starobogatov,
Mikhalevich, 1985). The ground for such elevation of their rank I’ve recorded earlier
in a set of works (beginning from 1980). In the brief conclusion: it is highly
specialized and evolutionary advanced group having multichambered deeply
differentiated mostly calcareous skeleton with the complex integrative systems unique
among Protists and striking for the unicellular level. The peculiarities of the
Foraminiferal reticulopodia, gametes, life cycles and nuclear apparatus (with the
polymeruzation of nuclei and genoms and the presence of nuclear dymorphism in
their more advanced groups) also well defines this phylum from all the other
protistan phyla.
The foraminiferal rRNA sequences differ from any other known rRNA sequences,
their SSU and rDNA sequences also confirm that Foraminifera represent a separate
monophiletic line of protists (Pawlowski et al., 1997, Pawlowski, 2000).
In the classifications composed under the first approach giving the predominant
significance to the shell wall composition and structure we meet a huge amount of
pairs of the agglutinated shells which are strikingly isomorphic analogue with one or
another kind of the forms with the secreted wall.
Only some of such bright examples are demonstrated in plate 2, I and 2, II:
1. beginning from the simpliest unilocular Lagenammina and Pygmaeoseistron,
Saccammina (very interesting Antarctic new species recently described by me,
Pronina and Nestell (Mikhalevich et all., 2000) and having needle-like projection at
its base) and Lagena with the similar projection, up to the uniserial
pseudochambered Hormosinella and Grigelis, Polychasmina and Geinitzina, the
morphotypes of the agglutinated Pseudopalmula with the unusual V-shape chambers
and of the calcareous secreted Kyphopixa, and at last in the forms having a peculiar
high-spire: the agglutinated Nouria and the some Polymorphinids, Abdulevia and
Paleopolymorphina(pl.2,I : 1 – 7) ;
2. these examples can be continued in the planospiral and trochospiral
pseudotwochambered forms with the long tubular second chamber (Ammodiscus
Spirillina, Arenoturrispirillina Trocholina) and advanced multichambered forms
probably of the same line( Alpillina – Chapmanina and some others) (pl. 1,pl. 2,I:8 –
3. they are very demonstrative in the shells with a few (usually 2) tubular chambers
per whorl both in their agglutinated and calcareous secreted isomorphs with different
miliolid types of coiling [ the irregular(Schlumbergerina Miliola) and regular
glomerate ( Dentostomina Quinqueloculina), sigmoid ( Sigmoilopsis Sigmella),
planospiral ( Spiroglutina – Spiroloculina) and combined miliolid types of coiling (
Pseudoflintina, early stage quinqueloculine with two chambers per whorl, later
planospiral with 3 chambers per whorl Flintina, , early stage triloculine with two
chambers per whorl, later planospiral with 3 chambers per whorl )] as well as in
Miliolid forms with wide and more numeral chambers per whorl (Trochamminoides -
. Hechtina)(pl. 2,II).
4. the similar agglutinated and calcareous analogues are well known in trochoid
(Trochamminidae – Rosalinidae(pl. 2), planospiral ( Haplophragmoididae
Nonionidae (pl. 1)) and derived from them monoaxial forms ( Ataxophragmiidae
some Buliminids(pl. 1)) having in principal another plan of the shell structure than
the previous examples (pl. 1, pl. 2,I: 12, pl.3: 7 – 12).
The rows of such examples can be many times multiplied.
Such isomorphic repetitive forms were considered evolving (rising) as a
product of evolutionary convergence. The phenomenon of the convergence is known
in the evolution of the different groups of organisms but, perhaps, in no other group
it is met so often as in Foraminifera. As if we have two nearly mirror reflections
differing only in their wall character. Why are they so strikingly similar ? The
isomorphic calcareous forms were divided (separated) into several taxa long
ago. Is it right to unite all their agglutinated analogues in one taxon, including all the
known foraminiferal morphotypes as it was previously in Textulariina ? Is it possible
to give them distinctive diagnose when the only common feature is the agglutinated
character of the wall ? (With the separation of the orders Astrorhizida, Lituolida and
Textulariida it is possible to give the distinctive diagnose to the first one, but two
others are still too wide ).
Trying to understand the whole Foraminiferal group I undertook the
comparative morphological analysis of all the main Foraminiferal morphotypes
evaluating the taxonomic weight of the main morphological features of the shell and
their wall (Mikhalevich, 1980, 1981). As a result of the analysis performed it turned
out that it is possible to single out main well outlined morphotaxonomic groups
regarded here as classes: Astrorhizata, Spirillinata, Miliolata, Nodosariata and
Rotaliata. Each of this classes have in their composition the subclasses with the
agglutinated forms (see above).
Inspite of the diversity of the forms in each class it’s possible to outline their
main features concerning: 1. number of chambers: 2.the characteristic chamber
form, 3. pathways of chamber formation resulting in chamber conjunction character
4. form of the shell and predominant mode of coiling, 5.form and position of the
main aperture, 6. the development of the inner apertural structures, 7. the
development of the integrative apertural systems, 8. presence or absence of the
additional apertures, 9. the presence of the additional skeletal plates, 10. presence or
absence of the integrative systems of the whole shell.
Let us look through the display (the presence and the character)
of all of these features (how all these features are expressed ) in
the following classes, each of them including the agglutinated subclasses as well.
1. number of chambers and 2.the characteristic chamber form:
in Astrorhizata shells unilocular, rarely pseudochambered, chambers mostly
circular, tubular or irregular in form (some of the representatives of this class may
occur belonging to another protistan groups as it was shown for Syringammina
(Tendal, 1979), Pelosina (Mikhalevich, Voronova, 1999) and for some other
in Spirillinata - shells pseudotwochambered with long tubular second chamber or
derived from them multichambered, often with the preserved initial tubular part (in
multichambered forms chambers isomorphic);
in Miliolata – shells predominantly multichambered, unilocular ones as an exception,
pseudotwochambered forms also exist, in multichambered shells chambers mostly
tubular, a few (2 -–3) per whorl, or wide, in some shells having wide chambers, the
initial tubular flexostyle often presenting;
in Nodosariata shells may be unilocular (in Lagenids, which I don’t consider as
secondary unilocular, as well as in their agglutinated analogues Saccamminids), but
mostly multichambered, chambers subsphaerical, isomorphic, often of special V-
shaped form not met in any other phyla;
in Rotaliata shells exclusively multichambered, chambers isomorphic, no tubular
chambers present.
Thus among the multichambered shells in Spirillinata and Miliolata we can see the
predominance of the tubular chambers, in Nodosariata – a presence of a special V-
shaped chambers), in Rotaliata predominance of isomorphic chambers (see
Hottinger, 2000). These features characteristic for each class are expressed in each of
the both subclasses of every of the classes in consideration.
3. Pathways of chamber formation (a. the pseudochambers - formed under
continuous growth, when already formed chamber lumen is subdivided, b. true
chambers – newly formed units as a result of additional growth). These two modes of
the chamber formation result in chamber conjunction character (Mikhalevich,
1981) : in Astrorhizata only the first way present, forming pseudochambers only, in
Spirillinata both are present, the first predominates, in Miliolata also both are
present, the second predominates, in Nodosariata first one very rare, mostly the
second one, in Rotaliata exclusively second one, forming true chambers, the first
one only as additional mode forming chamberlets within the chambers.
4. predominant mode of coiling in the pseudotwochambered and
multichambered shells resulting in the form of the shell :
in Spirillinata – irregular glomerate, planospiral (often combined with the glomerate
one), as well as a trochospiral one;
in Miliolata- predominantly irregular and regular glomerate and planospiral (often
combined), trochospiral one only as an exception (e.g. Fisherinella – such type of
coiling is so unusual for Miliolata, that I’ve checked the ultrustructure of its wall to
confirm its belonging to miliolids (Mikhalevich et al., 1988);
in Nodosariata – the predominant shell types are elongated uniserial and monoaxial,
sometimes biserial, planospiral also, a special poymorphinoid and
plectofrondicularian types occur, the trochoid ones absent;
in Rotaliata exclusevely plano- and trochospiral forms and derived from them
monoaxial (mostly bi- and triserial ones).
The predominance of one or another type of coiling is similar in the pairs of
subclasses within the class they belong.
The form and position of the main (initially terminal or areal or initially basal
pl. 4) and additional apertures, the development of the inner apertural structures
and of the integrative apertural systems is the most demonstrative and conclusive in
classes description (Mikhalevich, Debenay, 2001, pl-s 1 - 7 ) and may serve as key
elements to distinguish them .
5)position of the main apertures :
in Astrorhizana - aperture may be absent, may be of irregular position and form,
may be primary multiple, the single aperture in elongated forms is terminal
(Mikhalevich.,1995, Mikhalevich, Debenay, 2001, pl. 1 (2,5,6,8,13 16), pl. 1 of
the present article) ( just the same character of the aperture together with the
similarity of their shells in the representatives having the tectinous wall of the
subclass Lagynana (= Allogromiina) gives me ground to unite both in one taxon -
class Astrorhizata (Mikhalevich, 1999, Mikhalevich et al., 2000) (this is consistent
with the Loeblich and Tappan (1988) and my own previous position, when both taxa
where considered having the equal rank);
in Spirillinata - aperture is terminal, at the end of the tube, though in trochoid, and
especially in advanced multichambered calcareous shells it shows a tendency to be
on the umbilical side, in their more primitive agglutinated forms (subclass
Ammodiscana) it is always terminal in position ( Mikhalevich, Debenay, 2001, pl.1
(17 –24) and pl. 1, pl. 2,I of the present article);
in Miliolata– aperture is always terminal in their both subclasses
(Schlumbergerinana and Miliolana) (it is terminal and is not connected with the base
of the apertural face even in their coiled forms Fisherinella, Alveolinidae)
(Mikhalevich, Debenay, 2001, pl.2 and pl. 1, pl. 2,II, pl. 3: 3 - 11 of the present
article), ( mark the difference in position of the aperture in the trochospiral forms of
Miliolana (Fisherinella) and Textulariana (Trochammina) – pl. 3: 11a and 11b);
in Nodosariata aperture also is always terminal in their both subclasses
Hormosinana and Nodosariana (Mikhalevich, Debenay, 2001, pl.3 and pl. 1, 2, I: 1
7, pl. 3: 1 - 2 of the present article) ,
it is terminal even in their coiled forms (this circumstance facilitates the
distinguishing between for instance Lenticulininae and some involutely coiled
Rotaliina with the basal aperture), this fact makes it necessary to move the
agglutinated planospiral forms with the terminal aperture displaced to the angle of
the shell from Textulariana to the other corresponding taxa of Hormosinana;
in Rotaliata –– aperture is basal in position (at the base of the apertural face) (pl. 4:
2a – e, 3a) in both subclasses Textulariana and Rotaliana, at least initially, becoming
secondary terminal usually in some monoaxial forms (Mikhalevich, Debenay, 2001,
pl.4 and pl. 1, 2, I: 12, pl. 3: 7 - 12 of the present article), (the change of the
apertural position in this leneage is strictly connected with the profound change of
the whole plan of their shell architecture – pl.3, 11b).
6) a special types of the aperture and inner appertural structures:
in Astrorhizata inner apertural structures are not developed (the only inner
apertural structure mentioned in this group was entosolenian tube in the agglutinated
Ovammina, but basing ourselves on this fact (such tubes are characteristic for
Nodosariata) as well as on the morphological similarities of Saccaminids and
Lagenids as we have seen earlier – Saccamminids were moved from Astrorhizana to
Hormosinana ( Mikhalevich et al., 2000));
in Spirillinata no inner apertural structures are developed in their both subclasses
(though in their advanced calcareous multichambered forms rarely an external plates
of a special S – or T- form are present) ( Mikhalevich, Debenay, 2001, pl.1 (17 –24)
and pl. 1 of the present article);
in Miliolata in advanced forms the unique inner tooth and complex tooth-
structures derived from it are well developed in their both subclasses
Schlumbergerinana and Miliolana (Mikhalevich, Debenay, 2001, pl.2, 6 and pl. 1,
pl. 3: 3, 4 of the present article), such kind of the inner apertural apparatus is not
occur in other classes and in other agglutinated taxa.
in Nodosariata - a special types of the radial aperture (sometimes with inner radial
ribs), assymmetrically-fissured aperture, aperture with the inner entosolenian tube
and with additional ampula are present– the first three of the mentioned types occur
also in Hormosinana ( radial with inner ribs - in Nodosinum, assymmetrically-
fissured aperture - in Cuneata , aperture with the inner entosolenian tube - in
Ovammina) (Mikhalevich, Debenay, 2001, pl.3 and pl. 1, pl. 3: 1 - 2 of the present
article), thus here we can see again the profound similarity of the agglutinated and
calcareous analogues within the same class; these apertural types are not met in
any other group of agglutinated or calcareous foraminifera and well separate the
Hormosinana from Litoulids.
in Rotaliata – a very complex special inner apertural structures are often developed (
tooth-plates, valves, groves, tubes (of another kind than nodosariate entosolenian
ones) in elongated and trochoid forms) (Mikhalevich, Debenay, 2001, pl.5 (1 –9)
and pl.3:7 - 10 of the present article) not met in any other group. The perfect photoes
of Hottinger (Hottinger et al., 1993) of the Clavulina’s inner apertural plates similar
to that ones of some Buliminids does not leave any doubt of the close affinity of the
agglutinated and calcareous analogues of Rotaliats.
7. integrative apertural systems (only in some taxa with multichambered
representatives, having inner apertural structures):
in Miliolata – practically not developed;
in Nodosariata – purely developed only in some representatives of their calcareous
subclass Nodosariana (in Pleurostomella, Ellipsopolymorphyna (Mikhalevich,
Debenay, 2001, pl.3 (9, 21)), though the very elongated apertural neck in Ginesina
and elongated inner radial ribs in Nodosinum may promote to form the longitudinal
cytoplasmic flews;
in Rotaliata integrative apertural systems are developed strongly and widely
(Mikhalevich, Debenay, 2001, pl.5 (1 –9) and pl.3: 8 - 10 of the present article) and
though being more strongly developed in the calcareous subclass Rotaliana
nevertheless occur also in the agglutinated forms of the subclass Textulariana
(example given .Clavulina (Hottinger et al., 1993), pl. 3: 7a, 8a, 9a of the present
article), in the process of the further study such examples among the Textulariina
supposingly will be more multiple.
8. presence or absence of the additional apertures:
in Spirillinata sutural additional apertures along the spiral suture are met only in
their calcareous subclass Spirillinana;
in Miliolata and Nodosariata sutural additional apertures occur only as an
exceptions in their calcareous subclasses;
in Rotaliata sutural, periferal and umbilical additional apertures are widely
developed in their both subclasses, often in quite similar way (e.g. Toretammina
Schwancia, Balticammina - Discanomalina (pl. 3: 12a,b)).
9.additional skeletal plates:
well developed only in Rotaliata in their both subclasses, often representing the
mirror similar analogues (e.g. AsterotrochamminaDiscorbis (umbilical plates) (pl.
1), Arenagula Neocribrella (apertural plates).
10. the development of the integrative systems of the whole shell: canals are
developed in the pseudotwochambered forms with the long tubular second chamber
(Spirillinata) and in the multichambered forms (Rotaliata) only in the representatives
of their higher subclasses (pl. 1), stolons are met in both subclasses of Miliolata and
in both subclasses of Rotaliata.
I’d like to emphasise that all the taxonomic features in consideration are
evolutionary and functionally a very significant ones. Combining the sets of all the
enumerated features and also some other additional less significant ones it is
possible to give the distinctive diagnosis to each of these lineages while the previous
Textulariina included all the known foraminiferal morphotypes and the only
common feature for all of them was the agglutinated wall. The similarity of the
subclasses of the forms having agglutinated and calcareous wall within each class
was stressed during this display.
There are also some additional evidences of the unity of the agglutinated and
of their calcareous analogues within this groups:
11. The similarity of the tendencies of the onthogenetic and phylogenetic
development of the representatives of the pairs of the subclasses within each class.
For instance the development of the complex inner tooth from the previous simple
one in Miliolana and Schlumbergerinana (pl. 3: 3 - 5) , the change of the position of
the aperture from the basal at the initial stages into the terminal one at the final stages
in monoaxial forms of the subclasses Textulariana and Rotaliana (pl.4: 3).
12. The groups discussed above differ as well in the mode of growth and of
the formation of the new chamber. Though these processes are studied insufficiently
and unevenly in the different groups - the comparative analysis of the previous
works and our own observations permits to conclude that this mechanism has the
profound distinctions in the groups in consideration.
Within the Rotaliata the situation is just the diverse – we can see
the similarity in the process of the building of the new chamber in Textulariana
(Angell, 1990 on Trochammina inflata, Bender, 1992 on Textularia candeiana with
the mixed agglutinated-microgranular wall) and in Rotaliana (Angell, 1979 on
Rosalina floridana ): Their main steps are the following: 1. the exit of the veneer of
the reticulopodia through the aperture of the last chamber forming the cyst around
this last chamber (not around the whole test); 2. formation inside the cyst in front of
the aperture of the last chamber of the cavity having approximately the size of the
future chamber which defines the space for its building (predeterminating its future
form just at once); 3. formation of the outer organic layer (envelope = plasmatic
membrane ) around the cytoplasmic projection; 4. the deposition of the
biomineralized calcareous matter in the direction from the distal to the proximal part
of the chamber wall , the oriented disposition of the crystals forming the cylindrical
cavities of the pore system ( evidently there preexists the chemical “marking” of the
plasmatic membrane into the zones participating in the deposition of the calcareous
matter and the islands” in which such deposition is absent (the future pores)
(Labas, Mikhalevich, 1987); 5. formation of the organic lining of the pores; 6.
formation of the thin inner organic layer of the chamber after the cyst is destroyed
and then the increasing of this layer into the several layers during the subsequent
growth of the shell with the addition of the each new chamber.
The difference of the formation of the new chamber of the Rosalina and of the
other hyaline Rotaliata (opposite to Miliolata) is that the calcareous crystal units are
laid at the both sides of the POM which obviously corresponds to the “envelope” or
the “plasmatic membrane” in Textularia candeian . In the both cases the new
chamber is outlined at once and the organic and calcareous secretion are of
alternating sequence and are separated in time opposite to the process of the new
chamber formation in the representatives with the porcellaneous wall. In tubular
forms of Miliolana the new chamber is growing piece by piece (according to
Angell, 1980 stadying the Spiroloculina hyalina and to our own observations
in Massilina secans (Mikhalevich, 1999, Mikhalevich, Debenay, Stauff, in the
preparation for the publication). The process begins from the formation of the cyst
around the whole test, then the exit of a small amount of the cytoplasm through the
aperture. The chamber wall is build with the participation of this cytoplasm and the
reticulopodia not at once but piece by piece in the direction from the previous
aperture, opposite to it. The cytoplasm is transporting to the place of building two
kinds of the vacuoles – with the organic matter and with the secreted needle-crystals
formed in the Golgi apparatus (Hemleben et al., 1986). The calcareous needles are
embedded into the organic layer of the future wall ( not laid on it). The organic and
calcareous secretion are going simultaneously. Each subsequently built new wall
segment is then underlined by the thick inner membrane differing in its composition
and tightness from the organic matter in which the needles are embedded. When the
process of the building of the new chamber is finished the shell turns out and goes
out of the cyst. The cytoplasmic projection and the envelope are not formed in
tubular Miliolata ( In the process of building of the wide chambers of Peneroplis
(Wetmore, 1999) the anlage was observed but the mechanism of laying secreted and
organic material is supposingly the same as in tubular Miliolata.
The process of the chamber formation in hyaline radial Nodosriata type is not
The rest hyaline radial Spirillinata type differ from the other hyaline radial types
concerning the mode of the chamber growth - it goes (according to the literary data –
Loeblich, Tappan, 1988) as a gradual (continuous) lateral accretion along the edge
of the chamber wall, neither cyst nor envelope are not formed.
Thus the processes of the chamber formation differ in the majority of the classes in
13. The gradual change of the agglutinated shell wall into the characteristic
miliolidean one also shows the close affinity of the morphologically similar
representatives of the subclasses Schlumbergerinana and Miliolana ( In the
Dentostomina agglutinans long calcareous needles underlining the inner surface of the
shell and forming the tooth are combined with the thick agglutinated wall
(Mikhalevich et al., 1986, fig. 5 of the present article). Such combined shell wall with
the calcite needles forming very thin inner layer in another agglutinated species of the
Schlumbergerinana was observed also in Siphonaperta compta (Mikhalevich et al.,
1986) and in Schlumbergerina alveoliniformis and Sigmoihauerina bradyi
( Hottinger et al., 1993).
The absence of the distinct boundary between the agglutinated forms and the
microgranular calcitic ones among the other agglutinated groups, the variable amount
of the microgranular matter in their wall composition was also marked (Coleman,
1980; Plotnikova, Egorova, 1982; Toksvad, Hansen, 1983; Bender, Hemleben, 1988;
Bender,1992, Mikhalevich, 1995).The examples of the transition of the agglutinated
wall into hyaline radial one though very rare are nevertheless exist: in Sabaudia and
in some Orbitolinids the hyaline radial wall composes part of the agglutinated test. The
formation of pores during the evolution of the agglutinated foraminiferal
representatives obviously was also going gradually , the variable kinds of the
pseudopores could not be a stable feature. Some authors (Desai, Banner, 1987; Banner
et al., 1991) had showed the iterative change of the microgranular calcitic canaliculate
forms from the non canaliculate ones. It’s also possible that namely the presence of the
tight calcareous material in the wall with the closely packed particles makes it
necessary to communicate with the surrounding water through the pore system while
the organic cement of the wall permits the penetration of molecules without forming a
special openings (nevertheless irregular occasional openings between the sand
particles are present in such wall – fig. 6 a – d). The species of the same genus often
have the variable proportion of the sand/calcitic material in their shell wall as well as
the different kinds (morphotypes) of the organic cement in the organically cemented
genera. Thus it is hardly reasonable to create the new isomorphic pairs of genera
among the agglutinated groups such as Eggerelloides Eggerella, Karrerulina
Karreriella and so on. And from the taxonomic point of view the variable unstable
feature can hardly be of great use. More over, the unevenly investigated material -
only 13% of the known agglutinated genera (Bender, 1995) (and among them
Pelosina was included which does not belong to Foraminifera at all (Mikhalevich,
Voronova, 1999) does not permit to take these features as the ground for the
taxonomic constructions. For example, in the Dentostomina agglutinans which was
considered to be non canaliculate species some irregular pore-like openings were
unenspectedly discovered (fig. 5,7). In the Lituolida though identified like
noncanaliculate the character of the wall is not studied in the majority of the type
species and this feature is extrapolated on the basis of the morphological similarity. In
Placopsilina the canaliculi were described (Bender, 1995). In Prolixoplectidae only in
the 3 genera among the 9 composing the family the shellwall is surely marked as
noncanaliculate. The data of Loeblich and Tappan (1985) had shown that even in the
recent species of Dorothia the canaliculate character is not always clearly seen and
may be masked as a result of calcification. Pseudopores were discovered in Haddonia
(Coscinophragmatoidea) (Hottinger et al., 1993) considered previously
noncanaliculate, in noncanaliculate Ataxophragmiids Coskinolina and Coleiconus
have “radial pores”, canaliculate Parurgonina was placed in Lituolida,
noncanaliculate Indomarssonella and Cylindroclavulina were placed among
canaliculate forms. The pseudopores of Glaucoammina, “scattered and irregular”
could hardly be called as canaliculi and the genus should be moved into uniserial
Hormosinana according to its shell morphology. Somewhat similar character of the
pseudopore openings we observed in Ginesina (Hormosinidae) (fig. 6a). In
Cribratina (Astrorhizana) shell wall is also penetrated (fig. 6). These features of the
agglutinated wall and their diagnostic value requires further investigations. The
questions about the taxonomic significance of the wall structure versus the
morphology in larger agglutinated Foraminifera were also risen by Gooday and Smart,
The morphological structure of the shell though also variable serves as more stable and
conservative base in taxonomic purposes.
14. The affinity (close relationship) of each pair of subclasses within
classes Miliolata and Rotaliata is also supported by the data of the molecular analysis
performed by Dr. Pawlowski with co-authors (Fahrni et al., 1997, Pawlowski et
al.,1995, Pawlowski, 2000). This is just the case when the results of the
morphological and molecular study coincide.
15. The classification proposed here can also be supported by the biostratigraphic
record: usually in each philogenetic line in consideration the agglutinated forms
were found earlier than their secreted analogues (fig. 8: 2, 9, 10, 12 ). The
representatives of the subclasses Astrorhizana (class Astrorhizata) and
Ammodiscana(class Spirillinata) are known from Cb. In the case of Miliolata we
have not such evidence, may be due to the several reasons. One of them might be that
the agglutinated Miliolata (Schlumbergerinana) are too fragile . Another one that
it’s difficult to identify their initial planospiral and glomerate tubular forms from the
forms of the ancient Spirillinata as the primitive forms in both groups are not reach in
distinguishable features . May be just in this case the molecular methods may be of the
most use. Fahrni et al., 1997 analysing the molecular data had supposed the more
earlier beginning of this group than it was recorded. I also hypothesized their pre-
existence much earlier (Mikhalevich, 1992, 2000). I think we need to find their roots
in the previous Fusulinoida (Fusulinacea) which I consider to be the heterogenous
I hope I managed to show in this brief review the profound similarity of the basic
shell structures, the unity of each of the classes enumerated and the continuity of the
morphological features within each phyletic lineage (class) from the agglutinated to
the secreted calcareous groups though in their agglutinated groups many of the
features and especially the complex structures are usually less developed and less
expressed. They are not realised so fully due to the lower constructive potentials of
the agglutinated wall comparing with the calcareous one.
There are some consequences (=results) from the analysis performed:
1. As the morphological regularities are the same for the agglutinated and calcareous
forms within the separate class and as they differ more significantly between the
agglutinated forms of the different classes than between the agglutinated and
calcareous analogues of the single class I regard the different groups of the
agglutinated Foraminifera as the preceding step in the development of their calcareous
analogues, and hence the last ones - as their direct descendants. Therefore the
relations between the corresponding pairs of such groups are not convergent ones, but
the close affinity.
2. Thereby the different kinds of the calcareous secreted wall evolved in the
evolutionary development of the Foraminifera independently and in parallel in
different phyletic lines and at the different periods of the geological history ( the
Spirillinata’s calcareous type wall was formed in the Upper Devonian, Nodosariata’s –
earlier in Devonian, Rotaliata’s - in Trias).
3. Thus the taxonomical significance of the composition and the ultrastructure of the
Foraminiferal shell wall has though important nevertheless subordinate significance
comparing the shell morphology. In the classification presented it is characteristic for
the taxa of the subclass (not class) level (in class Astrorhizata Saidova, 1981 it is
tectinous in the subclasses Lagynana Mikhalevich, 1980, and in questionable
Ammoscalariana Mikhalevich, 1980 (having tectinous or partly tectinous wall but
multichambered test) and agglutinated and microgranular in the subclass
Astrorhizana, it is agglutinated and microgranular in the 4 rest subclasses with the
representatives having the agglutinated wall).
4. The former Textulariina represent heterogenous rather then monophyletic group.
5. The agglutinated Foraminifera (together with the tectinous ones) are composing
entirely the class Astrorhizata Saidova, 1981 and the lower subclasses Ammodiscana
Mikhalevich, 1980, Schlumbergerinana Mikhalevich 1992, Hormosinana Mikhalevich,
1992 and Textulariana Mikhalevich, 1980 of the classes Spirillinata Maslakova, 1990,
Miliolata Saidova, 1981, Nodosariata Mikhalevich, 1992 and Rotaliata Mikhalevich,
1980 accordingly.
6. This new classification eliminates the significant number of the cases of parallelism
and convergence of the previous systems, comes nearer to the phylogenetic
development of the group and gives more distinctive diagnoses based in the first turn
on the shell morphology.
7. The new approach to the Foraminiferal taxonomy together with the new
understanding of their evolution requires the profound revision of the previous
agglutinated taxa and to some extent shows the pathways for such revision. In a set of
articles and in the two newly published books (Mikhalevich, 1999, 2000) I began to
move in this direction (for instance separating the genuine Lagenids (such as
Collaniellids, Paleotextulariids) and some unilocular shells from the Fusulinid group.
The taxonomy of the subclasses Astrorhizana and Miliolana are elaborated by me up
to the generic level (Mikhalevich, 1988, 1995), the rest subclasses up to the family
level (Mikhalevich, 1992, 1998, 1999, 2000). But to elaborate the new up date
classification of all the agglutinated Foraminifera we have to unite the efforts of the
scientific collective.
The taxonomic classification of the agglutinated forms represented here is elaborated
unevenly as it was mentioned above. And this classification is not finished yet as the
large groups need thorough revision. Except the classification of the subclass
Astrorhizana given in detail the classification of the rest 4 subclasses is to some extent
only outlined (contoured) in the direction of the future investigations but showing
the possible preliminary (initial) way of their examination (review,
inspection). That’s why some of them are placed questionably (marked by ?).
The former Textulariacea and Fusulinacea are to my opinion the two large
heterogenous groups mostly requiring to be revised. The separation of the five main
phyletic foraminiferal lines (classes Astrorhizata, Spirillinata, Miliolata, Nodosariata,
Rotaliata) and of the regularities of the development of their main structures
(Mikhalevich, 1981,1992, 1998, 2000, Mikhalevich, Debenay, 2001) may help us to
bring to light their roots in the previous two groups. It is more easier to demarcate
(=differentiate) the complex multichambered forms with the complex
apertural structures but in the case of the unilocular or simple pseudotwochambered
forms with the second tubular chamber scarce in their features it is nearly impossible
and just in such cases we especially need the addition of the molecular methods. It was
suggested earlier (Loeblich, Tappan, 1988) that the Nodosariid, Spirillinid and
Miliolid forms may have evolved from some extinct Fusulinids. I think that more over
some of the former Fusulinacean groups are themselves the ancient representatives of
these three phyletic lines. It is clear in the case of the ancient Nodosariids (=Lagenids)
and of the more advanced Spirillinids, especially of those having the radial wall layer
(Mikhalevich, 1992, 2000).
The tubular pseudotwochambered forms such as the more primitive Tournayellids
with the undivided or only partly and irregularly subdivided tubular chamber may be
compared and formally may be attributed to the Spirillinids or to the Miliolids. But
evolutionary line Tournayellids- Endothytids Fusulinids is one well expressed and
documented united line more resembling Miliolids and I dare to assume that they may
represent a special ancient Miliolata’s branch, at least to be developed from the
common root as a separate branch having much in common. Some bases for such
unexpected hypothesis may be as follows:
1). The similar form of the test and the chambers in the primitive (compare
Endochernella and Baisalina(Pl. 9,I: 1) with short protrusions of the chamber roof
forming pseudochambers, Endothyranella Zoella, Tournayella Fisherina, the
spiral side of the Fisherinella) and complex advanced forms (some Pseudoendothyra
some Reticulogira (not endoskeleton), some Fusulinids some Alveolinids, some
Rhapydionids (Pseudedomia) (not endoskeleton)). The fusulinoid forms are met only
in the Fusulinid and Alveolinid phyletic lines.
2). The predominance of the glomerate or planospiral (or combined) coiling with the
often change of plan of coiling in both lines (I do not regard the Tetrataxacea
belonging to the Fusulinacean line.)
3). Many of the multichambered forms in both lines have chambers of more or less
distinct tubular character even in the planospiral evolute forms which are circular in
outline (e.g. Tournayella Fisherina, Dolosella) or may preserve the traces of such
character in more complex forms.
4). The terminal position of the aperture (Pl. 4: 1d, pl. 9, II). Though in such forms as
in Tournayella, Endothyranella and some other Tournayellids the aperture is marked
in their diagnosis as basal as a matter of fact it represents more or less wide circular
or semicircular open end of the tubular chamber and differs in principal from the true
basal aperture of Rotaliata (Mikhalevich, Debenay, 2001). The multiple circular
openings at the broad septal face of Fusulinids are slightly elevated above the septal
base as in Alveolinids .
5). In many cases the special character of septation, of the inner formations at the
chamber floor sreening the aperture and foramen are similar in form and position in
both lines (Pl. 9, I; 9,II) (the depositions at the chamber floor of Nodochernyshenella
and Bosniella (pl. 9, I: 2), Avesnella and Melathrokerion, Danubiella, Hechtina(Pl. 9,
II:4). In sections of recent Miliolids their flat teeth (flap) screening the aperture often
have the same triangular form at the base of the chamber (pl. 9,II:3), in some positions
it looks like a spine (pl.9,II:2d) strongly resembling the inner spine-like projection of
Globoendothyra and related forms (pl.9,II:2c). Such bottom depositions occur only in
these two groups.
6). The facts, that the special porcellaneos Miliolata’s wall can turn into the
microgranular one in ancient forms (e.g. in Triassic Ophtalmidium triski Janger the
microgranular grains look in LM as a dark layer( Gubenko, 1988). The same author
watched the microgranular structure of the wall in the Permian Agatammina pusilla
(Geinitz). Recristallization in living porcellaneous Archais angulatus was described by
Macintyre, Reid, 1998), of carboniferous and Permian Trepeilopsis by Loeblich,
Tappan, 1988. The recrystallization of the more ancient forms is the more possible.
Who knows if the three layered recrystallized miliolid wall couldn’t look somewhat
similar to the three layered fusulinacean one ? The miliolid wall of some recent
Antarctic species in the exemplars of large size at the adult stages becomes very thick
and differentiated into the three or four layers (Pl. 10): except the thin tightly packed
external and internal outer layers (comparable with the outer and inner tectoriumum) a
the inner thick friable layer of the needle crystals is pierced by the radially going units
of crystals with the longitudinal between them cavities widening at the upper part of
this layer..) and differing though not sharply from its lower part(compare with
fusulinoid wall). As a result the transverse section of the wall looks radial (similar to
fibtous radial character of the wall of many fusulinids). The radial character of the
miliolids wall could be seen even in the light microscope and I observed such radiality
in many large thick-walled Antarctic species (Pyrgo sarsii, P. peruviana, Pyrgoella
sphaera, Planispirinoides bucculentus and some others). These radially going cavities
are sometimes opened at the inner shell surface like pore openings (Pl. 10, 11).
Somewhat similar though more shorter and scarse radial “tubules” and pseudopore
openings were marked in Archaias angulatus (Fichtel et Moll) (Gudmundson, 1994).
The alveolar structure of the wall of some Miliolata (see AustrotrillinaLoeblich,
Tappan, 1988, pl. 362, fig. 10 –14, Pseudobroeckinella Ibid.,pl. 403, fig. 7) could
also be compared to some extent with the wall of some fusulinaceans. (The forms with
the typical vitreous hyaline radial layer I do not attribute to the Fusulinacea). The
unstable and variable character of porosity in both lines is also comparable and it
seems to be correlated with the thickness and the friability of the wall. Though the
majority of the Miliolata are imperforate for some of them straight or branching pore
canaliculi are marked (Milioliporinae, Kamuraninae, in Peneroplis and Nubeculina).
In this case (as well as in many others) the presence or the absence of the wall
openings could not serve as a feature having the taxonomic significance for the whole
group, it could be used only for the taxa of the lower level.
The thickness of the wall in Miliolids varies strongly in closely related species and
even in one exemplar during its growth, it is thinner and not radially arranged in the
yuanger chambers becoming thicker and radial in the adult ones. This tendency could
be observed in Fusulinids also. The thick inner layer of miliolid wall could look radial
through or has less structured lower part, the more tight wery thin inner and outer
layers are often destroyed and thus the wall varies from 4 to 3 or 2 layered one just as
it is the case with the Fusulinid wall.
Except the variable thickness of the wall in these two groups (Miliolids and
Fusulinids) the amount of the agglutinated material included into microgranular wall
also varies strongly.
7). Fahrni’s et al.,1997 supposition (see above) of the more earlier rise of Miliolata
also gives ground to search their roots in Fusulinoid group.
The main objectives against such hypothesis is the difference in the character of the
inner shell structure( the absence of homata and fluted septa in the Miliolata) . But it
should be noticed that in Miliolata the possibility of the laying of the floor deposits in
the last volutions is also developed and sometimes thise depositions have similar
dentate character (see Periloculina and Maklaya and Cancellina(pl. 9, I:4,5 ). The
similarity of screening (=shelding) deposits at the bottom of the chamber far back the
last septa in some Tournayellida, Endothyrida and some Miliolids was mentioned
above (Pl. 9, I; 9, II). The form of fusulinid chomata at both sides of the aperture could
be similar in form (as it is seen from the sections (pl. 9, I:6)) with weak depositions at
the sides of Pyrgo proloculus. The chomata are developed in different Fusulinid
groups to different extent.
On the base of the characters viewed above it is possible to suppose that Fusulinids
represent a special Miliolata line with a strong and peculiar direction of their inner
structures in advanced forms. In this case the Tournayellids with the agglutinated wall
ought to be included in the subclass Schlumbergerinana. Of course these hypothesis
has a preliminary character and needs further investigations.
The significant difficulties arise also to make distinction between the multichambered
agglutinated planospiral involute tests with the terminal aperture among which the
forms belonging to Miliolata (Schlumbergerinana) and Nodosariata (Hormosinana)
may exist, conventionally speaking the Peneropline and Lenticuline types (with the
simple, non radial aperture) among the agglutinated forms. The additional
morphological features, the molecular data and may be in some cases the wall
character could be of use in these studies.
Many genera need to be reinvestigated as their apertural structures and position of the
aperture at the initial stages of their development are not known yet.
The former Textulariacean forms included here in subclasses Schlumbergerinana and
Hormosinana are described as noncanaliculate. The forms isomorphic to higher
Rotaliata and included here in subclass Textulariana unite both canaliculate and
noncanaliculate taxa.
The endings of the superfamily names are accepted as oidea instead of -acea. The
last ones are more used in the Botanic nomenclature.
Having in mind all these circumstances the outline of the new classification of the
agglutinated forms elaborated to a different degree is given below.
Phylum Foraminifera d'Orbigny,1826
Class Astrorhizata Saidova, 1980
Subclass Astrorhizana Saidova, 1980
Shells unilocular, sometimes pseudocolonial or pseudochambered, of subsphaerical,
elongated, tubular, stellate, branching or other irregular form; wall 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 the tubes), or single in more advanced
forms (usually of definite form and position, may be slightly elevated on the neck,
may have a small lip); free-living or (very often) attached, Cm - Holocene.
Remarks. The taxa of the order and family level are separated on the basis of their test
form, presence or absence of the aperture, number of the apertures and their position,
mode of life (free, attached, pseudocolonial) and on the basis of the complication of
their shell wall and chamber lumen, as well as on the combination of these features.
Thus the forms having an aperture and lacking it, or singular and vise versa
pseudocolonial tests which were united previously in one family now are separated
into two different ones. The subclass includes many transitional forms whose features
permits to include them in Astrorhizana or with some remarks (proviso) into other
subclasses: thus the Vanhoeffenellidae, Causiinae and Amphifenestrellinae are
transitional from the subclass Lagynana, Ammovoluminidae to subclass
Ammodiscana, many of the Hippocrepinida to subclass Hormosinana. Some of the
microgranular forms included previously in the lower Fusulinacea are transferred to
Astrorhizana as the boundary between the agglutinated and microgranular wall is often
transitional. Thus in the Ammovertellininae were previously included forms both with
the agglutinated and microgranular wall (Gandiella, Pilammina) though in the
diagnosis of their higher taxon Ammodiscacea only the agglutinated character of the
wall was marked. Indeed, often it is not possible to separate the forms only on the
base of this feature. The most ancient typical Astrorhizana coexisted with their
microgranular analogues from Cambrian time, the known range of many of them was
significantly widened owing to the new data (Tyszka, 1997, Kaminski, this vol. and
many others). Komokiacea are not included here (their taxonomy is given in
Mikhalevich, 1995).
Order ASTRORHIZIDA Haeckel, 1894
Test free, single, tubulate or with tubes extending from central body; wall simple;
apertures at open ends of tubes, two or more.
Superfamily Astrorhizacea Brady, 1881
Test with the simple wall and chamber lumen not subdivided.
Family Vanhoeffenellidae Saidova, 1981
Test large, discoidal, fusiform or patelliform, with one, two or rarely more tubular
cylindrical outgrowths extending from it; wall agglutinated except the central capsule
with non agglutinated flexible tectinous area of organic wall at opposite sides of
central body. L. and M. Dev. , Holocene. Vanhoeffenella, Inhauris.*
* - Here and further the authors of the genera are given only in those published after
1988 year.
Family Astrorhizidae Brady, 1881
Test with more or less inflated central part and with tubular arms of differing length
extending from it, these arms sometimes branching, very often partly broken; number
of arms from 3 to 9 or 11; wall agglutinated throughout, usually stout. Ord – Hol.
Astrorhiza, Astrorhizoides, Cystingarhiza Bell, 1996, Cylindrammina Bell, 1996.
Remarks. The genus Clados Schroeder, Medioli et Scott, 1989 assigned by its authors
to Astrorhizinae belongs more probably to Komokiida.
Family Rhabdamminidae Brady, 1884
Test straight, long and unbranched or sparsely branched tube without inner septa, but
very often with constrictions, reflecting the growth periods, wall usually thick, from
smooth to roughly agglutinated, firm or friable; apertures at open ends of tubes.
Subfamily Bathysiphoninae Avnimelech, 1952
Test a free long tube of nearly equal diameter throughout; growth occurs at one end of
the test, the opposite end may be closed by debris and periodically discarded; wall
usually thick and firm. L. Cm, U. Tr., Hol. Bathysiphon, Bogdanowiczia, Nothia,
Platysolenites, Rhabdamminella, ? Silicotuba.
Remarks. The genus Astrorhizinula is removed from the subfamily and transferred to
Hippocrepinellidae as it has an oval test and several apertural openings at its restricted
Subfamily Rhabdammininae Brady, 1884
Test free, tubular, consisting of a single tube or a number of tubes (3, rarely 5)
extending from the center, tubes usually straight, of equal diameter throughout, more
rarely very slightly curved and tapering to the end; wall of different thickness, firmly
cemented, usually hard. Ord. Hol. Rhabdammina, Oculosiphon, Marsipella, Linea
Schroeder, Medioli et Scott, 1989.
Remarks. The genera Rhizammina and Testulorhiza whose representatives have
flexible and branching test I regard as belonging to Rhizammininae (order
Komokiida, incertae sedis).
Family Hippocrepinellidae Loeblich et Tappan, 1984, emend.
Test elongated, thickened to different degree in central part, externally sometimes with
transverse striation reflecting periodicity of growth; wall thick, finally grained, friable,
with large amounts of light cement; apertures as rather small subcircular openings at
constricted or semiclosed ends of shell, two or more. Ord. Hol. Hippocrepinella,
Amphitremoida, Crespinatella, Croneisella, Astrorhizinulla.
Remarks. The family differs from the closely related family Rhabdamminidae in the
shorter shell, tapered apertural end, and structure of the aperture. The composition of
the family is expanded by including the genera Amphitremoida, Crespinatella and
Croneisella transferred from Saccamminidae and the genus Astrorhizinulla from the
Order DENDROFRYIDA Haeckel, 1894, emend.
Test attached or free, single or pseudocolonial, of irregular and often of fanciful form,
strongly branched; wall simple or complicated by inner partitions or labyrinthic, with
different amounts of cement, more often friable, flexible; apertures open ends of
ramifications or apertures not fixed, looking as perforations of wall.
Remarks. The genus Arborammina Shires, Gooday et Jones, 1994 of the family
Arboramminidae Shires, Gooday et Jones, 1994 belongs more possibly to
Xenophiophoria as it has no apertural and chamber lumen and its body consists of
organic fibrils.
Superfamily Dendrophryoidea Haeckel, 1894
Test wall simple, test lumen not subdivided.
Family Dendrophryidae Haeckel, 1894
Test attached at base, with one or several tubular outgrowths extending upward from
the base; wall fragile, flexible when moist; apertures – open ends of tubular
ramifications or aperture absent.
Subfamily Dendrophryinae Haeckel, 1894
Test with the apertures at the open ends of the tubular ramifications. U. Cretaceous;
Pleistocene to Holocene. Dendrophrya, Psammatodendron, Saccodendron,
Globodendrina Plewes, Palmer et Haynes, 1993.
Remarks. The genera Saccodendron and Globodendrina are transferred from the
family Astrorhizidae as their representatives have strongly branching outgrowths.
Subfamily Spiculidendroninae subfam. nov.
Test with tubular branches without apertures. Holocene. Spiculidendron Ruetzler et
Richardson, 1996.
Family Notodendrodidae De Laca, Lipps et Hessler, 1980, emend.
Test free-living, buried in the mud, consisting of elongated tubular chamber branching
to different extent in its upper and lower part, the latter burried in ground, lower part
of the test may have circular turgidity from which multiple ramifies resembling
rhizoids are running; wall friable, flexible when moist; apertures – open ends of tubes.
Notodendrodes, Radicula.
Remarks. the genus Radicula is transferred from the family Astrorhizidae as its
representatives have a branched test buried in the ground. The family
Notodendrodidae was included earlier in Hippocrepinacea.
Family Dryorhizopsidae Loeblich et Tappan, 1984
Test entirely attached to substratum, branched tubes prostrate, sometimes extending
from central body; wall of fine sand grains; apertures circular, at ends of tubular
ramifies. U. Carboniferous – Holocene. Dryorhizopsis, Sagenina.
Superfamily Shizamminoidea Norvang, 1961
Test with inner cavity complicated by inner subdivisions or with labyrintic wall.
Family Shizamminidae Norvang, 1961
Test free, tubular or flattened, dichotomically branched or flabelliform and branched at
margins into numerous tubules, with inner cavity not subdivided; wall thick, hard,
with great amount of ferruginous cement, externally smooth and with transverse
wrinkles, labyrintic or cancellate inside, sometimes with small pseudopore openings at
surface; apertures – open ends of tubes. Holocene. Schizammina, Jullienella.
Family Halyphysematidae Loeblich et Tappan, 1984, nom. corrected.
Test attached, place of attachment spreading, discoidal, with raising from it conical
extension or erect tube branching at end, basal widening internally subdivided; wall
often with great amount of sponge spicules; aperture – open ends of tubes or at apex of
conus, often surrounded by clusters of spicules.Holocene. Halyphysema, Dendronina.
Family Diffusilinidae Loeblich et Tappan, 1961
Test attached, irregular in outline, high or flattened, sometimes with numerous tubular
outgrowths of wall or with septa forming small cellules of subquadrate or irregular
form; wall with moderate amount of cement; apertures at ends of peripheral tubes or
interstitial pseudopores at ends of sparse small pustules on test surface. Ordovic,
Holocene. Diffusulina, Kerionammina, Atelikamara.
Remarks. The genus Atelikamara is removed from the subfamily
Hemisphaerammininae as having an attached test of irregular form with incomplete
inner septa.
Order PSAMMOSPHAERIDA Haeckel, 1894
Test circular, oval, slightly elongated or hemispherical, free or attached, singular or
pseudocolonial; wall simple or complicated (labyrintic, cellular, with inner septa);
apertures numerous, single or absent.
Suborder STEGNAMMININA Mikhalevich, 2000
Test singular.
Superfamily Stegnamminoidea Moreman, 1930
Test with simple wall.
Family Stegnamminidae Moreman, 1930
Test free or attached, aperture absent.
Subfamily Amphifenestrellinae Mikhalevich, 1995
Test free; wall not fully agglutinated, with large naked area. Silurian, Holocene.
Amphifenestrella, Blastammina.
Remarks. The genus Amphifenestrella is removed from the subfamily
Vanhoeffenellinae on the basis of its test morphology (without tubular extensions open
at ends) and the absence of marked aperture. The genus Blastammina is excluded from
the subfamily Stegnammininae on the basis of its wall character with only sparce sand
Subfamily Stegnammininae Moreman,1930, emend.
Test free, circular, discoidal or slightly elongated, subcylindrical or nearly so,
sometimes with closed outgrowths; wall entirely agglutinated. Lower Cambrium
Upper Cretaceous. Stegnammina, Ceratammina, Raibosammina, Thekammina,
Anictosphaera, Bykovaeina, Pseudoastrorhiza, Storthosphaera, Thuramminoides,
Spiculosiphon, Gastroammina, Luekatiella Zhigulina, 1999.
Remarks. Although in the diagnosis of Raibosammina the possibility of branching of
its test is indicated, the specimens of the type species have an oval test without any
branches (Loeblich, Tappan, 1988, pl. 20, figs. 6,7). The genus is transferred from
the subfamily Thurammininae as lacking the aperture and some other genera from the
subfamily Psammosphaerinae as they have a single (not pseudocolonial) test.
Subfamily Haemisphaerammininae Loeblich et Tappan, 1961, emend.
Test attached, hemisphaerical; wall entirely agglutinated, connection with the
environment apparently through interstitial pseudopores. Ordovic – Holocene.
Hemisphaerammina, Fairliella(restored by Rauser-Chernousova et Reitlinger in
Vdovenko et al., 1993), Sorosphaerella.
Remarks. Only the genera with singular attached test without aperture are left in the
subfamily. The genus Sorosphaerella needs further investigations, it is possible that it
has pseudocolonial test and hence it should be placed in Telamminidae.
Family Colonamminidae Rauser-Chernousova et Reytlinger, 1993
Test free or attached, with single or multiple aperture.
Subfamily Causiinae Mikhalevich, 1995
Test planoconvex, circular in outline; wall organic, with small amount of agglutinated
material at peripheral margin of the test; aperture rounded, at centre of the concave
side of the test. Holocene. Causia.
Remarks. The subfamily differs from the other subfamilies of the family
Colonamminidae in the organic test wall.
Subfamily Colonammininae Rauser-Chernousova et Reytlinger, 1993
Test attached, hemispherical or elongated; wall entirely agglutinated, usually roughly
sandgrained; aperture single, at apex of chamber, sometimes on elevated neck. Silur –
Holocene. Colonammina, Jascotella, Nubeculariella.
Remarks. The genera Colonammina and Jascotella are excluded from the
Hemispaerammininae on the basis of presence of distinct aperture at the apex of the
shell, the genus Nubeculariella is is removed from the family Haliphysematidae ,
which is characterized by differing external structure of the test and by subdivision of
the inner lumen. Opposite to the authors of the family I left in it only representatives
with a single test and one aperture excluding Tholosina (with multiple aperture) and
Webbinelloidea (pseudocolonial), as well as Saccamminis whose representatives have
pseudochambered test with one commom aperture.
Subfamily Tholosininae Mikhalevich, 1995
Test attached, hemisperical, circular, oval or somewhat irregular in outline; wall
entirely agglutinated; aperture multiple (openings at base of the test attachement or at
ends of short outgrowths, extending from the place of attachement). Ordovic
Holocene. Tholosina, Iridia, Mesammina, ?Scyphocodon.
Remarks. All these genera are excluded from the subfamily Hemispaerammininae, in
which I have left only the genera with the distinct aperture, in distinction from the
present subfamily whose representatives have a multiple aperture.
Superfamily Crithioninoidea Goes, 1894, stat. n.
Test with labyrintic wall or chamber lumen subdivided by inner septa.
Family Crithionidae Goes, 1894
Test free or attached, with chamber lumen subdivided by inner septa.
Remarks. The family differs from the Oryctodermatidae in the presence of inner septa
subdividing the test lumen and in the absence of the labyrintic wall.
Subfamily Daitroninae Mikhalevich, 1995
Test free, discoidal or ovoidal; inner shell cavity subdivided by radial or vertical and
transverse semisepta; wall entirely agglutinated, finaly sandgrained; without distinct
aperture or aperture slit-like (Nephrosphaera). Upper Ordovic, Holocene. Daitrona,
Remarks. Subfamily differs from the subfamily Crithioninae in the test form and in the
free mode of life.
Subfamily Crithioninae Goes, 1894
Test attached, hemispherical; shell cavity subdivided to different extent by inner
asymmetrical septa; aperture at centre of upper part of test or at outstanding
outgrowth, or absent. Holocene. Crithionina, Pseudowebbinella, Verrucina.
Remarks. The former subfamily Crithioninae included representatives with free-living
as well as attached tests and tests with inner septa as well as tests with labyrintic wall.
Family Oryctodermatidae Saidova, 1981
= Subfam. Masonelinae Saidova, 1981 (as “Marsonellinae”), part.
Test free, circular or discoidal; wall thick, labyrintic, with system of regular or
irregular alveolar canaliculi radially extending from central cavity; canaliculi may be
opened at shell surface, there also may be gaps of irregular form in the thick friable
wall; no distinct aperture. Lower Carbon Holocene. Oryctoderma, Masonella,
Remarks. The genus Masonella is removed from Crithioninae.
Suborder PSAMMOSPHAERINA Haeckel, 1894
Test pseudocolonial or pseudomultichambered.
Family Psammosphaeridae Haeckel, 1894
Test pseudocolonial, without distinct aperture.
Subfamily Psammosphaerinae Haeckel, 1894
Test free, sometimes with interstitial pores between agglutinated particles. Middle
Cambrian – Holocene. Psammosphaera, Cellonina, Psammophax, Sorosphaera,
Remarks. The genera with singular test without aperture are transferred from the
former subfamily Psammosphaerinae to the subfamily Stegnammininae as well as the
newly described genus Luekatiella.
Subfamily Telammininae Loeblich et Tappan, 1985, emend.
Test attached, consisting of a series of circular, oval or hemisphaerical
pseudochambers, closely adjoining each other or loosely linked together by thin tubes
resembling stolons and forming rows or rather complex ramified net, as from one
pseudochamber may extend several stolons, linking it with the neighboring ones; wall
fine-sand grained. Silur Devon, Holocene. Telammina, Tumidotubus,
Metamorphina, ?Patellammina Bell, 1996, Webbinelloidea, Ropostrum Jonasson,
Schroeder-Adams, 1996.
Remarks. The subfamily is removed from the Hormosinacea, as its representatives
have no true apertures and represent pseudocolonial organism. The genus
Aggerostramen is excluded from the subfamily and placed here in family
Lacustrinellidae as its representatives are characterized as having an aperture
pseudocolonial organisms. The genus Patellammina is included conditionally as
according to its morphology its pseudocolonial tests are the sooner detached from the
surface and the basal side with the “aperture” is supposingly the side of attachment
(regarded as close to Webbinelloidea).
Family Polysaccamminidae Loeblich et Tappan, 1984
Test pseudomultichambered, with one or two apertures.
Subfamily Polysaccammininae Loeblich et Tappan, 1984
Test free, consisting of pseudochambers of circular or irregular form aligned in a
roughly uniserial, biserial or wholly irregular series; wall usually fine-grained with
considerable amount of cement, flexible, sometimes with a few sand grains attached
to surface; one aperture at end of last chamber, sometimes slightly elevated and
bordered by slight rim. Holocene. Polysaccammina, Goatopitigba.
Remarks. The uniserial tests resemble those of the representatives of the family
Hormosinidae, but differ from them in the absence of true chambers.
Subfamily Saccamminidinae Mikhalevich, 1995
Test attached, consisting of one row of hemisphaerical or oval pseudochambers; wall
of sand grains of medium size; one common aperture on the last pseudochamber,
sometimes at end of slightly produced neck. Upper Carbon. Saccamminis.
Remarks. The subfamily is separated from the Hemisphaerammininae on the basis of
the pseudochambered test having one commom aperture.
Subfamily Amphicervicinae Mikhalevich, 1995
Test attached, circular or oval in outline, consisting of 2-3 pseudochambers rapidly
enlarging in size and enveloped dorsally by the last hemisphaerical pseudochamber;
wall well cemented, of fine to medium size sand grains; two apertures at opposite
ends of last pseudochamber near place of attachment, somewhat produced. Lower
Silur. Amphicervicis.
Remarks. The subfamily is separated from Hemisphaerammininae at the base of its
pseudomultichambered embracing test and presence of apertures.
Family Lacustrinellidae Mikhalevich, 1995
Test pseudocolonial, attached, of circular, hemisphaerical or subangular form,
arranged in closely appressed groups or in a row, where they may be somewhat
distant from each other or linked by stolons or sponge spicules; wall sometimes with
a great amount of spicules or entirely of sand grains; every test with its own aperture
level flash with surface, at elevated papillae or at end of stolon. Middle Silur,
Holocene. Lacustrinella, Ammopemphix, Sorostomasphaera, Aggerostramen.
Remarks. The family is separated from the heterogenous subfamily
Hemisphaerammininae, as Lacustrinellidae are pseudoolonial and have an aperture.
The genus is transferred from the family Telamminidae earlier included in
Hormosinacea. The latter includes only representatives with true multichambered
Оrder Parathuramminida Mikhalevich, 1980
Test free or attached, single or pseudocolonial, of circular, elongated or irregular
form; wall microgranular, simple or complicated, perforated or imperforated; aperture
one, many or aperture absent. Silur – Perm.
Remarks. This order differ from the order Psammosphaerida including the suborder
Stegnaminina only in its wall structure. As I regard the step of the formation of the
microgranular wall following the agglutinated wall the order Parathuramminida is
transferred here from the previous Fusulinacea into Astrorhizata. Their revision is
not finished because of the pure preservation and unclear features of many of them.
For the detailed diagnoses see Mikhalevich, 1995.
Composition. Superfamily Parathuramminoidea E.V.Bykova, 1955:families
Archaesphaeridae Malakhova, 1956 with the subfamilies Archaesphaerinae (
Archaesphaera, Corbiella, Neoarchaesphaera, Quasiirregularina, Vicinisphaera,
Vulgarella), Insolentithecinae (Insolentitheca), Usloniinae (Uslonia, Bisphaera,
Cribrosphaeroides (= Parphia)), Eoammosphaeroidinae Mikhalevich, 1995
(Eoammosphaeroides, Arakaevella, Diplosphaerina) and Parathuramminidae
E.V.Bykova, 1955 with the subfamilies Parathurammininae (Parathurammina,
Cribrohemisphaeroides, Parathuramminites, Saltovskajina), Irregularininae
(Irregularina, Pachythurammina, Palachemonella), Chrysothurammininae (
Chrysothurammina, Salpingothurammina).
Superfamily Marginarioidea Loeblich et Tappan, 1986 (= Ivanovelloidea Chuvashov
et Yuferev, 1984) : families Marginaridae Loeblich et Tappan, 1986 with the
subfamilies Marginarinae (Marginara, Cordatella, Turkmeniella), Uralinellinae
(Uralinella, Sogdianina),
Tuberitinidae A.D.Miklukho-Maklay, 1958 with the subfamilies Tuberitininae
(Tuberitina, Bituberitina, Eotuberitina, Ivdelina, Mendipsia, Orientina, ?Tubeporella,
Tubeporina) and Hemithurammininae Mikhalevich, 1995 ( Hemithurammina,
Illigata), Auroriidae Loeblich et Tappan, 1986 (Auroria, Apertauroria),
Eovolutinidae Loeblich et Tappan, 1986 (Eovolutina, Biorbis), Rauserinidae Sabirov,
1987 (Rauserina, Baituganella, Petchorina).
Order HIPPOCREPINIDA Saidova, 1981
Test free or attached, pseudotwochambered, rarely pseudomultichambered,
commonly elongate; from the subspherical and distinct or scarcely marked
proloculus the second tubular pseudochamber extends upwards, it may be straight,
branched at end, curved or weakly coiled, or widened into a conus or funnel, it may
be undivided or subdivided by incomplete inner partitions, or may have external or
inner constrictions; wall agglutinated or microgranular; aperture terminal (wide or
tapering open end of tube), circular or oval; branched shells are accordingly with two
Superfamily Hippocrepinoidea Rhumbler, 1895
Test with undivided inner cavity.
Family Hippocrepinidae Rhumbler, 1895
Test free, closed and acute at aboral end; proloculus not clearly marked, much
narrower than the following widened part of test; wall agglutinated, often with
transverse constrictions, reflecting periodicity of growth; aperture a wide circular
open end of tube or a small opening at centre of apertural face.
Subfamily Hippocrepininae Rhumbler, 1895
= Hyperamminoididae Loeblich et Tappan, 1984 (part).
Test with closed and slightly inflated apertural end of tube, in transverse section
circular, oval or square with depressed sides; wall usually finely sandgrained, with
transverse constrictions externally; aperture – rather small circular or oval opening at
centre of apertural face. Upper Devon, Carbon, Perm, Holocene. Hippocrepina,
Hyperamminoides, Giraliarella, Pseudohyperammina.
Remarks. The genera whose representatives have a widely open aperture are
excluded from the subfamily. At the same time the genera Hyperamminoides,
Giraliarella and Pseudohyperammina are transferred from Hyperamminoididae on
the basis of similarity of their test and apertural structure with those of
In the heterogenous family Hyperamminoididae there were included ancient forms
(Devonian Cretaceous) of different structure. Inclusion of forms of different age
but of similar morphology in one subfamily appears to be justified as it is done in
Hyperammininae (Silurian – Holocene) and in many other subfamilies.
Subfamily Jaculellininae Mikhalevich, 1995 (nom. corrected)
= Hyperamminoididae Loeblich et Tappan, 1984 (part).
Test resembling stick, conus, or funnel, with widely open apertural end, circular in
transverse section; wall finely or coarsely agglutinated, with or without transverse
constrictions; aperture wide open end of test, sometimes with a small everted lip
(Sansabaina). Upper Ordovic, Silur, Devon, Carbon, Perm, Holocene. Jaculella,
Aciculella, Arenosiphon, Tasmanammina, Kechenotiske, Sansabaina.
Remarks. The subfamily is separated from the Hippocrepininae on the basis of
apertural structure. The genera Kechenotiske and Sansabaina are transferred from
the Hyperamminoididae. The position of the latter is not clear and needs further
Family Hyperamminidae Eimer et Fickert, 1899
Test free, with clearly marked bulbose proloculus; tubular pseudochamber extending
from it is of lesser diameter than proloculus; in the upper part it may slightly widen
or fork into two branches; wall agglutinated, microgranular or mixed; aperture an
open end of tube, branched tests with two apertures.
Subfamily Hyperammininae Eimer et Fickert, 1899
= Hyperamminoididae Loeblich et Tappan, 1984 (part).
Test with straight or widening unbranched tube extending from subspherical
widened proloculus; wall agglutinated; one aperture. Middle Silur Holocene.
Hyperammina, Areniconulus, Sacchararena.
Remarks. The genus Sacchararena is transferred from the Hyperamminoididae as it
has distinct proloculus.
Subfamily Earlandiinae Cummings, 1955
Test as in Hyperammininae, but differs in the microgranular wall. Silur Perm.
Earlandia, Aeolisaccus, Gigasbia.
Subfamily Saccorhizinae Eimer et Fickert, 1899
Test with straight or slightly curved tube extending from widened proloculus and
branching at end; wall agglutinated, often with sponge spicules; two apertures at end
tubes. Upper Silur – Holocene. Saccorhiza, Saccarena.
Remarks. The subfamily is restored as a separate taxon from the synonymy of
Family Ammovoluminidae Chernych, 1967
Test free, with distinct proloculus, second tubular pseudochamber sometimes
widened or rugged and of irregular form, it is curved and coiled in a loose spiral;
wall agglutinated; aperture at end of tube, circular, oval or irregular. Middle Silur,
Devon, Holocene. Ammovolumina, Hyperbathoides, Psammonyx, Serpenulina.
Remarks. The family is transferred from the order Ammodiiscida. It is intermidiate
between the two orders, but the irregular test form, short and often only slightly
curved tube more sharply differentiate it from the regular ammodiscid tests with
numerous volutions.
Superfamily Botellinoidea Chapman et Parr, 1936
= Caligelloidea Reytlinger, 1959
Test pseudomultichambered, with pseudochambers formed by incomplite septa or
wall constrictions.
Family Botellinidae Chapman et Parr, 1936
Test free, tubular, with slightly marked or indistinguishable proloculus, with obtuse
base and small inner projections of wall into chamber lumen; wall agglutinated,
sometimes with sponge spicules; aperture open end of tube, sometimes slightly
tapering. Holocene. Botellina, Protobotellina.
Remarks. The family is restored from the synonymy of Hippocrepinidae, its
representatives differ from that ones of the Hippocrepinidae and Hyperamminidae
(Loeblich, Tappan, 1988) in the peculiarities of their external and internal structure.
Family Hormosinellidae Rauser et Reytlinger, 1986
Test free, elongated, rocary-like, with elongated or wide pseudochambers formed by
wall constrictions; wall agglutinated; aperture terminal, circular. Middle Devon
Holocene. Hormosinella ( according to M. Kaminski(this book) Rothina and
Caudammina are the synonyms of Hormosinella, representing its broken chamber),
Remarks. The family is transferred from Hormosinida as having pseudochambers
instead of a true chambers.
Family Paratikhinellidae Loeblich et Tappan, 1984
Test free or attached, with markedly isolated proloculus and with second tubular or
widened pseudochamber, often of irregular form, sometimes bifurcated; inner cavity
subdivided by incomplite septa or constrictions of wall arranged irregulary, rarely
evenly; wall microgranular entirely or including agglutinated particles; aperture an
open end of tube, branching shells with two apertures.
Subfamily Caligellinae Reytlinger, 1959
Test attached, tubular part unbranched; septa irregular; wall microgranular; one
aperture. Upper Silur Lower Carbon. Caligella, Glubokoevella, Paracaligella,
Subfamily Saccorhininae Mikhalevich, 1995
Test attached, upper part of tubular pseudochamber bifurcated; septa irregular; wall
microgranular; two apertures. Devon. Saccorhina, Evlania.
Remarks. The subfamily differs from Saccorhizinae in the microgranular wall and
subdivided shell cavity. The genus Saccorhina was previously included in
Moravamminidae, Evlania in Caligellidae.
Subfamily Paratikhinellinae Loeblich et Tappan, 1984
Test free, tubular, unbranched, septa or wall constrictions regular; wall
microgranular; one aperture: open or tapering end of the tube. Middle Devon
Carbon. Paratikhinella, Saccamminopsis.
Class SPIRILLINATA Maslakova, 1990
Subclass AMMODISCANA Mikhalevich, 1980
Test pseudotwochambered, with a long tubular second chamber coiled around the
proloculus, in higher taxa may be with the tendency of transition to
multichamberedness usually with the preservation of the tubular character of the
chambers or pseudochambers, initial undivided part often preserved, types of coiling -
glomerate, planospiral, trochospiral, the predominant type of shells - flat,
subsphaerical, rarely patellinoid, sometimes irregular; wall agglutinated, usually with
the ferrugenous cement or microgranular, rarely tectinous and partly agglutinated;
aperture at the end of the tubular chamber, simple, without any inner structure;mostly
free living forms, more rarely attached. Cambrian - Holocene.
Order AMMODISCIDA Mikhalevich, 1980
Test with the second tubular chamber coiled planospirally or sometimes weakly
trochospirally or even in high trochoid spire in the free living forms, in attached forms
– uncoiled or irregularly coiled.
Ammodiscidae (Ammodiscus, Ammodiscoides, Arenoturrispirillina, Bifurcammina,
Hemidiscus, Rectoammodiscus Reitlinger, 1993, Spirillinoides, Spirosolenites, without
Agatamminoides), Pseudoammodiscidae from Fusulinacea(Pseudoammodiscus, ?
Brunsia, ?Brunsiella, ?Pseudoglomospira, Quasilituotuba, Turrispiroides, without
Warnantella), Tolypamminidae (Tolypammina, Ammodiscella, Ammodiscellites Resig
et Glenn, 1997, Ammolagena, Ammovertella, Hemidiscella, Saturnella Hedinger,
1993, Serpulopsis), Turritellellidae fam. reinstated and given stat. nov. (Turritellella,
Order AMMOVERTELLINIDA Mikhalevich, 1999
Test with the second tubular chamber coiled glomerately at least at the early stage
(Ammovertellinidae) in distinction of the Ammodiscids or with the sharp change of
the regular types of coiling (Usbekistaniidae).
Usbekistaniidae (Usbekistania, Flagrospira, without Turritellella and Repmanina),
Ammovertellinidae with Ammovertellininae - having the erected part
(Ammovertellina, Rectoglomospira), Glomospirellinae without erected part, subfamily
reinstated (Glomospirella, Agatamminoides from Ammodiscinae, Annectina,
Arenomeandrospira Jones et Wonders, 2000, Gandinella, Glomospira, Vostokovella),
and Pilammininae Urosevic, 1992 (Pilammina, Pilamminella, Rectopilammina
Urosevic, 1992) , Pseudolituotubidae from Fusulinacea (Pseudolituotuba,
Warnantella from Pseudoammodiscidae).
Order PLAGIORAPHIDA ord. nov.
Test with the chambers in high spire, following the proloculus, each chamber forming
an entire whorl or more with strongly oblique sutures between the chambers; wall
agglutinated; aperture circular, large, at the open end of the enrolled chamber. Middle
and Upper Trias. Plagioraphidae Mikhalevich, 1995 (Plagioraphe).
Class MILIOLATA Saidova, 1981
Subclass SCHLUMBERGERINANA Mikhalevich, 1992
Test pseudotwochambered, pseudomultichambered, multichambered, the
predominant type of coiling – irregular- and regular glomerate, planospiral,
trochospiral or biserial at the last stage rarely, as an exclusion; shell of the higher
representatives could be cyclical and fusulinoid in form; the piece of the tubular
chamber may be preserved around the proloculus; chambers predominantly tubular,
usually two per volution, in some forms more wide like in Peneroplis-similar forms
or sometimes subsphaerical, may be strongly embracing, in higher representatives
chamber number greatly increases , chamber lumen could be subdivided by the inner
sceletal elements, advanced forms may have integrative systems (stolons, tunnels);
wall agglutinated, microgranular or agglutinated and partly with the secretious
needles; aperture terminal, simple in the most primitive forms, in more complex
ones - with the outer (flap) or inner tooth of a special structure, sometimes aperture
secondarily multiple, additional apertures non developed; bottom forms, mostly free
living. Carbon - Holocene.
Remarks. In some ancient representatives wall may be microgranular as a result of
the diagenesis.
? Order TOURNAYELLIDA Hohenegger et Piller, 1973
Test with the second tubular chamber with the glomerate, planospiral or combined
coiling, often with the irregular alternating of the plan of coiling, with the following
subdivision of the tubular chamber lumen into rather irregular pseudochambers,
sometimes forming in the last volutions true chambers, usually of somewhat irregular
form and dimensions, bearing some tubular features, rarely biserial at the last part,
which may be erected; wall agglutinated or microgranular; aperture – an open end of
the tubular chamber. Middle Devon Holocene. Families ?Moravamminidae
(Moravammina, Kettnerammina, without Saccorhina)), Tournayellidae (the
composition according to Rauser-Chernousova et al., 1996, pp. 7 27 (= Ord.
Tournayellida of that book), including 4 families, 8 subfamilies, 46 genera),
Lituotubidae with Lituotubinae having later uncoiled part (Lituotuba) and
Trochamminoidinae Haynes et Nwabufo-Ene, 1998 (emend.) - without uncoiled part
(Paratrochamminoides, ?Conglophragmium, Trochamminoides, Sokotina Haynes et
Nwabufo-Ene, 1998).
Order SCHLUMBERGERINIDA Mikhalevich, 1980
Test multichambered, of one of the miliolid type of construction; wall agglutinated or combined with calcareous needles,
aperture often with a tooth. Cretaceous – Holocene. Families: Rzehakinidae with Rzehakininae ( Rzehakina, Ammoflintina,
Psamminopelta, Spirolocammina, Spirosigmoilinella, without Rothina(see Bubik,1997 and Kaminski, this book – the latter
considers it to be a broken part of Hormosinella shell (personal communication)) and also without Silicosigmoilina and genera
transferred into Miliammininae) and Miliammininae - subfam. reinstated (Miliammina, Birsteiniola, Silicomassilina, ?
Trilocularena), Schlumbergerinidae (Schlumbergerina, ?Tekkeina Farinacci et Yeniay, 1994 (if valid)) , Siphonapertidae
(Siphonaperta, Agglutinella, Ammomassilina, Dentostomina, Pseudoflintina), Sigmoilopsidae (Sigmoilopsis, Silicosigmoilina
from Rzehakininae ) , Spiroglutinidae (Spiroglutina)(the genera of the last 4 families from the former Miliolacea).
Order HAPLOPHRAGMIIDA Mikhalevich, 1992
Test multichambered, streptospirally coiled, at least initially (or in one of the
generations), evolute or involute, later may be uncoiled, aperture terminal or
elevated , single or multiple.
Superfamily Recurvoioidea Alekseychik-Mitskevich,1973
Test with a simple wall. Families: Cystamminidae fam. nov. test with a few
volutions and few chambers , subsphaerical or ovoid, strongly inflated(Cystammina,
Praecystammina, Saccamminoides), Recurvoididae (Recurvoides, Budashevella, ,
Thalmanammina (according Neagu et Platon, 1994 the aperture of this genus is areal
(terminal)), Ammobaculinidae with Ammobaculininae (Ammobaculinus,
Bulbobaculites) and Telatynellinae (Telatynella Gawor-Biedowa, 1987), Acupeinidae
(Acupeina, Navarella – transferred from Ammobaculinidae as having multiple
aperture), Plectorecurvoididae (Plectorecurvoides, Pokornyammina Neagu et Platon,
1994) (this family stands apart from the rest Recurvoioidea having biserially arranged
enrolled chambers and possibly deserves to be elevated to the order rank. But the
scrupulose work of Neagu et Platon, 1994 had shown the unstable mode of coiling in
Plectorecurvoides(“both planospiral and streptospiral” ) and unstable biseriality in
Pokornyammina (“biserial to larger uniserial”). Bubik (1995) also had showed the
variable irregular coiling of some species of Plectorecurvoides (including the type
species) as well as the terminal character of its aperture (judjing from the inner tubes
connecting the chambers seen in sections) and of the aperture of Recurvoides (Bubik,
2000). More over, some specimens of Thalmannammina (fam. Recurvoididae) look
occasionally as biserial ones (see Neagu et Platon, 1994). All of these
genera have terminal aperture (though sometimes it looks externally as basal
(interiomarginal) and the further investigation of their variability is necessary for the
taxonomic conclusions. They looks closely related from the morphological point of
Superfamily Haplophragmioidea Eimer et Fichert, 1899
Test with the alveolar wall. Middle and Upper Jurrassic. Families:
Haplophragmiidae with Haplophragmiinae(Haplophragmium) and
Streptocyclammininae subfam. nov. – test flattened, early stage streptospiral, at least
in one of the generations, later planospiral, with numerous low, broad, strongly
recurved chambers forming peneropline appearance, wall of imperforate outer layer
and fine subepidermal network, septa massive, with numerous apertures, at the later
stage a few irregular pillars may present (Streptocyclammina), aperture multiple,
openings covering entire apertural face (Streptocyclammina transferred from
Spirocyclinidae (former Loftusiacea) as having initial streptospiral coiling,
Alveosepta transferred from Hottingeritidae ), Mesoendothyridae
( Mesoendothyra, Audienusina), Hottingeritidae (Hottingerita) (two of the last
families transferred from the former Loftusiacea ).
Superfamily Labyrinthidomatidoidea Loeblich et Tappan, 1988
Test with the chamber lumen subdivided by the inner septa. Upper Cretaceous.
Family Labyrinthidomatidae(Labyrinthidoma, Bulbophragmium).
Order SPHAERAMMINIDA Mikhalevich et Kaminski, ord. nov.
Test multichambered, planispirally coiled, involute, with a few strongly overlapping
subsphaerical chambers; wall agglutinated, aperture terminal, areal, with tooth or
without it. Paleocene Holocene. Family Sphaeramminidae (subfamily
Sphaerammininae with a tooth (Sphaerammina, Ammosphaerulina, Canepaia) and
Praesphaerammininae Kaminski, subfam. nov. - without tooth (Praesphaerammina
Kaminski et Filipescu, 2000)
Order LITUOLIDA Lankester, 1885
Test multichambered, planospiral, at least initially, later may be uncoiled,
(trochospiral as an exception Nezzazatidae), chambers broad or subtriangular in
outline, more often with straight sutures, aperture terminal, without tooth.
Remarks. Forms with the clearly basal aperture, at least at the initial stage are
transferred from Lituolida to the new order Nautiloculinida (see below). Lituolids
included here are to strong degree isomorphic to some Peneroplids, Soritids and
Alveolinids. In the composition of the order Lituolida (Loeblich, Tappan, 1994 and
in some following classifications) the genera both having agglutinated and
microgranular wall and with or without canaliculi or some kind of the pore openings
were included in spite of the diagnosis of the order defining it as agglutinated and
lacking pores. It is impossible indeed to put the sharp boundary in many cases. More
often it is easier to distinguish the taxa if we base ourselves on the shell form and
position and character of the aperture.
Superfamily Lituolinoidea de Blainville, 1827
Test free, with the simple wall. Families: Lituolidae – with Ammobaculitinae (single
aperture)(Ammobaculites, Ammobacularia, Sculptobaculites, Simobaculites,
Ostiobaculites Bronniman, Whittaker, Zaninetti, 1992, ?Apostrophoides McNeil,
1997) and Lituolinae (as Ammobaculitinae, but multiple aperture)(Lituola,
Atactolituola, Bulbobuccicrenata, ?Kolchidina,Carasuella Neagu, 2000),
Buzasinidae fam. nov. – test planospiral entirely, involute, aperture areal, boarded by
a narrow lip, looking as transverse slit or circular (Buzasina, Ammosiphonia), ?
Mayncinidae (in some genera it is necessary to study the aperture at the initial stage)
(Mayncina, Biconcava, Comaliamma, Daxia,?Flabellocyclolina, Freixialina,
Gendrotella, ?Phenacophragma, Stomatostoecha, Hinogammina Neagu, 2000,
Paleomayncina Sepfontaine in Kaminski, 2000, (without Deuterospira - basal
aperture at the initial stage), Placopsilinidae (attached)(Placopsilininae -
Placopsilina, Acruliammina, ?Ammocibicides, ?Ammocibicoides, Lapillincola
Wilson, 1986, ?Subbdelloidina, ? Flatschkofella Rettori, Senowbari-Daryan, Zuhike,
1996); Adhaerentiinae ( Adhaerentia).
Superfamily Barkerinoidea Smout, 1956
Test with the chambers, internally subdivided by the transverse partitions
(Barkerinidae) or basally digitate plate of varied form. Shell wall microgranular or
microgranular-agglutinated. Lower Cretaceous. Family Barkerinidae (Barkerina),
??Nezzazatidae (seems to be the heterogenous group with fully secreted and perhaps
diagenetically changed shell wall) with the Biplanatinae subfam.nov.
planospirally enrolled (Biplanata, Merlingia, Demirina Ozcan, 1994) and
trochospirally enrolled Nezzazatinae (Nezzazata, Trochospira, Pyrenina, without
Nezzazatinella,which according Neagu, 1979 belongs to Pfenderinid line) and
Coixitinae (Coixites, ?Antalyna, Rabanitina) .
Superfamily Biokovinoidea Gusczic, 1977
Test free, entirely coiled or with later uncoiled part which may be straight or
peneropliform and reniform, with thickened complicated wall(may be with canaliculi
and keriotheca- and phrenotheca –like layers) and endoskeletal elements (pillars,
partitions, chomata-like structures), may be alveolar (Buccicrenatidae,
Choffatellidae, Spirocyclinidae) ; aperture terminal, single or multiple.
Families Charentiidae (Charentia, Ismailia, ?Karaisella, Melathrokerion,
Praekaraisella, Praepeneroplis, Bosniella from Biokovinidae), Lituoliporidae
(Lituolipora), Biokovinidae (Biokovina, without Chablasia), Labyrinthinidae
Sepfontaine, 1988 with Labyrinthininae (Labyrinthina), Planiseptinae Sepfontaine,
1988, nom. nudum (Planisepta Sepfontaine in Kaminski, 2000) and Levantinellinae
Fourcade, Mouty et Teherani, 1997 ( Levantinella Fourcade, Mouty et Teherani,
1997) , Buccicrenatidae (Buccicrenata, Evertycyclammina), Choffatellidae with
Chofatellinae ( Choffatella, Feurtillia, ?Paracyclammina, Pseudocyclammina,
Torinosuella, ?Redmondellina Banner, Whittaker, 1991, Ijdranella Bassoullet,
Boutakiout et Echarfaoui, 1999, Palaeocyclammina Bassoullet, Boutakiout et
Echarfaoui, 1999, Popovia, Sabellovoluta (two last genera from the
Alveolophragmiinae as having areal (not basal) aperture at the initial stage), without
Amijiella, Bramkampella and Rectocyclammina) and Pseudochoffatellinae
(Pseudochoffatella, Alzonella, Broeckinella, Dhrumella, Torremiroella),
Spirocyclinidae (Spirocyclina, Anchispirocyclina, Martiguesia, Pseudospirocyclina,
Reissella, Sornayina (without Quataria, Saudia, Thomasella Sirel, 1998,
Timidonella, Vania having cyclical or annular chambers, without Streptocyclammina
with the initial streptospiral coiling and without conical and initially trochospirally
coiled forms (Haurania, Spiraloconulus)).
Remarks. Family Buccicrenatidae is transferred from Loftusiacea, it differs from
Charentiidae in its alveolar wall, the position of the inner septa at the bottom of the
chambers is similar in both families as well as in Biokovinidae. I regard this line as
the continuation of the phyletic line of the planospiral Tournayellids and Endothyrids
(see Loeblich, Tappan, 1988, pl. 79 (8) and pl. 233(11); pl. 79 ( 11) and pl. 243
(16) ). Families Choffatellidae and Spirocyclinidae are also transferred from the
former Loftusiacea.
? Superfamily Coscinophragmatoidea Thalmann, 1951
Test attached, with variably coiled earlier part and irregular spreading, sometimes
branching later stage, wall complex (canaliculate, alveolar); aperture terminal, may
be cribrate. Family Haddoniidae (Haddonia, Stylolina – reinstated by Popescu et al.,
1998 from the synonimy of Lituola), Coscinophragmatidae (Coscinophragma,
Alpinophragmium, Bdelloidina, Goellipora Senowbari-Daryan, Zankl, 2000, ?
Order CYCLOLINIDA Mikhalevich, 1992
Test multichambered, initially planospiral or coiling oscillating, than Peneropliform
and later have cyclical or annular chambers; aperture multiple a rows of circular
openings at the apertural face.
Superfamily Cyclolinoidea Loeblich et Tappan, 1964
Test with the simple wall and undivided cavity. Lower and Upper Cretaceous.
Family Cyclolinidae (Cyclolina, Ammocycloloculina).
Superfamily Cyclopsinelloidea Loeblich et Tappan, 1984
Test with the chamber lumen subdivided by the pillars or septa, or both, advanced
forms with the stolons. Families Cyclopsinellidae (Cyclopsinella,
Mangashtia),Illerdorbidae (Ilerdorbis, Dohaia, Eclusia), Orbitopsellidae
(Orbitopsella, Cyclorbitopsella, Orbitammina).
Superfamily Vaniinoidea superfam. nov.
Test with subepidermal network of beams and rafters, and often with pillars.
Family Vaniidae fam. nov. test planospiral, with the cyclical or annular chambers
at the adult stage, wall with the subepidermal network of beams and rafters forming
alveolar wall and in some cases may form chamberlets, pillars often formed at the
later stages, aperture multiple, rows of openings along the peripheral margin; with
subfamilies Vaniinae subfam. nov.- without pillars( Vania, Quataria from
Spirocyclinidae, Balkhania, ? Montsechiana from Pseudochoffatellinae) and
Thomasellinae subfam. nov. - with pillars (Thomasella Sirel, 1998, Timidonella ,
Saudia – all the three from Spirocyclinidae) .
Remarks. The new family is separated from the former Loftusiacea (from
Spirocyclinidae and Cyclamminidae (Pseudochoffatellinae)) as having annular and
cyclical chambers at the adult stage. The superfamily differs from the Cyclolinoidea
in its complex wall and endoskeleton, from Cyclopsinelloidea in the character of the
wall (having subepidermal network).
Order LOFTUSIIDA ord. nov.
Test large, planospiral, axis of coiling elongate resulting in alveolinid and fusulinid
(aspect?)appearance, a short tubular chamber around proloculus may be preserved;
chamber interior with exoskeletal beams and rafters and endoskeletal pillars, canal-
like openings may be present; wall agglutinated and microgranular, with imperforate
epidermis and inner alveolar layer; aperture terminal at the initial stage, later
multiple – rows of rounded openings little above the base of apertural face. Low and
Upper Cretaceous.
Remarks. The affinity of these forms to Miliolata is confirmed not only by the
similarity of their outer (like Alveolinids) and inner shell structure (Reticulinella
even has preseptal canal) but also by the presence of the tubular chamber around
proloculus (Praereticulinella).
Family Loftusiidae (Loftusia, Praereticulinella, Reticulinella).
Class NODOSARIATA Mikhalevich, 1992
Subclass HORMOSINANA Mikhalevich, 1992
Shells unilocular to multichambered , predominant shell types - uniserial, high-
trochospiral (usually in a special polymorphinoid spire), planospiral (nearly
exceptionally involute), typical trochospiral type poorely developed; chambers often
subcircular, triangular, V-shaped, palmate, chevron-like and arcuate in uniserial part,
outer additional skeletal plates absent; wall agglutinated or microgranular, may be
complicated from the interior; aperture terminal even in the coiled forms, singular
(often displaced to the peripheral margin of the test) or multiple, rounded, oval, slit-
like, asymmetrically-fissured, may be with entosolenian tube, sometimes radial with
inner longitudinal ribs (Nodosinum, Haymanella) ( asymmetrically fissured and
Nodosariid kind of radial aperture, as well as entosolenian tube are unique and are
found only in this group), additional apertures absent; mostly free-living, bottom
forms, Ordovician - Holocene.
Remarks. Not only shell form (Lagena-like, Nodosaria-like, Marginulina -, Lingulina
-, Astacolus -, Kyphopixa-, Polymorphina-like etc. ) are analogous to some higher
Nodosariata, but also the unique structure of the aperture coincides in agglutinated
Hormosinana and secreted Nodosariana forms. I was expecting the existence of the
Lenticulina-like form with the aperture at the peripheral angle among the
agglutinated forms and luckily such form (Agardhella) was found and described by
Nagy et Basov (1998). By the way this example shows once more the incompleteness
of the geological record up to date. The only known exclusion of the position of the
aperture in cl. Nodosariata ( in their higher s/cl. Nodosariana) was the aperture of
Robuloides which was unclear and described as basal (or areal). But in pl. 437, fig. 4
of Loeblich &Tappan, 1988 it is definitely terminal and Prof. Neagu (personal
communication) also observed in his material terminal position of the Robuloides
Order SACCAMMINIDA Lankester, 1885
Test unilocular, of more or less regular form – sphaerical, subsphaerical, oval, slightly
elongated, with multiple or more often single aperture which may be at the neck or
with entosolenian tube, free living forms. U. Vendian - Holocene.
Remarks. The order transferred here from the subclass Astrorhizana (see Mikhalevich
et al., 2000) and is regarded as the preceding group for Lagenids.
Families Saccamminidae with Saccammininae ( Saccammina, Brachysiphon,
Hyperamminita, Lagenammina, Pilulinella, Placentammina, Psammophaga,
Sacculinella, Stomasphaera, Technitella, Marsupulinoides Bronnimann, 1988,
Saccamminella Bronnimann, Whittaker et Zaninetti, 1992, Titanotheka Gauche et
Sprechmann, 1999 if really belongs to Foraminifera ), Ovammininae subfam. nov.
aperture with the entosolenian tube (Ovammina), Pilulininae (Pilulina),
Thurammininae (Thurammina, Astrammina, Orbulinelloides, Ordovicina,
Order HORMOSINIDA Mikhalevich, 1980
Test multichambered, represents rectilinear or somewhat curved row of chambers
enlarging in size; wall often ferrugenous, may be pierced by canaliculi or alveolar;
aperture terminal, rounded, slitlike or cribrate, in one genus (Nodosinum) with inner
radial ribs, in forms with the ramifying last chamber two apertures. Ordovic
Remarks. The uniserial Dusenburyinidae are transferred into subclass Textulariana
(class Rotaliata) as having inner tooth similar with that one of Stilostomellidae.
Superfamily Hormosinoidea Haeckel, 1894
Test with the simple wall.
Families Hormosinidae with Hormosininae (Hormosina, Ginesina, Loeblichopsis,
Pseudonodosinella, Reophanus as having true chambers, Siliconodosaria) and
Nodosininae (Nodosinum, without Cribratinoides lacking inner ribs),
Aschemocellidae (Aschemocella, Kalamopsis, Calos Schroeder, Medioli et Scott,
1989 ), Reophacidae with Reophacinae (Reophax, Hormosinoides, Nodulina, without
Adelungia ), Polychasmininae Kaminski, 200.. in this book (Polychasmina), and
Bireophacinae subfam. nov. – test branching dichotomously with rounded apertures
at the end of each branch (Bireophax), Oxinoxisidae ( Oxinoxis) , Glaucoamminidae
(transferred from the former Textulariacea) (Glaucoammina), Cuneatidae (Cuneata,
Acostata Bronnimamm, Whittaker et Valleri, 1992, Cribratinoides,
Psammolingulina, Sulcophax, Warrentia), Kunklerinidae (Kunklerina, Leptohalysis
from Reophacinae chambers slightly differentiated in dimensions, numerous,
Scherochorella, Subreophax), Earlandinitidae (wall microgranular with agglutinated
particles)(Earlandinita, Darjella, Lugtonia).
Superfamily Cribratinoidea Loeblich et Tappan, 1984
Test with the complicated wall structure – with canaliculi or alveolar or labyrintic.
Families: Cribratinidae (Cribratina, Pseudotriplasia (from Pavonitininae, having
uniserial test), ? Haplostiche (inner character unknown but crenulate chambers
permit to suppose the complex wall), Thomasinellidae (Thomasinella, ?
Protoschista, Axicolumella Hercogova, 1988).
Test multichambered, often compressed, may be triangular or quadrangular in
transverse section, planospiral, at least initially, later usually elongated, uncoiled
into uniserial or rarely biserial part (Orectostominidae), chambers usually narrow,
often arched, sutures oblique, more often curved; wall agglutinated; aperture
terminal, tending to displacement to the peripheral end, rounded or more often ovate
or slit-like, may be cribrate. Jurassic – Holocene.
Remarks. The forms transferred from Lituolida differ from planospiral uncoiled
lituolid forms in their test and chamber form (usually flat, with elongated oblique or
arquate chambers, sometimes with limbate sutures) and in the form and position of
the aperture (often elongated and displaced to the peripheral margin), analogous in
their features and appearance to some higher secreted Nodosriata (some
Superfamily Ammomarginulinoidea Podobina, 1978
Test with the simple wall.
Families Ammomarginulinidae with Ammomarginulininae (Ammomarginulina,
Ammotium, ?Adelungia (transferred from Reophacidae as having the appearance as
that one arising from coiled form ), ?Discamminoides, Eratidus, ?Kutsevella,
Lamina), ?Haymanellinae subfam. nov. differs from other subfamilies of the
family in the pure development of the coiled part, “goulot”(chevron-like) chambers
with long bottled neck and radial aperture with longitudinal ribs projecting into
apertural opening (Haymanella Sirel, 1999), Flabellammininae (Flabellammina,
Ammopalmula, Pterammina, Triplasia) and ?Ammoastutinae (Ammoastuta, ?
Praeammoastuta), Bykoviellidae (fam . reinstaited by Kaminski, 200.. and here
given the status nov.) (differs from Vialovinae in which it was placed in synonimy in
its terminal instead of basal (interiomarginal) aperture) with Bykovaellinae
(Bykoviella, Sepetibaella from Jadammininae, having characteristic aperture near the
peripheral angle, ?Polskyammina (if not having basal aperture at the initial stage),
Ponceammininae Seiglie, 1991 (Ponceammina Seiglie, 1991) and Agardhellinae
subfam. nov. – test planospirally coiled entirely, without rectilinear part, aperture at
the peripheral angle of the test (Agardhella Nagy et Basov, 1998 ), Orectostominidae
fam.n. test with initial part coiled asymmetrically planospirally or in very low
trochospire similar to Bykoviella or Ponceammina but the rectilinear part is biserial,
aperture terminal throughout, a transverse slit with protruding lips, at the center of
the apertural face; the new family differs from the Bykoviellidae in its biserial
erected part, from Spiroplectamminidae in the terminal instead of basal position of
the aperture (Orectostomina transferred from the Spiroplectamminidae),
Lacroixinidae fam.n. – test biserial or loosely biserial with cuneate chambers at later
stage, tending to become uniserial, in microsphaeric form the residual chambers of
the unclear initial coiling may present (Lacroixina), wall agglutinated,
noncanaliculate, aperture terminal from the very beginning, with Lacroixininae
subfam. n. - the biserial part constitute the significant or the most part of the test,
aperture usually on a short neck (Lacroixina, Parvigenerina (both genera from the
Pseudobolivinidae which differ from the new family in having basal interiomarginal
aperture, Haghimashella Neagu et Neagu, 1995, Bicazammina Neagu et Neagu,
1995, Rashnovammina Neagu et Neagu, 1995, ?Plectinella from Textulariopsidae)
and Monotaleinae Bronnimann, Whittaker et Zaninetti, 1992 – iserial part short, most
of the test is uniserial, aperture terminal, circular, large, devoid of everted border
(Monotalea Bronnimann, Whittaker et Zaninetti, 1992, Trunculocavus Bronnimann
et Whittaker, 1993) .
Superfamily Flabellamminopsidoidea superfam. nov.
Test with the complicated alveolar wall
Family Flabellamminopsidae fam. nov. - test planospirally enrolled in the earlier
stage, then uncoiled, uncoiled part with arched septa, circular, flattened, triangular or
quadrangular in section; aperture terminal, at the center of uncoiled part in adult,
singular or multiple, with Flabellamminopsinae (rectilinear uncoiled part variously
flattened, triangulate or quadrate in section) (Flabellamminopsis) and ?Amijiellinae
Sepfontaine, 1988 ( (transferred from Hauraniidae Sepfontaine, 1988, diagnosis
changed) (Amijiella, Bramkampella, Rectocyclammina, ?Coprolithina, (from
Pernerinidae), ?Kastamonina Sirel, 1993 if having coiled part (most probably
planospiral) as is supposed in its diagnosis, otherwise uniserial and should be placed
into Cribratinoidea), Spiropsammiidae from Pavonitacea) (Spiropsammia).
Order NOURIIDA Mikhalevich, 1980
Test high trochospiral with the chambers in polymorphinoid arrangement or initial
part may be triserial or biserial later changing into uniserial ; aperture
terminal,simple, rounded or slitlike.
Remarks. The aperture of the initial chambers needs to be specified in many genera
of the order.
Superfamily Nouriidoidea Chapman et Parr, 1936
Test with the simple wall and chamber lumen.
Families Nouriidae with the longitudinally elongated chambers (transferred from the
former Textulariacea) (Nouria, Adbullaevia), Reophacellidae Mikhalevich &
Kaminski fam. nov. test high trochospire of polymorphinoid type, at least initially,
with reduced number of chambers in the terminal part of the shell, chambers
subsphaerical, inflated, with long narrow apertural tubules not visible externally,
running in the multichambered part along the side of the adjacent chamber close or
pressed to it, tubules visible only in sections, with Reophacellinae Mikhalevich &
Kaminski subfam. nov. tests with the number of chambers reducing from the
multiserial initial part to three or two in the latest part( Reophacella,
Falsogaudryinella, Uvigerinammina, ?Paragaudryina, ?Talimuella, ?
Paleogaudryina, ?Karrerulina, (initial aperture in 4 last genera supposingly terminal,
6 first genera from Verneuilinidae, the last one from Prolixoplectidae),
Gerochammina Neagu, 1990 with terminal aperture(the character of the inner
structure of, Falsogaudryinella and Uvigerinammina was shown by Kaminski and
Filipescu, 2000) and Pseudoreophaxinae Mikhalevich & Kaminski subfam. nov.
(with the type species Pseudoreophax cisovnicensis Geroch, 1961, non
Pseudoreophacinae Suleymanov, 1963 invalid according to Loeblich & Tappan,
1988 (with t.sp. Pseudoreophax marginulinae Suleymanov, 1963 (= Adelungia
Suleymanov, 1966)),
test with the uniserial latest part (Pseudoreophax, (from Verneuilinidae)), ?
Cylindroclavulininae subfam. reinstatedthe coiled part of the shell small, reduced,
at least in one of the generations, chambers wide, of “goulot” (chevron-shaped)
character (close to chevron-shaped) elongated into narrow usually long apertural
tube, which is unclear externally, visible only in sections, the most part of the shell is
uniserial, character of the inner “tooth” needs further study (Cylindroclavulina (from
Valvulinidae)), Barbourinellidae with ? Spiroplectinatinae (Spiroplectinata,
Gaudryinoides, Spiroplectina)(without Belorussiella with the basal aperture) and
Barbourinellinae (Barbourinella, ?Bermudezina, ?Latentoverneuilina (from
Verneuilininae) without Heterostomella wich have the basal aperture at the initial
stage) .
Superfamily Liebuselloidea Saidova, 1981, stat. n.
Test with the complicated wall or subdivided chamber lumen.
Family Liebusellidae (Liebusella, Cubanina, ?Kaeveria (from Pernerinidae ), ?
Guppiella from Textulariellidae, (without genera Jarvisella, Matanzia, Remesella
and Ruakituria having the basal aperture at least initially), Marieitidae (Hensonia,
Order PSEUDOPALMULIDA Mikhalevich, 1992
Test biserial, at least initially, may be with characteristic V-shaped chambers
(Pseudopalmulidae); wall microgranular or agglutinated; aperture terminal. Devon.
Family Pseudopalmulidae (Pseudopalmula).
Incertae Sedis: family Agglutisoleniidae fam. nov. – test with supposed
trochospiral initial coil, later uniserial, but may be it is uniserial throughout, uniserial
part with broad low inflated chambers narrowing at the base, with strongly
constricted sutures, wall thick, agglutinated, aperture terminal, on a short neck, with
inner tube, tubes of the adjacent chambers interconnected. U. Triassic
(Agglutisolenia, transferred from Pernerinidae).
Remarks. If the test is uniserial throughout the family belongs to Hormosinida,
otherwise – to Nouriida.
Class ROTALIATA Mikhalevich, 1980
Subclass TEXTULARIANA Mikhalevich, 1980
Test multichambered only, all regular types of chamber arrangement well presented
(glomerate type practically absent), mostly trochospiral, planospiral 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) and pillars till septa and septula (of the first and the second order)
in higher representatives; chambers usually isomorphic, chamber number could
increase to some hundreds; additional skeletal plates above the sutures and
umbilicus may present; wall agglutinated and microgranular, often with various
complication of the inner wall, which could be cancellate, alveolar, spongy, may
have pseudopores or sometimes canaliculi;; aperture basal, usually at the base of the
apertural face both in coiled and in some biserial and triserial monoaxial forms, in
the last ones could became secondarily terminal(areal, sieve-like), may have various
inner structures ( tooth-plates and open subcylindrical or closed cylindrical plates)
which could be interconnected and could form a single system, supplementary
apertures well developed (sutural, umbilical, peripheral); mostly free living, rarely
attached. … - Holocene.
Superorder DUSENBURYINOIDA Mikhalevich, 1980
?Order DUSENBURYINIDA Mikhalevich, 1980
Test uniserial, aperture with the inner tooth. Holocene. Family Dusenburyinidae
Test planospiral, in one case biserially planospiral, aperture basal (interio-marginal),
at least initially, suplementary apertures may present. Upper Trias – Holocene.
Remarks. The order differs from the order Lituolida in the character of its aperture
which is really basal (not terminal) at least at the early stages of the ontogenetic
development or rarely may be interio-areal, from the order Haplophragmiida in its
planospiral coiling. In some genera the initial position of their aperture needs to be
restudied (= elucidated).
Superfamily Nautiloculinoidea Loeblich et Tappan, 1985
Test with the simple wall and chamber interior.
Families Haplophragmoididae with Haplophragmoidinae (lacking supplementary
apertures) (Haplophragmoides, Asanospira, Cribrostomellus, Cribrostomoides,
Labrospira, Unitendina, Recurvoidella,?Debarina, ?Evolutinella, ?Gobbettia, ?
Trochamminita, ?Veleroninoides, ?Deuterospira (transferred from Mayncinidae as
having basal aperture), Nonionammina Neagu, 2000, without Ammosiphonina,
Buzasina) and Trematophragmoidinae subfam. n. having supplementary sutural
apertures (Trematophragmoides Bronnimann et Keij, 1986), Nautiloculinidae
(Nautiloculina, Murgeina).
Superfamily Alveolophragmioidea Shchedrina, 1936
Test with the complicated wall, more usually alveolar.
Families Alveolophragmiidae with Alveolophragmiinae (Alveolophragmium,
Reticulophragmium, Reticulophragmoides Gardstein et Kaminski, 1989,
Quasicyclammina, without Popovia and Sabellovoluta having terminal aperture from
the very beginning) and Hemicyclammininae (Hemicyclammina, Alveocyclammina,
without Flabellamminopsis), Discamminidae (Discammina, with the rest 3 genera
put into Ammoscalariidae Mikhalevich, 1982(subclass Lagynana with the tectinous
wall), Cyclamminidae (Cyclammina).
Order SPIROPLECTAMMINIDA Mikhalevich, 1992
Test elongated, may be widened at the upper part, consists of planospirally coiled
initial part and following uncoiled straight biserial part, sometimes changing into
uniserial one; aperture basal, interiomarginal, at least initially, but often in the adult
also, single or multiple.
Remarks. The planospiral part constitutes significant part of the test. From the former
Spiroplectamminacea only forms with the basal aperture (even if having it only
initially) are left. Many of the genera need to be restudied concerning the position of
their aperture at the initial stage.
Superfamily Spiroplectamminoidea Cushman, 1927
Test with the simple wall and chamber lumen not subdivided.
Family Spiroplectamminidae with Spiroplectammininae (Spiroplectammina, ?
Ammobaculoides, Bolivinopsis, Heterantyx, Textella Mikhalevich, 1979(=Palustrella
Bronnimann, Whittaker et Zaninetti, 1992), Quasispiroplectammina, ?Spiroplectella,
Spiroplectinella, without Orectostomina), Vulvulininae (Vulvulina, ?Ammospirata), ?
Morulaplectinae (coiling unclear – streptospiral ?) (Morulaplecta).
Superfamily Spirotextularioidea Saidova, 1975
Test with the inner septa or alveolar wall.
Families Spirotextulariidae with Spirotextulariinae (Spirotextularia, Septigerina),
Duquepsammininae Seiglie et Baker, 1987 (Duquepsammina Seiglie et Baker, 1987
– very close to Septigerina), Novalesiidae ( Novalesia), ?Ecougellidae (Ecougella), ?
Pavonitinidae (Pavonitina, Pavopsammia, ? Zotheculifida, without Pseudotriplasia).
Order TEXTULARIIDA Delage et Herouard, 1896
Test elongated, biserial, more often flattened, may be uniserial at the end, the initial
part often having the additional chamber resulting in pseudotriserial appearance;
aperture basal, at least initially, later may be terminal, single or multiple,
supplementary peripheral apertures may present.
Remarks. The forms lacking canaliculi are included here as separate families. Many of
the genera need to be restudied concerning the position of their aperture at the initial
Superfamily Textularioidea Ehrenberg, 1938
Test with the simple wall and undivided chamber lumen.
Families Pseudobolivinidae (Pseudobolivina (without Lacroixina and Parvigenerina
with the terminal aperture from the initial stages)) , Textulariopsidae with
Textulariopsinae (Textulariopsis, ?Aaptotoichus, ?Bimonilina, ?Haimasiella, ?
Minyaichme,?Pleurostomelloides, ?Trunculocavus (without Plectinella)) and
Kaminskiinae Neagu, 1999 (Kaminskia, Spirorutilus), Textulariidae with
Textulariinae (Textularia, Haeuslerella, Paravulvulina, Sahulia, Semivulvulina,
Tetragonostomina), Norvanganinae subfam. n.with supplementary peripheral
chamberlets open at the ends (Norvanganina), ?Siphotextulariinae ( Siphotextularia,
Karrerotextularia, Plecanium, Siphoscutula, Textulina) and Planctostomatinae
(Planctostoma, Olssonina, Poritextuaria), Bigenerinidae with Bigenerininae
subfam. reinstated (Bigenerina) and Cribrobigenerininae subfam. n. test with the
well developed uniserial part, aperture cribrate at the last stage (Cribrobigenerina),
Siphobigenerininae (transferred from Valvulinidae)(Siphobigenerina),
Textularioididae (Textularioides).
Superfamily Septotextularioidea Loeblich et Tappan, 1985
Test with the subdivided chamber lumen.
Families Septotextulariidae (Septotextularia, Colomita), Tawitawiidae (Tawitawia).
Order VERNEUILINIDA ord. nov.
Test high trochospiral throughout or only in the initial part, later part may have
increased or decreased number of chambers per whorl or may become uniserial or
cyclical, aperture basal at least in the initial part, later may become terminal, single
or multiple, some genera with inner apertural structures or supplementary peripheral
apertures. .
Remarks. Forms with the terminal aperture from the very beginning of the growth and
some structural Nodosariata’s features are transferred into subclass Hormosinana. In
family Conotrochaminidae the aperture of the initial stages needs to be checked.
Superfamily Verneuilinoidea Cushman, 1911
Test with the simple wall and undivided chamber lumen.
Families ?Conotrochamminidae (Conotrochammina, Alterammina Bronnimann et
Whittaker, 1988, Earlandammina Bronnimann et Whittaker, 1988, Resupinammina
Bronnimann et Whittaker, 1988, ?Tritaxis?Tritaxis), Prolixoplectidae with
Prolixoplectinae (Prolixoplecta, Arenogaudryina, Caronia Bronnimann, Whittaker et
Zaninetti, 1992, Convallina Mc-Neil, 1997, Danubia Neagu, 1997, Eomarssonella,
Kadriayina Al-Najdi, 1975 (reinstated by Kaminski, 2001), Magnesoina,
Neaguammina Kaminski, Holbourn et Geroch, 1997(differs from Gaudryinopsis in its
well developed triserial part (Kaminski et al., 1997), Orientalia, ?Plectina,
Praedorothia Desai et Banner, 1987, Protomarssonella Desai et Banner, 1987,
Riyadhella, Verneuilinella, without Karrerulina) and Gerochellinae Kaminski, 2003 in
this book (Gerochella Neagu, 1997), Piallinidae Rettori et Zaninetti, 1993 (Piallina
Rettori et Zaninetti, 1993), Verneuilinidae ( without Spiroplectinatinae and
Barbourinellinae) with Verneuilinoidinae (Verneuilinoides, Belorussiella (from
Spiroplectinatinae characterized by the terminal aperture), Flourensina,
Gaudryinopsis, Mooreinella, without Duotaxis transferred to Duotaxidae fam. n.,
without Eggerellina, Vialovella transferred to Globotextulariidae and without
Falsogaudryinella, Paleogaudryina, Paragaudryina, Pseudoreophax, Reophacella,
Talimuella, Uvigerinammina transferred to s/cl. Hormosinana) and Verneuilininae
(Verneuilina, Gaudryina, Gaudryinella, Paramigros Adb-Elsahfy et Ibragim, 1990,
Siphogaudryina, Heterostomella (from Barbourinellinae) (without Latentoverneuilina
and Pseudogaudryinella)), Globotextulariidae (Globotextularia, Cribroturretoides,
Gravellina, Rhumblerella, Tetrataxiella, Verneuilinulla and Eggerellina and
Vialovella from Verneuilinidae and Hagenowella and Sabulina from
Ataxophragmiinae), Tritaxidae (Tritaxia, Bitaxia).
Superfamily Crenaverneuilinoidea superfam. nov.
Test with subdivided chamber lumen.
Family Crenaverneuilinidae fam. n. test initially trochospiral, then triserial, with
intercameral sutures parallel to the axis of coiling or in usually triserial, rarely biserial
order , chambers with internal batresses, partitions in the marginal chamber zone,
lower chamber edge may be crenulate, wall agglutinated, aperture basal, arch- or loop-
shaped, at least initially. Family differs from Pernerinidae in the character of coiling
(not Ataxophragmium-like) and chamber direction not inclined to the axis of coiling,
from Textulariellidae in less regular conical form, inflated chambers and depressed
sutures (Crenaverneuilina, Hagenowina from Pernerinidae, Jarvisella, Matanzia,
Remesella, Ruakituria from Liebusellidae, Alveovalvulina and Hagenowinoides from
Superfamily Valvulinoidea Berthelin, 1880
Test with the simple wall and undivided chamber lumen.
Eggerellidae with Dorothininae (Dorothia, Arenodosaria, Bannerella, ?Marssonella,
?Connemarella Loeblich et Tappan, 1989), Minouxinae (Minouxia, Andersenia,
Tetraminouxia) and Eggerellinae (Eggerella, ?Eggerelloides, Eggerina, Karreriella,
Martinottiella, Mediamonella, Multifidella, Rudigaudryina), Pseudogaudryinidae
with Pseudogaudryininae (Pseudogaudryina, Clavulinoides, Clavulinopsis,
Hemlebenia Loeblich et Tappan, 1989, ?Paragaudryinella Popescu, 1998,
Pseudoclavulina, Pseudogaudryina, Valvoreussella) and Siphoniferoidinae
(Plotnikovina, Siphoniferoides), Valvulinidae (only forms with the external valvular
tooth (flap) are left in this family) with Valvulininae (Valvulina, Makarskiana,
Neoclavulina), Cribrobulimininae s/fam. n. with the valvular tooth cribrate at the
last stage (Cribrobulimina) and Vacuovalvulininae s/f. n. test conical, early part
trochospiral, with decreased or increased chamber number in the latest whorls, wall
agglutinated, canaliculate in the typical genus, aperture with external valvular tooth
(flap) occupying the center of the flat apertural side (Vacuovalvulina transferred from
Chrysalidinidae, pillars absent, ?Pityusina transferred from Ataxophragmiinae,
noncanaliculate, Redmondoides Banner, Simmons et Whittaker, 1991 from
Paravalvulininae Banner, Simmons et Whittaker, 1991), Clavulinidae Balakhmatova,
1973 - fam. reinstated here from Valvulinidae and given stat. n– differs from
Valvulinidae in having internal tooth, with Goesellinae s/f. n. – with short internal
tooth, teeth of the adjacent chambers not connected (Goesella, Gyrovalvulina, ?
Cribrogoesella (aperture multiple in adults)), Clavulininae Balakhmatova, 1973,
family reinstated here from Valvulinidae inner teeth of the successive chambers
form interconnected system running from aperture to foramen (Clavulina).
Superfamily Chrysalidininoidea d”Orbigny, 1839
Test with the chamber lumen subdivided by partitions or pillars.
Family Chrysalidinidae with ?Pseudodictiopsellinae Sepfontaine et De Matos, 1998
(Pseudodictiopsella Sepfontaine et De Matos, 1998 ), Tritaxilininae (Tritaxilina) ,
Chrysalidininae (Chrysalidina, ?Accordiella transferred by Kaminski from
Pfenderinidae, Dukhania, ?Pfendericonus, Praechrysalidina transferred by Kaminski
from Ataxophragmiinae, Pseudochrysalidina, without Parurgonia included by
Sepfontaine into Parurgoninidae Sepfontaine, 1988), Paravalvulininae Banner,
Simmons et Whittaker, 1991(Paravalvulina Sepfontaine, 1988, ?Indomarssonella, ?
Pseudomarssonella, ?Riyadhoides).
Incertae Sedis (to all the order): Siphovalvulina Sepfontaine, 1988.
Order TROCHAMMINIDA Saidova, 1981
Test low trochospiral, often temporary attached, usually with convex spiral and
concave or flat umbilical side, only few genera with convex umbilical and flat spiral
side, tendency to uncoil purely expressed, umbilical ends of the chambers often of a
special configuration (lobed, curved, elongated); aperture basal, at least initially, at
the base of the apertural face more often at the umbilical side, more rarely at the
peripheral one, later may be areal, single or multiple, supplementary apertures
(umbilical, sutural) well developed, or umbilicus may be opened; additional skeletal
plates occur.
Superfamily Trochamminoidea Schwager, 1877
Test with the simple wall and chamber lumen not subdivided.
Families Trochamminidae with subfamilies: Trochammininae (Trochammina,
Ammoanita Seiglie et Baker, 1987, Calyptammina Nagy et Basov V.A., 1998,
Ammoglobigerina, Asarotammina, Camurammina Bronnimann et Keij, 1986,
Globotrochamminopsis, Lepidoparatrochammina Bronnimann et Whittaker, 1986,
Lingulotrochammina Hercogova, 1987, Paratrochammina, Patellovalvulina,
Portatrochammina, ?Pseudoadercotryma, Trochamminopsis), Trochamminellinae
(Trochamminella, Atlantiella, Pseudotrochammina), Toretammininae Bronnimann,
1986 ( Toretammina Bronnimann, 1986 ), Zavadovskininae Bronnimann et Whittaker,
1988 (Zavadovskina Bronnimann et Whittaker, 1988), Arenoparrellinae
(Arenoparrella, Trochamminula), Jadammininae (Jadammina, ?Entzia),
Polystomammininae ( Polystomammina, Balticammina Bronnimann, Lutze et
Whittaker, 1989, Deuterammina, without Lepidodeuterammina), Vialoviidae
(Vialovia, Arenonionella), Rotaliamminidae (Rotaliammina, Siphotrochammina,
Tiphotrocha, Lepidodeuterammina (from Polystomammininae)), Adercotrymidae
Bronnimann et Whittaker, 1988 (Adercotryma, ?Insculptarenula),
Ammosphaeroidinidae (Ammosphaeroidina), Valvulamminidae (Valvulammina,
Arenagula, Discorinopsis).
Superfamily Remaneicioidea Loeblich et Tappan, 1964
Test with the chamber lumen subdivided by septula or by the infoldings of the wall.
Families Remaneicidae (Remaneica, Bruneica, Remaneicella (=
Septatrochammina)), Carterinidae (Carterina (=Zaninettia), Abyssotherma
Bronnimann, Van Dover et Whittaker, 1989), Asterotrochamminidae
(Asterotrochammina, Asteroparatrochammina), Dictyopsellidae (from the former
Ataxophragmiacea)(Dictyopsella, ?Abuhammadina Abd-Elsahfy et Ibrahim, 1990
(put by the authors into Choffatellinae, but having distinctly trochoid test),
Andamookia, Conorbinella, Dictiopselloides).
Order ATAXOPHRAGMIIDA Fursenko, 1958
Only forms with the typical strongly asymmetrical Ataxophragmiid spire usually
with elongated vertical chambers inclined to the previous whorls or conical shells of
Satorina, typical Coskinolina, Coskinon types also often with the initial part inclined
to the latest triserial or more usually uniserial part are left in this order (excluding the
other forms into the new order Verneuilinida). The phyletic line included here is a
special trochospiral line differing from Trochammina –like or Eggerella-like
trochospiral type in mostly high conical form with noninflated chambers,
development of the uniserial part and its special character, development of columella
or other kind of the integrative inner central structure and passages in the central
space of the shell (the low conical Dobrogelina and Nezzazatinella belong to the
same line having the inner central structure of the same Pfenderinid type as it was
shown by Neagu ( 1979)). It resembles strongly the Glomotrocholina (though
pseudotwochambered) Cylindrotrocholina (Involutinidae)- Tetrataxis -Patellina
Chapmanina Ferayina line, also with conical shells, noninflated chambers, similar
character of the uniserial part (if present) and of the inner structures (columella in
Patellina, hollow inner pillars are known in Pseudolituonella (Ataxophragmiida) and
in Angotia (Chapmaninidae ). The taxonomic elaboration of this line needs futher
In Ataxophragmiidae and Pernerinidae forms with the typical Ataxophragmium spire
and with the usual trochospire reduced to triserial one are separated here into
different families.
Superfamily Ataxophragmoidea Schwager, 1877
Test with the simple wall and undivided chamber lumen.
Families Duotaxidae fam.n. - test conical, circular in section, Textulariella-like, with
flattened apertural face, early part trochospiral, later biserial and may be uniserial,
chambers broad, nun inflated, wall agglutinated, smooth, aperture basal, a wide
horizontal slit at the base of the apertural face of the last chamber, may be sudivided
by the small median lobe; family differs from the representatives of Verneuilinidae
where it was previously placed in the character of the test and chamber shape
(Duotaxis), Ataxophragmiidae with Ataxophragmiinae (only genera having
Ataxophragmium-like largely involute spiral coiling with not visible early whorls and
with high chambers turned to the axis of coiling are preserved here)
(Ataxophragmium, Arenobulimina, ?Ataxorbignyna, without Hagenowella and
Sabulina (both transferred to Globotextulariidae) and without Pityusina and
Superfamily Textularielloidea Groenhagen et Luterbacher, 1966
Test with the subdivided chamber lumen or alveolar wall.
Families Pernerinidae with Pernerininae (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 tests lacking uniserial part
are preserved here) (Pernerina, Orbignyna, Voloshinoides, , ?Opertum, ?Anatoliella
without Agglutisolenia, Coprolithina and Kaeveria transferred to s/cl. Hormosinana
and without Crenaverneuilina, Hagenowina and Voloshinovella) and
Voloshinovellinae s/fam. n. - test with small initial distinctly trochospiral part , later
big rectilinear uniserial part inclined to trochospiral one, slightly widened to the end,
circular in section, rectilinear chambers subdivided marginally by vertical radial
partitions in the marginal zone of the chambers, partitions usually simple, rarely
bifurcating, wall agglutinated, solid, noncanaliculate, aperture basal in the uncoiled
stage, later areal, single or multiple - rounded openings in the central area; differs
from Pernerininae in its small coiled part, the most of the test formed by uniserial
rectilinear widening part giving to it subconical appearance, from Coskinolininae - in
its endoskeletal features: more simple and situated in the marginal part of the
chambers (Voloshinovella, Trochamijiella Athersuch, Banner et Simmons, 1992
( originally put into Biokovinidae but differs from it in its early trochoid coil)),
Textulariellidae (Textulariella, Cuneolinella, without Alveovalvulina,
Hagenowinoides transferred to Crenaverneuilinidae and Guppyella transferred to s/cl/
Hormosinana), Cuneolinidae without Sabaudiinae having hyaline radial layer of the
wall, with Cuneolininae (Cuneolina (in understanding of Neagu, 2000
Palaeolituonella, Pseudotextulariella, ?Vercorsella and Histerolina Neagu, 2000,
Scythiolina Neagu, 2000 two last genera with initial scarse planospiral part
according Neagu, 2000) and Montsaleviinae Zaninetti, Salvini-Bonnard, Charollais et
Decrouez, 1987 (Montsalevia Zaninetti, Salvini-Bonnard, Charollais et Decrouez,
1987), Dicyclinidae (Dicyclina), Pfenderinidae with Pseudopfenderininae (without
tunnels)(Pseudopfenderina, Pseudoeggerella), Paleopfenderininae ( Paleopfenderina
Sepfontaine, Conicopfenderina, Chablasia (transferred by Kaminski from
Biokovinidae), Satorina, Senderella, ?Steinkella (chambers subdivided), Pfenderininae
(Pfenderina, Drevennia, Pfenderinella, without Accordiella transferred by Kaminski
to Chrysalidininae), Dobrogelininae Mikhalevich, 1992 (Dobrogelina, Nezzazatinella)
and Kurnubiinae (Kurnubia, Gyroconulina, Praekurnubia, ?Conicokurnubia);
Coskinolinidae (Coskinolina, Coleiconus, Coskinon, Lituonelloides,
Pseudolituonella); Hauraniidae Septfontaine, 1988 (Haurania, ?Cymbriaella
Fugagnoli, 1999, Gutnicella, Meyendorfina, ?Plathyhaurania Bassoulet et Boutakiout,
1996, ?Socotraina Banner, Whittaker, Boudagher-Fadel et Samuel, 1997,
Spiraloconulus (from Spirocyclinidae), (without Amijiellinae); Parurgoninidae
Sepfontaine, 1988 (Parurgonia), Orbitolinidae with Dictyoconinae ( Dictyoconus,
Abrardia, Calveziconus, Campanellula, Carinoconus, Coskinolinoides, Cribellopsis,
Cushmania, Daviesiconus, Dictioconella, Fallotella, Falsurgonina,
Heterocoskinolina, Iraqia, Karsella Sirel, 1997, Orbitolinella, Orbitolinopsis,
Paleodictyoconus, Pseudorbitolina, Simplorbitolina, Urgonina, Valdanchella,
Verseyella), Praedictyorbitolininae (Praedictyorbitolina Schroeder, 1990,
Dictyorbitolina, Paracoskinolina) and Orbitolininae (Orbitolina, Alpillina,
Conicorbitolina, Eopalorbitolina, Eygalierina, Mesorbitolina, Naupliella, Neoiraqia,
Neoorbitolinopsis, Palorbitolina, Palorbitolinoides, Praeorbitolina,
Rectodictyoconus, Valserina).
I am deeply grateful to the Grzybowski Foundation for the support and invitation to
participate in the Sixth International Workshop on Agglutinated Foraminifera and
especially to Mike Kaminski for kindly providing original descriptions and literature
hardly available in my country. I would like also to sincerely thank him for
encouraging me to fulfill this work, for his help in preparing the english text and the
pictures. I also thank Theodor Neagu permitting me to work with his perfectly cured
collections, presenting me part of his excellent material and as well for the fruitful
discussions on the problems of the foraminiferal classification.
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Angell R.W. 1980. Test morphogenesis (chamber formation) in the fo-
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FOUNDATION (Editor M/A/ Kaminski)
... Members of the protistan Phylum Foraminifera (d'Orbigny, 1826) (phylum status was proposed by Mikhalevich, 2004 andPawlowski et al., 2013) are ubiquitous in marine environments. They are well preserved in Phanerozoic sediments and sedimentary rocks as a consequence of their organic, agglutinated, or calcareous shells (commonly called "tests"), making them among the most widely utilized organisms in earth and ocean sciences (e.g., Sen Gupta, 1999). ...
The Foraminifera are a diverse class of protists whose ubiquity in marine environments, small shells, and ease of collection have made them critical tools in bioindicator, bioassay, paleo-environmcntal, and paleoceanographic research. Despite the plethora of applications and accompanying literature on foraminifers, many aspects of their biology and ecology remain unexplored. One of these aspects is dormancy, a life-history strategy involving suspension of active life, arrested development, and reduced or suspended metabolic activity, mediated either by internal physiological factors (known as diapause) or exogenous factors (known as quiescence). Dormancy is a widespread adaptation, playing a role in the life cycles of a huge variety of organisms. Yet, despite anecdotal and circumstantial evidence, very little research has directly addressed this aspect of foraminiferal biology. The relatively recent discovery of cryptobiotic propagules has revealed a fundamental role for dormancy in the life cycles of foraminifers, most prominently for dispersal. Moreover, culture studies with environmental applications have shown that post-propagule quiescence (i.e., in juveniles, sub-adults and adults) may be a common response to environmental stressors, allowing rapid recovery of populations following disturbance or otherwise unfavorable conditions. A review of publications on foraminiferal biology revealed that observations indicating the potential for dormancy have been recorded for at least six decades, and that this potential is well represented throughout the class in a variety of forms, suggesting that dormancy may be a basic adaptation in the Foraminifera. If dormancy is as widespread as the literature suggests, its role in structuring foraminiferal assemblages and determining global distributions in the geological past, present, and future is fundamental. Further research into the mechanisms of dormancy will expand understanding of its role in foraminiferal life cycles, and provide new perspectives in the many fields that utilize and apply foraminiferal data.
... We use the classification system of higher foraminiferal taxa proposed by Cavalier-Smith (2002) and Mikhalevich (1998 Mikhalevich ( , 2000 Mikhalevich ( , 2004). Description and terminology mainly follow Gerke (1967), which was partially translated into English by Karavaeva, 1998] Order Nodosariida Calkins, 1926 [nom. ...
Full-text available
A latest Changhsingian (latest Permian) foraminiferal fauna composed of 19 species (belonging to 10 genera) was recovered from the Daxiakou section in Xingshan County, Hubei Province, South China. Compared to contemporaneous faunas in South China, the Daxiakou fauna displays unique features: (1) nodosariids are the dominant forms in abundance and diversity; and (2) the most abundant forms have elongate and flattened tests, and include Geinitzina , Howchinella , and Ichthyolaria . The predominance of flattened elongate tests and their smaller size suggest that these foraminifers possibly lived in an oxygen-depleted, deep-water environment. Seven new species, Nodosaria quinquecostata n. sp., Howchinella inflata n. sp., H. hubeiensis n. sp., H. complanata n. sp., H. xiangxiensis n. sp., Ichthyolaria celsa n. sp., and Pseudotristix elongata n. sp. are described.
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ABSTRACT - Owing to the intensive movement of the cytoplasm through the aperture where the communication with the environment is realised, this area is charged with a strong and variable functional burden. Additional skeletal structures have fundamental supporting functions along the course of the strong cytoplasmic stream and may be related to the compartmentalization and differentiation of the cytoplasm. In connection with these important functional roles, the structure of the aperture is one of the basic diagnostic features in foraminiferal taxonomy. The simplest and least diverse apertural types are found in the most ancient unilocular or pseudochambered representatives of the classes Lagynata and Astrorhizata. Their development from simple to complicated ones in the different classes (following the new foraminiferal macrosystem proposed by Mikhalevich) shows a significant number of parallelisms and convergences. In both the lower agglutinated groups and the higher calcareous members of the classes Spirillinata, Miliolata, Nodosariata and Rotaliata, the evolutionary trends of the apertures are similar within the same class, even if those of the lower groups are always structurally simpler and less diversified. These trends continue until all the possibilities of the pre-existing structures are exhausted. Then, new structures, sometimes affecting the whole cell organisation arise, and the possibilities of morphological changes multiply leading to evolutionary divergences.