The Paleozoic forams

Paleontology > Benthic Foraminifera > The Paleozoic forams

Introductory remarks: Paleozoic Foramifera

The fossil record indicates that the very first foraminifera (with a hard shell) was the unilocular simple agglutinated Allogromiina, which evolved into the calcareous agglutinated Textulariina during the Cambrian. While the Textualariina were a dominant group early in the Paleozoic, the Fusulinina became dominant later in the Paleozoic (from the Silurian).The Fusulinina are morphologically larger with compound, microgranular walls and complex internal structure and they constitute much of the Late Paleozoic shallow-marine limestone seen today (BouDagher-Fadel, 2008).

Because of their importance, the main focus of this sub-page is the suborder Fusulinina. Other suborders we will discuss here are the Lagenina, which evolved in the Silurian, the Miliolina and the early Involutinina. The lagenines, miliolines and involutinines were morphologically small and primitive throughout the Paleozoic, measuring between 0.4 and 1.3 mm in diameter. The evolution of the Paleozoic larger foraminifera is shown to the left (BouDagher-Fadel, 2008).

The suborder Fusulinina contains twelve superfamilies and 418 distinct genera (Loeblich and Tappan, 1988), most of which became highly specialized during the latter part of the Paleozoic. Rapid evolution facilitated large and diverse lineages of Fusulinina and they achieved a remarkable size for a single celled organism. By the end of the Permian, some forms of fusuline reached 15 cm in diameter (BouDagher-Fadel, 2008). Despite their dominating presence, complicated structure and large size, they were extinct by the end of the Paleozoic.

Below we present a brief description of each superfamily and family and include a description of select genera beneath each family. For more detailed taxonomic descriptions of all genera, see Loeblich and Tappan (1988) and Rauser-Chernousova et al. (1996). Additionally, please refer to the paper by M. BouDagher-Fadel (2008), from which this page is summarized, and references found there-in, especially regarding the individual descriptions for each family. Also, refer to the Miracle home-page, constructed by Dr. Marcelle K. BouDagher-Fadel of the University College of London (University College of London - MIRACLE).

For a list of key Suborders, Superfamilies and Families that are discussed in the following sections, please see the diagram below (after BouDagher-Fadel, 2008). This diagram, as well as the large diagrams in the following section can be viewed at a larger scale once the shadowbox has been activated by right-clicking on the diagram and opening it in a new tab (Google Chrome) or selecting the View Image function (IE).

SUBORDER FUSULININA

Test: homogeneously microgranular primary test made of low-magnesium calcite, in which the crystal units have no optical alignment.
Wall: advanced forms have two or more alignment
Geologic Range: Silurian to Triassic

Superfamily PARATHURAMMINOIDEA

Test: unilocular, globular or tubular
Wall: simple calcareoius microgranular or with granulo-fibrous layers.
Geologic range: Early Silurian to Permian. Live free or attached.

Family Archaesphaerida

Test: One or more globular to elongate chambers
Wall: No aparent aperture
Geologic range: Late Silurian to Early Permian

Diplosphaerina

Test: Two chambered, with small proloculus enveloped within a larger chamber

Geologic range: Middle Devonian to Tournaisian

Family Parathuramminidae

Test: globular or irregular
Wall: simple calcareous agglutinated wall with multiple aperture at end of tubular neck
Geologic range: Late Silurian to Carboniferous

Parathurammina

Test: Globular with aperature at ends of numerous tubular protuberances

Geologic range: Late Silurian to Carboniferous

Family Chrysothuramminidae

Test: globular to irregular in outline
Aperture: at the end of tubular projections
Geologic range: Middle Devonian to Carboniferous

Chrysothurammina (B)

Test: Single subspherical chamber

Aperture: on a neck-like protrusion.

Geologic range: Middle Devonian to Carboniferous

Family Ivanovellidae

Wall: thick calcareous wall with two layers; a thin dark compact granular inner layer and a thick grey radially fibrous outer layer.
Geologic range: Late Silurian to early Carboniferous

Elenella (B)

The outer surface has tiny projections;

Aperture: none

Geologic range: Middle Silurian to Carboniferous

Family Marginaridae

Wall: three layered; inner and outer layers are dark and the middle layer is light gray with canals running through it, which form projections outside the wall
Geologic Range: Middle to Late Devonian

Family Uralinellidae

Wall: small, unilocular with three layers. The median layer is thick and clear and the outer layers dark.
Aperture: a multiple apertural neck is projected from the test
Geologic range: Middle Devonian to Early Carboniferous

Sogdianina

Aperture: very prominent apertural necks

Geologic range: Carboniferous

Family Auroriidae

Test: ovate.
Wall: two layered, a thin inner finely porous dark layer and a thick outer canaliculated dark layer
Geologic range: Middle to Late Devonian.

Auroria

Test: irregularly globular

Geologic range: Middle to late Devonian

Family Usloniidae

Test: globular with no distinct aperture
Geologic range: Middle Devonian to Early Carboniferous

Parphia

Test: spherical to ovate

Wall: dark and evenly porous

Geologic range: Late Devonian

Family Eovolutinidae

Test: globular
Wall: microgranular with single distinct aperture
Geologic range: Late Silurian to Late Devonian

Eovolutina

Test: tiny with proloculus completely surrounded by second chamber

Geologic range: Late Silurian to Late Devonian

Family Tuberitinidae

Test: simple, attached, made of one or more globular chambers
Wall: calcareous, microgranular
Geologic range:Silurian to Permian

Draffania

Test: Flask-shaped tuberitinids

Aperture: at the end of an elongate neck

Geologic range: Tournaisian to Visean

Eotuberitina

Test: hemispherical with basal disc

Wall: microgranular, coarsely perforate

Geologic range: Middle Devonian to Middle Carboniferous

Paratuberitina

Test: highly hemispherical with basal disc

Wall: microgranular, coarsely perforate

Geologic range: Carboniferous (Visean to Moscovian)

Tuberitina

Test: Shows bulbous chambers in an arcuate or straight series

Geologic range: Late Carboniferous to Late Permian

Tubeporina

Test:hemispherical with basal disc and three-layered wall; a hyaline layer between two microgranular layers

Wall: coarsely perforate

Geologic range: Middle Devonian to Carboniferous

Superfamily EARLANDIOIDEA

Test: free, non-septate with a globular first chamber and a straight tubular second chamber
Geologic range: Late Silurian to Early Triassic

Family Earlandiidae

Chamber: single, free
Geologic range: Late Silurian to Early Triassic

Earlandia

Test: free, elongate, composed of globular proloculus followed by an undivided straight tubular chamber

Wall: calcareous microgranular

Geologic range: Late Silurian to Early Triassic

Family Pseudoammodiscidae

Test: small with a simple aperture and globular proloculus followed by planispiral, trochospiral or streptospirally coiled, undivided second chamber
Geologic range: Devonian to Permian

Brunsia

Test: proloculus is followed by an early streptospiral coil and later a planispirally enrolled tubular chamber

Geologic range: Devonian to Carboniferous

Brunsiella

Test: globular proloculus is followed by undivided second chamber, early whorls streptospiral, later ones planispiral and evolute

Geologic range: Carboniferous to Early Permian

Family Pseudolituotubidae

Test: attached
Chamber: single enrolled with simple terminal aperture
Wall: calcareous microgranular
Geologic range: Early Carboniferous to Late Carboniferous

Superfamily ARCHAEDISCOIDEA

Test: small, consisting of a proloculus followed by an enrolled chamber
Wall: one or more layers
Geologic range: Early Carboniferous to Late Carboniferous

Family Archaediscidae

Test: discoidal test composed of a proloculus followed by a stretospirally enrolled second chamber
Wall: dark inner granular layer and a clear radially fibrous or granulo-fibrous outer layer
Geologic range: Early Carboniferous to Late Carboniferous

Archaediscus

Test: free streptospiral, with an undivided tubular second chamber and thickened wall

Geologic range: Carboniferous

Asteroachaediscus

Test: small, lenticular with sigmoidal coiling, stellate and with occluded chamber cavity

Geologic range: Carboniferous

hemiarchaediscus

Test: small, discoidal with glomospiral early whorl, later becoming planispiral; chamber lumen remains open

Geologic range: Carboniferous

Planoarchaediscus

Test: small discoidal with glomospiral early stage, followed by a planispiral adult; chamber lumen remains open

Geologic range: Carboniferous

Propermodiscus

Test: small, lenticular wtih glomospiral early whorls and a planispiral, involute adult

Geologic range: Carboniferous

Family Lasiodiscidae

Test: discoidal to conical. The radial-fibrous outer layer is mainly concentrated in the umbilical region where it may for pillars, or a series of tubercles on the surface of one side of the test
Geologic range: Early Carboniferous to Permian

Howchinia

Test: conical with septal bridges. The umbilical region is filled with fibrous calcite-forming pillars

Geologic range: Carboniferous

Superfamily NODOSINELLOIDA

The Nodosinelloidea evolved from the Parathuramminoidea via the free Moravamminoidea. This evolution of the families is shown graphically below. Please refer to Figure 1 (below) to trace the evolution of the specific family and their genera.

Test: Free with more than one distinct chamber, partially septate or fully septate
Wall: simple microgranular, evolving into forms with an extra fibrous inner layer
Geologic range: Late Silurian to Permian

Family Earlandinitidae

Test: simple microgranular
Geologic range: Late Devonian to Late Carboniferous
The Earlandinitidae evolved from the Caligellidae by becoming elongate uniserial, semiseptate.
Genera: Earlandiinita. Subseptate with a terminal round aperature; Carboniferous

Family Nodosinellidae

Test: uniserial test developed in the Late Carboniferous
Wall: double-layered with an inner fibrous layer and an outer microgranular layer
Geologic range: late Silurian to Permian
The Nodosinellidae evolved from the Earlandinidae
Genera: Nodosinella. Free and fully septate test; Permian

Superfamily COLANIELLAOIDEA

Test: uniserial and fully septate
Chamber: strongly overlapping with partitions
Wall: an outer vitreous layer and an inner granular layer
Geologic range: Late Devonian to Late Permian

Family Colaniellidae

Test: strong internal partitions or septa
Geologic range: Late Devonian to Late Permian
Genera: Colaniella. Elongate and uniserial, having a calcareous wall with a fibrous structure and a terminal radiate aperture. Late Permian.

Superfamily PTYCHOCLADIOIDEA

Test: No distinct apertures; attached
Wall: microgrnular, banded with transverse tubuli
Geologic range: Late Devonian to Carboniferous

Family Ptychocladiidae

Test: uniserial and attached
Geologic range: Late Devonian to Late Carboniferous
Genera: Ptychocladia. Calcarous microgranular two layered wall with no distinct aperture. Late Pennsylvanian.

Superfamily Palaeotextularioidea

Test: biserial or uniserial
Wall: microgranular calcareous with single or multiple aperatures (in later stages)
Geologic range: Middle Devonian to Permian

Family Semitextulariidae

Test: biserial, flattened but often becoming monoserial with broad chambers; fully septate
Geologic range: Devonian to Pennsylvanian
Genera: Koskinotextularia. Elongate, biserial test with a single layered wall and single aperture in early stages but later becoming cribrate. Early Carboniferous to Late Carboniferous.

Figure 1. Schematic morphological evolution of the Fusulinina (and Lagenina) from the Allogromiina (BouDagher-Fadel, 2008)

Family Palaeotextulariidae

The palaeotextulariidae evolved from simple agglutinated froms in the Devonian (Cummings, 1956). Please refer to the figure below to trace the evolution of the palaeotextularioidea.

Test: dark granular calcareous outer wall and inner clear to yellowish 'fibrous' layer with stacks of granules perpendicular to the surface; resembles Textulariidae
Geologic range: Early Carboniferous to Permian

Palaeotextularia

Test: biserial

Wall: thick with a low slit (aperture) opening at the base of the final chamber

Geologic range: Carboniferous to Permian

Deckerellina

Test: biserial

Aperture: in the early stage is a slow slit but the adult aperture is two parallel slits

Geologic range: Carboniferous

Deckerella

Test: biserial in early stage, later uniserial and rectilinear

aperture: a low slit in th eearly stage, later terminal with two parallel selits separated by a a partition

Geologic range: Carboniferous to Permian

Cribrostomum

Test: biserial

aperture: in the early part is basal, in the last two whorls becoming areal, multiple and cribrate

Geologic range: Carboniferous to Permian

Climacammina

Test: biserial followed by a uniserial stage

Aperture: in the early stage is at the base of the last chamber, later becoming areal, multiple and cribrate

Geologic range: Carboniferous to Permian

Family Biseriamminidae

Test: In early stages the test is biserial but transitions to a planar test in later stages
Wall: microgranular with one or more layers
Geologic range: Carboniferous to Permian
Genera: Biseriella. the early stage is tightly coiled and later has an open helicoid spire. Carboniferous to Permian

Superfamily TETRATAXOIDEA

Test: conical with secondary internal partitions

Wall: microgranular with one or two layers

Geologic range: Carboniferous to Permian

Family Tetrataxidae

Test: conical leaving an open central umbilicus where the chambers are partially overlapping
Wall: calcareous microgranular and in two layers
Geologic range: early Carboniferous to Late Carboniferous

Tetrataxis

Test: outer layer dark and microgranular; inner layer is light and fibrous

Geologic range: Carboniferous

Family Pseudotaxidae

Wall: calcareous, microgranular and single layered
Geologic range: Tournaisian to Bashkirian
Genera: Vissariotaxis. Conical with trochospirally coiled chambers and wide aperture. Carboniferous.

Family Valvulinellidae

Test: conical with subdivided chambers
Wall: single layer, microgranular
Geologic range: early to late Carboniferous

Valvulinella

Test: chambers are separated by numerous vertical pillars and one or two horizontal ones

Geologic range: Carboniferous

Family Abadehellidae

Test: Representatives of this family differ from the Valvulinellidae as they have two double-layered walls instead of a single-layered wall
Geologic range: Late Permian
Genera: Abadehella. Conical with numerous whorls and subdivided chambers by regularly spaced pillars. Late permian

Superfamily MORAVAMMINOIDEA

Test: Simple free or attached consisting of a proloculus and a rectinilinear second chamber that was subseptate
Wall: simple
Geologic range: Evolved from the Parathuramminoidea in the Silurian but died out in the Early Carboniferous

Family Caligellidae

Test: attached and subseptate
Geologic range: Late Silurian to Tournaisian
Genus: Paracaligella. Subspherical proloculus followed by an irregular tubular chamber partially divided by incipient septa. Devonian to Carboniferous.

Family Moravamminidae

Test: attached and irregularly septate with an enrolled rectilinear part.
Geologic range: Middle to Late Devonian

Family Paratikhinellidae

Test: free, elongate, subseptate
Wall: simple microgranular
Geologic range: Middle Devonian to Carboniferous

Superfamily TOURNAYELLOIDEA

Test: the proloculus is followed by a planispiral or streptospiral non-septate to fully septate
Wall: made of microgranular calcite evolving into forms with a thick outer layer (tectum) and an inner granulo-fibrous layer
Geologic range: Late Devonian to Carboniferous

Family Tournayellidae

Test: ealry streptospiral or trochospiral coiling, becoming planispiral in adult stages
Aperture: simple and open at the end of a tubular chamber
Geologic range: Devonian to Carboniferous

The Tournayellidae are divided into two lineages, which evolved in the Devonian and followed parallel lines of evolution. the first lineage consists of simple planispiral, non-septate forms with an initial proloculus and an aperture at the end of an uncoiled tube. These forms developed a compound wall and rudimentary septa in the Late tournaisian to almost complete septa in the Late Visian. the second lineage is made of streptospiral tubular forms which gradually developed rudimentary septa also in the late Tournaisian, Please see the diagram below which describes the evolution of the Tournayellidae (BouDagher-Fadel, 2008).

Schematic evolution of the Tournayellidae

Family Palaeospiroplectamminidae

Test: initially streptospiral then planispiral and in later stages becomes biserial
Geologic range: Late Devonian to Carboniferous

Superfamily ENDOTHYROIDEA

Test: streptospiral with many chambers followed by a rectilinear stage in some forms
Wall: microgranular and calcareous, but some forms evolved two to three distinct layers
Geologic range: Late Devonian to Triassic

Family Endothyridae

Test: small, multilocular, enrolled with evolute lanispiral coiling with well developed septa in early whorls or initially streptospiral to planispiral characterized by the development of secondary deposits of cacite on the chamber floor
Aperture: low equatorial, advance forms may become uniserial with a cribrate terminal aperture
Geologic range: Devonian to Permian
Genera examples: Endothyra, Endothyranella, Endothyranopsis, Mikhailovella, Planoendothyra, Plectogyra, Spinoendothyra

Schematic evolution of the Endothyridae

Family Bradyinida

Test: planispiral with few chambers
Wall: microgranular calcareous, perforate with distinct radial lamellae with supplementary septal pores opening into the septal aperture
Aperture: primary aperture is cribrate
Geologic range: Carboniferous

Superfamily FUSULINOIDEA

Test: large, varies from spherical to fusiform; the development of seondary deposits of calcite is prominent in most forms but the numerous chambers are subdivided by folds or septula
Geologic range: Middle Devonian to Permian

There are over 100 genera recognize belonging to this superfamily. The Fusulinoidea have special morphological diagnostic characteristics that make this family unique. There are five types of wall structures that characterize the Fusulinoidea. The first four types of wall characterize the Ozawainellidae, Staffellidae and the Fusulinida. The last type of wall belong to the Schwagerinidae and Verbeekinidae. Please refer to the figure and legend below:

Form A: two-layered walls, an inner clear fibrous light layer (early tectorium) and a dark granulous outer layer (tectum)
Form B: Three-layered walls, the late and early tectorium surrounding the dark layer of the tectum
Form C: Four-layered walls, wth the late and early tectorium surrounding the tectum and a glossy layer called the diaphanotheca
Form D: Three-layered walls, where the late tecctorium disappears and they are left with the tecturm, diaphoanotheca and the early tectroium
Form E: Composed of a tectum and a glossy layer with alveoles running through it

Five types of wall structure recognized in the Fusulinoidea

Key recognizable features of the Fusulinoidea

The Fusulinoidea was divided into eight families, based on wall structure, presence or absence of the chomata and the degree of septal folding in the test. As stated, there are over 100 genera identified. Please refer to the works of BouDagher-Fadel (2008), Loeblich and Tappan (1964; 1988), Haynes (1981) and references therein for descriptions of these important families, subfamilies and genera belonging to the Fusulinoidea.

Family Loeblichiidae
Family Ozawainellidae
Family Schubertellidae
Family Fusulinidae
Family Schwagerinidae
Family Staffellidae
Family Verbeekinidae
Family Neoshwagerinidae

Schematic diagram of the most advanced forms of fusulines

SUBORDER LAGENINA

Test: Members of this suborder have walls that are monolamellar, with or without secondary laminations, with radiating calcite crystals which have the crystallographic c-axis perpendicular to the surface.
Wall: Only advanced forms have secondary lamination. Reiss proposed an extensive reclassification of lamellar forms and Gronlund and Hansen (1976) proposed new terminology for lamellarity;

atelo-monolamellar- primitive lagenines, no secondary lamellarity
plesio-monolamellar- secondary lamellarity envolops some but not all chambers
ortho-monolamellar- primary wall of each chamber secondarily envelops all previous chambers
poly-monolamellar- entire test is covered by a second secondary lamella

Superfamily ROBULOIDOIDEA

Test: typical of the Lagenina but without secondary lamellar or with slight lamination in younger taxa
Geologic range: Late Silurian to Early Cretaceous

Family Syzraniidae

Test: two chambered, elongate, composed of a proloculus and an undivided or subseptate tubular chamber partitioned to varying degrees by internal theickenings of the wall
Wall: calcareous with possibly two layers
Geologic range: Late Silurian to Late Permian

Superfamily GEINITZINOIDEA

Test: uniserial and appear similar to the Nodosinelloidea, but the microgranular layer is the innner dark layer while the radially fibrous layer is an outer layer
Wall: advanced forms have secondary lamellarity
Geologic range: late Devonian to Middle Triassic

Family Geinitzinidae

Test: free and uniserial evolving from species that are rounded in cross-section to species that are compressed with a terminal aperture
Geologic range: Late Devonian to Late Permian
Genera: Geinitzina. Uniserial elongate, laterally compressed test exhibiting in transverse view two planes of bilateral symmetry and an oval aperture in the center of the apertural face. Permian.

Family Pachyphloiidae

Test: free, uniserial compressed

Wall: simple calcareous microgranular but with secondary thickenings on both sides of the test

Geologic range: Permian

Pachyphloiidae

Test: globular proloculus followed by a uniserial succession of laterally compressed, strongly overlapping chambers

Wall: plesio or ortho-monolamellar with thin dark inner layer and thick hyaline outer layerand terminal, oval, radiate aperture

Geologic range: Early to Late Permian

Family Protonodosariidae

Test: This family includes the uniserial Lagenina with atelo-monolamellar or plesio-monolamellar wall structure consisting of radial-fibrous
Geologic range: Late Carboniferous to Late Permian
Genera: Protonodosaria. Elongate test with horizontal straight sutures and round aperture. Late Carboniferous to Late Permian.

Family Ichtyolariidae

Test: elongate uniserial with a single-layered wall. May show secondary lamination.
Aperture: simple, terminal
Geologic range: Early Permian to Early Cretaceous

Family Robuloididae

Test: uniserial, enrolled
Wall: atelo-monolamellar
Aperture: terminal
Geologic range: Late Permian
Genera: Calvezina. Proloculous followed by five or more uniserial chambers, weakly trochospirally enrolled.The chamber is slightly compressed laterally and size increases from hemispherical to lobate. Late Permian.

SUBORDER INVOLUTININA

This suborder includes all forms with an enrolled second chamber. Walls are aragonitic but commonly recrystallized to give a homogeneous microgranular structure. They have an umbilical region with pillar-like structures on one or both sides of the test and range from Early Permian to Late Cretaceous.

Superfamily INVOLUTINOIDEA

This superfamily comprises forms consising of a proloculus followed by a planispiral to trochospiral enrolled trubular second chamber. Early Permian to Late Cretaceous. This superfamily is discussed in more detail in the following section on Triassic forams.

SUBORDER MILIOLINA

Test: porcellaneous and imperforate, which are made of high-magnesium calcite with fine randomly oriented crystals
Geologic range: Carboniferous to Holocene

Superfamily CORNUSPIROIDEA

Test: free or attached and composed of a globular proloculus followed by a tubular enrolled chamber. The coiling is planispiral or trochospiral, evolute or involute.
Aperture: simple, at the end of the tube
Geologic range: Early Carboniferous to Holocene

Family Hemigordiidae

Test: Trochospiral in early stages, later becoming streptospiral
Geologic range: Carboniferous
Genera: Hemigordius; Neohemigordius

Family Hemigordiopsidae

Test: early streptospiral stage, later planispiral
Geologic range: Early Carboniferous to Holocene
Genera: Hemigodiopsis; Shanita

References

BouDagher-Fadel, M.K., 2008. Evolution and Geological Significance of Larger Benthic Foraminifera. Developments in Paleontology andstratigraphy, v. 21, 540p.

Gronlund, H., Hansen, H.J., 1976. Scanning electron microscopy of some recent and fossil nodosariid foraminifera. Bull. Geol. Soc. Denmark 25, 121–134.

Haynes, J.R., 1981. Foraminifera. MacMillan, London, 433p.

Loeblich, A.R., Jr., Tappan, H., 1964. Protista 2, Sarcodina, chiefly ‘‘Thecamoebians’’ and Foraminiferida. In: Moore, R.C. (Ed.), Treatise on Invertebrate Paleontology, University of Kansas Press, Kansas, Part C, Vols. 1 and 2.

Loeblich, A.R., Jr., Tappan, H., 1988. Foraminiferal Genera and their Classification. Van Nostrand Reinhold, New York, two volumes, 2047pp.

Rauser-Chernousova, D.M., Bensh, F.R., Vdovenko, M.V., Gibshman, N.B., Leven, E.Ya., Lipina, O.A., Reitlinger, E.A., Solovieva, M.N., Chedia, I.O., 1996. Guidebook on the Systematics of Foraminifers of Paleozoic. Nauka Publishing House (In Russian), Academy of Sciences of Russia.

** Page generated by Kerry McCarney-Castle; text summarized from BouDagher-Fadel (2008)**