Carnets Geol. 19 (3)

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[1. Introduction] [2. Historical background: Authors having cited the species]
[3. The different genera assigned to the species "Triloculina laevigata d'Orbigny, 1826" in Fornasini, 1905]
[4. The synonymy Pseudotriloculina-Sinuloculina in Loeblich and Tappan (1987) ...]
[5. The fossil form and the Recent one: only one and same species?] [Bibliographic references] and ... [Plates]

The "avatars" of Triloculina laevigata Orbigny, 1826,
generic and specific attribution

Armelle Poignant

5, rue Léon Bloy, 92260 Fontenay-aux-Roses (France)

Published online in final form (pdf) on February 4, 2019
DOI 10.4267/2042/69756
[Editor: Bruno Granier; language editor: Stephen Eagar]

Click here to download the PDF version!


The purpose of this study is to clarify the problem posed by Triloculina laevigata Orbigny, 1826, nom. nud., described and figured by Fornasini in 1905. An other species, also called Triloculina laevigata was created by Bornemann in 1855 and Orbigny's specific name became invalid. This invalidity was only recently assessed and numerous authors still use the original assignation. Consequently, it seems necessary to erect a new name. After listing the various names, revising the subgenus and genera attributed to Triloculina laevigata, and comparing fossil and recent specimens, the name: Affinetrina alcidi Lévy et al., 1992a, is confirmed.


• Foraminifera;
• systematics;
Triloculina laevigata Orbigny, 1826;
• subgenus Quinqueloculina (Pseudotriloculina) Cherif, 1970;
• genus Affinetrina Luczkowska, 1972;
• genus Sinuloculina Luczkowska, 1972


Poignant A. (2018).- The "avatars" of Triloculina laevigata Orbigny, 1826, generic and specific attribution.- Carnets Geol., Madrid, vol. 19, no. 3, p. 35-46.


Les "avatars" de Triloculina laevigata Orbigny, 1826, attribution générique et spécifique.- Ce travail a pour but de clarifier le problème posé par Triloculina laevigata Orbigny, 1826, nom. nud., figurée et décrite par Fornasini en 1905. Mais cette validation devient un homonyme secondaire, en raison de la description par Bornemann en 1855 d'une autre Triloculina laevigata. Cette invalidation n'a été reconnue que tardivement et plusieurs auteurs récents utilisent encore l'appellation originelle. Après la révision du sous-genre et des genres attribués à Triloculina laevigata et la comparaison entre les formes fossiles et actuelles, l'appellation Affinetrina alcidi Lévy et al., 1992a, est confirmée.


• Foraminifères ;
• systématique ;
Triloculina laevigata Orbigny, 1826 ;
• sous-genre Quinqueloculina (Pseudotriloculina) Cherif, 1970 ;
• genre Affinetrina Luczkowska, 1972 ;
• genre Sinuloculina Luczkowska, 1972

1. Introduction

In 1826, Orbigny proposed a species name: Triloculina laevigata in the "Tableau méthodique de la classe des Céphalopodes (nº 15, p. 300)" with only this location: "Habite la Méditerranée". This species was illustrated in the "Planches inédites" with 3 drawings (both sides and aperture with a long and bifid tooth) and the legend: "fossile à Valognes (W France), Éocène". His collection housed in the Muséum National d'Histoire Naturelle (MNHN) and includes a slide (MNHN.F.FO631) containing only 3 damaged specimens from Mediterranean. But the name "laevigata" remained invalid since it is without a description and figures. In 1905, Fornasini gave a good illustration but in his description, he figured a round aperture. Anyway, T. laevigata remains an homonym of Bornemann's species, described in 1855 and called Triloculina laevigata, but that is in fact a Miliolinella. Moreover, Terquem (1878, 1882) made a bad interpretation of Orbigny's species, describing it with a round aperture. So, the subsequent authors have described 2 different species according to Orbigny (see §2.1) or Terquem (see §2.2). In 1970, Cherif erected a new subgenus Quinqueloculina (Pseudotriloculina) laevigata for Orbigny's species. This subgenus became a genus: Pseudotriloculina in Loeblich and Tappan, 1987, with a non valid type-species name! The result was a total imbroglio.

It seemed significant to study the synonymy established by Loeblich and Tappan (1987) between Quinqueloculina (Pseudotriloculina) Cherif, 1970, and Sinuloculina Luczkowska, 1972, and also the genus Affinetrina Luczkowska, 1972, since T. laevigata has been referred to these 3 genera. Luczkowska (1972) worked substancially on the Miliolidae of the Miocene of Poland; she has outstandingly studied the internal structure and measured the angles between chambers during their ontogeny.

The aim of the present work is to bring out the various opinions of the authors, discuss the different generic attributions of Orbigny's species, and confirm the identity of the fossil and Recent specimens.

2. Historical background: Authors having cited the species

The main characters used by the authors is the shape of the aperture and of the tooth. These differences allow one to distinguish the two groups of authors: the first one chose the elongate aperture and tooth according to Orbigny (see §2.1, the second one chose the round aperture and short tooth according to Terquem (see §2.2).

2.1) the elongate aperture and tooth according to Orbigny

1826Triloculina laevigata d'Orbigny, nom. nud.

1893Triloculina laevigata d'Orbigny, Schlumberger (Pl. 1, figs. 45-47)

Schlumberger sees Mediterranean specimens. He shows 3 pictures of external morphology and 2 sections of A and B organisms (p. 206, Figs. 9-10). His species is unquestionably that of Orbigny: elongated aperture with a long tooth. He gives a short description of it.

1905Triloculina laevigata d'Orbigny, Fornasini (Pl. 1, fig. 10)

Fornasini worked on the unpublished plates of Orbigny. He reproduces Orbigny's drawings, clarifies the location of the species and its age, and gives a description: "T testa oblonga, laevigata, antice truncata, postice et circiter rotundata, loculis elongatis, leniter arcuatis inflatis, suturis angustis, apertura rotundata, dente apice bifurcata instructa". This publication describes and figures T. laevigata for the first time, giving the original drawing and a description (not quite accurate since the aperture is not round, but elongated).

19271928Triloculina laevigata d'Orbigny, Cushman

Cushman (1927, p. 32; 1928, p. 181), working on the classification of foraminifers, reproduced the figures of Schlumberger (1893): the external view (3 sides) and the section of A form. No comments are made.

1947Triloculina laevigata d'Orbigny, Lys and Sigal

They draw up some forms concerning the non-published plates of Orbigny. Their 'index card' (no. 977) reproduces the original drawing of T. laevigata. The provenance is: "Méditerranée, Belle-Île, et l'Océan Atlantique et Valognes (Manche), Castel-Arquato (Italie)". They indicate that the type of the MNHN specimens comes from Mediterranean. Unfortunately their work was not published.

1952Triloculina laevigata d'Orbigny, Bogdanovich

This author figures two drawings of Fornasini (1905, Fig. 10 and 10a) and Schlumberger's section (1893).

1958Triloculina laevigata d'Orbigny, J. and Y. Le Calvez (Pl. 6, figs. 63-64)

They point out the presence of the species in the Bay of Villefranche, Mediterranean (their Fig. 62 shows a round aperture !). They consider that Terquem has misinterpreted T. laevigata and that his figures are not accurate.

1969Quinqueloculina laevigata (d'Orbigny), Bogdanovich

The author studied the internal structure of some Triloculines (see further).

1970Triloculina lecalvezae Kaasschieter, Y. Le Calvez (Pl. 1, fig. 3)

Y. Le Calvez figures T. lecalvezae (p. 51-52, Pl. 7, fig. 1) in the Eocene of the Parisian Basin (Chaussy), but it is not Kaasschieter's species. Her specimen has an elongated aperture, a long and bifid tooth, and is identical to Orbigny's specimen from Valognes. She reports it in several Eocene localities of Parisian Basin (among them the Eocene "falunière" of Grignon, Yvelines, western Parisian Basin) where the present author found specimens (see Pl. 1, figs. 8-9), as well as in the Oligocene. The figures of Y. Le Calvez are the first since Fornasini (1905) that exactly shows the Eocene species of Orbigny, in spite of which she refers them to Terquem (1878), although earlier (J. et Y. Le Calvez, 1958) she considered that Terquem had misinterpreted T. laevigata (see below).

1970Quinqueloculina (Pseudotriloculina) laevigata (d'Orbigny), Cherif (Pl. 9, fig. 2; Pl. 21, fig. 2, fide Schlumberger)

When studying the Miliolacea from the western coast of Naxos, Cherif erected the subgenus Quinqueloculina (Pseudotriloculina) with T. laevigata Orbigny fide Schlumberger (1893) as type-species. His figures do not differ from Schlumberger and it is likely the same species. Diagnosis of the sub-genus and relative comments will be treated further.

1973Quinqueloculina (Pseudotriloculina) laevigata (d'Orbigny), Cherif (Pl. 10, fig. 6; Pl. 14, fig. 4, 10; Pl. 5, fig. 10)

Cherif repeated the description of the sub-genus with some new figures, but without any modification.

1986Triloculina laevigata d'Orbigny, Cabioch et al. (Pl. 3, fig. 3)

These authors point out and figure T. laevigata in the Quaternary of fringing reefs in Southern New-Caledonia, but they give Terquem reference.

1992aAffinetrina alcidi nom. nov. pro Triloculina laevigata d'Orbigny, 1826, Lévy et al. (Pl. 4 , figs. 5-6)

When studying the Mediterranean foraminifers of the Cretan shore and mainly specimens of Miliolacea with quinqueloculine and triloculine arrangement of the chambers, the present author and her coauthors have described (p. 123-124, Pl. 3, figs. 3-4) Affinetrina alcidi and placed into its synonymy Triloculina laevigata in Orbigny (1826), Schlumberger (1893), Fornasini (1905), Triloculina lecalvezae in Y. Le Calvez (1970), and Quinqueloculina (Pseudotriloculina) laevigata in Cherif (1970, 1973).

1992bAffinetrina alcidi Lévy et al. (p. 121, Pl. 2, figs. 9-10

1995Affinetrina alcidi Lévy et al., Lévy et al.

The species occurs in the Recent deposits of the archipelago Fernando de Noronha, north Brasil.

2.2) the round aperture and short tooth according to Terquem

1878Triloculina laevigata d'Orbigny, Terquem (Figs. 20a-21b)

Terquem considers he has found again T. laevigata in the Pliocene of Rhodes island. It cannot be the same species because the aperture of Terquem is round whereas that of Orbigny is elongated.

1882Triloculina laevigata d'Orbigny, Terquem (Pl. 17, figs. 22-23)

Terquem identified Orbigny's species again in the Eocene of Paris region. It is the same as his species from Rhodes.

1961Triloculina lecalvezae Kaasschieter nom. nov. pro Triloculina laevigata, 1826, nom. nud.

Kaasschieter was probably the first to recognize invalidity of Orbigny's species. Consequently, he introduced a new species name that dedicated to Y. Le Calvez. Unfortunately, the specimen he chose is different from Orbigny's one because the aperture is round. He put into the synonymy the original Orbigny's species, figured by Fornasini (1905), as well as that of Terquem (1878). He did not describe his species. Kaasschieter's specimen (Lutetian of Damery, Parisian Basin) is a different species.

1972Sinuloculina laevigata (d'Orbigny, 1878), Luczkowska (p. 365, no figure)

This genus will be examined further. It is obvious that Luczkowska chose Terquem reference, as it is indicated by the date of validity of the species.

1987Pseudotriloculina lecalvezae (Kaasschieter), Loeblich and Tappan (p. 342-343, Pl. 352, figs. 13-14)

In their treatise, Loeblich and Tappan raised to genus the Cherif's subgenus Pseudotriloculina but, doing so, they made an error: they said that the subgenus was Triloculina (Pseudotriloculina) instead of Quinqueloculina (Pseudotriloculina). They chose to validate Orbigny's species Triloculina lecalvezae Kaasschieter with Terquem figures from the Pliocene of Rhodes island, which are different (see supra) from Orbigny's species. They put into the synonymy Pseudotriloculina and Sinuloculina Luczkowska, 1972. This problem will be treated further.

Table 1 summarizes the various assignments of Triloculina laevigata Orbigny, 1826

Table 1: The various assignments of Triloculina laevigata Orbigny, 1826.

  Triloculina laevigata Orbigny, 1826, nom. nud.  
  Citations and illustrations. Eocene-Oligocene
Valognes (Manche), Paris Basin
  Triloculina laevigata Orbigny.- Fornasini, 1905
invalid, junior synonym of Triloculina laevigata Bornemann, 1855
  Triloculina lecalvezae Kaasschieter.- Y. Le Calvez, 1970
species name invalid
  Affinetrina alcidi Lévy et al.- Poignant, the present work  
Wrong assignments
Not Orbigny's species

Citations in various areas and epochs
Triloculina laevigata Orbigny.- Terquem, 1878
Pliocene, Rhodes Island
Triloculina laevigata Orbigny.- Schlumberger, 1893
Triloculina lecalvezae Kaasschieter, 1961
Eocene, Paris Basin
Triloculina laevigata Orbigny.- Cushman, 1927, 1928
Schlumberger's figures
Pseudotriloculina lecalvezae (Kaasschieter, 1961)
Loeblich & Tappan, 1987
Pliocene, Rhodes Island
Triloculina laevigata Orbigny.- Lys and Sigal, 1947
Pliocene (Castel-Arquato), Mediterranean
  Triloculina laevigata pro parte Orbigny.- J. et Y. Le Calvez, 1958
  Triloculina lecalvezae Kaasschieter.- Y. Le Calvez, 1970
Paris Basin
  Quinqueloculina (Pseudotriloculina) laevigata Orbigny.- 
, 1970, 1973
  Triloculina laevigata Orbigny.- Cabioch et al., 1986
Quaternary, New-Caledonia
  Affinetrina alcidi Lévy et al., 1992a, 1992b, 1995
  Affinetrina alcidi Lévy et al., 1992a.- 
, the present work

3. The different genera assigned to the species "Triloculina laevigata d'Orbigny, 1826" in Fornasini, 1905

In the literature the species is found with 5 assignements to genera and subgenus: Triloculina, Quinqueloculina (Pseudotriloculina), Sinuloculina, Affinetrina, Pseudotriloculina. We will see that every diagnosis, except Fornasini's one, takes into account the internal structure that is essential for a reliable determination.

Bogdanovich (1952) described the internal structures of Triloculina. He showed a figure (p. 12, Fig. 5) of T. laevigata from Schlumberger with a quinqueloculine structure and 3 chambers visible.

In 1969, he writes (p. 352-353): "However, cases where the number of chambers seen outside the tests of many quinqueloculine species decreases to 4 and even 3 are quite common. This phenomenon is caused by a rapid decrease of thickness of chambers as the test grows, and thus by their involution. As a result, the last three chambers cover the earlier ones – the fifth and also not seldom the fourth chambers (...)". The author proposed to name this structure as "cryptoquinqueloculine" and to regard it as a modification of the true quinqueloculine structure of the Miliolidae.

Luczkowska (1972) made an extremely interesting work on the Miliolidae of the Polish Miocene, later accompanied by an abundant illustration in 1974. Effectively, she made numerous sections and measured the angles between chambers during the ontogeny in particular for the genera Triloculina and two new genera that she erected: Affinetrina and Sinuloculina. Her measurements of angles particularly concerns the B generation. Among others, she used 2 terms corresponding to 2 types of coiling:

Genera Triloculina and Quinqueloculina

According to Loeblich and Tappan (1987), Triloculina is initially cryptoquinqueloculine at least in microspheric generation (Pl. 3 , fig. 2, from Schlumberger, 1893), with 3 chambers visible externally (Pl. 2, figs. 1-2). Quinqueloculina has earlier chambers in quinqueloculine arrangement (Pl. 3 , fig. 1), with 5 chambers visible externally.

Subgenus Quinqueloculina (Pseudotriloculina) Cherif

(Pl. 1 , fig. 5)

1970Quinqueloculina (Pseudotriloculina) Cherif, vol. 142, no. 1, p. 112.

Type species: Triloculina laevigata Orbigny, 1826, emend Schlumberger, 1893.

Original diagnosis: "Gehause frei, Kammernordnung quinqueloculin, Äquatorial-Windung involut, von aussen 3 Kammern sichtbar. Wand porzellanartig, hochglänzend. Mündung mit oder ohne Hals, mit oder ohne Lippe, einfacher oder bifider Zahn.
Beziehungen: Quinqueloculina (Pseudotriloculina) n. subgen., unterscheidet von Triloculina d'Orbigny durch ihre quinqueloculine Kammeraordnung. Sie unterscheidet sich von Quinqueloculina (Quinqueloculina) d'Orbigny durch ihre involute Äquatorialwindung, die nur 3 Kammern von auben sehen läbt."

Cherif (1973) repeated the diagnosis of his sub-genus without any modifications and added a few figures.

Cherif distinguished Pseudotriloculina from Triloculina Orbigny (T. trigonula type) by the quinqueloculine coiling of the first genus, which is incorrect since Loeblich and Tappan (1987) mention a quinqueloculine coiling in Triloculina, at least in microspheric forms. Moreover, Cherif's section of Triloculina does not show the initial quinqueloculine coiling (this work: Pl. 3 , fig. 5 C). Cherif wanted to name the particular structure he saw in Triloculina laevigata Orbigny (see his description) and consequently created the sub-genus called Quinqueloculina (Pseudotriloculina). This structure was called "cryptoquinqueloculine" by Bogdanovich, 1969 (see supra).

Cherif (1970) found the species in the Mediterranean (Naxos coasts, Greece); consequently, he determined his species according to Schlumberger (1893) whose specimens came from the Mediterranean. He quoted Fornasini but not Terquem (1878). He knew the work of Kaasschieter (1961) but he did not take into account the new name of T. laevigata, probably because Kaasschieter's figure does not concern the Orbigny's species (see supra). In his synonymy, apart Fornasini, he only mentions authors having observed the species in Recent environments, i.e., Martinotti (1920) and Wiesner (1923).

Genus Affinetrina Luczkowska

(Pl. 1 , figs. 4, 6; Pl. 2 , figs. 8-9, 11-12; Pl. 4 , figs. 5-6)

1972Affinetrina n.gen. – Luczkowska, p. 368, Pl. 13, figs. 4-5.

Type species: Triloculina planciana Orbigny, 1839

Diagnosis: "Test with an internal structure as in Triloculina, but with an high slit-like aperture nearly filled with a long, slender tooth, slightly enlarged at the end; three chambers visible externally".

Affinetrina mainly differs from Triloculina in having an elongated aperture and a long tooth. The genus was recognized by Loeblich and Tappan (1987, p. 337), who report Luczkowska's diagnosis.

In fact, the section of the micro generation of Affinetrina planciana given by Luczkowska is more complex. It shows a kryptoquinqueloculine initial part and a pseudotriloculine mature stage (see Fig. 11, p. 358).

Lévy et al. (1992a, 1992b) put Triloculina laevigata fide Orbigny in Schlumberger (1893), Fornasini (1905), Y. Le Calvez (1970), Cherif (1970), in the genus Affinetrina.

Genus Sinuloculina Luczkowska

(Pl. 2 , figs. 3-7, 10)

1972Sinuloculina n.gen. – Luczkowska, p. 370, Pl. 12, figs. 4-5.

Type species: Biloculina cyclostoma Reuss, 1850

Diagnosis: "Test initially with kryptoquinqueloculine chamber arrangement, later chambers added in planes of coiling increasing irregularly to 180° or more, so that alternating chambers form the sinusoid curve in sections; three proloculus sizes are distinguished; chambers without floor, broadly overlapping the preceding chamber and giving externally triloculine or biloculine appearance, aperture large, rounded or oval, usually bordered with a thick rim, filled with a thick, bifid tooth, protruding over the aperture edge".

The type species only shows 2 chambers in external view.

Luczkowska referred several species to her new genus like Triloculina consobrina Orbigny (see section, Pl. 3 , fig. 1). Of course, Luczkowska did not know Cherif's study.

4. The synonymy Pseudotriloculina-Sinuloculina in Loeblich and Tappan (1987) - 
Differences between both genera – 
Validity of the genus Pseudotriloculina
Specific assignment of Triloculina laevigata

The diagnosis of the sub-genus Quinqueloculina (Pseudotriloculina) Cherif, which was erected as a genus by Loeblich et Tappan, is essentially that of Luczkowska's genus, as far as the internal structure is concerned.

Loeblich and Tappan figure the genus Pseudotriloculina (Pl. 352) with Reuss species: Biloculina cyclostoma, type species of Sinuloculina Luczkowska, but this species only shows 2 external chambers, which is not mentioned in Cherif's diagnosis. Moreover, they also illustrate Pseudotriloculina lecalvezae with Terquem's figures, showing a rounded aperture and without elongated and protruding tooth, in contradiction with Schlumberger (1893), Fornasini (1905), Cherif (1970), and Y. Le Calvez (1970).

Loeblich and Tappan recognize the "pseudotriloculine" structure described by Luczkowska and the genus Pseudotriloculina of Cherif the type-species of which displays a kryptoquinqueloculine and a pseudotriloculine mature stage structure according to Luczkowska (see below).

We draw attention that something important is lacking in Cherif's diagnosis; his description of the external morphology is almost totally insignificant: "Mündung mit oder ohne Hals, mit oder ohne Lippe, einfacher oder bifider Zahn". In contrast Luczkowska's diagnosis of Sinuloculina is much more significant: "aperture large, rounded, usually bordered with a thick rim, filled with a thick, bifid tooth, protruding over the aperture edge".

The external morphology has important since it allows one to observe from the outside the number of chambers on each side, the presence or lack of a peristome, and the shape of the aperture and tooth. For example, Triloculina has a sub-triangular to triangular outside in oral view (Pl. 2 , figs. 1-2) and Sinuloculina an oval one (Pl. 2 , figs. 4-5, 7); the aperture can be rounded, oval, elongated (Pl. 2 , figs. 1-3, 6, 10); the tooth can be thick, thin, short, elongated, bifid, even ring-shaped (Pl. 2 , figs. 1-3, 6, 10). The genus Sinuloculina allows one to identify it more often, without making a section; moreover, its type species possesses only 2 visible chambers.

On the other hand, Loeblich and Tappan (1987) admit the genus Affinetrina Luczkowska, 1972, which shows an elongate aperture and a long and bifid tooth, protruding over the apertural rim (see supra).

Finally, the present author adheres to the opinion she previously held with her coauthors (Lévy et al., 1992a, 1992b) placing "Triloculina laevigata" Orbigny (fide Schlumberger), provided with a slit-like aperture and an elongate tooth, in the genus Affinetrina. The section of Triloculina laevigata is identical to the one of T. planciana Orbigny, 1839, type species of the genus Affinetrina. For Luczkowska (1972, p. 358, Fig. 11c), A. planciana has a kryptoquinqueloculine section, then pseudotriloculine, i.e., between the 3 chambers of the last coil: 2 angles > 130° and 1 < 90°. The section of Triloculina planciana in Luczkowska (Pl. 4 , fig. 1) is quite similar with Schlumberger's sections for T. laevigata (Pl. 4 , figs. 3-4).

Consequently, a question must be considered: what are the implications for the genus Pseudotriloculina if it loses its type-species, i.e., Pseudotriloculina laevigata, that possesses the external and internal characters of the genus Affinetrina? In any case, the present author decided to stop using it.

As for the specific assignment of Triloculina laevigata, neither laevigata Orbigny can be used, having been invalidated by Bornemann, nor lecalvezae Kaasschieter, because Kaasschieter's figure has a rounded aperture. Only "alcidi" (Lévy et al., 1992a), which was erected for a Recent species and put into synonymy with "Triloculina laevigata" sensu Orbigny, is left.

5. The fossil form and the Recent one: only one and same species?

Finally, is the Recent form the same as the fossil one ?

Firstly, Orbigny who described the species in the Eocene also reports it from the Recent. The fossil form has been found in the Eocene and Oligocene in the following localities:

In the Eocene-Oligocene, it seems to have only been observed in the Franco-Belgian Basin and Eocene of Manche; it has never been recorded in Aquitaine or from sediments of Miocene age. Lys and Sigal (1947) found it in sediments of Pliocene age at Castel-Gandolfo (Italy, Emilia).

However, the Recent form is quite widespread. Orbigny (unpublished plates) indicated it at Belle-Île, Atlantic Ocean, Mediterranean, and the type specimen, stored at the MNHN, comes from the Mediterranean. Schlumberger (1893) studied it in the Mediterranean Sea and a mount of the Schlumberger collection, i.e., a microfossil slide, with 16 specimens coming from Marseille (bryozoa zone) has been observed at the MNHN. Two mounts of the J. and Y. Le Calvez Mediterranean collection (no. 191-192) have also been observed; they contain several well preserved specimens of T. laevigata.

Cherif (1970, 1973) studied it in Greece whereas the present author and her coauthors were investigating in Crete (Lévy et al., 1992a, 1992b). In both cases, the Recent form was placed in the synonymy with the fossil form. Cabioch et al. (1986) observed and figured it in the Quaternary of fringing reef of New-Caledonia, but they take the Terquem reference. It was also found in the Fernando de Noronha archipel (Northeastern Brasil) by Lévy et al. (1995).

The present author studied both forms but, as it has been said above, the fossil specimens are rare compared to the Recent ones, even if Y. Le Calvez found it in many exposures of the Paris Basin (see above). As far as the external morphology is concerned, no noteworthy differences can be seen; the specimens observed in the Eocene of Grignon: those by Y. Le Calvez one (1970) and the ones in the present work are identical to the Mediterranean or NeW Caledonian forms of the mentioned authors. In addition, the Eocene specimen of Orbigny, figured by Fornasini, shows no special features except for its typical tooth.

Therefore, it is difficult to distinguish the two different species since Orbigny saw one species only. Revets (1994) writes: "d'Orbigny was a naturalist of high repute as the result of his careful observations and his keen eye for detail". Affinetrina alcidi is perhaps rare to very rare in Eocene-Oligocene and Miocene and Pliocene too and abundant in Recent.

In conclusion, there would very probably exist only one and the same species: Affinetrina alcidi Lévy et al., 1992a.


I am deeply indebted at first to the MNHN for the facilities given for the examination of the various collections and unpublished Orbigny's plate; without its assistance, I could not have carried out my work. I particularly thank Annachiara Bartolini (MNHN) for her comments, suggestions and her help in the realisation of the illustration; this paper owes a lot to her. First, I highly thank one of the revisers who make a very serious study of my work and also Bruno Cahuzac, Stephen Eagar, Florent Falize, Bruce Hayward, Irina Konenkova, Jean-Pierre Margerel, Rainer Müller, Jean-Michel Pacaud, Christian Rupp, Huguette Salles-Astier, Karoly Sztrakos, Marie-Thérèse Vénec-Peyré, for their help in various fields: comments, discussions, pictures, translations, drawings, photocopies, and linguistic improvements.

Bibliographic references

Bogdanovich A.K. (1952).- Miliolidae et Peneroplidae. Foraminifères fossiles de l'URSS.- Travaux de l'Institut de Recherches scientifiques des Mines de l'URSS (n.s.), Moscou, vol. 64, 338 p. [French translation].

Bogdanovich A.K. (1969).- To the revision of Miliolidae with Quinqueloculine and Triloculine structure of tests.- Annales de la Société géologique de Pologne, Kraków, vol. 39, no. 1-3, p. 351-360.

Bornemann J.G. (1855).- Die mikroskopische Fauna des Septarienthones von Hermsdorf bei Berlin.- Zeitschrift der Deutschen Geologischen Gesellschaft, Berlin, Band 7, p. 307-371. URL:

Cabioch G., Anglada R. & Babinot J.-F. (1986).- Microfaunes et paléoenvironnements des récifs frangeants quaternaires de Mamié et Ricaudy (Nouvelle-Calédonie).- Cahiers de Micropaléontologie, Paris, vol. 1, no. 1-2, p. 5-36.

Cherif O.H. (1970).- Die Miliolacea der Westküste von Naxos (Griechenland) und ihre lebensbereiche.- Lehrstuhl für Erdölgeologie Technische Universität Clausthal, 176 p.

Cherif O.H. (1973).- Zur Klassifierung der Gattung Quinqueloculina (Foraminifera).- Neues Jahrburg Geologie Paläontologie, Abhandlungen, Stuttgart, Band 142, Heft 1, p. 73-96.

Colom G. (1942).- Una contribución al conocimiento de los Foraminíferos de la Bahía de Palma de Mallorca.- Instituto Español de Oceanografía, Notas y Resumenes, Madrid, vol. 2, no. 108, 53 p.

Cushman J.A. (1927).- An outline of a re-classification of the foraminifera.- Contributions from the Cushman Laboratory for Foraminiferal Research, Sharon, MA, vol. 3, 105 p.

Cushman J.A. (1928).- Foraminifera their classification and economic use.- Special Publications Cushman Laboratory for Foraminiferal Research, Sharon, MA, vol. 1, 401 p.

Ellis B.F. & Messina A. (1940).- Catalogue of Foraminifera (+ supplements).- The American Museum of Natural History, New-York.

Fornasini C. (1905).- Illustrazione di specie orbignyane di Miliolidi institute nel 1826.- Memorie delle Reale Accademie degli Scienze dell' Istituto di Bologna (Serie VI), t. II, p. 59-70 (4 Pls.). URL:

Kaasschieter J.P.H. (1961).- Foraminifera of the Eocene of Belgium.- Institut royal des Sciences naturelles de Belgique, Mémoires, Bruxelles, no. 147, p. 1-259.

Le Calvez J. & Le Calvez Y. (1958).- Répartition des Foraminifères dans la Baie de Villefranche. I. Miliolidae.- Annales de l'Institut océanographique, Monaco, t. XXXV, fasc. 3, p. 160-226.

Le Calvez Y. (1970).- Contribution à l'étude des Foraminifères paléogènes du Bassin de Paris.- Cahiers de Paléontologie, Paris, 326 p.

Lévy A., Mathieu R., Poignant A. & Rosset-Moulinier M. (1991).- Morphologie et morphométrie de foraminifères benthiques (Miliolacea) du Cénozoïque à l'Actuel.- Geobios, Villeurbanne, vol. 24, suppl. 1, p. 97-104.

Lévy A., Mathieu R., Poignant A. & Rosset-Moulinier M. (1992a).- Foraminifères à arrangement quinquéloculin et triloculin (Miliolacea) de Méditerranée.- Revue de Paléobiologie, Genève, vol. 11, fasc. 1, p. 111-135.

Lévy A., Mathieu R., Poignant A. & Rosset-Moulinier M. (1992b).- The Miliolacea with Quinqueloculina and Triloculina morphology from the Mediterranean.- Benthos'90, Tokai University Press (Japan), p. 117-125.

Lévy A., Mathieu R., Poignant A., Rosset-Moulinier M. & Ambroise D. (1995).- Benthic foraminifera from the Fernando de Noronha Archipelago (northern Brazil).- Marine Micropaleontology, vol. 26, no. 1-4, p. 89-97.

Loeblich A.R. & Tappan H. (1987).- Foraminiferal genera and their classification.- Van Nostrand Reinhold Comp., New-York, 970 p.

Luczkowska E. (1972).- Miliolidae (Foraminiferida) from Miocene of Poland. Part I. Revision of the classification.- Acta Palaeontologica Polonica, Warszawa, vol. 17, no. 3, p. 341-377.

Luczkowska E. (1974).- Miliolidae (Foraminiferida) from Miocene of Poland. Part II. Biostratigraphy, Palaeontology and systematics.- Acta Palaeontologica Polonica, Warszawa, vol. 19, no. 1, p. 1-169.

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Plate 1: 1. Triloculina laevigata Orbigny. Unpublished Orbigny's drawing in Fornasini, 1905. Eocene, Valognes (Manche). X no mentioned. 2. Triloculina laevigata Orbigny, in Schlumberger, 1893, Pl. 1, figs. 45-47, X 40. 3. Triloculina lecalvezae Kaasschieter, in Y. Le Calvez, 1970, Pl. 7, fig. 1. Eocene, Chaussy (Paris Basin). Side view: X 30, apertural view: X 140. 4, 6. Affinetrina alcidi Lévy et al., Eocene, "falunière" of Grignon (Yvelines, Paris Basin). X 50. 5. Triloculina laevigata Orbigny, in Cherif, 1970, Pl. 9, fig. 2a,b. Recent, Naxos. Side view: X 50; apertural view: X 200. Graphical scale = 5 cm.

Pl. 1
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Plate 2: 1. Triloculina trigonula (Lamarck). Recent, Sicily. X 80. 2. Triloculina gibba Orbigny. Recent, Cyprus. X 80. 3, 7. Sinuloculina cyclostoma (Reuss). Recent, Sicily, Cagliari. X 65. 4, 10. Sinuloculina williamsoni (Terquem). Recent, English Channel. 4: X 60; 10: X 300 (ring-shaped tooth). 5-6. Sinuloculina consobrina (Orbigny). Recent, Crete. 5: X 65; 6: X 95. 8-9. Affinetrina bermudezi (Acosta). Recent, Tel Aviv, Israel. 8: X 70; 9: X 100. 11-12. Affinetrina eburnea (Orbigny). Recent, Cyprus. 11: Side view: X 80; 12: X 120 (Figures 3, 6, 7, 8, 9, 11, 12, excerpts from Lévy et al., 1992a). Graphical scale = 5 cm.

Pl. 2
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Plate 3: 1. Quinqueloculina boueana Orbigny. Section of a microspheric form. Aquitanian, Le Plantat, Northern Aquitaine. X 225. 2. Triloculina schreiberiana Orbigny. A: Section of a megalospheric form; B: section of a microspheric form, in Schlumberger, 1893, p. 204, Fig. 5, A form; Fig. 6: B form. Recent, Mediterranean. X 66. 3. Sinuloculina consobrina (Orbigny), section of a megalospheric form. Burdigalian, Pont-Pourquey, Northern Aquitaine. X 200. 4. Sinuloculina cyclostoma (Reuss). Section of a microspheric form, in Luczkowska, 1972, p. 358, Fig. 11a. Miocene, Weglinek, Poland. X 1300. 5. A - Section of Quinqueloculina; B – Section of Pseudotriloculina (the last coiling has 3 chambers because the youngest chamber hides the 4th chamber); C - Section of Triloculina, in Cherif, 1970, Pl. VI, figs. 5-7. Recent Mediterranean. X no mentioned (Figure 3 excerpt from Lévy et al., 1991). Graphical scale = 5 cm.

Pl. 3
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Plate 4: 1. Affinetrina planciana (Orbigny). Section of a microspheric form, in Luczkowska, 1972, p. 358, Fig. 11c. Miocene, Gliwice st., Poland. X 800. 2. Triloculina laevigata Orbigny. a: b form; b: a form. Sections in Cherif, 1970, Pl. VII, X no mentioned. 3. Triloculina laevigata Orbigny, section of Schlumberger (MNHN) with addition of angular measures. Recent, Mediterranean. 4. Triloculina laevigata Orbigny. Section of a microspheric form, in Schlumberger, p. 206, Fig. 10, X 100. 5-6. Affinetrina alcidi Lévy et al. 5: side view, X 30; 6: apertural view, X 40. Recent, Crete (Figures 5-6, excerpt from Lévy et al., 1992b). Graphical scale = 5 cm.

Pl. 4
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The present author could examine the original sample of Orbigny (stored at the MNHN) but she only found a few badly preserved specimens.