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ACTA GEOLOGICA HISPANICA,
v.
3 1 (1996),
no'
1-3,
p.
183-187 (Pub. 1998)
Morphostructure and paleobiology of Mesogean orthophragminids(Discocyclininidae and Orbitoclypeidae, Foraminifera)
CARLES FERRANDEZ-CAÑADELL
Departament d'Estratigrafia i Paleontologia, Facultat de Geologia, Universitat de Barcelona,Martí Franqués s/n, 08071-Barcelona (Spain).
cferran@natura.geo.ub.es
Orthophragminids are Paleocene-Eocene bilamellar-perforate larger foraminifers characterized by a lenti-cular test with an equatorial layer of rectangular cham-berlets arranged in concentric rings and lateralchamberlets on either side. Usually called "discocy-clinids", orthophragrninids include genera from two fa-milies: Discocyclinidae and Orbitoclypeidae.This paper presents a detailed study of the mor-phostructure of the four Mesogean orthophragminid ge-nera: the Discocyclinidae
Discocyclina
and
Nem-kovella,
and the Orbitoclypeidae
Orbitoclypeus
and
Asrerocyclina
using exceptionally well-preservedspecimens from the early Cuisian sites of Gan and Bos-darros (southern France). Following the morphos-tructural study, a revision of the systematics and a study
sf
the ontogeny, the functional morphology, themorphogenesis and biocalcification of the test, and thephylogeny and evolution of the group are undertaken.
Morphostructure
Three-dimensional models reflecting the lamellarconstruction, the arrangement of chambers and cham-berlets, and the structural elements (pores, stolons,apertures, "pillars", etc) were built for each genus(Fig. 1).
Lamellar construction:
One of the main results of themorphostnictural study was the identification of the tridi-mensional lamellar construction of the test in al1 four ge-nera studied. The model obtained differs from those des-cnbed in other foraminiferal groups. The test is built fromadditive chambers. Each chamber is delimited by an outerlamella, which forms a new annulus and which comple-tely covers the previous test. The annuli are subdivided bybox-like independent inner lamellae, one for eachequatorial rectangular chamberlet. On the lateral surfaces,the involute outer lamellae form lateral chamberlets,which are intemally covered by inner lamella.
Chamber and chamberlet:
Given these cha-racteristics of the lamellar construction, and taking thedefinition of chamber as al1 the structures formed in onegrowth step (Hottinger, 1978), in orthophragminids achamber is delimited by an outer lamella and comprisessevera1 geometrically separated elements: an
annulus
ofrectangular
equatorial chamberlets
plus a number of
lateral chamberlets,
scattered on the lateral surfaces.The term "chamber" must, thus, be used only formegalospheric embryonic chambers and rnicrosphencproloculus and initial spiral chambers.
Stolons and apertures:
The chambers andchamberlets are connected by cylindrical passages calledstolons. Four types of stolon were differentiated:
radial
 
lsolated chamberlets (orbitoidal growth)
Figure 1
-
Summary o€ he main morphostructural characters in Mesogean orthophragminids Orthophragminids comprise two families, Discocyclinidaeand Asterocyclinidae, that are differentiated according to their microspheric nepiont. Within the Discocyclinidae, Discocyclina is differentiated fromNemkovella by the presence of annular stolons in the former. The Orbitoclypeidae also lack annular stolons; only in Asterocyclina the annuli are lo-cally enlarged and subdivided into equatorial chamberlets axially superposed.
 
stolons
(those connecting equatorial chamberlets of adja-cent annuli),
annular stolons
(those connecting equa-torial chamberlets of the same annulus),
oblique stolons
(those connecting equatorial and lateral chamberlets) and
interlateral stolons
(those interconnecting lateral cham-berlets). The stolons are arranged in stolon systems. Theradial stolon system mns radially from the centre of thetest towards the penphery within the equatorial plane,and ends in the
equatorial apertures
in the periphery ofthe test. The annular stolons form annular circuits withineach annulus crossing the septula. The interlateral stolonsystem forms a three-dimensional network in the laterallayers, connected to the annuli by the oblique stolons,and opening to the exterior by the
lateral apertures
in thelateral surfaces of the test. The equatorial, oblique andlateral apertures become stolons when a new chamber isformed, thus extending the stolon systems.
Crystalline cones:
Anew term, crystalline cone is pro-posed, instead of "pillar" (which is also used in aggluti-nated and porcellaneous forarninifers), for the imperfo-rate conical structures formed by the superposition ofthickened imperforate sectors of the involute outer la-mellae. The crystalline cones originate as hemisphericalgranules on the lateral walls of embryonic chambers andequatorial and lateral chamberlets. The preferential for-mation on the equatorial chamberlets produces the con-centric arrangement of granules on the lateral surfacesfound in several orthophragminid species.
Embryonic chambers:
In al1 four genera the embryo isbilocular, with a subspherical protoconch and a kidney-shaped deuteroconch, of a size and shape characteristic foreach species. The two embryonic chambers are connectedby the
protoconchal stolon.
The deuteroconch is connec-ted to each equatorial chamberlet of the first annulus by a
deuteroconchal equatorial stolon,
and to the first lateralchamberlets by the
deuteroconchal lateral stolons.Microspheric forms:
Two models of ontogeny werecharactenzed (Fig.
l),
differentiated into three stages: spi-ral, transitional, and neanic. In Discocyclinidae, the cham-bers of the spiral stage are simple and do not form a thic-kened wall, while those of the transitional stage arefalciform and subdivided into chamberlets by septula ofinner lamella, becoming progressively larger until theyform a complete annulus. In the Orbitoclypeidae, the spi-ral chambers form a thickened spiral wall and have stellarstructure, with the ventral side subdivided into a mainchamberlet and a stellar chamberlet (Fig.
2),
whereas thetransitional stage follows the orbitoidal growth model.
Morphogenesis and biocalcification
A
model for the morphogenesis of the test isproposed. Each new chamber is formed from previousapertures. ~onse~uently,he shape of each new chamberand the number of chamberlets within it are determinedby the number and arrangement of apertures in theprevious test. In megalospheric forms, the first annulusafter the embryo is formed from the deuteroconchalequatorial apertures, and the subsequent new annuli fromthe equatonal apertures in the previous one. The lateralchamberlets are formed either from the deuteroconchallateral apertures in the embryo, from the oblique
Figure
2.-
Equatorial section
of
the microsphenc nepiont of Asterocyclina and Orbitoclypeus showing the ventral (A) and dorsal(B)halves. Each cham-ber is subdivided in its ventral half into a main chamberlet and a stellar chamberlet. This stellar structure indicates a relationship with theAsterigerinacea.
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