Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02)

Illustrated glossary of terms used in foraminiferal research

Lukas HOTTINGER1

Summary: An illustrated glossary of terms used in the analysis of the shells of recent and fossil foraminifera supplemented by a rigorous selection of terms that facilitate an understanding of their biology and their use in ecology and biostratigraphy. The glossary includes some 650 entries illustrated by 83 - often composite – figures many of which are stereographs or 3D models. A taxonomic index lists the 140 taxa illustrated. Key Words: Architecture, comparative anatomy, Foraminifera, hierarchy of terms, glossary, micropaleontology

Citation: HOTTINGER L. (2006).- Illustrated glossary of terms used in foraminiferal research.- Carnets de Géologie / Notebooks on Geology, Brest, Memoir 2006/02 (CG2006_M02) Abrégé : Glossaire illustré de termes utilisés dans la recherche sur les Foraminifères.- Ce glossaire des termes utilisé dans l’analyse de l’architecture des tests récents et fossiles de Foraminifères, complété par une sélection rigoureuse de ces termes, facilite la compréhension de leur biologie et leur utilisation en écologie et en biostratigraphie. Il comporte près de 650 entrées accompagnées par 83 figures, souvent composites, dont de nombreuses images stéréographiques ou modélisations en 3D. Un index taxonomique répertorie les 140 taxons illustrés. Mots-Clefs : Anatomie comparée, architecture, Foraminifères, glossaire, hérarchie des termes, micropaléontologie Ubersicht: Illustriertes Glossar der Fachbegriffe in der Foraminiferenforschung.- Ein Glossar von Begriffen zur Analyse der Architektur von rezenten und fossilen Foraminiferenschalen, erweitert mit einer strengen Auslese von Begriffen, die das Verständnis der Biologie sowie der Ökologie und der Biostratigraphie der Foraminiferen fördern. Das Glossar umfasst um die 650 Einträge welche mit 83 meist zusammengesetzten Figuren illustriert werden. Darunter sind viele Modellzeichnungen räumlicher Strukturen oder Bilder von Skulpturen der Schalenarchitektur. Die 140 abgebildeten Taxa werden in einem alphabetischen Verzeichnis aufgeführt. Stichwort: Foraminiferen, Glossar, Hierarchie der Begriffe, Mikropaläontologie, Schalenarchitektur, Vergleichende Anatomie

terminology cannot be avoided. Introduction Nature and significance of specialized Most distinctive morphological characters terms have been named through the coinage of Diagnoses and descriptive texts of taxa specialized terms. They may be based only on demand the use of a specialized terminology to geometrical features in the second or third embrace in a reasonably short and manageable dimension, or may include in their concept text the complex morphology of any taxon. biological processes such as growth or Pictures of the object described are necessary reproduction. For example: what is a to supplement its description in words. In many "chamber"? and what is a "chamberlet"? in cases however the "description" is restricted to radial foraminifera (DROOGER, 1993). the specification of diagnostic features, often According to a concept based on the geometry without an indication of their location in the of the cavities in the shell, neighboring lumina illustration or photograph. An ideal description in an annular cycle that are not in direct requires that a reasonable number of communication are called "chambers". A specimens per taxon be illustrated at the same concept including growth would consider that all scale and with the same orientation. This cavities in the annulus have been formed at the procedure helps the eye to distinguish, because same instant and would call them of the repetition of diagnostic characters, those "chamberlets". All cavities of the annulus would features that define the taxon from intraspecific form a chamberlet cycle and correspond to an variations. Desirable descriptive texts support undivided annular chamber. Thus, terms based this process by pointing out what to look for. on common concepts help to avoid comparing But no matter how well illustrated such a "apples and pears". They are the basis for the "description" may be, the use of an explicit comparative anatomy of foraminiferal shell

1 Museum of Natural History, CH 4001 Basel (Switzerland)

1 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02) structures introduced by DOUVILLÉ (1906). accuracy; differences in the methods used to prepare samples for examination (HOTTINGER, Accurate description of foraminiferal 1978). In some cases, the terms are based on architecture began in the second half of the 19th the geometry of shapes as they appear in a century (CARPENTER, PARKER & JONES, 1862) particular orientation of a section rather than on but the biological meaning of many their three-dimensional geometry. The morphological features of the foraminiferal shell employment of such terms may be misleading was discovered only in the second half of the and is not recommended. 20th century. Thereafter a considerable evolution in terminology has taken place. To The terms listed in the following Glossary day, the electronic treatment of data plays an are accompanied [in square brackets] by ever-increasing role in science. As yet, the alternative terms and (in parentheses) by terms machine is not able to recognize the context in considered to be synonymous, partially which these terms are used. synonymous, unclear or to be avoided for other reasons. Where in my own experience the In contrast to Linnean taxonomy, where the translation of a term into French or German is principle of priority contributes enormously to difficult, the translation is indicated in the stability of the taxonomic system, an (parentheses). The plural of terms coined in introduction of priority to morphological terms Latin, such as foramen, are indicated by: , pl. would block all progress in understanding the foramina. Biological terms are selectively complex morphology of the shells, their included in the number considered essential to morphogenesis and their biological, functional an understanding of the foraminifers and the and evolutionary meaning. However, wherever biological significance of their shell structure. it is possible and meaningful the use of the Some specific ecological terms are defined and most widely recognized term is recommended discussed to facilitate understanding of the in order to stabilize, as far as possible, and to increasingly abundant literature concerning the standardize diagnoses and descriptions. ecology and biogeography of foraminifera. There is a major parallelism in the terms Current discussions on the "Paleonet" list used to describe fusulinids of the Late Paleozoic [http://jerwood.nhm.ac.uk/archives/paleonet/] on one hand and of larger nonlamellar- show that the presentation of some basic terms imperforate benthics from the Mesozoic and used in taxonomy, biostratigraphy and Cenozoic on the other. This parallelism was evolutionary theory may be helpful. generated by the traditional specialization of The taxa of a generic or higher level used to the respective researchers; so far, very few portray the meaning of a term or to illustrate have been active in both fields. Although the the range of its application are from LOEBLICH concept of a suborder Fusulinina separating and TAPPAN (1987). For taxa that are not a most of the Late Paleozoic benthic foraminifera generotype, additional specific references are from the later clades is supported by the largest given in the text. Clicking in the alphabetic biological turnover in Earth History at the end index brings up illustrations of the taxon in of Permian times, there is no reason to use question that may help to verify an discrete morphological terms with regard to identification. major divisions of geological time, for throughout the Phanerozoic many The hierarchy of basic architectural characteristics of th shell are common to all elements in benthic foraminifera foraminifera. The present glossary may help to unify some of the parallel terms to a common Using basic concepts of biomineralisation usage. and wall construction (HOTTINGER, 1986; BENDER, 1989) that include the lamellar theory Current terminology used by various authors of SMOUT (1954) and HANSEN (1999), in describing the morphological features of the chamberwise growth, reproduction, foraminiferal test is far from being uniform and comparative anatomy (DOUVILLÉ, 1906; varies greatly from case to case (compare e.g. REICHEL, 1936-1937; HOTTINGER, 1978), and REISS, 1963; HOTTINGER, 1967, 1978; functional morphology including the demands of HOTTINGER et alii, 1993; LOEBLICH & TAPPAN, symbiosis with unicellular algae (HOTTINGER, 1964, 1987). The reasons for the coinage of 1997), a hierarchy of the most important several terms for the same feature are in most morphological terms can be formulated. On cases not at all formal, but due to one or more each hierarchical level, the terms for wall of several factors: differences of opinion with elements, cavities and connections between regard to the significance of a named element them are specific. They should not be used out as it relates to homologies and analogies; of their hierarchical context. differences in the degree of observational

2 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02)

1. Elements not modifying the shape of the living protoplast: 1.1. wall textures indicative of one of the several processes of biomineralization 1.1.1. agglutinated walls with noncalcareous cements binding the grains 1.1.2. intracellular biomineralization 1.1.2.1 calcareous cements binding agglutinated grains . 1.1.2.2 porcelaneous walls . 1.1.3. extracellular biomineralization: lamellar wall 1.2. wall textures produced by discrete arrangement and/or composition of constituents of the wall 1.2.1. wall polarity outside-inside 1.2.1.1 transition gradual, no layers: tectum, epiderm, pavement, endoskeletal . textures (in basal layers for example). 1.2.1.2 transition abrupt, wall layered, produced by a single or double-sided organic . template: inner and/or outer lamellae (lining) 1.2.2. differentiation of repetitive elements perpendicular to wall surface: parapores, pits, keriotheca, pseudokeriothecal textures, pores, pore chimneys. Opening of pores at wall surface: pore mouth; internal polygonal modification housing symbionts: eggholders. 1.3. ornaments produced by repetitive differentiation of external wall parts or layers: ribs, spines, pseudospines, spikes, pustules, beads, papillae, piles. 2. Elements modifying the shape of the living cell by chamber-wise growth: 2.1. incorporation of frontal shell walls including aperture(s) into the shell by adding a new compartment to the previous shell, a new chamber (with its aperture). First chamber: proloculus; second chamber (if not differentiated from later ones): deuteroloculus. Added cavity: chamber lumen; separating wall: septum; connection between successive chamber lumina: intercameral foramen, if tubular and multiple: stolon, if generated by resorption and wide open: tunnel. 2.2. subdivision of chamber lumen by deformation of chamber walls: fluting (including cuniculi) if frontal; retral lobes or processes if affecting the posterior chamber wall. 2.3. subdivision by circular invaginations of lateral parts of the chamber wall by calyces or similar structures in expanse chambers. 2.4. generation of multiple chamber lumina within a single growth step, enclosed by the same primary wall: chamberlet cycle. 3. Generation of shell shape (cylindrical, conical, globular, fusiform, lenticular) by chamber shape and arrangement: 3.1. serial disposition of chambers: uniserial, biserial, triserial. 3.2. spiral disposition of chambers: streptospiral, planispiral, trochospiral, multiple spirals. 3.3. milioline chamber arrangement. 3.4. by modification of chamber shape: chevron-shaped, involute, evolute, concentric (in two dimensions): annular, concentric (in three dimensions): spherical-concentric. 4. Elements modifying the shape of the protoplast in the chamber lumen: 4.1. elements independent of protoplasmic flux as revealed by patterns in subsequent foramenal disposition (arrangement of foraminal axes): exoskeleton. Cavities: compartments such as alveoles, subepidermal networks, alcoves; separating walls: beams (perpendicular to septum), rafters (parallel to septum); connecting open chamber lumen between exoskeletal cavities: passages. 4.2. elements reflecting protoplasmic flux by conforming to foramenal features

3 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02)

and patterns: endoskeleton. Cavities: compartments, if more or less closed: chamberlets; separating shell elements: if free standing and circular in section: pillar, if forming a wall perpendicular to primary chamber wall: septulum, if a free standing wall: hemiseptulum; if covering the previous septal face only: septal flap; if covering at the chamber bottom previous septal faces and/or surfaces of previous shell whorls: basal layer; connecting open chamber lumen between compartments: passages. 5. Shell cavitites produced by (lamellar) growth covering interlocular space: supplemental skeleton. Cavities: if tubular: canals, if more or less isometric: supplemental chamberlets 5.1 cavities produced by separation of the frontal wall of the previous and the proximal wall of the subsequent chamber: intraseptal interlocular space, may be subdivided into intraseptal canals 5.2 cavities produced by a space between subsequent whorls of a spiral array of chambers: spiral interlocular space, often restricted to a more or less tubular spiral canal. Opening of canals at shell surface: orifice. 5.3. cavities produced between subsequent whorls of spiral chambers and structures filling partially the umbilical space: umbilical interlocular spaces or canals. 5.4 cavities produced by infolds of outer lamellae over peripheral parts of spiral chambers or chamberlet cycles: marginal interlocular spaces (marginal cord, marginal crest, canalicular spines). 6. Branching canalicular cavities running obliquely through subsequent, regularly perforate outer lamellae: trabeculae. 7. Embryonic apparatus different in architecture and/or structure as compared to adult parts of the shell: 7.1. first or first two or three septa straight, uncurved, different from the adult: bi-, tri- or quadriconchs. 7.2. tubular extension of protoconch foramen (flexostyle) eventually feeding a particular deuteroconch with multiple apertures (vestibule). 7.3. concentric deuteroconch enclosing an often poorly calcified protoconch with an exoskeleton: sphaeroconch. 7.4. hemispheric deuteroconch with exoskeleton (supraembryonic chamber) feeding a cycle of nepionic chamberlets with exoskeleton only (subembryonic chamber). 7.5. multilamellar envelope of embryo enclosing several chambers and feeding a particular cycle of first (nepionic) chamberlets (corona, with auxiliary and adauxiliary chamberlets).

4 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02)

also: brood chambers, Fig. 24. Glossary A agglutinated - shell texture characterized by components gathered in the ambient A - form – see gamont, schizont. environment and bound by organic or biomineralized cements produced by the cell. abaxial - directed away from or far from Particles may be selected according to size and the shell axis. shape to form a closefitting mosaic. Often, the aboral – directed away from or in a agglutination in the external and internal parts position opposite to the apertural [oral] end of of the wall is differentiated but there are no shell. sharp boundaries (see also wall polarity). Fig. 6. accessory aperture (infralaminal aperture, auct.) - aperture that does not lead alar prolongation - winglike extension of directly into the main chamber lumen, but umbilico-lateral portions of involute chambers extends beneath accessory structures (e.g. in on lateral surfaces of previous whorls in bullae). Fig. 1. lenticular tests. May be meandering. Fig. 7. acervuline - chambers in irregular, alcove - a blind compartment of a chamber clustered arrangement, as in Acervulina. lumen delimited by beams and the lateral chamber wall, as in Orbitopsella. Fig. 72. acicular - needle-shaped. allopatric speciation - emergence of acicular spine - see spine. species by geographic isolation of populations actin – cytoplasmic protein producing in its for a time sufficient to alter the genome of the polymerized form short, contractile fibers. If populations involved to a mutual reproductive positioned below the cell wall, they may form incompatibility. stellar arrays. alternating arrangement of shell acuminate - tapering, i.e. getting thinner compartments – Shell compartments or pointed; conical. (chambers or chamberlets) of successive growth stages arranged on alternating radii of acute - shape with acute or sharp angles. the shell. Fig. 8. adapertural depression (periapertural Remarks:. The alternating arrangement of the depression; cavity; apertural fissure, sulcus, shell cavities is a very fundamental and auct.) - space formed by a toothplate that widespread pattern of the foraminiferal separates it partly or completely from the main architecture. It occurs in all three dimensions: chamber lumen. Interconnected adapertural the first, linear dimension (= biserial), the depressions produce a canal. Fig. 2. second, planar dimension, in annular- concentric shells, in particular in the main adauxiliary chamberlet - chamberlet layers of orbitoidiform groups, and in the third, arising from a single radial stolon (additional spatial dimension, in shells with spherical with respect to the apertures between proto- concentric growth, where it forms chessboard and deuteroconch) in the embryonic wall of patterns at the surface of the globular shells. orbitoidiform shells. Fig. 4. alternation of generations - see life adaxial - directed towards or positioned cycle. Fig. 5. near the shell axis. alveolar layer - layer of alveoles in lateral adventitious - produced by or with the chamber walls forming diagnostic exoskeletal help of foreign particles such as those forming structures lacking a differentiation into beams an agglutinated test. and rafters as well as polygonal subepidermal advolute chamber arrangement - in those patterns; distinct from subepidermal, spirally coiled forms where the chamber polygonal networks or keriothecal textures. lumina of a whorl cover laterally those of the See exoskeleton (Fig. 45) and keriotheca (Fig. preceding whorl to a considerable extent, but 62). not entirely, on one or both sides. Fig. 3. alveole (alveolus, pl. alveoli) - a recess of Compare: evolute, varying depth in lateral walls, coated by the agamogony - asexual reproduction within organic lining, blindly ending with a rounded the reproductive cycle, from the first mitotic contour below an epiderm or some equivalent division of the zygote to meiosis. Fig. 5. outer layer of the wall and opening into the chamber lumen. May branch towards the outer agamont - specimen grown from the part of the wall, each generation of branches zygote, producing either gamonts or schizonts forming layers within the wall. Fig. 9. in an asexual process involving apogamous Remarks: The term alveole is used here nuclear divisions and/or meiosis. Foraminiferal exclusively for exoskeletal structures, i.e. for agamonts, produced by a sexual reproduction, subepidermal, tiny compartments of the are called microspheric (B - form). Fig. 5. See chamber cavity coated by an organic lining.

5 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02)

Alveoles are supposedly filled with chamber system" as in advanced Eulepidina). See also: plasm. Alveoles must be distinguished from preseptal passage. paraporous or parakeriothecal cavities that are annulus - ring-shaped chamber, which an adjunct of wall texture. Like true pores, may be subdivided, or ring-shaped cycle of these cavities are (by definititon) neither filled chamberlets. See chamber arrangement, Fig. with living chamber plasm nor coated by the 37. organic lining. The simultaneous presence of both alveolar structures and keriothecal wall anterior - directed to or positioned near or texture in the fusulinid Verbeekina and its on frontal part of chamber, usually including relatives, the combination of an alveolar the main aperture, distal in respect to direction exoskeleton with a paraporous external wall in of growth. Dicyclina or with a bilamellar perforate wall in Fabiania supports a consequent restriction of antetheca - apertural face in fusulinids. the term's use to exoskeletal structures, never apertural axis – the shell axis defined by to textures. a placement of foramina in a single line. See Layers of alveoles coating the lateral chamber milioline coiling, Fig. 68. wall are present in various agglutinated groups of which Cyclammina is the most prominent apertural chamberlet – the cavity in while Everticyclammina is an early extinct preseptal position below the radiate aperture representative of an exoskeletal layer of in Lenticulina and related forms. The lamellar exclusively undivided, shallow alveoles. The nature of the wall between the main chamber Neogene group of Textulariella CUSHMAN and the apertural chamberlet remains unclear (GRÖNHAGEN & LUTERBACHER, 1966) has at present. branching alveoles. Among the porcelaneous apertural face – that surface of the foraminifera with alveoles, Austrotrillina is a chamber-wall that contains the main cameral prominent group. In this genus, species with aperture. See also face, septal face, umbilical deep and branching alveoles - A. howchini face, Fig. 48. (SCHLUMBERGER) - are said to evolve from earlier forms with layers of shallow, undivided apertural flange - see lip. alveoles (ADAMS, 1968). The term alveole is apertural lip - see lip. also used for rows of blind recesses in postseptal position over supplementary apertural plate [basal plate] - a plate-like apertures in the previous septal face as in structure that restricts the base of an Subalveolina or Bullalveolina. We do not yet interiomarginal aperture and restricting the know, how to interpret (in terms of exo- or latter. Fig. 10. endoskeletal elements) these alveoles nor apertural tooth - see milioline tooth and what might be their biological meaning. valvular tooth. analogous – similarity in function and, aperture – the primary opening of the where shape is functional, also in shape, of foraminiferal shell cavity towards the ambient independent ontogenetic origin and hence of environment. May be covered by subsequent different evolutionary origin. See also chambers and thus transformed into a homologous, homomorphous. foramen. May be masked (see mask). May be annular arrangement - arrangement of single or multiple. Fig. 48 (See also cameral concentric annular chambers. aperture; foliar aperture; supplementary aperture; labial aperture). annular canal - free space in preseptal Remarks: We insist here on restricting the use position between endo- and exoskeletal of the term "aperture" in chambered shells to structures in spirocyclinids. the ultimate opening of the last chamber Remarks: Unrelated to canal systems in cavity into the ambient environment. When a lamellar perforate foraminifera. The term new chamber is added to the previous one as should not be used but replaced by annular a main process of growth, the aperture is passage. transformed into a means of communication annular chamber - ring-shaped chamber. between successive chamber lumina and thus May be subdivided as in Cycloclypeus. its function changes. The transformed aperture is called a (intercameral) foramen. Its annular passage [annular stolon] - morphology may change through selective annular, open space in preseptal position, may resorption of its margins. Often, the last be single, in the equatorial plane, as in chamber of the shell is not preserved because Sorites, or double, in lateral position, as in the of its thin walls (see lamellation) and the archaiasines. In orbitoidiform foraminifera the aperture is lost. adjacent lumina of a chamberlet cycle may be connected by a pair of tubiform lateral apex - initial portion of trochospiral or passages in postseptal position (lateral annular conical test. stolons, as in Discocyclina, Fig. 41 A) or by apical - referring to the initial part of a several layers of annular stolons ("six-stolon

6 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02) trochospiral or conical test. foraminifers. May be double in embryos with a bilateral symmetry or multiple, when the apogamous – reproducing according to the protoconch and the deuteroconch have mode of apogamy. additional equatorial stolons. Together with the apogamy – the process of reproduction adauxiliary chamberlets, they form a corona. where the offspring has the same number of Fig. 4; Fig. 36. chromosomes as the parent cell. auxiliary tunnel – the coalescence of arborescent – in permanently attached several subsequent cuniculi as in shells, a tree-like, branching growth pattern. Polydiexodina. Fig. 11. axial filling - secondary deposits in narrow areal - positioned within the apertural face, spaces around the axial columella in fusulinids neither at its base nor at the shell margin. and pfenderinids. areal aperture - cameral aperture in a axial plate – see umbilical plate, cover distal wall, not at a suture. May be single or plate. multiple. Fig. 12. Remarks: This term, introduced for Ammonia by CIFELLI (1962), was substituted arenaceous - see agglutinated. erroneously by PARVATI (1971) by "umbilical areolate – the chamber wall is subdivided plate" but in fact is a cover plate. Continued into more or less equal surfaces (areoli) as in use of the term "axial plate" would add to the Homotrema. Fig. 11. considerable confusion in the current concepts of rotaliid umbilical architecture (see also LÉVY asexual reproduction - a mode of et alii, 1986). reproduction where the genome of a single individual is duplicated. axial section – a slice bisecting the test in a plane coinciding with the axis of coiling and astral fissure - see folium, foliar aperture, intersecting the proloculus. Compare foliar suture. equatorial section. Fig. 15. astral furrow - see foliar suture. axial septulum - an exoskeletal structural astral lobe - see folium. element parallel to the septum in verbeekinid Remarks: CARPENTER et alii (1862), BRADY fusulinids and consequently corresponding to (1884) and in particular DAVIES (1932) in his the rafter in the exoskeletons of other basic monograph on Rotalia trochidiformis agglutinated and porcelaneous foraminifera. used the term astral lobe (Fig. 13) to axis of coiling – an imaginary line around designate what is currently called a folium. In which a spiral test is coiled. Fig. 7 & Fig. 68. spite of their similar ventral-adumbilical See also chamber arrangement. position astral lobes must be distinguished from stellar chamberlets (as in B Amphisteginidae) that are separated from their B - form – see agamont. corresponding main chamber by a complete wall. However, functional similarities may not balloon chamber – a hemi- to sub- be excluded. spherical chamber surrounding a float- chamber in the pseudoplanktic stage of some attachment – the permanent fixation of a benthic foraminifera; provided with multiple shell onto its substrate. See also surface of openings for extrusion of gametes. Fig. 16. attachment, arborescent growth, encrusting growth, Fig. 60. basal - at the base or parallel and proximate to the base of a structural element attics (French: mansardes) – in or of a complete architecture. porcelaneous shells the outermost lateral or Remarks:. The term is occasionally used for abaxial layer of chamberlets in a multi-layered interiomarginal apertures or foramina at the endoskeleton, distinguished from less lateral base of the apertural face adjacent to the or adaxial ones by the comparatively small previous whorl (in contrast to areal) or for the caliber of the tubiform chamberlets, as in base of a cone in conical shells including Alveolinella. Fig. 14. sections in or parallel to the base of the cone. autogamy – a process of sexual Fig. 80 B. See also basal layer. reproduction where amoeboid gametes from basal disc - the thickened but porous the same gamont mate (in foraminifera within extension of the median layer in the proximal the mother shell) to form a zygote. May be pore cavity that is part of the pore plug. Fig. combined with ogamy. 75 A-B. auxiliary chamber [or auxiliary basal flap – in milioline forms an chamberlet] – a chamber or a chamberlet fed interiomarginal, spathulate, more or less by a stolon positioned in the suture between excavated infold of the distal wall projecting protoconch and deuteroconch in orbitoidiform into the aperture and restricting it. Fig. 17.

7 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02)

endoskeletal elements such as septula, Remarks: The basal flap in miliolids seems to particularly in verbeekinids, orbitolinids and be an extension of the basal layer detached cuneolinids. May occur as unique exoskeletal from the previous whorl and extending into the element (Orbitopsella, Fig. 72) or in large aperture. It may be a homologue of the combination with minor, shorter exoskeletal milioline tooth. elements producing a subepidermal polygonal network (Spirocyclinidae, Fig. 45). basal layer - in imperforate foraminifera: Remarks: DAVIES (1930) distinguished the parts of a chamber wall coating the "primary" or "major" (1939) partitions in the previous coil and/or the septal face of the description of orbitolinids in opposition to previous chamber. As they are not in contact "secondary" or "minor" elements. HENSON with the ambient environment, these portions (1948) separated "subepidermal plates" from of the wall lack the differentiations in the main partitions (the latter being of texture of the outer wall. Basal layers endoskeletal nature and therefore to be called represent or are part of the endoskeleton, and septula nowadays; compare HENSON's fig. 7, may be sculptured by ribs in the chamber reproduced here as Fig. 19). HENSON's (1948) lumen or form endoskeletal elements reaching subepidermal plates, synonymous with the ceiling of the chamber, i.e. pillars or SILVESTRI's "trabecole perpendicolari" (1932), septula. Fig. 18. were subdivided again into "transverse" and Remarks: The basal layer is a very important "parallel" partitions corresponding to DAVIES' element of the architecture of imperforate major and minor elements. HENSON's general foraminifera. In spiral-involute and miliolid term "subepidermal partition" may include shells it may become much thicker than the main partitions (= septula), transverse external chamber wall (flosculinisation); in partitions (= beams) and parallel partitions (= fusiform shells it may form a double columella rafters), where septula and beams fuse to reaching both poles of coiling. Thickened basal produce (mostly radial) chamber layers often have numerous tubular passages compartments. In order to distinguish of irregular shape connecting successive partitions according to their origin, HOTTINGER chamber lumina, with (edomiids) or without (1967) introduced particular terms for (elongate alveolinids) intermediate preseptal partitions exclusively of exoskeletal origin, i.e. spaces. In our view the basal layer is "poutre" for major and "poutrelle" for minor homologous with the "basal skeleton" of partitions, translated into English in 1978 as fusulinids (chomata and derived structural "beam" and "rafter" (Balken and Bälkchen in elements). Where tubular chamberlets are German). For illustration see exoskeleton. vertically superposed, the basal layer may form regular floors parallel to the chamber biconcave - test having both sides concave roof, as in elongate Praealveolina. (in coiled forms). basal lobe - a finger-like extension of a biconch - protoconch and deuteroconch chamber wall at the spiral or septal suture (in together, if separated by a straight septum, the absence of an interlocular space). (See thus differing in shape from later, curved ones. retral lobe and compare ponticulus). The straight septal wall suggests that hydrostatic pressure in protoconch and basal skeleton - see chomata. deuteroconch was equal as a morphogenetic basement – cavities below a main control before the wall was calcified. Thus, the chamberlet layer, developing polewards in two first chambers were formed together and elongate alveolinids, delimited by floors and represent a single growth stage similar to an septula, opening into a preseptal space and embryonic apparatus. connected vertically by shafts, as in Remarks: The status of a biconch as a Praealveolina. Fig. 70. particular growth stage is supported by RÖTTGER's (1974) observation that biconchs in bead – a small, rounded to hemispherical nummulitids are formed after hatching, i.e. protuberance on the surface of lamellar shells, after the naked embryo is squeezed out of the forming strings along septal, septular or narrow canal orifices. In such biconchs, the hemiseptular sutures. Usually imperforate or second chamber is used to keep the few poorly perforate. To be distinguished from symbionts transmitted from the mother shell papilla. Fig. 73. in their first stage of procreation within the beading – strings of beads along linear new generation of their hosts. Other embryos shell elements such as sutures. To be calcify their shell with straight septa beween distinguished from costellae. the first two, three or even four chambers within the mother shell and will leave it during beam – an exoskeletal main partition of the hatching process by dissolving chamber the chamber lumen, perpendicular to the walls of their mother. chamber septum and to the lateral chamber wall. In discoidal shells often separated from biconvex - test having both sides convex an endoskeleton by an empty space in the (in coiled forms). chamber (annular passage). May fuse with

8 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02)

bifid - divided into two branches. See also whorl and covering primary, main or milioline teeth. supplementary apertures. May have marginal accessory apertures. Present in planktic biformed - change in chamber foraminifers only. Fig. 1. arrangement during ontogeny, for example from coiled to uncoiled or triserial to bi- or buttress - see pillar. uniserial. C bilamellar wall - in perforate foraminifera a chamber wall formed primarily by two calyx, pl. calyces (pillar-pore) – a funnel- mineralized layers (outer and inner lamellae) shaped, perforate invagination of lateral wall on either side of a primary organic sheet, the supporting expanse chambers over a wide area median layer. See also lamellar wall, outer as in Miniacina. Fig. 25. lamella, inner lamella. cameral [chamber] aperture - single or bilateral - having two equal sides by multiple opening in a chamber-wall that allow mirror symmetry. the communication of a main chamber lumen with the ambient environment. It may later be bilocular - said of an embryonic apparatus converted partially or entirely into an having two chambers differing in size and intercameral foramen, unless that foramen has shape from the following ones. been formed secondarily by resorption. The position of apertures on the apertural face biloculine - see milioline coiling. Fig. 68. (interiomarginal, marginal, areal, umbilical, bipartitor – a bridge-like structure, extraumbilical, terminal) and persistent extending posteriorly from an umbilical plate, patterns in multiple apertures (see crossing the preceding aperture and attached endoskeleton) are of fundamental taxonomic to the adjacent coil, thus cutting off the value. See also supplementary aperture; foliar intercameral foramen from the opening into a aperture, tunnel, mask. Fig. 48. primary umbilical canal. Fig. 21. canal systems - a term collectively and biserial - trochospiral chamber broadly applied to those interconnected spaces arrangement with about 180° between of the foraminiferal test, that are primarily or consecutive chambers, thus producing two secondarily separated from the main chamber rows of chambers. See serial disposition of lumina, but with which they may communicate chambers. Fig. 37.6-7. in one or in successive whorls by openings other than intercameral foramina, the so- biumbilicate - spiral test having umbilici called loop-holes. Canal-systems contain on both sides. Fig. 22. functional microtubular ectoplasm and biumbonate - having an umbo on both represent bypasses of spiral main chamber sides of the test. lumina so that ectoplasm deep inside the early parts of the test connects directly with blades - plate-like, strongly protruding, extrathalamous rhizopodial ectoplasm. The short or long costae. Fig. 23 A. spaces forming canal-systems are delimited by blueprinting – a morphogenetic process: a number of discrete elements of the test: repetitive induction of shape by preexisting umbilical plate, cover plate joined to a shapes, such as linear angularities or foramenal plate, sealing plate, toothplate, shoulders on a previous chamber that induce septal flap, folia, previous coil, as well as by the position of the suture of the following consecutive outer lamellae. See also: primary chamber, or of areal reliefs by protrudent pile and secondary umbilical spiral canals; heads on which subsequent lamellae are intraseptal interlocular spaces; grooves; thickened. enveloping canals; funnels; cord, crest. Fig. 26. boss – see umbo. Compare: plug; pile. canalicular structure – a structural brood chamber - chamber(s) or pattern produced by repeated modes of canal chamberlet cycle(s) with enlarged cavities that disposition in the foraminiferal shell. See also: house the offspring before hatching. The canal system. enlarged chamber cavities may be produced by partial resorption of shell material, in canaliculate crest - see crest. particular of the endoskeleton and the septa. canaliculate [canaliferous] spine – a To date, brood chambers have been observed spine- or club-shaped to arborescent radial exclusively in agamonts (microspheric structure composed of consecutive outer specimens). Fig. 24. lamellae enclosing canals (supplemental buccal aperture - see funnel. skeleton). May contain spikes. Compare pseudospine; spine. Fig. 27. buccal ring - see chomata. cancellate - having honeycomb-like bulla – a blister-like test element surface ridges as ornament. Fig. 28. extending over the umbilicus of the ultimate

9 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02)

carina [keel] – a peripheral thickening of annular passages. However, they are always the shell. May be doubled in Cretaceous connected indirectly by the retrovert stolons planktic foraminifera. In bilamellar feeding the lateral chamberlets. Fig. 36. foraminifera formed exclusively by the outer chessboard pattern – a chamberlet lamella and therefore imperforate. Fig. 29. arrangement in superposed chamberlet layers cell envelope in foraminifera - all in alternating positions resembling those of a elements covering the living cell body of a chessboard: in one layer, the position of the foraminifer delimited by its plasmalemma. chamberlets corresponds to the black fields, in the following layer to the white fields, as in cell wall – biomineralized cover of living Sphaerogypsina. The outline of the cell. chamberlets may be quadratic, polygonal or central complex – see reticular zone. somewhat irregular. This three-dimensional Adaxial cone in conical agglutinated forms array corresponds to the annular chamberlet where radial septula are fused to an irregular arrangement, that is alternating radial pattern of meshes. In contrast to radial zone positions in the second dimension. Fig. 8 F-H. and marginal zone. Fig. 20 & Fig. 71. chevron shape – the inverse of V- shaped. chamber [loculus] – the space and its See also: equitant. enclosing biomineralized walls formed at one chirality – the proportion of right-hand to instar, i.e. during a single step of nepionic and left-hand coiling in a population of trochospiral ephebic growth. See also: chamber lumen, organisms, as seen from the spiral (dorsal) chamberlet. side. See dextral and sinistral coilings. chamber arrangement – the pattern of chloroplast (plastid) – the organelle of a disposition of chambers in a shell. Fig. 37. plant cell (in foraminifera endosymbiontic chamber lobe - see folium. algae) that are responsible for photosynthesis, with ultrastructures (thylacoids, pyrenoid) of chamber (or chamberlet) lumen – a shell different pigmentation diagnostic for several cavity filled with protoplasm (usually of an algal groups. Fig. 30. endoplasmic nature except in newly formed chambers), coated by an organic lining, the chloroplast husbandry – a particular primary cell envelope. By definition, chamber symbiotic relationship in which chloroplasts of lumina communicate exclusively through ingested algal food cells are kept in the host intercameral foramina and/or multiple stolon protoplasm in a state of partial functionality. systems. The term "chamber lumen" is used in Fig. 30. particular to distinguish between the "inside" choma (pl. chomata; German: Basalreifen) and "outside" of a shell, i.e. between intra- – a dense deposit on previous whorl and interlocular spaces. The latter are never constituting the chamber floor, and in papered with an organic lining. Within the fusulinids forming a pair of parallel ridges, boundaries of intraspecific variability the each extending from a tunnel margin to the volume of chamber lumina may depend on the previous one. May extend progressively seasonal availability of food and be polewards over the entire chamber floor in independent of wall thickness which in lamellar staffellids, much like a basal layer in fusiform foraminifera is dependent on water turbulence. porcelaneous shells. Fig. 31. chamber passage - see passage. chomatal pores – porelike texture of chamber wall(s) - skeletal elements chomata in higher fusulinids as continuation of formed at one and the same instar, enclosing pore-like textural elements in the keriotheca or the corresponding chamber and coated by an parakeriotheca below the tectum of the inner organic lining. previous whorl. Fig. 66. chamberlets - segments or subdivisions of circumproloculus – a deuteroconch that a chamber, produced during the same instar. in some advanced fusulinids envelops the See also: subsidiary chamberlets; cycle of megalosphere to a large extent. chamberlets; foliar chamberlet; lateral clavate - club-shaped. chamberlets; stellar chamberlet. climax – end of an ecological succession, chamberlet cycles – the chamberlets when an equilibrium with the long term forming an annulus within one instar. The ecological conditions is reached. annuli may produce a single layer thus forming a disc, as in Planorbulinella, or in a lenticular closing chamber – a symmetric shell be comprised of a main chamberlet layer interauxiliary chamber. See chamber with lateral layers of chamberlets or lateral arrangement in orbitoidiform foraminifera. Fig. expanse chambers, as in orbitoidiform 36. architecture. The chamberlets of an coalescence – the fusion of separate, orbitoidiform that cycle in alternating radial elongated parts. positions may or may not be connected by

10 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02)

coenocline (community gradient) – the when describing the relation between the distribution of populations of species in diameter of the embryonic apparatus and the ecological gradients. number of chamberlets in the immediately succeeding growth stage. See also: cohort – a demoecological term for a periembryonic chamberlets. group of individuals conditioned in their temporal development by a common event in cosmopolitan - Occurring all over the the ambient environment. In foraminifera: the world where ever there is a suitable habitat, individuals produced by a more or less the contrary of endemic. synchronous hatching triggered by seasonal costae - raised ribs or ridges on test change, and without reference to their mode of surface. See also: striae, blades. Fig. 23 B. origin, i.e. agamonts or schizonts which differ greatly in length of life. costate - having costae. coil - see whorl. costellae – a particular ornament on the chamber wall produced by more or less coiling axis - see axis of coiling. elongate ridges formed by an aligned fusion of columella – the solid, trochospiral pustules produced by secondary lamellation. structure formed by the basal walls of spiral chambers coalescing around the coiling axis, Remarks: The term was introduced to describe as in many gastropod shells, or symmetrically the particular ornament of Costellagerina, a poleward in planispiral - fusiform shells. May rugoglobigerinid from the late Cretaceous. It be thickened by secondary deposits or by an might be useful to extend the term as a extensive thickening of a basal layer in the description of the radial ornamentation of the polar realm of the shell (alveolinids). Fig. 32. ventral chamber walls in smaller benthics indicating ogamy (pairing of two gamonts compartment – the lumen of the shell venter against venter), as in Glabratellina. Fig. housing chamber protoplasm, communicating 23 C-D. with other lumina in the shell by constricted passage-ways. Supposedly functions as a cotype – see type. reactor receptacle for cell metabolism. A counterseptum – a kind of lower lip of an chamber may constitute a compartment or be interiomarginal - basal aperture appearing in subdivided into chamberlets that are units of a appropriate sections as a forward directed compartment. Compartments may have hook below the foramen and glued to the specific shapes. Tubular and isometric shapes previous shell whorl, as in Eulinderina do not occur together in the same taxon and guayabalensis (NUTTALL) or Amphistegina may reflect different types of bioreactions. The lopeztrigoi PALMER (BUTTERLIN, 1987). The volume of the compartment remains hook is said to develop into "complete septal comparatively stable throughout growth but walls" in Helicostegina. Fig. 33. may be modified by the availability of food Remarks: The term was introduced by BARKER during the seasonal cycle. On the other hand, and GRIMSDALE (1936) for proximal portions the number of compartments produced during of the main chamber septum below the main one growth step may increase during foramen in A. lopeztrigoi. The hook may be ontogeny. Narrow passage-ways between identical with the gutter in Old World compartments, stolons or foramina, may be amphisteginids. The nature of the "complete", temporarily closed by organic plugs, thus helicosteginid countersepta remains to be isolating the compartment's protoplasm from analysed the more so as the relation between other compartments and in particular from the "countersepta" and stellar septula in the New one containing the nucleus. World amphisteginids is not clear. congeneric - belonging to the same genus. cover plate [umbilical cover plate] conspecific - belonging to the same (retroparies, pars auct.) - a more or less species. folded, imperforate extension of the septal flap into the preceding chamber through the contrefort – a lateral thickening of the intercameral foramen, separating - in the inner wall layer against the base of the preceding chamber - the main chamber lumen chamber lumen that thins or vanishes in the from a foliar chamberlet. Usually attached to a equatorial plane, as in some archaediscids. preceding foramenal plate. It is a secondary May be homologous with basal layer. feature, never present in the ultimate chamber convolute - see evolute. and thus not homologous with a primary foramenal plate or umbilical plate. Compare corona - first cycle of chamberlets also: sealing plate. Fig. 34. enveloping an embryonal apparatus completely, at least in one plane of cribrate - perforated by multiple holes. sectionning, as in Discocyclina. Fig. 41 A. Should be used exclusively for numerous and small multiple apertures. See also Remarks: The term may be useful in biometry trematophore.

11 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02)

cruciform - cross-shaped (aperture). tendency to produce a more or less complete fusion of the pillars into a median shell wall. crystalline cone – see pile of lamellae; Fig. 47 F. compare papilla. cyst – a temporary cover enveloping part cubiculum (pl. cubicula) – a lateral shell of the shell, the whole shell or several cavitiy in spiral and annular-concentric shells individuals, consisting of adventitious material with a main chamber- or chamberlet- layer. bound together by organic material or slime,

as a protection of chamber building or of Remarks: This term, introduced by BANNER reproduction processes. Accumulations of and HODGKINSON (1991) and exemplified by refused food particles in front of the face at Spiroclypeus, "orbitoid, miogypsine and other, the margin of discoidal shells are called three-layered rotalines" is a substitute for feeding cysts. what is traditionally called lateral chamberlets in larger foraminifera (compare FERRANDEZ, cytoplasm - protoplasm, excluding the 1999). Apparently, BANNER and HODGKINSON nucleus. mean by "true" lateral chamberlets in heterosteginids their alar prolongations D subdivided into chamberlets. They show them dendritic - see arborescent. only in axial or transverse sections where the distinction between septal and secondary deuteroconch – the second chamber in an subdivision of shell cavitiesis is difficult to embryonic apparatus. See also: sphaeroconch. carry out. I suspect these "lateral chamberlet" Fig. 38. to be undivided alar prolongations cut at an deuteroloculus – the second chamber in a oblique angle in respect to the median line of shell without embryonic differentiation, i.e. the alae. See also: lateral chamberlet, lacking an embryonic apparatus. supplemental chamberlet. dextral coiling – a clockwise direction of cuneate, cuneiform - wedge-shaped, as coiling, as seen from the spiral side. in Cuneolina. diaphanotheca – a light-colored or cuniculus (pl. cuniculi; cunicular passage) translucent wall layer immediately below the - in fusulinids a transverse passage at the tectum in fusulinids. Fig. 39. chamber base, parallel to the tunnel, connecting the lumina of alternating chambers. dimorphism - coexistence of two discrete It is produced by the fusion of opposed folds in morphotypes representing different adjacent septa that lose contact with the generations in the life cycle of a single species. previous whorl at their point of fusion. The They are expressed in the adult growth stages geometry of the apertural face in cunicular and/or in the protoconch and in the following fusulinid shells remains unclear. Fig. 35. nepionic chambers. The protoconch diameter is large (megalospheric = A - form) when the cupule – radial, rounded ridges on the agamont's protoplasm is distributed (including shell face of conical shells resulting from an eventual symbionts) among the cloned arrangement of slightly inflated radial offspring. The protoconch diameter is small chamberlets alternating in radial position in (microspheric = B - form) when the gamont's subsequent cycles. gametes fuse pairwise to form a zygote whith Remarks: the term "cupule" means "cup" in no protoplasm or symbionts from the mother. French and English and refers to the cup- If there is dimorphism in the adult shell, the B shaped sections perpendicular to an elongate - form becomes larger than the A - form. The barrel vault forming the radial chamberlets as compartments of the microspheric initial in Orbitolinopsis, Orbitolinella or Halkyardia. phases are small. It takes many growth steps cycle of chamberlets - multiple segments to reach the initial shell size of the of a main chamber lumen, produced at one megalospheric generation. Reaching the adult and the same instar. Fig. 36. oversize of the microspheric generation demands numerous additional instars. cyclical arrangement - arrangement of Consequently, the dimorphism of foraminiferal cyclical chamberlets in one plane or in generations reflects different life times and concentric layers. Fig. 8 C-H & Fig. 37. thus different strategies of life within the same cyclical chamberlet - one of the cyclical species: the microspheric generation is segments of the main chamber-lumen adapted to the permanent basic low-level produced at one and the same instar. Fig. 36. carrying capacity of an oligotrophic and warm environment, while the megalospheric cyclopsinellid structure - arrangement of generation with its short life time adapts to endoskeletal elements as in Cyclopsinella: the both spring and eventual autumn seasonal arrangement of endoskeletal pillars following peaks of carrying capacity. See also: odd the pattern of the radial stolon axes pairs. Nepionts with large megalospheres may superposed in a radial position in neighbouring have a particular architecture different from stolon planes. This particular pattern has a that of the adult (see megalospheric

12 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02) apparatus). In complex life cycles, a third without spinose projections at the junction (schizontic) generation may produce with adjacent pores. Harbours symbionts in megalospheric shells with a morphology order to enhance the gas exchange of the slightly different from that of the gamonts. See symbiont with the ambient environment of the also: trimorphism. Fig. 5 & Fig. 70. host. Fig. 30 and Fig. 40. diploid – a genome with paired elongation index - in alveolinids and chromosomes, one coming from the father, other fusiform or otherwise elongate shells, one from the mother. See also: haploid. the ratio of the length of the axis of coiling to the diameter at the equator. Fig. 78. disclimax – an ecological succession truncated by periodical ecological events, as embryo(n) [embryonic apparatus] – the for instance the destructive turnover of hard group of chambers including protoconch and substrates by winter storms. deuteroconch (nucleoconch), and in some genera a flexostyle, that differs in size, shape distal - farthest from the proloculus in and arrangement from subsequent chambers. direction of growth. Fig. 41 A. distal chamber wall - farthest, in embryonic – the earliest growth stage in direction of growth, chamber wall. foraminiferal ontogeny, usually distinguished distal face - outer surface of distal from later stages by an abrupt change in shell chamber-wall. See also: face. architecture, commonly with thickened walls indicating a longer period of cessation in distinctly radial texture – the appearance growth as frequent in the megalospheric under crossed nicols of test wall fragment generation of larger K-strategists (bi- and producing a dark cross of interference and triconchs; embryonic apparatus). concentric colour rings. Corresponds to fibrous ultrastructure. embryonic apparatus - see embryo(n). dorsal - the side of a free, flattened embryonic pseudochamber organism turned away from its substrate, as ("Zwischenkammer" in GRELL, 1954) – a opposed to ventral. See also: spiral side; hemispherical cavity between the umbilical side. deuteroconch and a third chamber seen on the Remarks: REVETS (1994) rightly points out dorsal side of the shell as hemicircular that the terms dorsal and ventral, beyond their protrusion. specific meaning in the architecture of Remarks: The "true", i.e. lamellar nature of vertebrates, refer to the orientation of a this cavity and its connection with the regular flattened organism in respect to its substrate. spiral chambers seen in Rotaliellidae (see also In foraminifera, benthic, free shells live with PAVLOWSKI et alii, 1992) is not yet known. their apertural face on their substrate in order encrusting growth – the mode of growth to gather food. Permanently attached shells, of permanently attached shells covering large usually filter feeders, are living with their surfaces by keeping low and close to it, apertures turned away from the substrate. accelerating the occupation of a surface by They must have free apertures in order to add forming annular expanse chambers (as in new chambers during growth. In most free, certain species of Gypsina), in competition trochospiral forms, the spiral side is in a with rhodophyceans for instance. See also dorsal, the umbilical side in a ventral position. arborescent growth, surface of attachment. Nepionic, spiral stages in permanently attached forms (Planogypsina (Cibicididae) for endemic - occurring in restricted instance) show by the position of their geographic areas; the contrary of foramina in the early spiral whorl that their cosmopolitan. Compare: vicarious species. surface of attachment is on their spiral (dorsal) endoplasm – the central part of side. However, in current uage the terms protoplasm containing the nucleus or nuclei dorsal-ventral and spiral-umbilical respectively and in which the major metabolic processes may not always be synonymous. take place. E endoskeleton - localized thickenings on ectoplasm – the microtubular outer zone the inner surface of the chamber wall that of cytoplasm, that also forms the pseudopodia. partly or totally subdivide the main chamber They have many functions for movement, lumen in the lee of protoplasmic streams catching and transport of food, removal of according to a pattern produced by the excretory products and of gametes from the arrangement of intercameral foramina in mother shell, in sensory tasks, in gas successive septa. Plate-like elements exchange, as well as in the processes of test (septula), usually perpendicular to the septum, formation. See also: canal systems. may form more or less complete partitions touching the lateral walls or fusing with egg-holder - internal pore mouth, elements of the exoskeleton. Discontinuous, enlarged to a cup with polygonal rims, with or columnar partitions are called pillars (or

13 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02) interseptal pillars). A third type of the way to clarify, by comparisons, the endoskeleton is produced by layers of shell significance of true toothplate structures. deposited on the chamber floor and coating entosolenian tube [endosolen] – a tube- the previously exposed outer shell surface like internal skeletal structure extending from completely (basal layer) or partially (chomata the aperture in a proximal direction. Fig. 43. and parachomata of fusulinids). In different taxa, the three endoskeletal types may occur enveloping canals (intramural cavities, alone or in varying combinations. Fig. 47 & auct.) - more or less tubular spaces parallel Fig. 63. Often, endoskeletal elements appear with the test surface formed within lateral only in the course of ontogeny, usually later chamber walls and communicating with than exoskeletal elements. In agglutinated intraseptal interlocular spaces. The enveloping shells endoskeletal elements include the canals are produced by (non-adhering) septum and may be recognized by remarkably imperforate portions of outer lamellae, that coarse and irregularly shaped particles that also cover partly the intraseptal spaces and obscure the genetically fixed pattern in are fold into these spaces, leaving on both contrast to the more ordered exoskeletal sides of this "flying cover" alternating rows of elements of the same specimen. Whether the openings for ectoplasmic flow over grooves in toothplates and their equivalents in lamellar the perforate wall, situated between perforate foraminifera and the secondary septa imperforate inflational ridges (feathering). Fig. produced by folds of the inner lamella in 44 & Fig. 65. orthophragminid lamellar architecture are homologous equivalents of the endoskeletal ephebic - "adult", i.e. a post-nepionic structures of non-lamellar-imperforate growth stage during which the features foraminifera remains an open question. characterizing the shell architecture remain constant. The ephebic stage may grade into a Remarks: The term endoskeleton was later gerontic stage where the architectural introduced by H. DOUVILLÉ (1906, p. 593 and features risk becoming irregular, or are altered 602) in a key paper comparing the anatomy of to form brood chambers, as is often observed imperforate fusiform shells, i.e. fusulinids, in the microspheric generation of larger, loftusiids and alveolinids. DOUVILLÉ had complex foraminifera. See also: gerontic already recognized the close morphological stage. relationship between apertural and epiderm – the thin, outermost coat of a endoskeletal patterns. In his monograph on foraminiferal non-lamellar, imperforate alveolinids, M. REICHEL (1936-1937) adopted chamber wall if it differs in texture from that of the term endoskeleton to designate the the inner layers. Present in all exoskeletons structural elements subdividing the chamber in consisting of a polygonal network as in contrast to a so-called exoskeleton comprising orbitolinids (s.str.) and spirocyclinids. May be the lateral and frontal chamber walls including homologous with the tectum of fusulinids. Fig. the apertural face. The strict correspondence 45. between the pattern of distribution of the foramina on the septal face and the patterns epiembryonic - see subembryonic. produced by the endoskeletal elements (in the epiphyte – Organism living on vegetal alveolinids by the septula) were clearly substrates. Larger porcelaneous foraminifera demonstrated by models of the shell cavities prefer seagrass leaves to algal thalli because and their connections through the septum as if the leaf-hairs (trichomes) give them hold on they were a cast of the shell cavities their surface. Fig. 42. (REICHEL, 1936-1937, fig. 27). These patterns are still used today as diagnostic features for epitheca - biomineralized deposits on inner the definition of alveolinid genera. chamber surface in fusulinids, comprising HOTTINGER (1967) modified and extended tectorium and chomata. (1978) DOUVILLÉ's term to include all equator(ial) – the peripheral line in the structures subdividing the chamber lumen and median plane, perpendicular to the axis of linked to the patterns of intralocular planispiral coiling or to radial symmetry in protoplasmic streaming in contrast to chamber arrangement. exoskeletal partitions that are not affected by such patterns. Thus, the originally descriptive equatorial aperture – an interiomarginal term is expanded to include a functional primary chamber aperture in spiral tests, meaning and extended to all corresponding straddling the equatorial periphery. features in imperforate shells. In some equatorial chambers or chamberlets - lamellar-perforate foraminifera, comparatively see main equatorial chamberlets. rare structures (such as the hollow pillars in Chapmanina) correspond in shape and position equatorial crest – in discocyclinids the to the definition of endoskeletal features. equatorial thickening of the inner lamella of There is no reason to interpret their function the equatorial main chamberlets. Fig. 41. otherwise. So such features may be called endoskeletal without hesitation. They may lead equatorial section – a slice of the test in

14 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02) the equatorial plane. Fig. 37.13. considered at that time to be mainly the expression of a particular "behaviour" in the equitant (in German "reitende" Kammer) - diagenetic process. Nowadays, we know that uniserial arrangement of chevron-shaped this particular kind of differentiation in the chambers, more or less embracing the flanks porcelaneous wall is textural and corresponds of the preceeding ones, as in Flabellina. to the general differentiation of all lamellar- Foraminiferal chambers of this type, usually perforate and non-lamellar walls into inner and with a single, terminal aperture, are commonly outer layers. two-dimensional, as in Flabellinella, rarely In agglutinated foraminifera, there are often three-dimentional, as in the Permian true structural elements to be classified in Colaniella. Biological significane unclear. Fig. separate categories in much the same 49. geometric way as REICHEL conceived it, for eucaryote – a cell with a complete, they carpet the internal surfaces of the free membrane-coated nucleus housing a set of lateral chamber walls (exoskeleton) or follow single (haploid) or double (diploid) the foraminal pattern in the septa chromosomes. (endoskeleton). In discoidal agglutinated shells, exoskeleton and endoskeleton are euphotic zone - the water layers in which separated by open spaces running parallel to light penetration permits the photosynthesis of the septum, the annular passages. In eucaryotic cells, to about 140 m depth in uniserial-conical constructions, the main radial pristine water bodies (blue deserts). partitions of a chamber may be continuous, evolute chamber arrangement - in spirally extending from the shell center to the coiled foraminifera where - due to chamber periphery of the chamber and comprise an shape - the chamber lumina of a coil do not inner, endoskeletal and an outer, exoskeletal cover laterally those of the preceding coil. Fig. part (as in Orbitolina). Thus, REICHEL's (1936- 7. Compare involute. 1937) term exoskeleton was conserved but its definition restricted to true structures as exocytosis – expulsion of fluids and solids opposed to the also exclusively structural from the cytoplasm into the ambient elements of the endoskeleton (HOTTINGER, environment through the cell membrane. 1967, 1978). Therefore, the endoskeleton as exoskeleton - localized thickenings on the conceived here comprises septal structures inner surface of the chamber's outer walls that only when such elements consist of blind- subdivide the chamber lumen into blindly ending recesses carpeting also the lateral ending compartments. They form geometric chamber walls (Hottingerita) or when more patterns which are independent in number and complex structures derive from such a feature direction from those determined by (Alveosepta). BANNER's "hypodermis" (1966, protoplasmic circulation through foraminal according to the English manuscript) is not systems, i.e. endoskeletal patterns. synonymous with "exoskeleton" but a general Exoskeletal elements may consist simply of term for all kinds of alveolar layers, including partitions (beams) perpendicular to the the pigeon-holes of the polygonal septum and to the lateral chamber wall, that subepidermal network, but excluding simpler produce simple alcoves, or of a tapestry of "pseudalveolar" structures as in Orbignya or alveoles of various kinds coating the internal Voloskinovella. As the distinction of polygonal surface of the outer chamber walls. Two main networks and various alveolar structures is of alveolar types may be distinguished: 1) great taxonomic and stratigraphic importance, branched or unbranched alveoles with a blind we do not recommend the use of the term ending of rounded outline below the external hypodermis. wall surface: the partitions lack a expanse chamber (dome-shaped differentiation into beams and rafters and 2) chamber, auct.) – a chamber extending over a pigeon-holes with a blind ending polygonal in wide area above a previous chamber and outline below an abaxial, that is produced by adhering to it in a vermiculate or reticulate partitions differentiated into beams and pattern. Fig. 46. rafters. Fig. 45. Exoskeletal elements may be very deep, subdividing the chamber almost extrathalamous - situated outside the completely as in Dictyopsella. test. Remarks: M. REICHEL (1936-1937) introduced extraumbilical aperture - an aperture in the term "expressis verbis" as a contrary of the primary chamber of coiled shells situated DOUVILLÉ's "endoskeleton" for alveolinids. He at the interior margin but not connected with thus designated the free, outer chamber walls the umbilicus. including the frontal wall with its apertures as exoskeletal, all internal partitions (septula, extraumbilical-umbilical aperture – an basal layer and chamber floors) as aperture located on the interior margin of the endoskeletal. This concept was supported by primary chamber of a coiled shell that extends an obvious differentiation of the wall texture in from the umbilicus to the periphery. the external cover of the chamber walls

15 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02)

F intensity in order to avoid photoinhibition. Fig. 30 B. feathered – ornamented by numerous parallel grooves extending in more or less flange (French: collerette) – a thin, fragile perpendicular direction from sutural furrow or marginal portion of an orbitoidiform shell fissure onto shoulders of ventral chamber where the main chamberlet layer is exposed walls, as in many rotaliids. Fig. 50. for lack of lateral chamberlets. Fig. 52. face – a differentiated part of the shell flexostyle (French: goulot) - in surface delimited by modification of shell porcelaneous foraminifera an eaves-like shape, of wall texture or of ornamentation, extension of the proloculus wall that forms a where single or multiple primary apertures space, u-shaped in section, over part of the and/or orifices of interlocular spaces are proloculus. Fig. 41. grouped to form a functional unit. Fig. 48. float-chamber – a large hemi- to

subspherical chamber that facilitates Remarks: Sofar, the term "face" has been floatation, found in the pseudoplanktic stage of used in combination with a modifyer like some benthic smaller foraminifera. Occurs apertural face or septal face, that concern only inside a so-called balloon chamber. Fig. 16. a single chamber. In addition, we introduce here the generalized term "face" to designate floor (French: plancher; German: Boden) - that portion of the shell surface which by its Part of endoskeleton: wall parallel to basal morphological differentiation indicates a layer separating superposed, regular particular function. This surface may be chamberlet layers in alveolinids. Fig. 14 & Fig. delimited by breaks in the topography of the 70 G-H. shell surface, such as an angular periphery in Remarks: Advances in the comparative trochospiral shells that delimit a more or less anatomy of alveolinids and miliolids requires flattened base to the conical shape of the that superposed regular layers of chamberlets shell, or by a particular wall texture, the lack separated by floors (= planchers, REICHEL, of perforation for instance, and/or by a 1936-1937) as in Praealveolina or Alveolinella discrete ornamentation such as radial be distinguished from irregular, tubular, (Discorbinoides) or parallel-linear supplementary chamberlets present in a (Scarificatina) ornaments over spiral-umbilical thickened basal layer as in Alveolina or surfaces. Faces are directed towards the Subalveolina. Both regular supplementary substrate in motile benthics, away from the chamberlets and irregular, tubular chamberlets substrate in permanently sessile, arborescent appear as soon as the fusiform shell reaches a forms or are opposite each other in paired specific minimum elongation (HOTTINGER, shells ready for plastogamy. The grouping of 1962). In particular the irregular chamberlets apertures and/or orifices on the face may of Alveolina appear during ontogeny, at first in indicate its autecological functions. very low numbers, in shells where the polar thickening of the basal layer has produced an feeding cyst – pads or rolls of the remains elongation of 1.4 : 1 in axial length versus of digested food that accumulate temporarily equatorial diameter. A relationship between in front of apertural faces of benthic elongation index and polar structure is foraminifera. May contain identifyable particles observed also in the parallel phyletic lineages. such as coccoliths or diatom frustules However, the difference between regular indicating elements of the diet of the supplementary chamberlets and irregular- foraminifer. tubular passages in the polar columella is as fistulose chamberlet (marginal diagnostic as all other structural features. chamberlet; peripheral chamberlet auct.) - in agglutinated foraminifera the space produced When HAMAOUI and FOURCADE (1973) were through separation either peripherally or revising the axially compressed relatives of the laterally of part of the chamber from the main classical alveolinids, they described the basal chamber-lumen by a paraporous partition. May layer pierced by tubular supplementary occur in species with either solid or paraporous passages as "central thickening". REICHEL walls. In the latter case, the partition is a (1984, fig. 3, p. 530) established the continuation of the paraporous inner wall-layer homology of the thickened basal layer in the beneath the outward turning outer, solid layer, polar area of Subalveolina with the "central the pavement. Fig. 51. thickening" of Rhapydionia. Chubbina was described by ROBINSON (1968) as having flabelliform - fan-shaped. tubular passages in a basal layer, while DE flagellum – a tubular extension of the cell, CASTRO established (1990) Pseudochubbina as reinforced with microtubuli and anchored by the taxon for a compressed alveolinid "with centrioles deep inside the protoplast of floors". In published illustrations, the Chubbina dinophycean symbionts. The short flagellae are endoskeleton rather resembles that of an used for locomotion in the lacunar system of advanced Praealveolina with floors while the host as a device for regulating irradiation Pseudochubbina has, particularly in adult

16 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02) growth stages, very irregular tubular passages outline and often texturally differentiated in a massive basal layer. Thus, the unfortunate (porosity). The limit between the main lateral current use of the term floor is synonymous chamber wall and the folium may be marked with basal layer. A detailed structural analysis by a short, posterior indentation or "notch" of laterally compressed alveolinids (FLEURY & and/or by an umbilical plate-suture. An FOURCADE, 1990) will be necessary to opening (foliar aperture) is always present distinguish precisely the two kinds of between the anterior margin of a folium and endoskeleton. the adjacent previous coil. In addition, umbilical and/or posterior openings may be flosculinisation – a conspicuous present, depending upon whether the folia are thickening of the basal layer in the equatorial free or attached by their tips or along their zone of miliolid and alveolinid shells. Fig. 18. posterior margin. In some genera such as

Asterorotalia, the folia may extend onto the Remarks: The term was derived from the preceding chamber and be attached to it, ancient generic name Flosculina STACHE by partially covering intraseptal interlocular SCHWAGER in 1883. Flosculinisation occurs not spaces wherever present. A folium is only in alveolinids but in other, unrelated composed of the same layers as those forming porcelaneous genera as well. It may be the wall of the main chamber. See also: foliar restricted to a particular period of ontogenesis, chamberlet; foliar aperture. Fig. 53. or sometimes to two successive stages, and be combined with a polar thickening of the basal foramen [intercameral foramen] (plural: layer. foramina) – the opening or openings that allow communication between the lumina of fluting (septal fluting) - folding of the consecutive chambers and provide passage for septa at their base in fusiform shells, as in functional endoplasm. May be primary, hence fusulinids. The folds in consecutive septa formed by an initial cameral aperture, or oppose each other and may fuse at their secondary, i.e. formed by resorption of masks bases, thus loosing contact with the surface of or other parts of the septum (tunnel). Cameral the previous whorl that is the chamber floor apertures converted into intercameral and consequently producing passageways foramina may be modified in shape by parallel to the tunnel, the so-called cuniculi. resorption or through restriction by attachment Fig. 35. of a toothplate, foramenal plate or umbilical foliar aperture (labial aperture, pars plate. See also: stolon. auct.) – the primary opening of a folium to the foramenal axis - an axis common to two exterior in an interiomarginal anterior, or more intercameral foramina or stolons in posterior or umbilical (axial) position at the subsequent septa; may form regular patterns. folium borders; it lies between the main Fig. 47 & Fig. 80. chamber wall and the folium (foliar slit; astral fissure); or in the folium itself. A foliar foramenal disposition – the pattern aperture may be a continuation of the primary generated by a regular spacial disposition of cameral aperture or separate from it, but it is foramina on septal faces. Fig. 47 & Fig. 80. never converted into an intercameral foramen. foramenal plate (toothplate; paries It may lead to funnels when covered by proximus, pars auct.) - basically a primary secondary lamellae. Fig. 53. infold or "inpush" in the direction of growth of foliar chamberlet – the part of a chamber the posterio-lateral chamber wall at a sutural delimited by a folium. A foliar chamberlet may notch, and attached to a single intercameral be continuous with the main chamber, and is foramen. A foramenal plate is a detached separated primarily (and partly or more continuation of a septal flap that may or may completely) from the main chamber by a not be connected with a cover plate in the foramenal plate, by an umbilical plate, or - previous chamber. A foramenal plate separates secondarily - by a cover plate or a sealing to some degree the main chamber lumen from plate. A foliar chamberlet communicates with the lumen of the foliar chamberlet and the the exterior through foliar apertures and with chamber plasm from the ectoplasm in the the lumen of its own main chamber through interlocular spaces. Compare: umbilical plate; openings in the umbilical plate or cover plate bipartitor. Fig. 34. (when present) or at its margin. Depending fore-court – a deuteroconch with multiple upon the geometry of the test, a foliar apertures covering the outer flexostyle chamberlet may be converted into part of a opening and parts or all of the wall of the secondary spiral umbilical canal as part of the megalosphere in complex soritid embryos. umbilical cavity and communicates with Introduced by LEHMANN (1961) under the intraseptal interlocular spaces. Fig. 53. German term "Vorhof". Fig. 41. folium (lip; tenon; umbilical flap; astral fossette – the opening to the exterior of lobe, pars auct.) - in spiral lamellar an intraseptal interlocular space which is foraminifera an axial-umbilical portion of the subdivided by ponticuli with retral processes. lateral chamber wall, generally triangular in

17 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02)

Fig. 54. a climatic belt around the globe and any one of its subdivisions, such as the photic zone within four-stolon system (or pattern) – a the tropical oceans. The time period governing pattern of the stolon arrangement of arcuate a GCM unit is of geological dimensions. chamberlet cycles that alternate in radial Disrupting geological events which produce the position from one cycle to the next and are boundaries of the larger chronostratigraphic connected by 2 oblique stolons to both the units in the geological time scale often preceding and following cycles. truncate the GCM cycles. foveolate - see pits, pitted. glomerulus – a nepiont characterized by funnel (vertical canal) – a tubular streptospiral coiling (prior to fixation of a (interlocular) space more or less normal to the coiling axis) in fusulinids, alveolinids and many test surface produced by secondary lamination smaller imperforate foraminifera, often over several whorls. Funnels originate from the exclusively in the microspheric generation. Fig. margins of sutural canals or fossettes, or of 57. foliar apertures communicating with a spiral- Golgi body [Golgi apparatus] – a cell umbilical canal. Vertical canals persist in organelle consisting of a stack of flattened secondarily laminated parts of the test as long vacuolar cavities (cisternae). In the as they are not covered by later chambers. foraminiferal cell, the Golgi body produces the Fig. 55. high-magnesium calcite needles which, after fusiform – the outer shape of a shell exocytosis, are used for bonding grains in the similar to a spindle, with its polar ends more agglutinated shell or for the construction of or less tapered (see shell shape, Fig. 37.13). porcelaneous walls. Fig. 56. In fusulinids, loftusias, alveolinids, Fusarchaias granular texture - see jagged granular and Boreloides, it is produced by planispiral- and mosaic granular texture. involute growth when the length of successive chambers in axial direction increases more granule – see bead, papilla. rapidly than the half circumference of the groove – an elongated depression corresponding equator of the shell. Note the produced by external local thinning of the shortening of the pathways through a fusiform outer lamella. It is deepened by persistent shell in polar direction for up to one order of thinning ot the secondary laminations and a magnitude as compared to the equatorial path concurrent thickening of adjacent inflational and see also polar torsion. costae or ridges that are usually present. The G grooves reflect the flow of pseudopodial protoplasm extruding from interlocular spaces gametogamy – a process of sexual (feathering) or along the shell's periphery reproduction in which gamonts release their (cord). They my be covered or partially closed flagellate gametes into the free water column, by secondary lamellation in later stages of where they mate. growth and converted into canals. Fig. 50 & gamogony - sexual reproduction within Fig. 64 A. the reproductive cycle, from meiosis to gutter - in Amphisteginidae: the free end fecundation, generating the zygote. of the stellar septulum, folded anteriorly at an gamont – a specimen producing gametes angle to the axis of coiling. May be identical in the process of reproduction irrespective of with the counterseptum in New World its involvment in meiosis (diploid gamonts) or amphisteginids. Compare foramenal plate. Fig. not (haploid gamonts). Foraminiferal gamonts, 33. produced by asexual reproduction are H megalospheric (A - form). hamulus - see tooth. This term has been gamontogamy – process of sexual coined for endothyrids. reproduction where two gamonts form a nuptial cyst in which, under its protection, the haploid – having a reduced set (half of the amoeboid gametes from the two gamonts may total number) of chromosomes in the gamontic mate. generation as compared to the full set (diploid) in the agamontic generation. Fig. 5. gerontic (growth stage) – the ultimate growth stage distinguished from ephebic hemiseptula - in Amphisteginidae: the (adult) growth stages by a reduction in rates infolds of the inner lamella partially of growth and usually many structural subdividing a chamber. Fig. 58. irregularities. hemiseptular suture – the line of global community maturation (GCM) - adherence of hemiseptula to a lateral wall. gradual change in the composition of a hispid - covered with minute pustules or community by evolutionary and coevolutionary pseudospines. processes in a global ecological realm such as

18 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02)

holotype – see type. crystals formed initially on both sides of the primary organic sheet, crystals that some homologous – equivalent morphological authors include in the so-called median layer. features of common evolutionary origin. May Fig. 75. have different shapes and functions. inner organic lining (IOL) - see organic homomorphous - identical or similar in lining. Fig. 75. morphology but of different phylogenetic origin. instar - a single step in the discontinuous growth process of most foraminifera. It is homonym – the same name for different reflected by the formation of one chamber of taxa. Each taxon must have one name and one the shell (or of a cycle of chamberlets). See type. also: chamber; chamberlet cycle; lamellation. host – an organism housing symbionts. Remarks: The use of this term in husband, husbandry - see chloroplast foraminiferology is contested, mainly by husbandry. biologists. The Cambridge Dictionary of Biology hypodermis - exoskeletal structures below (WALKER P.M.B., ed., 1990) defines instar as an epiderm. See: polygonal network; alveoles. "the form assumed by an insect during a particular (ontogenic) stadium" (such as pupa, hypodigm (Latin: hypodigma) – at a imago etc.). Others have used the term for the species level all specimens that a taxonomist period of existence of marine arthropods considers to belong to that one unit. The between moults (ecdysis). Transferred to hypodigm also includes the specimens of foraminifera, the formal definition would mean previous workers as listed in the synonymy. the stages of growth commonly designated as See also type. embryonic, nepionic or ephebic, each I characterized by the addition of several or numerous chambers. However, because in imperforate - lacking pores or parapores. foraminifera each growth-step modifyes the form of the shell and can be individually incisional ornamentation - pattern of identified in every specimen by counting the primarily thinned portions of outer lamella number of chambers, I see no reason to reject deepened by secondary thin lamination, the already traditional and most convenient usually adjacent to inflational ornaments. See use of this term as substitute for growth-step. also: grooves. Thus, an instar of a lamellar foraminifer index of elongation - see elongation includes whatever the shell produces by the index. same growth-step, i.e. the walls of the a chamber or of a chamberlet cycle and the indistinctly radial texture - appearance respective outer lamella that covers all the under crossed nicols of extinction in the center exposed surfaces of the previous shell and of a fragment of test with other extinctions participates in the construction of surficial irregularly distributed. Corresponds to a ornements including piles, umbilical plugs and bundle-shaped ultrastructure. umbos. See also: supplemental skeleton, inflational ornamentation – a pattern of where the distinction of successive growth- primarily thickened, often imperforate areas of steps is difficult. outer lamella becoming additionally thickened interauxiliary chamber(s) - by secondary lamination. See also: pustules; periembryonic chambers that lack direct beadings; pseudospines; costae; carina. communication with the embryo lumen. They inflational pillar - see pile of lamellae. form the periembryonic spirals in miogypsinids. inframarginal sulcus - see infundibulum. intercameral foramen - see foramen. infundibulum [inframarginal sulcus] (scrobis septalis, pars auct.) – a distinct interio-areal aperture - aperture situated proximally directed infold in a distal chamber near the base of the distal wall, but not at the wall located beneath the periphery on the suture with the preceding coil. See also: umbilical side of trochospiral forms. See also: aperture. marginal prolongation. Fig. 61. interiomarginal aperture - aperture inner lamella (inner lining, inner situated at suture between the distal wall and calcareous layer, pars auct.) - the inner preceding coil. mineralized part of the primary bilamellar interlamellar organic sheets - sheets of chamber wall between an inner organic lining organic material occurring between and a primary organic sheet. Made up mainly consecutive secondary outer lamellae. Fig. 75 of stacked calcitic platelets or of aragonitic C. prisms. In calcitic foraminifera it comprises also the inner array of large, paired rhombic interlamellar space – a space formed

19 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02) between successive outer lamellas or by flying intraumbilical - see umbilical. covers of primary, bilamellar walls. These involute chamber arrangement - in spaces may be inflated to become a spirally coiled forms where - due to chamber supplemental chamberlet, fed by canal orifices, shape - the lumina of the chambers in one coil not by apertures nor stolons, and hence are cover laterally those of the preceding coil. See independent of the arrangement of primary also alar prolongations. Fig. 7. chambers. Seen in particular in Siderolites, Pellatispira and consorts. See also: isodiscodine - see biconch. supplemental skeleton. Fig. 65. J interlocular space (lacuna, pars auct.) - a space formed as a consequence of a deeply jagged-granular texture - appearance sunken suture between consecutive chamber under crossed nicols of a fragment of the test walls or between consecutive coils. See: wall showing minutely granulated crystals with intraseptal and spiral interlocular spaces. Fig. irregularly shaped sutures. The crystals usually 26 A. have a more or less uniform yellowish colour in polarized light. This texture indicates an interpore ridges - the external residual or intricate ultrastructure. built-up test material in the areas between large pores. Fig. 28. joist - see beam. interradius (pl. interadii) - in orbitoidiform juvenarium - see nepiont. architecture: sector of undifferentiated K equatorial layer inbetween radii. keel – see carina. interseptal - located between consecutive septa. keriotheca – an alveolar, honeycomblike structure of the spirotheca in advanced interseptum (in Amphisteginids for fusulinids, consisting of an "upper", extern and instance) – the partial subdivision of a spiral a "lower" intern "layer" produced by a split of chamber by infolds of the inner lamella in a the alveoli into narrower subunits below a direction that is more or less parallel to the tectum. Fig. 62. septum. Remarks: The terms "upper" and "lower" layer Remarks: The term is impropriate. Structural of keriothecal structures refer to the traditional elements subdividing spiral chambers in orientation of single chambers or wall parts in lamellar shells are septula, not septa. Use specialized papers. They should be replaced by instead hemiseptulum. the terms extern and intern when referring interseptal pillars [pillars] (lamelliform to their position within the shell, quite buttresses) - in porcelaneous and agglutinated independently of the orientation of the species: the multiple columnar projections illustrations. between consecutive septa and parallel to The term "keriotheca" is restricted here to protoplasmic flow. Pillars are part of the structures with two layers of alveoles. Alveolar endoskeleton. To be distinguished from piles of layers with uniform radial elements, as in lamellae. Fig. 47 E-G & Fig. 72. Verbeekinid fusulinids, must have another designation in view of the importance of these intradermal plate - see beam; septulum. strucural differences in higher systematics (see In order to avoid ambiguity tis term should not pseudokeriotheca, parakeriotheca). be used. knob - see boss, umbo. intraseptal interlocular space - the interlocular space formed between the L posterior bilamellar wall of a chamber and the labial aperture - see foliar aperture. distal bilamellar wall of the preceding chamber, as a result of a deeply sunken labyrinthic – a spacial disposition of suture. Intraseptal spaces may be open to the subdivided chamber cavities without regularity exterior along their margins either or pattern. In most cases however, the continuously or through openings between labyrinthic disposition is only apparent, in one points of marginal adherence of consecutive instance due to the strong curvatures of lateral chamber walls. See also: sutural consecutive septa in both axial and equatorial canals; fossettes. Fig. 44. directions, so intersections with structural elements in oriented sections of the shell are intraseptular space - space formed in the oblique, as in spiral Anchispirocyclina. wall (septulum) between adjacent subsidiary chamberlets of the same instar, produced by lamellation – a mode of wall formation by lateral infolding of the chamber wall. May extracellular biomineralization on organic contain canals communicating with an sheets (templates) shaped - for the chamber intraseptal canal system. under construction - by temporary ectoplasmic strands similar to brush borders of other cells. intrathalamous - situated inside the test. The external organic coat of previously

20 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02) formed, exposed hardparts may form an passages in anchispirocyclinids. See also: additional template for the deposit of an outer passage. lamella. In his monograph on rotaliids, SMOUT latero-marginal aperture – a primary (1954) formulated for the first time this very aperture situated on a lateral umbilical wall of basic principle of wall formation that was coiled tests, in a subperipheral position. subsequently refined and modified by HANSEN (1999). life cycle - most eucariotic, free-living cells reproduce asexually but shift from time to time lamellar wall – a test-wall consisting of to sexual reproduction in order to avoid layers of calcite or aragonite formed at degeneration (MULLER's ratchet, 1964). In consecutive instars and covering the exposed many foraminifera the alternation of sexually surfaces of the test already formed. This wall and asexually produced generations is generally possesses true pores produced in documented in the morphology of the shell by consecutive lamellae by so-called blueprints. dimorphism or trimorphism (see: gamont, Most lamellar genera are bilamellar and some schizont, megalospheric generation, agamont primarily multilamellar. and microspheric generation). In foraminifera, lamelliform buttress - see beam if there are many different types of life-cycles exoskeletal, septulum or pillar if endoskeletal. (LEE et alii, 1991; GOLDSTEIN, 1999). They differ in particular by the position of the Remarks: HENSON (1948) introduced this term nuclear reduction division (abbreviated R!) for chamber partitions leaning against free within the cycle. Planktic foraminifera are outer (mostly lateral) walls as suggested by interpreted as reproducing exclusively by the meaning of the term in architecture. sexual processes. See also: alternation of Buttress is by and large a synonym for generations. Fig. 5. subepidermal partition. HOTTINGER (1967) limbate – a descriptive term for the showed the distinction of endo- and thickened, more or less prominent sutures on exoskeletal structural elements to be of the test surface at the boundaries of taxonomic and phylogenetic relevance and chambers. Fig. 77 F. defined terms that are specific to each category of partitions. There is no reason to lip [apertural lip] – an everted, in lamellar continue the use of the term buttress. Where a forms imperforate extension of the chamber neutral and purely descriptive term may be wall along an elongate cameral aperture. May helpful in a case difficult to interpret, it is be narrow or broad, small or large. See also: sufficient to speak about "partitions". phialine lip; rim. Compare: folium. lamination [secondary lamination] - loculus - chamber. layering of test-walls due to superposition of consecutively deposited outer lamellae on loop-hole – a small connection between exposed outer shell surface. See also: lamellar chamber lumen and interlocular space. May be wall. located at the periphery (leading into a sulcus in nummulitids), at the base of the chamber, lateral canals - see sutural canals. as in elphidiids, or in front or in back of umbilical plates or cover plates, as in rotaliids. lateral chamber wall – that portion of the Fig. 34. main chamber wall never converted into part Remarks: The "septal passage" of PARVATI of a septum; more or less clearly separated (1971) is a loop-hole in pre- or post-septal from the chamber periphery or peripheral wall. position. As the term "passage" is now lateral chamberlets (cubicula) - in reserved for connections between cavities of orbitoidiform architecture: chamberlets of the same chamber (or instar), the term "septal roughly isometric or irregular to vermicular passage" leads to confusion and is replaced outline, layer-wise arranged in a chessboard here by pre- or post-septal loop-hole. Loop- pattern. They cover both lateral surfaces of holes function as backdoors for the extrusion orbitoidiform shells. Retral stolons connect the of ectoplasm when the chamberplasm retracts main chamberlets in the median chamberlet to inner chambers during a perturbation in the layer with the adjacent lateral layers of ambient environment. chamberlets. Consecutive layers of lateral lumen - see chamber (or chamberlet) chamberlets are connected with oblique lumen. stolons. The lateral chamberlets of one layer may be connected between themselves by M more or less restricted passages. There may be gradual transitions between lateral maerl - coarse-grained sediment consisting chamberlet layers and expanse chambers. of shells, coral fragments and other skeletal debris, that form extensive carpets on the sea lateral openings – a gap for intracameral floor where the components are bound by communication in neoschwagerinid septula, at living algal filaments and/or coralline red algal the junction of exoskeleton and endoskeleton crusts, and consequently resist erosion by and hence homologous to the lateral annular rapid bottom currents. May be inhabited by

21 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02) normally epiphytic larger foraminifera. margin produced by a marginal canal sytem, as in Pellatispira. It may be overgrown by main chamber lumen – a complete or supplemental chamberlets either on the lateral segmented chamber cavity communicating flanks alone (Biplanispira) or on all sides of the with the preceding and succeeding main shell (Vacuolispira). Fig. 65. chamber lumina through primary or secondary intercameral foramina. In certain lamellar marginal prolongation (tectum) - in a foraminifera the main lumina are separated trochospiral test, a distally directed partly or completely from a foliar or stellar prolongation of the spiro-marginal portion of a chamberlet, as well as from canal systems by chamber, leading the spiral sutures to be much a foramenal plate, toothplate, umbilical plate, more inclined on the spiral side than on the cover plate, or sealing plate. umbilical one. Fig. 61. Remarks: This feature has also been called main chamber wall – that portion of the tectum, a term preoccupied in fusulinids by the walls of a test enclosing a main chamber outermost layer of the chamber wall in the lumen. chamber roof. Therefore the term tectum is main chamberlet layer - in orbitoidiform not acceptable as an alternative to "marginal architecture: cycles of chamberlets in the prolongation". median (equatorial) plane of the shell, marginal trough – a circular depression commonly annular, that are distinguished from between the marginal and central part of the a lateral chamberlet layer (if present) by their chamber face in uniserial-conical foraminifera; smaller size and more regular structure. A it may be the site of a circular row of marginal flange in orbitoidiform shells exhibits the apertures. peripheral parts of a main chamberlet layer that lacks the cover of lateral chamberlets. The marginal ridge – a circular ridge in the last cycle of main chamberlets bears the discoidal chamber produced by a marginal apertural face of the shell. Fig. 52. trough, that marks the boundary between the marginal and the central chamber lumen, and main partition - in Orbitolina: (radial) often the limit between exo- and endoskeletal septulum. elements of the shell. marginal apertures – a single row of marginal zone – the marginal portion of apertures on the shell margin, distinguishable the discoidal chamber in conical shells or of from areal multiple apertures by their oblique- the lateral parts of an annular chamber in radial direction, as in Tertiary conical discoidal-evolute shells. It often houses an agglutinated forms or in Marginopora. Single exoskeleton. In uniserial-conical shells the marginal apertures in lamellar-perforates, as marginal zone is usually separated from a in almaenids, may be closed by a secondary central area of the chamber by a furrow, the lamellation already present in the penultimate trough. The furrow is produced by a recess of chamber. the marginal zone for third or half of the marginal canal system - enveloping chamber height (in the direction of growth). In canals grouped at the periphery of the shell, discoidal-annular shells, a similar recess often extended in radial direction, as in produces a pair of circular marginal troughs Pseudosiderolites, in contrast to cords, where framing the apertural face, as in Marginopora. the peripheral orientation of the canals is See also radial zone, reticular zone. Fig. 20 & emphasized. Fig. 65. Fig. 71 G. Remarks: HOTTINGER (1978) used this term marginoporid structure – a three- as a synonym of marginal cord. It seems dimensional arrangement of endoskeletal preferable now to distinguish between elements as in extant Marginopora: apertural predominantly radial and peripheral systems. axes oblique with respect to radial direction, The latter are never modified to include overcrossing in neighbouring stolon planes and canaliculate spines. See also crest. alternating in radial position from one stolon marginal chamberlet - see fistulose plane to the next. Endoskeleton may consist of chamberlet. septula or pillars. marginal cord – a thickened shell margin mask - mineralized element(s) of the test produced by a peripheral system of numerous that obstruct a primary cameral aperture; longitudinal anastomosing grooves and resorbed in subsequent growth stages. Fig. 77 adjacent imperforate elongated ridges and B. islands of an inflational ornament type. maturo-evolute – planispiral-involute grooves are closed by secondary lamination shells tending to become evolute in mature and converted to an anastomosing bundle of growth stages. peripheral tubular canals. The marginal cord is Remarks: Term introduced by BANNER and the diagnostic feature of the family HODGKINSON (1991), exemplified by Nummulitidae. Fig. 64. Heterostegina depressa. However, in that marginal crest – the thickened shell species and many other nummulitids, the

22 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02) degree of involuteness changes in relation to generations. the depth at which the individual lived. microspheric - in dimorphic species: a Microspheric specimens tend toward maturo- test having a small proloculus or microsphere; evoluteness because their ontogeny is much commonly an agamont. longer than that of the permanently involute megalospheric generation of the same species. microstriae - minute longitudinal, usually See also: evolute and involute in lamellar- anastomosing ridges on the surface of perforate foraminifera. Fig. 7. porcelaneous test. Most are visible only under high magnification, especially by Scanning meandrine – a tortuous, winding path of Electron Microscopy. SEM. linear features, in particular the septal sutures of long alar prolongations or supplemental microtubule - polymerized heterodimers chambers, as in some advanced agamonts of of alpha and beta tubulin in helicoids that form nummulitids, miscellaneids, meandropsines long, straight cylinders reinforcing the and archaiasines. pseudopodial ectoplasm of the pseudopods and the plasmatic content of interlocular spaces, median layer - in bilamellar foraminifera a i.e. canal systems. May depolymerize and form term applied to both the spongy primary paracrystals in the chamber plasm. The organic sheet and the distinctly larger crystals paracrystals are believed to be the reservoir formed initially in vesicles and occurring as that feeds the polymerization when pairs on either side of the primary organic pseudopods expand through an orifice into the sheet. See also: outer lamella; inner lamella. ambient environment. Fig. 67. Fig. 75 A. milioline - the taxa or the characteristics median section – a slice in the central of the suborder Miliolina. sagittal position normal to the axis of coiling. milioline coiling - in porcelaneous megalosphere – the large proloculus in di- foraminifera: bilocular coils where all terminal or trimorphic species; a defining characteristic apertures are positioned on one common axis for gamonts and schizonts in contrast to the (apertural axis). The axis of coiling is normal microsphere of the agamont. See also: to the apertural axis and is rotated so that alternation of generations. Fig. 5. several descrete angles exist between the megalospheric - in dimorphic species: median planes of consecutive chambers: these test having a large proloculus or are 72° (quinqueloculine), 120° (triloculine) or megalosphere; commonly a gamont or 180° (spiroloculine or biloculine). If the schizont. chambers are positioned so that in a section normal to the apertural axis they form an S- meiosis [reduction division] – step in the shaped curve, their arrangement is called process of cell division, during which the sigmoiline. See also: streptospiral. Fig. 37.14- chromosomes of the parent cell are reduced 17 & Fig. 68. from a double to a simple set (from diploid to haploid) for each daughter cell. See milioline tooth/teeth - one or more reproduction cycle: Fig. 5. inward projections of the inner portion of the chamber-wall into the aperture of milioline metabolism – The sum of biochemical species. May be bar-like, spatulate, bifid, or activity in a living organism. In perforate anvil-, T-, Y-, anchor-, spur-, scoop- or spoon- foraminifera the degree of its magnitude may shaped. A single tooth is always present be reflected to some extent by the diameter primarily in an interiomarginal position; it may and density of pores in the shell walls in be accompanied by additional teeth that relation to the volume of the chamber lumina. project from the opposite margin of the microgranular – a wall texture: under the aperture, from the chamber roof or from the optical microscope a granular appearence of lateral wall. See also: trematophore. walls composed of calcareous elements. More mitochondrion – the cytoplasmic stable and resistant to diagenetic organelle of a cell delimited by an outer recrystallization than porcelaneous walls. In membrane and enveloping a folded inner fusulinids and pfenderinids, the microgranular membrane (cristae); it is responsible mainly wall never encases agglutinated grains. In for the respiration of the cell. In oxygen- Mesozoic imperforates, agglutinated textures depleted environments, the foraminiferal may be replaced by microgranular material if mitochondria may assemble below the inner no grains are available in the ambient pore mouths thus documenting the role of environment. foraminiferal pores in the exchange of small microsphere – the small proloculus of the molecules with the ambient environment. Fig. agamont in di- or trimorphic species, in 75 B. contrast to the megalosphere of the gamonts mitosis – the "normal" cell division, where and schizonts. Microspheres never have each daughter cell inherits the gamut of the multiple apertures nor structural chromosomes of the the parent cell. complications. See also: alternation of

23 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02)

monolamellar – a perforate wall nucleus - the cell organelle that contains consisting of one lamina, the outer lamella the genomic material organized in only, lacking both a median layer and an inner chromosomes and that is bounded by a double lamella. The outer lamella may cover the membrane. In foraminifera, a single, larger exposed parts of earlier formed shell somatic nucleus that may be polyploid controls completely or partially. the metabolism of the cell. It degenerates when, at the start of the meiotic process, monothalamous [unilocular] – a shell several smaller, generative nuclei divide prior consisting of a single chamber. to reproduction. mosaic granular texture - under crossed O nicols fragments of the test wall are shown to be composed of large sutured calcite crystals oblique section – a slice through a test with finely serrated margins. Corresponds to cut neither parallel nor normal to its axis. It clumpy ultrastructure. may be centered, that is show the proloculus, or noncentered. Fig. 83. multiple spirals – Plani- and trochospiral shells may produce supplementary spirals odd pairs or associations (Français: growing at the same rhythm as the primary associations dépariées) - common associations spirals, in nummulitids indicated by of two or more foraminiferal species that simultaneous deviation from mean volume exhibit an identical or a closely related accretion rates during ontogeny. These architecture in their adult form but differ deviations are interpreted as seasonal effects. markedly in their adult size. In most cases, the Multiple spirals are known in alveolinids difference in the size of the adults is matched (Multispirina), possibly in meandropsinids by a corresponding, size-dependant (Fallotia), in nummulitids and in rotaliids architecture in the megalospheric embryo, i.e. (Dictyokathina, Dictyoconoides). Fig. 55. a large embryonal apparatus in the larger form, versus a simple, more or less murica – see pseudospine. undifferentiated megalosphere in the odd murus reflectus - see umbilical plate. partner. These partnerships are usually restricted to one or two pairings of species. An N example of a recent odd pair is the frequent neanic – a postnepionic growth stage with association of Amphisorus hemprichii with the the architecture of an adult shell. May be odd partner Sorites orbiculus. They have the either a synonym for the adult or ephebic same habitat on seagrass leaves but reproduce growth stage or designate an early phase of at different times in the seasonal cycle the adult stage. Should be used only where (HOTTINGER, 1999). An example of a fossil there is a qualitative delimitation to later adult odd pair is given in Fig. 70. stages, for instance the transition from spiral odd partner - the smaller representative to flabellar or from flabellar to annular growth. of an odd association. Fig. 37. oral - apertural. See: aperture, foramen. neck – a tubiform extension in terminal position of the aperture, as in uvigerinids. orbitoid(al) architecture – an arrangement of chambers like those of in nepionic – the juvenile stage immediately orbitoids s.l., i.e. an annular series of after an embryonic and preceding an ephebic chamberlets form a sagittal, equatorial main stage. chamberlet layer covered on both lateral nepionic acceleration – the reduction of surfaces by corresponding cycles of lateral nepionic chambers in geologic time within a chamberlets or expanse chambers. phyletic line of genera with either a spiral or a orbitoid(al) growth – an arrangement of radial architecture. chambers like those of orbitoids s.l.: a layer of nepiont – the growth-stage following the annular chamberlets alternating in radial embryonic stage and different in architecture position in successive cycles, each chamberlet from the adult stage. communicating in a diagonal direction with its neighbour in the previous and following annuli. nonlamellar - chamber walls lacking a Fig. 36. lamellar texture, as in most agglutinating and porcelaneous forms. orbitolinid structure – an arrangement of endoskeletal and exoskeletal elements in notch (French: encoche) – a single space like that of Orbitolina and some of its indentation of the proximal chamber wall in a closest relatives; i.e. a radial subdivision of the sutural position. It marks the limit between a discoidal chambers by septula that fuse with spiral main chamber and its folium and may an exoskeletal polygonal network, where the extend into an internal infold. See also: septula adjust to radial rows of apertures with foramenal plate. Fig. 34. crosswise oblique stolon axes. Fig. 71. nucleoconch - see embryonic apparatus. orbitolitid structure – an arrangement of

24 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02) endoskeletal elements as in Orbitolites: the transmission electron microscope. It apertural axes oblique in respect to the radial performs one or several, well defined direction in the discoidal shell, overcrossing functions. See in particular: nucleus, and superposed in radial positions in mitochondria, Golgi apparatus, chloroplast, neighbouring stolon planes. So far, only vacuoles, rhizopodia. septular (i.e. continuous) endoskeletal orifice – any unspecified opening in the elements are known to follow the orbitolitid test such as apertures or the mouths of a pattern of stolon axes. Orbitolinid structure is canal system. The term is to be restricted to a basically the same as orbitolitid structure, but functional meaning, i.e. to those openings it produces radial subdivisions within a where protoplasm extrudes from the shell. discoidal chamber whereas the orbitolitid structure produces subdivsions of an annular ornamentation – the patterns formed at chamber. Fig. 47 H. the shell surface that are generated by a regular modification of wall textures orbitopsellid structure - arrangement of (perforation, pitting), of the thickness of outer endoskeletal elements as in Orbitopsella: the portions of the wall that generates ribs, beads, arrangement of endoskeletal pillars follows the papillae, pustules, pseudospines, etc., or of a pattern of the radial stolon axes that alternate combination of both. See also: inflational in radial position from one stolon plane to the ornamentation; incisional ornamentation; next. Exoskeletal elements consist of beams textural ornamentation. only. Fig. 72. outer lamella – the mineralized layer of organic lining – an organic cell envelope the primary wall in bilamellar foraminifera, on said to consist of mucopolysaccharides, located the outer side of the primary organic sheet. At between the plasmalemma and the its contact with the primary organic sheet is biomineralized cell envelope. It covers the the outer array of distinctly larger paired protoplasmic cell body in the chamber lumina crystals (included by some authors in the so- and the connecting cavities inbetween them called median layer) succeeded by the stacks (foramina, stolons), but never the interlocular of calcite platelets or pseudohexagonal spaces. Whether it occurs in chamberlet aragonite prisms that form the major part of cavities of the supplemental skeleton is the outer lamella, and is completed by the unknown at present. May be discontinuous or outermost thin layer of blocky columnar extremly thin over pore mouths and/or in the crystals that comprise the veneer. Fig. 44 & ultimate and penultimate chambers, and Fig. 75 A. commonly thickens in the direction of earlier growth stages. May be involved in stolon P plugging. May be resorbed together with the biomineralized wall when brood chambers are papilla [pl. papillae] – a small, rounded, formed. The organic lining resists dissolution poorly or non-perforate protuberance, single or of the biomineralized shell by acidic attack, multiple, on the outer surface of perforate maintains the shape of the protoplasmic body chamber or chamberlet walls, produced by and is capable of remineralizing its shell when local inflation of outer lamellae and linked to a the ambient environment returns to normal. conical outward spreading of the pores. Fig. However, the role of the organic lining in 73. biomineralization has to be investigated parachomata - chomata that supplement further. Fig. 69 & Fig. 75. the primary pair of ridges in the equatorial Remarks: There is no reason to abandon the zone of the shell laterally polewards; regularly traditional designation of organic lining (as intercalated between supplementary tunnels used by LOEBLICH & TAPPAN, 1987) in favour up to the polar end of the chamber as in of "inner organic lining" (IOL) as introduced by Pseudodoliolina. May fuse with the beams of ANDERSON and BÉ (1978). The eventual use of an exoskeleton in order to form complete an "outer organic layer" called for by the IOL partitions in the chamber, as in that might be applied either to the outer neoschwagerinids. Fig. 74. organic cover of the biomineralized shell or to the temporary organic envelope that in some parafossette – an opening between species protects the process of chamber bifurcating ponticuli and the margin of the formation would add confusion in the preceding chamber wall. They communicate description of the several layers of the cell with intraseptal interlocular spaces and envelope and their specific functions. fossettes. Fig. 54. Moreover, the term "inner organic lining" is parakeriotheca – a single layer of easily confused with the term "inner lining" uniform, radial, closely spaced and more or which is equivalent to "inner lamella", the less parallel cavities in the spirotheca of inner calcified lamella of the primary wall in advanced fusulinids, covered by a tectum. perforate foraminifera.

organelle – a morphologically distinct, Remarks: This term is introduced here to named unit in the cell plasm, usually visible in emphasize the significance of the difference

25 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02) between one- and two-layered keriothecas. exoskeleton and endoskeleton, as in The single layer is interpreted as a wall texture verbeekinids. In peneropliform to concentric similar to pores and parapores, lacking a architectures, the passages may be plasmatic filling and serving to facilitate gas semiannular-annular, in a preseptal position, exchange through the shell. The interpretation as in Sorites, or paired in lateral positions, of the "alveoles" as pore-like textural elements separating exoskeleton from endoskeleton, as derives from their similarities to the in Anchispirocyclina or Orbitopsella. Fig. 72 & construction of pores: like the lamellar Fig. 79. constructions in perforate foraminifera, the pavement - in agglutinated foraminifera chomatopores are constructed by replicating the outer layer of solid wall covering an inner the pore cavity from one layer to the next paraporous layer. Fig. 51. without morphogenetic intervention by the protoplast. penultimate chamber - chamber before the last in an individual. parapores (canaliculi; pseudopores, pars auct.) - in agglutinated foraminifera: straight perforate - referring usually to walls to tortuous tubular spaces, round to polygonal possessing true pores. Where the term is in section, more or less normal to the test applied to walls possessing parapores it should surface, coated and closed off internally by the be replaced by "paraporous". organic lining. May be branching and anastomosing and - usually - restricted to the perforation pattern – the pattern of inner wall-layer, thus ending blindly beneath distribution of external pore mouths on an outer solid "pavement". No sieve-plate lamellar shell surfaces, either combined with present. Parapores may be laterally or totally apart from the ornamental sculptures interconnected. They may lead into irregular of the shell. cavities or fistulose chamberlets between a periapertural depression - see paraporous wall-layer and the pavement. The adapertural depression. partition between a main chamber lumen and a fistulose chamberlet, wherever present, is periembryonic chamberlets [corona] always paraporous. Compare: pores; pits. Fig. (periembryonic cells) - all chamberlets in 6 & Fig. 51 E. contact with an embryonic apparatus. In orbitoidiform architecture the term is restricted paries proximus – an integrative term to the chamberlets of the equatorial main layer designating the proximal, septal chamber wall and is a synonym of "corona". In orbitolinids and the various extensions produced the periembryonic chamberlets designate the independently by the inner lamella at the forth stage of growth consisting of an annular bottom of the chamber or at its adaxial chamber subdivided into chamberlets. The fifth umbilical region, including also structures in chamber may be called nepionic, and is the previous chamber such as umbilical cover annular or discoidal, radially subdivided by plates. septula following the constraints of crosswise- Remarks: this term was introduced by LÉVY et oblique stolon axes disposed in radial rows. alii (1979, p. 68) in order to support the Fig. 41 I-L. revision of discorbid foraminifera. It integrates Remarks: Introduced by DOUGLAS (1960), the the following terms currently in use: septal term periembryonic originally included flap, proximal wall, foramenal plate, abaxial subembryonic chambers, without analyzing in cover plate and umbilical plate. It is a partial detail the complicated structures of this part of equivalent of HOFKER's (1951) toothplate. This the test. Today, much weight is given to the term also is too broad to be helpful in distinction between periembryonic and distinguishing the various structures that are subembryonic chambers (SCHROEDER, 1962; diagnostic for defining the genera in several ARNAUD-VANNEAU, 1980). See also: auxiliary families. HOFKER's idea (1956) was to place all chamber, adauxiliary chamberlets. ("tooth"-) plate-bearing taxa into a common group, the "foraminifera dentata". Despite this, peripheral chamberlet - see fistulose the paper of LÉVY et alii (1979) is most chamberlet. important for it constitutes the basis of peristome [peristomal lip, peristomal rim] LOEBLICH and TAPPAN's conception of the – a raised rim or tube around an aperture or Discorbidae (1987) in present-day foramen. systematics. phenetic – morphological (in the context partitional pore(s) - see passage(s). of evolutionary theory). passage – the means of communication phialine lip - lip on apertural neck. between adjacent compartments of the same chamber; it is an opening that may be sited photoinhibition - diminishing rates of below the frontal chamber wall (preseptal), photosynthesis under stronger irradiation than after the septal wall (postseptal) as in necessary for optimal rates. alveolinids, or at the fusion between photosynthesis – reduction of

26 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02) carbondioxide to carbonhydrates powered by plastogamic plate – a plate-like structure energy from solar irradiation absorbed by covering the umbilicus in plastogamic chlorophyll pigments. specimens of some benthic foraminifera. Fig. 48 J. phototropy – orientation of organism towards (positive phototropy) or away from plastogamy – two gamonts form a pair by (negative phototropy) sunlight by growth or joining their faces and exchanging gametes active movement. within a common shell lumen where fecundation takes place. Zygotes are hatched phrenotheca - thin, calcified partitions from the paired shells to form an agamont that divide chambers irregularly in various embryo. The mother shells are then discarded directions, as in Pseudofusulina. (see ERSKIAN & LIPPS, 1987). The faces of pigeon-hole – an alveolar cavity in the the shells are decorated by numerous rows of exoskeletal polygonal network that expands at small pustules (costellae) in a radial pattern its blind end below the epiderm into a independent of the chamber arrangement. Fig. polygonal shape. Its opening into the chamber 48. cavity is delimited by longer beams plate suture - line marking the trace of (perpendicular to the septum) and by shorter the adherence of the umbilical plate to the rafters (parallel to the septum). Fig. 45. lateral chamber wall. pile (inflational pillar, cristalline cone auct.) plectogyral - see streptospiral. - superposed lamellar thickenings (pustules) on lateral walls or folia in consecutive whorls plesiotype – see type. or consecutive chamberlet layers, that produce plug [umbilical plug] – an expanding pile of an aspect of pillar-like structures in the thickened lamellae in axial position in an sections of shells. Compare: interseptal pillar. umbilicus or in an umbilical bowl. May be See also blueprinting. single, compound and/or canaliculate. Fig. 77. pilintradermal plate - see beam. pole (of shell) - in subspherical to fusiform pillar - see interseptal pillar. A pillar is not shells the point where the tips of involute a pile! chambers in a planispiral whorl meet the axis of coiling. Fig. 83. See also: polar torsion. pillar-pore - see calyx. polarity (of chamber wall) – a textural pioneering – an early stage of community differentiation of outer and inner portions in maturation initiated in a totally or almost non-lamellar primary chamber walls. See also: empty habitat invaded by immigrants agglutination, basal layer, flosculinisation. specialized for rapidly occupying empty spaces with their unused resources. See also: polar torsion - helicoidal torsion of septa community maturation. and septal sutures at the poles of fusiform larger foraminifera (mainly in fusulinids and pits (pseudopores; punctuations, pars. elongate alveolinids), enforced by the auct.) - in porcelaneous foraminifera, and in broadening of the apertural face and usually spirillinid and some unilocular genera: small linked to a polar multiplication of apertures. cavities in the wall opening at the external surface of the shell, that are rounded to oval in polygonal subepidermal network - section and tubular to conical in shape. They exoskeletal structure formed by a layer of penetrate the shell surface at normal and always undivided, deep, tubular recesses in oblique angles to depths that vary according chamber walls, ending blindly below a thin, genus. Sometimes they occur in several tiers often transparent epiderm or similar structure. and commonly anastomose thus resembling They generate a polygonal pattern at their the parapores of agglutinating foraminifera but distal end and open proximally into the opening to the exterior. The term "pitted wall" chamber lumen with rounded mouths that are is often used to describe the texture of the test slightly restricted between lateral partitions of of planktic, non-spinose foraminifera the chamber differentiated in beams and possessing distinct external pore-funnels or rafters. Fig. 47 E-G. "pore-pits". Remarks: In foraminifera with greatly inflated chambers like Bradyina or Gyroconulina, the planispiral chamber arrangement – beams may produce a polygonal pattern of chambers arranged in whorls where the rate of their own rather than form a row of partitions translation (net rate of movement along the perpendicular to the septum. growth axis to the net rate of movement away Introduced by H. DOUVILLÉ in 1906 under the from the axis) is zero. The spiral and umbilical name "réseau sous-épidermique", the term sides of the test are identical and symmetrical was used mainly in the description of with regard to the plane of bilateral symmetry. orbitolinids where already in 1930 DAVIES Fig. 37.1-3 & 5. distinguished major and minor partitions. In plastid – see chloroplast. the later Anglosaxon literature, these details were not considered as important: COX (1937)

27 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02) described the Loftusia exoskeleton as "alveolar conical agglutinated foraminifera. In the layer" and HENSON (1947) preferred to use ontogeny of specimens belonging to one of the the general term "subepidermal partition" for numerous genera involved no such series of all elements subdividing the chamber lumen exoskeletal structural complication has been into "pidgeon-holes" or "subepidermal cells". observed. In my view, the question remains The general term "subepidermal partition" was unresolved (for discussion see HOTTINGER, extended by HENSON himself and by later 1978, table 1, p. 255). authors to any kind of lateral chamber ponticulus, pl. ponticuli – a bridge of partition, thus depriving the term of any lateral wall spanning an intraseptal interlocular significance for studies of comparative or space. It may be massive or hollow. If hollow, functional anatomy. For this reason, we it covers a retral chamber process. Compare: recommend dropping the use of the term basal lobe; retral lobe; fossette. Fig. 54. "subepidermal partition". HENSON's term "pigeon-holes" rather than "subepidermal polythalamous (multithalamous, cells" is appropriate to distinguish these from multilocular) – a shell consisting of numerous ordinary alveoles or alcoves. chambers. A polygonal network has not been found in any living foraminifer. Therefore, that the recesses polyvalent individuals (polyvalent tests, were coated by the organic lining can not be twins or triplets) - individuals with two or more directly confirmed. The interpretation of the megalospheric embryos belonging probably to polygonal network as an exoskeletal structure the same clone, with a common late growth rather than as a particular kind of wall texture stage. Accidental association, not related to (as HENSON, 1947, had already pointed out) is gamontogamy. supported by their common combination: porcelaneous test wall - composed of lamellar Fabiania and its relatives exhibit a optically cryptocrystalline lathes and rods or perforate epiderm covering their subepidermal needles of calcite produced in Golgi vesicles network, advanced Cuneolina and Dicyclina within the protoplast and transported through have a paraporous epiderm and many the cell wall by exocytosis to the site of wall fusulinids bearing a parakeriotheca combine construction. Rods arranged randomly, lathes this wall texture with an exoskeleton. The arranged in a tile-roof pattern and forming the extremely thin, often transparent epiderm in outer wall-layer. Wall imperforate, but may agglutinated foraminifera suggests, that the possess pits. polygonal network is a device to keep symbionts exposed to light and in the pore – a minute tubular perforation immediate vicinity of the location where gas traversing a lamellar chamber wall, coated exchange through the shell should be internally by an organic sheathe. Subdivided enhanced by particular, porous textures. by organic discs ("pore plate" auct.) and closed off internally by the inner organic lining. It may be of taxonomical importance at a The latter may fuse with the basal disc supra-generic level to distinguish at least two corresponding to the median layer and form an kinds of polygonal networks. The first one to organic pore plug. The size and shape of the appear in the Mesozoic (Haurania and external and internal pore openings may be Amijiella, Middle Lias) is comparatively coarse identical (rounded to elongated) or dissimilar, and deep. There is probably no clear when symbionts are positioned in egg-holders differentiation of an epiderm, just a thin outer below the pore mouths for gas exchange. wall covering the polygonal ends of the blind Ultrathinsections of living benthic foraminifera recesses. The differentiation of beams and that show in the TEM some cytoplasm in pores rafters does not seem to be very orderly, even are considered to be artefacts due to imperfect in evolute, fan-shaped to discoidal shells such preparation. Compare: parapores; pits. Fig. 75 as Timidonella or Alzonella. The second group pore-chimney – an enlarged pore-cavity that appeared not much later with in a secondarily laminated wall, grouping 2-4 Pseudocyclammina liasica, has finer meshes smaller pores of the primary chamber wall. and a clearly differentiated, extremely thin 178/5,7. epiderm. This type characterizes most late Mesozoic conical foraminifera as well as the pore-fields - local concentrations of pores peneropliform spirocyclinids. in certain areas on the surface of the chamber The question arises: is there an evolutionary wall. Fig. 75 F-G. series of exoskeletonal development from pore-funnel [pore-pit] – the externally simple to complex, such as Praekurnubia (Late enlarged outlet of a pore (in planktic Middle Jurassic) with simple beams as foraminifera). exoskeleton, through Kurnubia (Early Upper Jurassic) with a comparatively simple pore-pit – see pore-funnel. polygonal network, to Rectokurnubia (Late pore plate [pore sieve-plate] – a minute, Upper Jurassic) with a deep and more complex microperforated, more or less calcified disk polygonal network. Similar evolutionary trends located in the pore tubulus at the level of a have been proposed for reticulinellids and

28 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02) distinct constriction which reflects the position or less tubular or flattened space between the of the primary organic sheet (median layer). umbilical plates and the wall of the preceding Fig. 75 A. coil or between plates, folia and the preceding coil; or between toothplates and the preceding pore plug – an organic structure plugging coil. Fig. 26. a pore funnel. It is produced by the coalescence of the organic cell envelope and progenitor – a direct ancestor. the organic median layer over the pore lumen. progressive chamber – a chamber with a May be partially calcified by minutely supplementary, retrovert aperture giving rise perforated platelets called abaxialpore sieve to a supplementary series of chambers (in plates. Fig. 75 A. architectural types with multiple spirals or in preseptal passage (preseptal canal, orbitoidiform growth following the annular passage auct.) - in porcelaneous development of spiral nepionts. fusiform shells an elongate, undivided space proloculus – the initial chamber of a beneath the septal wall containing the foraminiferal test without nepionic apertures. Fig. 79. differentiation. Usually. a proloculus has a primary - belonging to the last formed, i.e. spherical outline and a single aperture. ultimate chamber. protheca – the free chamber wall of primary aperture - see main cameral fusulinids composed of a tectum and a aperture; supplementary aperture; accessory diaphanotheca. aperture. protoconch – the first chamber of a test primary chamber - see chamber. with an embryonic apparatus or in which a deuteroconch is differentiated. In most primary foramen - see aperture. dimorphic larger foraminifera, the microspheric primary organic membrane (POM) - see generation has a proloculus, the megalospheric primary organic sheet and median layer. generation a protoconch or megalosphere. Fig. 41. Remarks: This term (BÉ et alii, 1980; protoforamen – an aperture or ANDERSON & LEE, 1991) is misleading and intercameral foramen to which a toothplate is should not be used at all in foraminiferology attatched. for the following reasons: The median layer is part of an outer, biomineralized cell envelope protoplasm – the living matter comprising and has nothing in common with the primary the cell-body. cell membrane (plasmalemma) of the protopore - comparatively narrow pores or foraminifer; neither their biochemistry nor parapores believed to represent early their geometry are comparable (see phylogenetic features. HOTTINGER & DREHER, 1974; LEUTENEGGER, 1977). Moreover, the acronym POM is used in proximal - nearer to the proloculus, oceanography for "particulate organic matter" contrary to the direction of growth. in the water column. Foraminiferal organic linings or its fragments are known to be one of proximal [posterior] chamber wall – the many components of particulate organic wall separating a chamber from the preceding matter, especially below the lysocline. The use one, formed by a septal flap or by a basal of this term or its acronym may cause layer, or (in part) by a strongly inflected confusion. lateral wall that together with the distal wall of the preceding chamber produces an intraseptal primary organic sheet (primary organic space. membrane auct.) - sheet of spongy organic material between outer and inner lamellae in pseudalveolar - having a simple bilamellar foraminifera. In calcitic foraminifera, exoskeletal structure consisting of alcoves, as the primary organic sheet is usually bounded the agglutinated test of Orbignya or Cubanina. by rhombic, paired crystals which are different pseudokeriotheca – a texture of external in size and shape from the stacks of platelets chamber walls in Mesozoic and later forming the major part of the mineralized agglutinated foraminifers consisting of layers. These crystals are interpreted as the uniform, parallel, radial elements covered by nucleation sites for the biomineralization of some kind of tectum. The interpretation of both inner and outer primary lamellae on an these textural elements as pore-like cavities in organic template. Some authors include them the shell and their delimitation from the in the so-called median layer. usually much larger and often more irregular primary plates - in orbitolinids: see parapores is not clear a present. exoskeleton. The term comprises beams and pseudoplanktic – a life habit: benthic rafters. forms emigrate into the planktic realm prior to primary spiral-umbilical canal – a more reproduction, a strategy to enhance the

29 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02) dispersion of a population. Pseudoplanktic radial zone – the annular zone in the foraminifera produce as a penultimate growth discoidal chambers of uniserial cones, where step a float chamber followed by a balloon radial septula subdivide the chamber into chamber with multiple apertures to release the radial compartments. See also marginal zone, hatching (Fig. 16). As fossils and in recent reticular zone. Fig. 20 & Fig. 71. sediments, pseudoplanktic foraminiferal shells radiate aperture – a single aperture, in are found in comparatively small numbers terminal or margino-terminal position with disseminated over all the ecological gradients radially directed, slitlike or pointed extensions. of the photic zone. The margins of radial apertures may fuse and pseudopodia - semipermanent or thereby subdivide the aperture, as in many temporary extrathalamous ectoplasmic nodosariaceans. projections. Foraminiferal pseudopodia form a radius (pl. radii) - in orbitoidiform reticular network (reticulopodia) reinforced by architecture: a multiplication of the main microtubuli. chamberlet layer in four or five sectors of the pseudopore - see parapore, pit. equatorial plane (as in Asterocyclina) that produces star-shaped tests. pseudorbitoid layer - see marginal canal system. rafter – a minor exoskeletal partition of the chamber lumen parallel to the chamber pseudospine (murica; spine, pars auct.) - septum and perpendicular to beams and the a pointed conical, or elongated spine-like, lateral chamber wall. Together with beams usually solid but sometimes hollow, inflational produces a subepidermal polygonal network. ornament feature. Compare: spine; acicular Fig. 47. spine; canaliculate spine. Remarks: The term rafter is a translation into pseudospinose - possessing English of DOUVILLÉ's term "poutrelle" (1906) pseudospines. restricted here however to minor elements, the major ones being called "poutres" in French pseudoumbilicus (false umbilicus, (or beams in English). DOUVILLÉ did not pars.auct.) - externally visible, extra-axial, distinguish major and minor partitions in his narrow or wide, cup-shaped space between an "réseau polygonal" or polygonal subepidermal infolded distal wall below main cameral network at that time. aperture and the adjacent coil, mimicking an umbilicus. Usually leads into an uncovered ramp – a linear surface in sections that spiral umbilical canal. Compare: umbilicus; traverse successive chambers with an umbilical bowl; umbilical depression. endoskeleton defined by crosswise-oblique stolon axes. The ramp effect is produced by punctate (punctuation) - see pits. the stolons alternating their inclination in pustule [tubercle; papilla] – a successive stolon planes of discoidal shells hemispherical to subconical inflational (Orbitolites) or cone mantels of uniserial cones protuberance of the outer lamella. See also: (Orbitolina). Fig. 47 H & Fig. 71. pseudospine. Fig. 73. rectilinear chamber arrangement - pycnotheca – in fusulinids: a uniform, chambers in a straight line. dense part of the septal wall below the tectum, reduction division - see meiosis. wedged between the keriotheca of two successive chambers. regeneration – repair of the shell after injury, first by closing chamber cavities and pylome – see aperture. subsequently by locally accelerated growth, pyrenoid – in foraminiferal symbionts a until a specific outer shape of the shell is more dense body rich in proteins in the chloroplast, or less perfectly restored. Shell fragments often surrounded by storage carbohydrates without an embryo may regenerate. This fact (starch). Seen in the transmission electron confirms that the nucleus of the cell moves microscope the shape of pyrenoids may be away from the center of the shell during diagnostic for the differentiation of groups of ontogeny. Fig. 78. algal endosymbionts. Fig. 30. reniform - kidney-shaped. Q residual pillar – an endoskeletal pillar quinqueloculine - see milioline coiling. supporting a frontal wall that spans a large preseptal space, frequent in trematophores. R Where septula and floors are interrupted by a preseptal space, residual pillars may remain radial – the direction from pole or axis between (horizontal) preseptal passages and toward any part of circumference of the test. (vertical) shafts in support of the frontal radial texture - see distinctly radial, chamber wall, as in elongate Praealveolina. indistinctly radial texture. Fig. 70 G & Fig. 82.

30 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02)

respiration – exchange of oxygen and agamont reproducing either by another carbon dioxide between an organism and its apogamous nuclear division and cytotomy (i.e. ambient environment. by distributing the mother protoplasm among the offspring) or undergoing meiosis. Because reticular zone – the center of discoidal the foraminiferal schizonts are produced by chambers in uniserial cones, where septula cytotomy, they are megalospheric (A - forms). narrowing towards the center fuse into a Fig. 5. reticular network in to minmize the volume of chamberlet cavities. See also marginal zone, scrobis septalis - see infundibulum. radial zone. Fig. 20 & Fig. 71 G-H. sealing plate (sealing-off plate; cover reticulate - having ornamental or other plate, pars auct.) - a thin plate, secondarily features arranged in a network. See: plugging the opening in an umbilical plate cancellate. which - primarily - provides communication between a main chamber lumen and a foliar reticulopodia - see pseudopods, chamberlet. Never present in the ultimate microtubules. chamber. Compare: cover plate. retral lobes - finger-like, hollow secondary - belonging to the penultimate extensions of the proximal chamber-wall (in or to earlier chambers if they differ from the the absence of an interlocular space). See last one. also: ponticuli; retral processes; basal lobes. secondary apertures - see supplementary retral processes - finger-like proximally apertures. directed extensions of the chamber lumen covered by ponticuli present at the margins of secondary lamination - see lamination. an intraseptal interlocular space. Fig. 54. secondary passage - see interseptal, retral [retrovert] stolon – in orbitoidiform interlocular space. foraminifera a stolon connecting main secondary septum - see septulum. chamberlet lumina with lateral chamberlets. It is positioned in the median plane of the secondary septulum - in chamberlet parallel to the shell axis and is neoschwagerinids: see rafter. directed backwards, feeding the lateral chamberlets corresponding to its cycle. Fig. secondary spiral umbilical canal - 36. tubular to flattened space comprized between cover plates and the lateral wall of the retroparies – see cover plate, umbilical adjoining previous coil. cover plate. selliform – the deformation of a discoidal retrovert foramen – a second, primary shell into a shape like a horse's saddel. foramen located at the base of the marginal Common in advanced Discocyclina ("D. sella") chamber suture, opening in a proximal but also to be observed in Orbitoides, direction and giving rise to orbitoidal growth Somalina or Eulepidina. Selliform orbitoidal after a spiral nepionic stage or to multiple shells produce characteristic sections (Fig. 76). spirals. Compare: supplementary apertures. Whether this deformation is a specific taxonomic character or a functional response reversed trochoid chamber to bottom currents remains an open question. arrangement - trochospiral arrangement in which the spiral side more involute than the septal face – that surface of a chamber- umbilical one. wall to be converted into a septum at a subsequent instar. rhizopodia - bifurcating and anastomosing pseudopodia, reinforced by microtubules. septal filaments - sutures of alar chamber extensions in involute nummulitids, often rim [peristomal rim] – the thickened meandrine. margin of an aperture. See also: lip. septal flap (paries proximus, pars auct.) – S that part of the inner lamella that covers the saddle - the distally directed, u-shaped preceding septal face. By its adherence to the embayment between retral lobes. septal face, the septal flap produces a trilamellar septum in a primarily bilamellar sagittal section – a slice through the test foraminifer. It may extend into an umbilical normal to the axis of coiling and passing plate, a foramenal plate, a bipartitor, a cover through the proloculus. plate or a toothplate. Fig. 53 F & Fig. 65 I-J. salients - rudiments of septa left over after the excavation of cuniculi (in fusulinids). Remarks: The term septal flap was used earlier for all parts of the proximal chamber sarcode - see protoplasm. wall, LÉVY's paries proximus (LÉVY et alii, schizont - apogamic offspring of an 1979). Here we restrict it to those areas of the

31 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02) proximal chamber wall that are glued to the "septulum" which is inconsistent with the face of the previous frontal chamber wall. This structural interpretation of all other area may be minimized to a narrow imperforate foraminifera. In French papers, hemicircular band around an interiomarginal where the orbitolinid mode of growth was foramen or reduced to sectors extending from correctly recognized as a uniserial stack of the foramen over the face of the previous chambers, we find the simultaneous use of chamber either in equatorial or in dorsal septum (for the true septum), cloison for direction. Free parts of the proximal chamber endoskeletal radial main partitions and wall are covered by outer lamellas like all free cloisonette (for beams in the exoskeleton). outer surfaces of the lamellar shell. The open septum (French: cloison) – a wall space between frontal and proximal walls of separating two consecutive main chamber subsequent chambers is called intraseptal lumina, i.e. the portion of the free chamber interlocular space. The line of adherence of the wall that is covered by subsequent chambers septal flap on the previous frontal chamber and thus incorporated in the architecture of wall delimits the intraseptal interlocular space the shell as a partition between successive in proximal direction and is in fact a deeply main chamber lumina. The connection sunken cameral suture. between them is assured by one or many septal fluting - see fluting. openings in the septum (intercameral foramina, stolon systems) that are in most septal foramen - see intercameral cases converted primary apertures. When foramen. multiple chamberlets form simultaneously the septal passage - in rotaliids: provides septum may consist of many discrete parts communication between the main the chamber acting as partitions between the lumina of lumen and the spiral canal. See: loop-hole. successive chamberlet cycles (not of neighbouring chamberlets). septal pore - primary small multiple apertures irregularly distributed on the sere (or series) – the complete sequence of apertural face (antetheca) of fusulinids. Fig. biocoenoses in a succession, from pioneering 66. stages to climax. septal suture – the line of adherence of a serial disposition of chambers (uniserial, chamber to the previous one. biserial, triserial etc.) – the regular arrangement of a small number of chambers in septular suture – the line marking the a trochospiral shell that will produce one, two, position of a septulum below a lateral chamber three or more rows of chambers in a regularly wall. superposed sequence in successive whorls. septulum (French: cloisonette) - Fig. 37 Endoskeletal, wall-like partition extending from sessile - permanently attached, usually the lateral wall into the chamber lumen, with the attachment surface on the dorsal dividing it into compartments (chamberlets). (spiral) side of trochospiral shells. Also In imperforate forms, these partitions are designates a sedentary life habit. produced by local thickening of the inner part of the shell wall. Their disposition has a close shaft – a preseptal space vertically relation to the arrangement of the foramenal connecting superposed preseptal passages in axes. In lamellar-perforate species, the septula elongate alveolinids. Fig. 70 G-H are produced by folded inner lamellas. Fig. 71. sieve plate – a calcified disk with minute

perforations closing the pore as a continuation Remarks: In describing alveolinid structures, of the median layer that separates primary there has never been any difficulty in inner and outer lamellas. Fig. 75 A. distinguishing between a septum (closing off a chamber) and a septulum (partitioning the sigmoid - s-shaped. chamber into chamberlets) ever since sigmoiline - see milioline coiling. CARPENTER (1862), DOUVILLÉ (1906) and REICHEL (1936-1937). In orbitolinids however, sinistral coiling - counterclockwise there is considerable confusion. CARPENTER direction of coiling as viewed from the spiral (1862) described "Patellina" (= Orbitolina) side. lenticularis with a "large chamber layer". This term corresponds in modern terminology to a sipho - term broadly applied both to single, discoidal (or annular) chamber strongly folded buliminid toothplates and to separated from the next "layer" by a septum. entosolenian tubes. R. SCHROEDER kept up the idea of the six - stolon system – the manner in which chamber layer to at least 1973 but abandoned each equatorial ogival or spatulate chamberlet it in his 1975 paper. The concept of a chamber of a main chamberlet layer is connected to its layer in orbitolinids leads to the interpretation adjacent chamberlets, in the same cycle by of the orbitolinid radial partition as "septum" annular passages and to the neighbouring two and of secondary, exoskeletal partitions as a chamberlets in the following and in the

32 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02) previous cycle by crosswise oblique stolons. spiral aperture - interiomarginal aperture The pattern may be duplicated or multiplied in along a spiral suture. Usually supplementary, successive stolon planes parallel to the not converted into a foramen. equatorial plane. spiral canals - see primary and secondary skeleton – all structural elements that umbilical spiral canals. supplement the primary chamber walls in spiral fissure - deep, circular, umbilical shaping permanently the protoplast. The three space separating ventral chamber tips or folia basic skeleton types, endoskeleton, from an umbilical plug, as in Ammonia. Fig. exoskeleton and supplemental skeleton may 77. occur in all possible combinations that together with chamber shape and chamber spiral interlocular space - space formed arrangement determine the architecture of the between adjacent coils along deeply sunken shell. The term should not be used as a spiral suture. See also: intraseptal interlocular synonym of test or of shell both of which space. Fig. 26. designate the total biomineralized cell envelope. Fig. 63. spiral side - that side of the test in trochospiral forms which contains the socculus (pl. socculi) - in porcelaneous proloculus. See also: dorsal. foraminifera low reliefs on the basal layer that do not touch the chamber roof. May form the spiral suture [whorl suture] – the line of pedestal-like base of pillars. On previous adhesion of adjacent whorls in spiral shells. apertural faces, low ridges may connect spiroconvex – a trochspiral shell with a neighbouring pillars or septula (Amphisorus). convex spiral side and a flattened to concave Ribs on basal layers may support pillars as in umbilical side. lacazinids. Socculi are a primarily internal feature of endoskeletal nature. They have to spiroloculine - see milioline coiling. be distinguished from ornamental elements on spirotheca – the free outer wall of the surface of chambers in previous whorls fusiform larger foraminifera, in particular of that have been covered by a chamber lumen fusulinids, that constitutes the chamber roof of the next whorl, such as the plugs or ridges and forms a spiral in equatorial section. on the apertural face in Amphistegina. Deposits on the chamber floor (tectorium, sphaeroconch - in agglutinated larger basal layer) of the next whorl are often foraminifera a spherical deuteroconch included in the term. The term's equivalent in enveloping a thin-walled, often poorly calcified discoidal-involute planispiral shells would be megalosphere. Usually possesses an "spiral sheet". exoskeleton, never an endoskeleton. Fig. 41. stellar chamberlet – an umbilical closed spicular wall – a test composed of segment of the chamber separated from the (secreted) fusiform calcite spicules. main chamber lumen by a folded inner lamella, the stellar septulum. Communicates with its spike - minute, conical to elongate spine- own main chamber lumen through a gap like projection on surfaces of the external wall between septulum and the adjacent coil and of lamellar foraminifera, not thickened by with the preceding chamberlet through the secondary lamination. Spikes occur often on umbilical part of the preceding foramen. Fig. the walls deeply inside interlocular spaces, as 48 K & M & Fig. 77 D. in the fossettes of larger elphidiids, to fend off larger particles such as diatom frustules stellate - star-shaped. transported with the food into the canal stolon – a tubular opening in a chamber system by the pseudopods. wall whose length is greater than its diameter, spine [acicular spine] – a calcite rod forming an intercameral foramen that permits normal to the test surface, thin, round, communication between consecutive chambers triangular to triradiate in section, running or between cyclical or subsidiary chamberlets through a hole in the outer lamella of planktic of one or two consecutive instars. Fig. 47 A-D. foraminifera. Arises apparently from the stolon axis - the axis common to stolons median layer of the chamber wall. At its base when they are aligned in succeeding septa. it is surrounded by a more or less conical mound, the spine base. Acicular spines are stolon plane - plane defined by stolons shed during the planktonic life cycle, when that are regularly arranged in layers. Fig. 80. descending in the water column prior to stolon system – the geometric disposition reproduction. of stolons in regular patterns. spine-base - see spine. stratophenetics – the reconstruction of spinose - possessing true, acicular spines phylogenetic relationships based on (in planktic foraminifera). See also: morphological (phenetic) similarity on one pseudospinose. hand and on the other by the time

33 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02) relationships provided by biostratigraphic any kind of structural element subdividing ranges, in contrast to cladistics. The most external (lateral) parts of chamber lumen. May common procedure in the construction of the be of exoskeletal (beams and rafters) or phyletic lineages of foraminifera. For backup endoskeletal (septula) nature. theory see GINGERICH, 1990. subsidiary chamberlets (secondary streptospiral arrangement - coiled in chamberlets, auct.) - subdivisions of main successively changing planes, like a ball of chamber lumen by folded inner lamella with twine. See also: milioline coiling.. Fig. 37.10- the primary organic sheet between folds, or by 11. septula. See also: stellar chamberlets. striae - thin costae. succession [ecological succession] – a gradual change over time, in any one area, of striate - having striae. the composition of the community through structure - of foraminiferal shells: a three- interspecific competition and coexistence, from dimensional design that defines the the arrival of pioneers in an empty habitat to a morphology of chamber cavities as patterns mature, equilibrated community, termed that are repeated in successive chambers or climax, when equilibrium with long-continued, chamberlet cycles. stable environmental conditions has been Remarks: It is recommended that the term achieved. Periodic disturbances in the "structure" be employed in a very precise and environment may shorten the length of the somewhat restricted way, i.e. that it not be succession to early phases of the process used to denote patterns unrelated to the (disclimax). shaping of chamber lumina, the wall sulcus – a peripheral infold of primary "textures". The term "architecture" should be chamber-wall, always imperforate. May or may regarded as having a broader connotation, the not have radial passages between the combination of textural and structural design underlying chamber-lumen and the ambient with chamber shape and arrangement in the environment or the interlocular space. May or entire test. "Architecture" means the complete may not be covered by additional marginal set of complex but highly diagnostic characters structures, such as a marginal cord. Fig. 7 F-G defining the taxa on the generic level. & Fig. 64. style – a massive, imperforate columnar Remarks: REVETS (1989, 1993) uses the term structure between lateral walls supporting sulcus for the adapertural depression between expanse chambers that occupy a wide area, as the apertural rim and the attached part of the in Homotrema. toothplate. This depression is positioned near or in the axis, not at the periphery of the spiral subembryonic chamberlets – shell, and should therefore be distinguished chamberlets that are produced in a mono- or from the nummulitid sulcus by a separate plurilocular third stage in the growth of term, the "adapertural depression". megalospheric embryos of agglutinated conical foraminifera. They are located below the supplemental foramen – an orifice proloculus in the cone axis and are subdivided produced by a strongly folded toothplate within by structural elements of uncertain, probably a protoforamen. May be completely discrete exoskeletal origin. Fig. 41. from the protoforamen, as in Remarks: J. HOFKER Jr (1963, p. 211, fig. 14) Siphogenerinoides. interpreted the life habit of an orbitolinid as supplemental skeleton – the imperforate face upward, i.e. the cone apex downward, refolds and flying covers produced by outer buried in the sediment. Therefore, he called lamellae that cover and/or restrict interlocular the third growth-stage in Orbitolina s.str. spaces to form enveloping canal systems, "epiembryonic". The confusion up-down is canaliculate spines, marginal crests and complicated by a supposed error in the legend marginal cords and the perforate chamberlets of fig. 2 in J. HOFKER Jr (1966), where that are fed exclusively by canal orifices and deuteroconch and epiembryonic chambers are that overgrow in more or less regular layers or reversed by comparison with his 1963 paper. tires the canalicular structures. The cavities of Today, all foraminifera with an extensive the supplemental skeleton can not be assigned apertural face covered by numerous apertures to particular stages of growth because there is live with their apertural face towards the no direct connection with the orderly cameral substrate. Undoubtedly, the same is true for system of shell cavities. In shells with an conical foraminifera in general. The term extensive supplementary skeleton, the cameral epiembryonic is therefore to be avoided. cavity system is reduced to neanic or even DOUGLAS (1960) did not distinguish between nepionic stages. Fig. 65. sub- and periembryonic chambers, both Remarks: Introduced by CARPENTER (1862) representing the third growth stage. See also: long before lamellar theory was developed, periembryonic and supraembryonic. and as now refined here, the term is a subepidermal partition (-plates, - welcome complement to the terms exoskeleton lamellae) - unspecified, descriptive terms for and endoskeleton. Both of these terms

34 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02) subdivide the chamber lumen while the shell on its substrate. "supplemental skeleton" structurizes sutural canals - openings to the exterior extralocular, "outer" space. It is meant to be of an intraseptal interlocular space whose used as generic term regrouping all forms that margins are partly closed by the local adhesion have canaliferous enveloping, marginal and of consecutive chamber walls. See also: pseudospinose structures like Siderolites, fossettes. Pellatispira, Calcarina and their allies along with all types of marginal cords (linked to a sutural supplementary apertures - single sulcus) as in Sulcoperculina and their additions to primary apertures in ventral or derivatives, Ranikothalia and all nummulitids dorsal sutural position that are not s.str. transformed into an intercameral foramen at the next instar. See also: supplementary supplementary aperture(s) (secondary apertures. aperture(s), pars. auct.) - primarily formed openings either in the apertural face suture – the line of adhesion of chamber ("apertural pores") or (slit-like) in a sutural wall(s) to the previously formed test. position, always in addition to a main cameral aperture. Sutural supplementary apertures are symbiont – an organism living together not converted into intercameral foramina with or within an other organism to the benefit because of their position and thus apparently of both. do not serve for passage of functional symbiosis - in foraminifera: algal cells endoplasm between chambers. The same living (as symbionts) within the foraminiferal seems to be true of some multiple cytoplasm in a mutualistic relationship with supplementary apertures, which - although their host. The symbionts actively situated in the septum - may be absent in photosythesize and reproduce asexually in the earlier chambers and/or may be plugged in host cell. They are engaged in recycling part at a subsequent instar. See also: nutrients. They live either in vacuoles of the accessory apertures. host cytoplasm and are displaced passively by supplementary chamberlet - a cavity in the host's protoplasmic streaming, or are the supplementary skeleton that is bounded by found in the lacunar system of the host cell a bilamellar, perforate wall in the direction of within which they may move actively using the ambient environment of the shell when it their shortened flagella to regulate the amount forms. It may be overgrown in later of their irradiation by sunlight so as to avoid ontogenetic stages by subsequent outer photoinhibition. During the asexual lamellas or by additional elements of the reproduction of the host, each offspring supplementary skeleton. Fig. 63 A & Fig. 65 F. inherits a small number of symbionts from the mother cell. But after sexual reproduction, the supplementary spirals – see multiple foraminiferal zygote must take up symbionts spirals. from its ambient environment. See also: chloroplast husbandry. Fig. 30. supraembryonic area – in advanced orbitolinids a circular area at the apex of the sympatric - inhabiting a common area or shell above the megalosphere, formed by a largely overlapping areas of distribution. subdivided annular deuteroconch. Compare: allopatric. supraembryonic chamber – a more or synonym – a different name for any one less hemispherical deuteroconch in apical taxon. May be invalidated by the priority of the position, embracing from above an often valid name (junior synonym). The names of incompletely calcified protoconch bearing erroneously identified other valid taxa also exoskeletal elements, as in Orbitolina s.str. appeare in synonymies. Fig. 41. Remarks: This term was introduced by synonymy list – a list in their order of DOUGLAS (1960) together with the term time of their publication of bibliographic "periembryonic" for the third (and partially references to previous descriptions and/or forth) growth stage, below the proloculus, named illustrations of the taxon considered implying a life position face downward. identical with the taxon being treated and thus SCHROEDER (1962, 1973) and later ARNAUD- a part of the hypodigm. VANNEAU (1980) attributed much phylogenetic Remarks:. The synonymy list is an weight to a distinction between supra- and indispensable instrument in taxonomic work periembryonic chambers. justifying the identification of taxa and eventually the creation of new ones. A surface of attachment – in permanently synonymy list should refer to the type attached (sessile) shells the surface that is description and in addition reflect the worker's fixed to and casts the substrate. Tiny own opinion on the taxon, which may be supplementary sutural apertures in the surface supplementd where necessary by particular of attachment may indicate that the cell comments. In most cases completeness of produces some kind of organic glue to stabilize synonymy lists is of secondary importance.

35 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02)

Copying synonymy lists of previous authors is chemical composition of test-walls. Compare: useless; they may be cited as a block ("with texture. synonymy") if the researcher agrees with the test-structure – any repeated pattern of opinion of the previous author in all cases. the elements that subdivide chamber lumina. Disagreements may be listed explicitly under the heading "non". Doubts or approximations test [wall]-texture - pattern of of the identity may be expressed by the terms arrangement of crystallites, agglutinated "affinis" (abbreviated aff.) and "confer" (cf.). grains, organic matter, pores, lamination or The former means "near to but not identical", layering. the latter expresses some doubts about the identification, useful in cases where not all textural ornamentation – the pattern diagnostic characters can be identified. generated by a regular grouping of pores or Qualified synonymy lists help to enhance and other textural elements on the surface of a deepen the concept of the taxon under shell. Fig. 75 F-G. consideration, and expand its documentation. thylacoid – the membrane-bounded, much The synonymy list facilitates the establishment compressed sac ocurring alone or associated in of the taxons' range in morphological stacks in the chloroplasts of vegetal cells and variation, space and time and permits factual in particular in the symbionts of the correction of a previous author's work, thus foraminifera. Traps sunlight as the source of avoiding unnecessary emotions. energy for the synthesis of sugars. Fig. 30. syntype – see type. tongue - see toothplate. T tooth (pl. teeth, French: dents) - inward projection(s) of the inner portion of the tectorium - in fusulinids: a slightly chamber wall into the aperture. This structural transparent internal layer of the shell that lines element is a continuation of the basal layer the chamber walls and is covered by the and may be more or less modified but not opaque tectum in the external, spiral wall. May suppressed when the aperture is transformed be combined with a diaphanotheca. into an intercameral foramen. Teeth growing tectum – in fusulinids: a thin, dense outer out from the basal layer may be (extern) layer of the spirotheca (spiral outer complemented by local thickenings of the inner wall), homologous in position to an epiderm portion of the free, marginal chamber walls. but possibly produced by a discrete shell- building process. May bear tiny, pore-like gaps Remarks: Teeth may be defined as discrete permitting replication of parakeriothecal endoskeletal elements restricted to the elements like chomatal pores in a superposed apertural area. The relationships between shell layer. See also: marginal prolongation. teeth and pillars (in particular those supporting a trematophore) on one hand, to valvular template – a sheet of protein substances teeth in agglutinated and to toothplates in on which biomineralization initiates. It governs lamellar-perforate foraminifera on the other is the pattern, shape and size of the a close one and may be transitional. True biomineralized chamber wall. During chamber teeth must be distinguished from tooth-shaped growth, the template is put in position by the masks in Borelis obstructing the main brush-like pseudopodia prior to the apertures but resorbed totally in intercameral biomineralization of the wall. See also: median foramina. See also: milioline tooth. layer. toothplate (sipho; central pillar, pars teratological – a pathological alteration of auct.) - a contorted plate running from an shell morphology, for example a loss of control intercameral foramen to an aperture, and that maintains axes of coiling or bilateral attached to both. It may be shaped to form a symmetry consistant. These aberrations are single, double or spiral fold (or "tongue") with common in gerontic growth stages or after a free, often serrated distal end and distally temporary extreme environmental conditions protruding into the aperture. A toothplate in tidal pools. separates partly or completely the main terminal - positioned at the distal end of a chamber lumen from an axial space linear structure or of an elongate chamber. (adapertural depression) in post-embryonic stages. It protrudes with a free edge distally test – the shell or skeletal components of a and adaxially into the aperture. Interconnected foraminifer. The test may be composed of a toothplates in low-trochospiral umbilicate variety of materials: secreted, agglutinated or shells may produce a primary spiral canal. A in combination. toothplate is never associated with a foliar or test-architecture – the spatial stellar chamberlet. The use of the term in low- arrangement of chambers, their subdivisions trochospiral rotaliid forms is under discussion and their connections. (see REVETS, 1993). test-composition – the mineralogical and topotype – see type.

36 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02)

trabecules [trabeculae] - imperforate shell represent haploid gamonts reproducing material extending from an imperforate sutural sexually. Thus, three different phenotypes zone into the perforate lateral chamber-wall would represent a (trimorphic) species. See and housing oblique, ramified trabecular also: alternation of generations. Fig. 5. canals opening between the pores on the triserial - chamber arrangement in a surface of the lateral chamber wall (Fig. 81). trochospire with three chambers per coil,

hence with about 120° between the median Remarks: Trabecules are the result of a planes of consecutive chambers. See serial deviation of pores from their parallel paths in disposition of chambers. the lateral chamber wall, to create - in a direction perpendicular to the septum - a V- tritoconch - the third chamber of the shaped zone of imperforate wall without megalospheric embryo of Miniacina. It has inflational deformation of the outer lamellae. multiple apertures that feed a more or less In contrast to ordinary elements of concentric chamberlet cycle of the nepionic ornamentation, they house a trabecular canal stage of growth. with a diameter only slightly larger than that of the pores, and therefore difficult to see in trochospiral arrangement - chamber fossil material. The trabecular canals take off arrangement in whorls or coils where the rate from the lateral intraseptal canal in a proximal of translation (net rate of movement along the and/or distal direction and represent therefore growth axis to the net rate of movement away an oblique extension of the interlocular space - from the axis) is more than zero. Spiral and without folding of the septal flap as in umbilical sides are dissimilar. May be involute Planoperculina - into the stack of outer or evolute on either the spiral or the umbilical lamellas. So far, trabeculae have been side. See also: reversed trochoid. observed only in some genera of the tube pillars - hollow pillars formed by a Nummulitinae, in particular in Nummulites, folded septal flap, as in Chapmanina. whereas they are absent in Assilina. tubercle - see pustule. transverse septulum - see beam. tuberculate, papillate, pustulate - covered trematophore – a sieve constituting the with tubercles, papillae or pustules. See also: face of many porcelaneous larger foraminifera, pustule. in miliolids produced by the coalescence of teeth, covering a large preseptal space. May tubulin – a globular protein molecule be supported by residual pillars. This forming the subunits to be polymerized to construction is in contrast to the multiple microtubules. apertures produced by the coalescence of tubulopore – a pore opening at the end of peristomal rims, as in Coscinospira. Fig. 82. a conical or tubular projection. trichome – leaf hair (of seagrasses for tubulospines, tubulospinate - hollow instance). May be casted by the chamberlet pseudospines. The cavity is a linear extension walls of foraminiferal epiphytes growing over of the chamber lumen that ends blindly below the leaf surface to strengthen the adhesion of the tip of the pseudospines. Caution! Many the epiphyte to its substrate. Fig. 42 pseudospines have been erroneously triconch – the first three chambers in a interpreted as hollow (as in Asterorotalia megalospheric generation separated by plane, pulchella). uncurved septa. These are shaped by an tunnel – An intercameral foramen in an equilibrated hydrostatic pressure, probably interiomarginal-basal position bordered by during a single instar. May be enveloped by endoskeletal structures narrowing the common secondary lamellas, for example in communication between the open chamber Planorbulinella. Fig. 60 B. lumen and the spiral space extending through Remarks: DROOGER (1993) calls this feature a the successive tunnel foramina. The tunnel is tritoconch, a term which, however, is produced by resorption of parts of an apertural preoccupied for the third chamber in face and/or of an apertural mask. May be megalospheric Miniacina. See also: biconch. multiplied to form a single row of basal triloculine - see milioline coiling. foramina in fusiform shells. Fig. 31, Fig. 32 A- D & Fig. 66. trimorphism – a morphologic Remarks: Introduced originally for fusulinids, differentiation of the megalospheric generation where the tunnel is bordered by chomata and in A1 and A2. A1 has a comparatively small may be multiplied - together with the megalosphere and reaches larger adult shell endoskeletal elements, the parachomata - the sizes than A2. The A1 shells are interpreted term has been extended to pfenderinids (HOFKER, 1968) as representing diploid (HOTTINGER, 1978). In this family the tunnel schizonts generated by the microspheric is bordered by being incised in a columellar agamont. The A2 shells, after the reduction endoskeleton and may be multiplied as division of the reproductive nuclei in A1, would multiple, parallel incisions in such complex

37 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02) genera as Sanderella. In addition, the term is U extended here to nummulitids where the tunnel is bordered by a pair of umbilical plates ultimate chamber – the last chamber in much the same position as the chomata in formed in an individual. fusulinids. In nummulitids, no multiplication of umbilical [intraumbilical] aperture – a tunnels has been observed so far. primary aperture of a chamber leading into an type – in taxonomy: the specimen or taxon umbilicus. of next lower rank designated to be always umbilical bowl (pseudoumbilicus, pars included in the respective taxon and to be auct.) - a deep, wide or narrow conical space excluded from neighbouring taxa, in what ever in axial position formed between inner way that taxon might be defined and/or umbilical chamber walls, wherever the latter delimited by subsequent authors. Several are separated from the outer umbilical walls by kinds of types are distingushed: holotype: the a distinct edge or shoulder. Compare: single specimen showing all characteristics umbilicus; pseudoumbilicus. Fig. 12 C. considered relevant to its identification and segregation at the time of its designation; umbilical canal system - umbilical syntypes: several specimens of the type interlocular space transformed into tubular population that together show all characters cavities by various skeletal elements and considered relevant at time of designation; chamber wall extensions (folia). Commonly cotype: an additional type specimen in modified by local resorption to create a support of the holotype; paratypes: network of communications between the specimens designated as such in addition to a tubular cavities. See also: spiral canal; funnel. holotype and considered important for the umbilical cavity - the axial complex of definition of the species' variability; topotype: interconnected passageways delimited by axial all specimens of the type population; chamber walls, inner umbilical walls, folia, lectotype: a single specimen selected from a foramenal plates, and cover plates. Includes series of paratypes, if a holotype has not been thus the umbilical canal systems. May be designated; neotype: a specimen, selected if restricted by piles or plugs and communicates possible from the original population, to be with the exterior through foliar apertures or designated as type if the original holotype has vertical canals. been lost; plesiotype: illustrated specimen used for justification of an identified taxon or umbilical depression (umbilicus, pars in support of a redescription of a valid taxon auct.) - a closed depression in axial position and deposited in a public collection; formed by the curvature of the umbilical generotype: type species of a genus. chamber-walls in the same coil. Compare: Remarks: Types play an important role in the umbilicus; pseudoumbilicus). revision of fossil taxa. The type specimens of umbilical flap - rotaliellid extension of the valid species must be deposited in a collection umbilical chamber wall deliminating a narrow accessible to the public. However, in most umbilicus and covering an umbilical aperture. cases, the type specimens are not available for The anterior margin may be glued to the further study by any invasive method (coating umbilical wall of chambers in the previous for Scanning Electron Microscopy, sectionning whorl. for structural and geochemical analysis etc.). Remarks: As the term "umbilical flap" was Therefore revision of a species must be used earlier as a synonym of "umbilical plate" focussed on topotypes. Consequently we in rotaliids, its revival for umbilical chamber recommend, in the proposal of new taxa, that wall extensions in Rotaliellidae (by not only the types but also as many topotypes PAVLOWSKI et alii, 1992, fig. 1) leads to as possible be deposited for further reference. confusion and should be abandoned. Instead, Fossil specimens considered to represent the umbilical chamber wall extensions in original (type-) population come from the Rotaliellidae should be compared to structures same field sample or from the same bed at a in planktic foraminifera with comparably open, particular (type-) locality, according to true umbilici and so be designated with circumstances. The use of syntypes may be corresponding terms. justified when the taxon is based exclusively on specimens in cemented rock studied by thin umbilical plate (foramenal plate; umbilical sections. In this case, some diagnostic flap; murus reflectus; toothplate; paries features may appear only in sections of proximus, pars auct.) - a more or less specific orientation. Otherwise, the use of the contorted plate-like test element, extending various kinds of types is to be limited to the between distal and proximal chamber walls curating of collections in museums. All and joined to both, attached to the specimens considered by an author to belong intercameral foramen and to the main to a taxon are part of the hypodigm, including aperture, but not protruding into the latter. the specimens of previous authors listed in the Separates the main chamber lumen from a synonymy. primary umbilical-spiral canal. Between plate and adjacent coil or within the plate itself an

38 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02) opening provides connection between chamber crystallites, an integral constituent of the outer and foliar chamberlet, wherever present.This lamella in bilamellar foraminifera. opening may remain open in all chambers or it venter – that part of free benthic shells, may be closed in all but the ultimate chamber particularly if flattened, facing the substrate. by a sealing plate. An umbilical plate may be See also face and surface of attachment. single or composed of two symmetrical branches in some planispiral genera, thereby ventral - the side of a flattened organism producing one or two umbilical-spiral canals turned to its substrate, as opposed to dorsal. between plate and adjacent coil. Fig. 7 F-G, Secondarily flattened, almost planispiral or Fig. 54 H & Fig. 63 D. slightly reversed-spiral, involute shells like Daviesina salsa (DAVIES et PINFOLD) or D. umbilical plug (-pile, -mass, umbonal langhami SMOUT reveal their trochospiral plug) – a pile of lamellae forming a solid, more phyletic origin by an asymmetric, ventral or less free-standing plug in the center of the position of the main cameral foramen and of umbilicus, often separated from foliar tips by a the umbilical plate. See also: umbilical side; spiral fissure. Fig. 77. spiral side; remarks to dorsal. umbilical primary aperture - see vertical canals (oblique canals, pars auct.) umbilical aperture. – see funnel. umbilical shoulder - see umbilical bowl. vestibule (Vorhof in German) – a umbilical side - in trochospiral tests the deuteroconch that embraces a protoconch side opposite to the spiral one. See also: including its wide-open flexostyle with a ventral. hemicylindrical to almost cylindrical frontal wall bearing numerous apertures, as in umbilical teeth - triangular modifications Amphisorus and Marginopora (LEHMANN, of the lip over umbilical apertures, as in 1961). Globoquadrina. vicarious species - closely related, even umbilicate - possessing a true umbilicus sister species, occupying identical or very on one or both sides (biumbilicate). similar niches ("ecological substitutes") in umbilicoconvex - in trochospiral shells: separate regions. spiral side flattened to concave, umbilical side vortex - a helicoidal extension of many convex. Compare: spiroconvex. consecutive alar prolongations spirally twisted umbilicus – the axial space in spiral around the coiling axis of a planispiral- foraminifera communicating directly through lenticular shell. There are transitions to apertures with surrounding main chamber meandrine structures. See also: polar torsion, lumina or foliar chamberlets. May be open or the equivalent in fusiform shells. restricted by an umbilical plug. (Compare W umbilical bowl; pseudoumbilicus; umbilical cavity; umbilical depression). whorl [coil] – in a spiral test, a single turn or volution through 360°. umbo (central pillar, Zentralpfeiler of German authors on nummulitids) – an Z expanding pile of thickened lamellae in an axial position in involute or orbitoidal zygote – a diploid cell resulting from fusion foraminifera. An umbo is never associated with of two (haploid) gametes in sexual an open umbilicus or with spiral umbilical reproduction. The biomineralized envelope of canals. See also: pile; plug. Fig. 77. the zygote is called a microsphere. See also: life cycle. unilocular (monolocular, monothalamous) - single- chambered. Acknowledgments uniserial - chambers arranged in a single This glossary has a long history going back row. Compare: biserial; triserial. See serial to international courses on larger foraminifera disposition of chambers. held during the 1990's as part of the European project COMMETT. At that time, Z. REISS V (* 1917 - † 1996) suggested the use of the vacuolar system (in Monolepidorbis) - see rich illustrations of the monograph on the lateral chamberlets. recent foraminiferal fauna of the Gulf of Aqaba (HOTTINGER et alii, 1993) as a support of the valvular tooth - in agglutinated glossary attached to this work. foraminifera: a flap-like extension from the With the help of V. SCHEURING (Basel) this distal margin of a main aperture, partly glossary was amplified to include fossil taxa for restricting it. subsequent COMMETT courses without veneer - outermost array of more or less illustrations. Most kindly, J. LIPPS put the blocky or columnar calcite or aragonite unillustrated glossary on the web in Berkeley.

39 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02)

The staff and many participants in the Téthys: l’Urgonien du Vercors septentrional COMMETT courses took an active part in et de la Chartreuse (Alpes occidentales).- discussing the meaning of terms significant in Géologie Alpine, Grenoble, Mémoire 11, 3 analyses of the comparative anatomy of the vols., 876 p. + 115 pls. foraminiferan shell, namely A. ARNAUD BANNER F.T. (1966).- Morphology, (Grenoble), D. BASSI (Ferrara), E. CAUS classification and stratigraphic significance (Barcelona), K. DROBNE (Ljubljana), C. of the Spirocyclinidae.- Voprosy FERRÀNDEZ (Barcelona), H. FORKE (Berlin), Mikropaleontologii, Moskva, N° 10, p. 201- M. LANGER (Bonn), U. LEPPIG (Freiburg i. 224 + 20 pls. (in Russian) Br.), J. PIGNATTI (Rome), J. REICH (Basel), BANNER F.T. & HODGKINSON R.L. (1991).- A E. VECCHIO (Naples) and E. VILLA OTERO revision of the foraminiferal subfamily (Oviedo). Heterostegininae.- Revista Española de The illustrations of the glossary presented here Micropaleontologia, Madrid, vol. XXIII, N° include a certain number of unpublished 2, p. 101-140. plasticine sculptures made by M. REICHEL BARKER R.W. & GRIMSDALE T.F. (1936).- A (* 1896 - † 1984) in the nineteen-forties and contribution to the phylogeny of the which since that time have been used for orbitoidal Foraminifera, with descriptions of instruction. The TEM graphs, mostly new forms from the Eocene of Mexico.- unpublished, are the work of S. REBER- Journal of Paleontology, London, vol. 10, N° LEUTENEGGER (Sissach). The SEM graphs 4, p. 231-247. were made in the SEM lab of the University of BÉ A.W.H., HEMLEBEN C., ANDERSON O.R. & Basel, at the time directed by R. SPINDLER M. (1980).- Pore structures in GUGGENHEIM (Basel). Most fossils used to planktonic Foraminifera.- Journal of illustrate this glossary were either donated to foraminiferal Research, Lawrence, vol. 10, the Geological Institute by F. ALLEMANN, M. N° 2, p. 117-128. CHATTON, Z. REISS and M. WANNIER were BENDER H. (1989).- Gehäuseaufbau, collected by the staff of the Geological Gehäusegenese und Biologie Institute after 1940 when courses on agglutinierender Foraminiferen (Sarcodina: micropaleontology began at the University. Textulariina).- Jahrbuch der Geologischen Most of this material is now deposited in the Bundesanstalt, Wien, vol. 132, N° 2, p. Museum of Natural History in Basel. 259-347. With trying efforts N. SANDER reviewed and BENDER H. (1995).- Test structure and polished the English text while B. GRANIER classification in agglutinated Foraminifera. edited the electronic version and contributed In: KAMINSKI M.A., GEROCH S. & many good ideas as to the presentation of the GASINSKI M.A. (eds.): Proceedings of the illustrations. D. BASSI and an anonymous Fourth International Workshop on reviewer gave their opinion on the texts. My Agglutinated Foraminifera.- Grzybowski sincere thanks to all of them, and also to the Foundation, Krakow, Special Publication 3, many students to whose questions I had no p. 27-70. immediate answer. BILLMAN H., HOTTINGER L. & OESTERLE H. (1980).- Neogene to Recent Rotaliid References Foraminifera from the Indopacific Ocean; ADAMS C.G. (1968).- A revision of the their canal system, their classification and foraminiferal genus Austrotrillina PARR.- their stratigraphic use.- Schweizerische Bulletin of the British Museum (Natural Paläontologische Abhandlungen, Basel, N° History), London, (Geology series), vol. 16, 101, p. 71-113. N° 2, p. 71-97. BRADY H.B. (1884).- Report on the AGUILAR M., BERNAUS J.M., CAUS E. & Foraminifera dredged by H.M.S. Challenger HOTTINGER L. (2002).- Lepidorbitoides during the years 1873-1876.- Reports of minima DOUVILLÉ from Mexico, a the Scientific Results of the Voyage of foraminiferal index fossil for the H.M.S. Challenger (Zoology), 9, 814 p. Campanian.- Journal of foraminiferal CARPENTER W.B., PARKER W.K. & JONES T.R. Research, Lawrence, vol. 32, N° 2, p. 126– (1862).- Introduction to the study of 134. Foraminifera.- Ray Society, London, 319 p. ANDERSON O.R. & BÉ A.W.H. (1978).- Recent + 22 pls. advances in foraminiferal fine structure CIFELLI R. (1962).- Morphology and structure research.- In: HEDLEY R.H. & ADAMS C.G. of Ammonia beccarii (LINNÉ).- (eds.), Foraminifera. Volume 3.- Academic Contributions from the Cushman Press, London, p. 121-202. Foundation for Foraminiferal Research, New ANDERSON O.R. & LEE J.J. (eds.) (1991).- York, vol. 13, p. 119-126. Biology of Foraminifera.- Academic Press, COX P.T. (1937).- The genus Loftusia in South London, 368 p. Western Iran.- Eclogae geologicae ARNAUD-VANNEAU A. (1980).- Helvetiae, Basel, vol. 30, N° 2, p. 431-450. Micropaléontologie, paleoécologie et DAVIES L.M. (1930).- The genus Dictyoconus sédimentologie d'une plate-forme and its allies: a review of the group, carbonatée de la marge passive de la together with a description of three new

40 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02)

species from the lower Eocene beds of der polythalamen Foraminifere Rotaliella northern Baluchistan.- Transactions of the heterocaryotica.- Archiv für Protistenkunde, Royal Society of Edinburgh, vol. 56, p. 485- Jena, vol. 100, N° 2, p. 211-235. 505. GRÖNHAGEN D. & LUTERBACHER H.-P. DAVIES L.M. (1932).- The genera (1966).- Beobachtungen an den Dictyoconoides NUTTALL, Lockhartia nov. Foraminiferen-Gattungen and Rotalia LAMARCK: Their type species, Pseudotextulariella und Textulariella sowie generic differences and fundamental verwandten Formen.- Eclogae geologicae distinction from the Dictyoconus group Helvetiae, Basel, vol. 59, N° 1, p. 235-246. forms.- Transactions of the Royal Society of HAMAOUI M. & FOURCADE É. (1973).- Révision Edinburgh, vol. 57, p. 397-428. des Rhapydionininae (Alveolinidae, DAVIES L.M. (1939).- An early Dictyoconus Foraminifères).- Bulletin du Centre de and the genus Orbitolina: their Recherche Pau - SNPA, vol. 7, N° 2, p. 361- contemporaneity, structural distinction and 435. respective natural allies.- Transactions of HANSEN H.J. (1999).- Shell construction in the Royal Society of Edinburgh, vol. 59, p. modern calcareous Foraminifera.- In: SEN 773-790. GUPTA B.K. (ed.), Modern Foraminifera.- DE CASTRO P. (1990).- Osservazioni Kluwer Academic Publishers, Dortrecht, p. paleontologiche sul Cretacio della località- 57-70. tipo di Raadshovenia salentina e su HENSON F.R.S. (1947).- New Trochamminidæ Pseudochubbina n. gen.- Quaderni dell' and Verneuilinidæ from the Middle East.- accademia pontaniana, Napoli, N° 10, 116 The Annals and Magazine of natural History, p. + 41 pls. London, (Ser. 11), vol. XIV, N° 117, p. DOUGLAS R.C. (1960).-. The foraminiferal 605-630 + 5 pls. genus Orbitolina in North America.- HENSON F.R.S. (1948).- Larger imperforate Geological Survey Professional Paper, Foraminifera of south-western Asia. Washington, 333, 52 p. + 14 pls. Families Lituolidae, Orbitolinidae and DOUVILLE H. (1906).- Évolution et Meandrosipinidae.- British Museum (Natural enchaînements des Foraminifères.- Bulletin History), London, 127 p. + 16 pls. de la Société géologique de France, Paris, HOFKER J. (1951).- The Foraminifera of the (4éme série), tome 6ème, p. 588-602 + pl. Siboga expedition. Part III. In: Siboga- XVIII. Expeditie.- Uitkomsten op Zoologisch, DROOGER C.W. (1993).- Radial Foraminifera; Botanisch, Oceanographisch en Geologisch morphometrics and evolution.- Gebied, Leiden, Monographie IVa, 513 p. Verhandelingen der Koninklijke Nederlandse HOFKER J. (1956).- Foraminifera dentata. Akademie van Wetenschappen, Afd. Foraminifera of Santa Cruz and Thatch- Natuurkunde, Eerste Reeks, Amsterdam, Island Virginia-Archipelago West-Indies.- deel 41, 242 p. Skrifter udgivet af Universitetets Zoologiske ERSKIAN M.G. & LIPPS J.H. (1987).- Museum, Copenhagen, XV, 237 p. Population dynamics of the foraminiferan HOFKER J. (1968).- Studies of Foraminifer. Glabratella ornatissima (CUSHMAN) in Part I. General problems- Publicaties van Northern California.- Journal of het Natuurhistorisch Genootschap in foraminiferal Research, Lawrence, vol. 17, Limburg, Maastricht, Reeks XVII, Aflevring N° 3, p. 240-256. 1-2, 135 p. FERRÀNDEZ I CAÑADELL C. (1999).- HOFKER J. jr (1963).- Studies on the genus Morfoestructura i paleobiologia dels Orbitolina (Foraminiferida).- Leidse Ortofragmínidos de la Mesogea Geologische Mededelingen, Leiden, vol. 29, (Discocyclinidae i Orbitocyclinidae, p. 181–253 + 23 pls. Foraminifera).- Arxius de les Seccions de HOFKER J. jr (1966).- Studies on the family Ciències, Barcelona, (Secció de Ciències i Orbitolinae.- Palaeontographica, Stuttgart, Tecnologia), CXXI, 339 p. (Abteilung A), Band 126, Lieferung 1-2, p. FLEURY J.J. & FOURCADE É. (1990).- La 1-34 + 10 pls. superfamille Alveolinacea (Foraminifères): HOTTINGER L. (1962).- Recherches sur les Systématique et essai d'interprétation Alvéolines du Paléocène et de l'Éocène.- phylogénétique.- Revue de Schweizerische Paläontologische Micropaléontologie, Paris, vol. 33, N° 3-4, Abhandlungen, Basel, N° 75/76, 243 p. + p. 241-268. 18 pls. GINGERICH P.D. (1990).- Stratophenetics. In: HOTTINGER L. (1967).- Foraminifères BRIGGS D.E.G. & CROWTHER P.R. (eds.), imperforés du Mésozoïque marocain.- Notes Palaeobiology. A synthesis.- Blackwell et Mémoires du Service géologique, Rabat, Scientific Publications, Oxford, p. 437-441. N° 209, p. 5-168 GOLDSTEIN S.T. (1999).- Foraminifera, a HOTTINGER L. (1977).- Foraminifères biological overview. In: SEN GUPTA B.K. operculiniformes.- Mémoires du Muséum (ed.), Modern Foraminifera.- Kluwer National d'Histoire Naturelle, Paris, (Série Academic Publishers, Dortrecht, p. 37-55. C, Sciences de la terre), t. XL, 159 p. + 66 GRELL K.H. (1954).- Der generationswechsel pls.

41 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02)

HOTTINGER L. (1978).- Comparative anatomy LEVY A., MATHIEU R., POIGNANT A. & of elementary shell structures in selected ROSSET-MOULINIER M. (1986).- larger Foraminifera. In: HEDLEY R.H. & Discorbidae and Rotaliidae: a classification ADAMS C.G. (eds.), Foraminifera. Volume to be revised.- Journal of foraminiferal 3.- Academic Press, London, p. 203-266. Research, Lawrence, vol. 16, N° 1, p. 63– HOTTINGER L. (1986).- Construction, structure 70. and function of foraminiferal shells. In: LOEBLICH A.R. & TAPPAN H. (1964).- LEADBEATER B. & RIDING R. (eds.), Sarcodina chiefly "thecamoebians" and Biomineralisation in lower plants and Foraminiferida. – In: MOORE R.C. (ed.), animals.- Systematics Association Special Treatise on Invertebrate Paleontology, Part Publications Series, N° 30, London, p. 219- C, Protista 2.- Geological Society of 235. America & University of Kansas Press, HOTTINGER L. (1997).- Shallow benthic Lawrence, 900 p. foraminiferal assemblages as signals for LOEBLICH A. & TAPPAN H. (1987).- depth of their deposition and their Foraminiferal genera and their limitations.- Bulletin de la Société classification.- Van Nostrand Reinhold, New géologique de France, Paris, vol. 168, N° 4, York, vol. 1, 970 p.; vol. 2, 212 p. + 847 p. 491-505. pls. HOTTINGER L. (1999).- "Odd partnership", a MULLER H.J. (1964).- The relation of particular size relation between close recombination to mutational advance.- species of larger foraminifera, with an Mutation Research, Amsterdam, vol. 1, N° emendation of an outstandingly odd 1, p. 2-9. partner, Glomalveolina delicatissima PARVATI S. (1971).- A study of some rotaliid (SMOUT, 1954), Middle Eocene.- Eclogae Foraminifera I.- Koninklijke Nederlandse geologicae Helvetiae, Basel, vol. 92, N° 3, Akademie van Wetenschappen, p. 385-393. Proceedings, Amsterdam, (series B), vol. HOTTINGER L. (2000).- Functional morphology 74, p. 1-26. of benthic foraminiferal shells, envelopes of PAVLOWSKI J., ZANINETTI L. & LEE J. cells beyond measure. In: LEE J.J. & (1992).- Systematic revision of tiny HALLOCK P. (eds.), Advances in biology of heterocaryotic foraminifera described by Foraminifera.- Micropaleontology, New Karl GRELL in the years 1954-1979.- Revue York, vol. 46, suppl. 1, p. 57-86. de Paléobiologie, Genève, vol. 11, N° 2, p. HOTTINGER L. & DREHER D. (1974).- 385–395. Differentiation of protoplasm in REICHEL M. (1933).- Sur une alvéoline Nummulitidae (Foraminifera) from Elat, Red cénomanienne du Bassin du Beausset.- Sea.- Marine Biology, Heidelberg, vol. 25, Eclogae geologicae Helvetiae, Basel, vol. N° 1, p. 41-61. 26, N° 2, p. 269-280. HOTTINGER L., HALICZ E. & REISS Z. (1993).- REICHEL M. (1936-1937).- Étude sur les Recent Foraminiferida from the Gulf of Alvéolines.- Mémoires de la Société Aqaba, Red Sea.- Opera SAZU, Ljubljana, paléontologique Suisse, Bâle, vols. LVII & 33, 179 p. + 230 pls. LIX, 147 p. + 11 pls. HOTTINGER L. & LEUTENEGGER S. (1980).- REICHEL M. (1984).- Le crible apertural de The structure of calcarinid Foraminifera.- Rhapydionina liburnica STACHE from the Schweizerische Paläontologische Vremski-Britof Maastrichtian, Yugoslavia. Abhandlungen, Basel, N° 101, p. 115-151. In: Benthos'83, 2nd International LEE J.J., FABER W.W., ANDERSON O.R. & Symposium on Benthic Foraminifera (Pau, PAWLOWSKI J. (1991).- Life cycle in 1983).- Bulletin des Centres de Recherches Foraminifera. In: LEE J.J & O.R. ANDERSON Exploration-Production Elf-Aquitaine, Pau, (eds.): Biology of the Foraminifera.- Mémoire 6, p. 525-532. Academic Press, London, p. 285–343. REISS Z. (1963).- Reclassification of perforate LEHMANN R. (1961).- Strukturanalyse einiger Foraminifera.- Bulletin of the Geological Gattungen der Subfamilie Orbitolitinae.- Survey of Israel, Jerusalem, N° 35, 111 p. Eclogae geologicae Helvetiae, Basel, vol. + 8 pls. 54, N° 2, p. 597-667. REVETS S.A. (1989).- Structure and LEUTENEGGER S. (1977).- Ultrastructure de comparative anatomy of the toothplate in Foraminifères perforés et imperforés ainsi the Buliminacea (Foraminiferida).- Journal que de leur symbiotes.- Cahiers de of Micropaleontology, London, vol. 8, part Micropaléontologie, Paris, fasc. 3-1977, 52 1, p. 23-36. p. + 54 pls. REVETS S. (1993).- The foraminiferal LEVY A., MATHIEU R., POIGNANT A., ROSSET- toothplate, a review.- Journal of MOULINIER M. & & ROUVILLOIS A. (1979).- Micropaleontology, London, vol. 12, part 2, Révision de quelques genres de la famille p. 155-168. Discorbidae (Foraminiferida) fondée sur REVETS S. (1994).- Spirals: their orientation l'observation de leur architecture interne.- and description.- Journal of Revue de Micropaléontologie, Paris, vol. 32, Micropaleontology, London, vol. 13, p. 54. N° 2, p. 66-88. ROBINSON E. (1968).- Some larger

42 Carnets de Géologie / Notebooks on Geology - Memoir 2006/02 (CG2006_M02)

Foraminifera from the Eocene Limestones trends in Orbitolinas.- Revista Española de at Red Gal Ring, Jamaica.- Verhandlungen Micropaleontologia, Madrid, Numero Naturforschende Gesellschaft in Basel, vol. Especial (1975), p. 117-128. 84, N° 1, p. 281-292. SCHWAGER C. (1883).- Die Foraminiferen aus RÖTTGER (1974).- Larger Foramiifera: den Eocaenablagerungen der lybischen Reproduction and early stages of Wüste und Aegyptens.- Palaeontographica, development in Heterostegina depressa.- Stuttgart, vol. 30, p. 79-154 + 6 pls. Marine Biology, Heidelberg, vol. 26, p. 5 – SILVESTRI A. (1932).- Foraminiferi del 12. Cretaceo della Somalia.- Paleontographica SCHRÖDER R. (1962).- Orbitolinen des Italica, Pisa, vol. XXXII (1931, Nuova Serie, Cenomans Südwesteuropas.- vol. II), p. 143-204. Paläontologische Zeitschrift, Stuttgart, Band SMOUT A.H. (1954).- Lower Tertiary 36, N° 3-4, p. 171-202 + pls. 20-21. Foraminifera of the Qatar Peninsula.- SCHROEDER R. (1973).- El corte de Aulet British Museum (Natural History), London, (Prov. de Huesca) Evolucion de las 96 p. Orbitolinas en el limite del Cretaceo inferior VAN HINTE J.E. (1966).- Orbitoides from the superior. In: XIII Coloquio Europeo de Campanian type section. Part I.- Koninklijke Micropaleontologia España 1973.- Nederlandse Akademie van Enadimsa, Madrid, p. 141-149. Wetenschappen, Proceedings, Amsterdam, SCHROEDER R. (1975).- General evolutionary (Series B), vol. 69, N° 1, p. 79-110.

Figure 1: Accessory aperture in bulla of Globigerinita glutinata (EGGER). SEM graph from HOTTINGER et alii, 1993. aca: accessory aperture; bu: bulla; ch: ordinary spiral chamber.

43