Carnets Geol. 16 (7)  

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Contents

[Introduction] [Material studied] [Discussion] [Systematics (B.G.)]
[Age of the stratum typicum] [The Brazilian morphotypes] [Conclusions]
[Bibliographic references] [Table] and ... [Plates]


On the fossil alga Marinella lugeoni Pfender, 1939, nom. cons.,
and its seven unfortunate avatars

Revision of the Juliette Pfender Collection. Part 2.
Revision of the Jesse Harlan Johnson Collection. Part 2

Bruno Granier

"Cátedra Franco-Brasileira no Estado de São Paulo 2015", UNESP - Universidade Estadual Paulista, Center for Geosciences Applied to Petroleum (UNESPetro), Caixa Postal 178, Av. 24 A, no. 1515, Bela Vista, CEP13506-900 - Rio Claro - SP (Brazil)
Dépt. STU, Fac. Sci. Tech., UBO, CS 93837, F-29238 Brest (France)

Department of Ecology and Evolutionary Biology, The University of Kansas, 1200 Sunnyside Avenue, Lawrence, Kansas 66045 (USA)

Dimas Dias-Brito

UNESP - Universidade Estadual Paulista, Center for Geosciences Applied to Petroleum (UNESPetro) & Departamento de Geologia Aplicada, Caixa Postal 178, Av. 24 A, no. 1515, Bela Vista, CEP13506-900 - Rio Claro - SP (Brazil)

Published online in final form (pdf) on April 14, 2016
[Editor: Michel Moullade; technical editor: Bruno Granier; language editor: Phil Salvador]

Click here to download the PDF version!

Abstract

A review of eight lookalike fossil species led to their being synonymized. Although Marinella lugeoni Pfender, 1939, is not the senior synonym, it is proposed to ascribe it the status of a "nomen conservandum". The age of its type-locality in Spain is Late Jurassic, not Early Jurassic. We also document small Marinella lumps found in Albian-Cenomanian strata of Brazil.

Key-words

Rhodogorgonales; Elianellaceae; Solenoporacea; Marinella.

Citation

Granier B. & Dias-Brito D. (2016).- On the fossil alga Marinella lugeoni Pfender, 1939, nom. cons., and its seven unfortunate avatars. Revision of the Juliette Pfender Collection. Part 2. Revision of the Jesse Harlan Johnson Collection. Part 2.- Carnets Geol., Madrid, vol. 16, no. 7, p. 231-245.

Résumé

À propos de l'algue fossile Marinella lugeoni Pfender, 1939, nom. cons., et ses septs avatars malheureux. Révision de la Collection Juliette Pfender. 2e partie. Révision de la Collection Jesse Harlan Johnson. 2e partie.- Le réexamen de huit espèces fossiles très semblables se conclut  sur leur mise en synonymie. Bien que Marinella lugeoni Pfender, 1939, ne soit pas le synonyme senior, nous proposons de lui attribuer le statut de "nomen conservandum". L'âge de sa localité-type en Espagne est Jurassique supérieur et non Jurassique inférieur. Enfin, nous consignons la découverte de petits nodules à Marinella dans des couches albo-cénomaniennes du Brésil.

Mots-clefs

Rhodogorgonales ; Elianellaceae ; Solenoporacea ; Marinella.


Introduction

Marinella lugeoni Pfender, 1939, is a fossil "calcareous alga" familiar to those people working on Upper Jurassic and Lower-"middle" Cretaceous limestones. It is not our intention to duplicate here the observations of earlier researchers on this nodular, sometimes encrusting alga, and it is particularly not our intention to duplicate the detailed information published by Barattolo and del Re (1984), supplemented by Leinfelder and Werner (1993), but we intend to present a summary of our understanding of the delimitation of this species: seven authors contributed to introduce one genus and eight species that were all synonymized (Johnson contributed twice). Marinella lugeoni was first found in Albian-Cenomanian strata of South Atlantic marginal basins in Angola (Romanes, 1916), and later in Brazil (Maury, 1937), before Pfender (1939) reported its occurrence in Jurassic strata from Spain. To complete this study, we reexamine the age of its stratum typicum in Spain and describe a particular morphotype commonly found in our Brazilian material.

Material studied

The first author (B.G.) examined material from the Juliette Pfender Collection in Paris, from the Jesse Harlan Johnson Collection in Washington D.C. and from the UNESPetro Collection in Rio Claro (São Paulo).

When revising the type-material of Marinella lugeoni Pfender, 1939, hosted at the "Laboratoire de Micropaléontologie" of the "Université Pierre et Marie Curie" (Paris, France), Barattolo and del Re (1984) had the opportunity to examine four thin sections, "probably those used by Pfender for the diagnosis of the genus" (the first author found only three of them), and ten additional thin sections "made in 1967" (the first author found only eight of them). In the meantime, some material (four thin sections) was lost (!). The eleven remaining thin sections (Fig. 1 ) will be relocated and deposited with a "PC" label at the "Herbier Cryptogamique, Département Systématique et Évolution, Muséum National d'Histoire Naturelle" (Paris, France) with the rest of the Juliette Pfender Collection. Two rock samples (Pl. 1 ) complete this partial inventory.

With respect to Johnson's material, hosted by the Smithsonian Institution, the first author managed to locate four key thin sections (Fig. 2 ) with USNM registration numbers D992-a-843 and D992-a-844 (Johnson, 1961), 42547 (Johnson & Kaska, 1965) and 42606 (Johnson, 1968), but not a fifth one with the USNM number 42467 (Johnson, 1965).

Fig. 1
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Figure 1: The 11 thin sections remaining of the Juliette Pfender Collection. The lectotype (defined here: Fig. 4 ; Pfender, 1939, Pl. II, fig. 1) is in the last thin section (the third from the left) of the first row. [All photos with 1 cm scale bar]

Fig. 2
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Figure 2: The 4 thin sections of the Jesse Harlan Johnson Collection:
A) D992 a844 / 2070 L 3441 Jur / Shell Co, Navarro Co, Texas;
B) D992 a843 / 2069 L-3441 Jur. / Shell Co; Navarro Co, Tex.;
A-B) both with Marinella lugeoni according to Johnson, 1961;
C) USNM 42606 with a Lithophyllum ? sp. or a Lithothamnium ? sp. according to Johnson, 1968;
D) USNM 42547 A-D with the type of Lithothamnium ? primitiva Johnson & Kaska, 1965. [All photos with 1 cm scale bar]

Discussion

It is easy today (half a century later) to blame the pioneers in paleophycology for their lack of scientific rigor. For instance:

However, knowledge on modern and fossil red algae has significantly grown since that time. In addition, access to information is getting easier.

Fig. 3
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Figure 3: Marinella lugeoni in thin section USNM no. 42547 of the Jesse Harlan Johnson Collection as:
A) Lithothamnium ? primitiva n. sp. (Johnson & Kaska, 1965, Pl. 6, fig. 1);
B) Marinella lugeoni (op. cit., Pl. 6, fig. 2);
C) Girvanella minuta (op. cit., Pl. 30, fig. 1). [All photos with 500 µm scale bar]

Early paleophycologists studying calcareous red algae and related forms were commonly relying on the external morphology and related features (measurements) to discriminate species. However, modern studies on the living and fossil forms (Woelkerling et al., 1993) have clearly demonstrated that these characters are unreliable due to the overlaps observed between discrete species and to the wide range of variation recorded within a single species.

Another set of key parameters used to split species apart consisted of measurements: the width and height of the thallus, as well as the diameter of the "branches" (all three of which are connected to morphology and therefore irrelevant for taxonomy), the height of cell rows or of the zones comprised between successive constrictions and the diameter of the filaments. Should we plot on a graph the last two parameters for the eight species, we would get a figure with overlapping clusters. For instance, according to Pfender (1939), the filament diameters and the zone heights are respectively 6-9 and 25-40 µm in her species whereas, according to Barattolo & del Re (1984), they are respectively 2-10 and 20-40 in Marinella yugoslavica Maslov, 1962 (a nomen nudum).

In conclusion, these eight species should be synonymized because external morphology and related features (measurements) cannot be used (Woelkerling et al., 1993) and because the remaining measurements (zone heights and filament diameters) fail to differentiate species from each others.

Marinella lugeoni is ascribed to the Family Elianellaceae Granier in Granier & Dias-Brito, 2016 (in replacement of the Family Solenoporaceae Pia, 1927), that groups those extinct "calcareous algae" sharing some morphological features:

A) they are either encrusting or form free structures;

B) they have a framework made of erected filaments, more or less densely juxtaposed, with common to rare horizontal partitions (i.e., they are pluricellular algae), defining a more or less preserved lattice-network with columns and rows. Horizontal partitions have been documented in Marinella by Barattolo & del Re (1984), Leinfelder & Werner (1993), and Bucur (1994);

C) their mineralization is intracellular, centripetal (starting from the cell walls inwards), and consists of elongated fibrous calcite (probably high-Mg calcite) crystals, another feature that justifies their ascription to the calcifying red algae;

D) they are deprived of fossilized reproductive organs (which were possibly external). The "sporangia" reported by Golonka (1970) or Leinfelder & Werner (1993) in Marinella are probably microborings sensu Granier (2014), which was the conclusion of Barattolo & del Re (1984).

This family was tentatively assigned to the Order Rhodogorgonales Fredericq & Norris, 1995, a sister group to the Sporolithales (Granier & Dias-Brito, 2016).

Systematics (B.G.)

Division Rhodophyta

Class Florideophyceae Cronquist, 1960

Subclass Corallinophycidae Le Gall & Saunders, 2007

Order ? Rhodogorgonales Fredericq & Norris, 1995

Family Elianellaceae Granier in Granier & Dias-Brito, 2016

(formerly Family Solenoporaceae Pia, 1927)

Genus Marinella Pfender, 1939

Type: Marinella lugeoni Pfender, 1939, p. 215-216, Pl. II, figs. 1-2. Lectotype (defined here): Fig. 4 ; Pfender, 1939, Pl. II, fig. 1.

Fig. 4
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Figure 4: Lectotype (defined here) of Marinella lugeoni Pfender, 1939 (op. cit., Pl. II, fig. 1). Small rounded structures up to 100 µm in diameter are microborings. Thin section of the Juliette Pfender Collection with labels "2" and "Marinella". [Scale bar 500 µm]

Marinella lugeoni Pfender, 1939, nomen conservandum

(Figs. 3.A-C , 4 , 5.A-C , 6 , 7 , 8 , 9 ; Pl. 2 , figs. A-K)

Synonymy list (not exhaustive):

1916

Lithothamnion angolense n. sp.- Romanes, p. 582-583, Pl., figs. 2 [here Fig. 5.A ], 3 & 4 [here Fig. 5.B ]
Lectotype (defined here): Romanes, 1916, Pl., fig. 4 [here Fig. 5.B ]

non

1916

Lithothamnion angolense n. sp., "conceptacles".- Romanes, p. 582-583, Pl., figs. 5-6

1928

Lithothamnium (?) sp.- Yabe & Toyama, p. 150-151, Pl. XIX, fig. 5 pars

1937

Lithothamnium ? regonis n. sp.- Howe in Maury, p. 45, Pl. XXVII, figs. 2-4; Pl. XXVIII, figs. 2-3

1939

Marinella lugeoni nov. gen. nov. sp.- Pfender, p. 215-216, Pl. I, fig. 1 pars; Pl. II, figs. 1-2
Lectotype (defined here): Pfender, 1939, Pl. II, fig. 1

1959

Lithophyllum (?) shebae sp. nov.- Elliott, p. 220 & 222, Pl. 1, fig. 7

1961

Marinella lugeoni.- Johnson, p. 147-148, Pl. 31, figs. 1 [here Fig. 6 , USNM D992 a843] - 2 [here Fig. 7 , USNM D992 a844]

nom. nud.

1961

Archaeolithothamnium somensis n. sp.- Endo, p. 56, Pl. 3, figs. 1-2; Pl. 4, figs. 1-4; Pl. 17, fig. 1 [Note: according to its author, the "Holotype" consists of the "Slide no. 675" with two figures (Pl. 4, figs. 2-3) assigned to it]
Lectotype (defined here): Endo, 1961, Pl. 4, fig. 2

nom. nud.

1962

Marinella yugoslavica sp. nov.- Maslov, p. 189-192, Fig. 127.a-e; Pl. XXXIV, figs. 1-5 [Note: according to its author, the holotype is the (?) sample "no. A 1"; the figures consist of a rock sample (x2), three specimens in thin section (x10) and an enlargement (x200)]
Lectotype (defined here): Maslov, 1962, Pl. XXXIV, fig. 2

1964

Marinella lugeoni.- Johnson, p. 25, Pl. 2, fig. 10; Pl. 23, figs. 1 (excerpt from Pfender, 1939, Pl. II, fig. 2) & 3 (excerpt from Pfender, 1939, Pl. II, fig. 1)

1964

Marinella lugeoni.- Johnson, Pl. 23, fig. 2 pars (excerpt from Pfender, 1939, Pl. I, fig. 1)

1964

Archaeolithothamnium somensis.- Johnson, Pl. 40, figs. 1 (excerpt from Endo, 1961, Pl. 3, fig. 1) & 2 (excerpt from Endo, 1961, Pl. 3, fig. 2)

nom. nud.

1965

Lithothamnium ? venezuelaensis n. sp.- Johnson, p. 719, Pl. 89, figs. 1-3 [Note: according to its author, the figures correspond to "cotypic sections"]
Lectotype (defined here): Johnson, 1965, Pl. 31, fig. 1

1965

Marinella lugeoni.- Imaizumi, p. 57-60, Pl. 11, figs. 7-17; Pl. 12, figs. 1-9; Pl. 13, figs. 1 (excerpt from Johnson, 1964, Pl. 2, fig. 10) & 2-4; Pl. 14, figs. 1-2

1965

Lithothamnium ? primitiva n. sp.- Johnson & Kaska, p. 30-31, Pl. 6, fig. 1 [here Fig. 3.A , USNM 42457]

1965

Marinella lugeoni.- Johnson & Kaska, p. 74, Pl. 6, fig. 2 [here Fig. 3.B , USNM 42547]

1965

Girvanella minuta.- Johnson & Kaska, p. 96, Pl. 30, fig. 1 [here Fig. 3.C , USNM 42547]

1968

Lithothamnium ? sp.- Johnson, p. 7

1968

Lithophyllum ? sp.- Johnson, Pl. 1, fig. 1 [here Fig. 8 , USNM 42606]

1968

Marinella lugeoni.- Bouroullec & Deloffre, p. 218-219, Pl. 1, figs. 7-9

1969

Lithothamnium ? primitiva.- Johnson, p. 11, Pl. 3, fig. 1 = Pl. 4, fig. 1 (excerpt from  Johnson & Kaska, 1965, Pl. 6, fig. 2 [here Fig. 3.B , USNM 42547])

1969

Lithothamnium (?) shebae.- Johnson, p. 11, Pl. 3, fig. 1 (excerpt from Elliott, 1959, Pl. 1, fig. 7)

1969

Marinella lugeoni.- Johnson, p. 34, Pl. 4, fig. 2 (excerpt from Johnson & Kaska, 1965, Pl. 6, fig. 2 [here Fig. 3.B , USNM 42547]); Pl. 21, figs. 1 (excerpt from Pfender, 1939, Pl. II, fig. 2), 2 pars (excerpt from Pfender, 1939, Pl. I, fig. 1) & 3 (excerpt from Pfender, 1939, Pl. II, fig. 1); Pl. 22, figs. 1 (excerpt from Imaizumi, 1965, Pl. 13, fig. 2) & 2 (excerpt from Imaizumi, 1965, Pl. 13, fig. 1)

nom. nud.

1969

Lithothamnium ? venezuelaensis.- Johnson, p. 13, Pl. 5, figs. 1 (excerpt from Johnson, 1965, Pl. 89, fig. 2), 2 (excerpt from Johnson, 1965, Pl. 89, fig. 1) & 3 (excerpt from Johnson, 1965, Pl. 89, fig. 3)

1970

Marinella lugeoni.- Golonka, p. 82-84, Photos 8-9

1971

Lithophyllum (?) shebae.- Basson & Edgell, p. 426 & 428, Pl. 7, figs. 1-2

1982

Marinella lugeoni.- Bengtson & Berthou, Pl. 4, fig. 5

1984

Marinella lugeoni.- Barattolo & del Re, p. 205-207 & 212-214, Fig. 1; Pl. I, figs. 1-2; Pl. II, figs. 1-2

nom. nud.

1984

Marinella yugoslavica.- Barattolo & del Re, p. 207-208 & 215-223, Figs. 3-6; Pl. III, figs. 1-5; Pl. IV, figs. 1-2; Pl. V, figs. 1-2; Pl. VI, figs. 1-4; Pl. VII, figs. 1-2; Pl. VIII, figs. 1-3; Pl. IX, figs. 1-4; Pl. X, fig. 1

nom. nud.

 ? 1984

Marinella yugoslavica.- Barattolo & del Re, p. 224-225, Pl. XI, figs. 1-3; Pl. XII, figs. 1-2; Pl. XIII, figs. 1-3

1987

Marinella lugeoni.- Granier, p. 266, Pl. 49, figs. a-d & f

1991

Lithothamnium angolense.- Granier et al., p. 173-175, Pl. 3, fig. 2

1991

Marinella lugeoni.- Granier et al., p. 175, Pl. 5, fig. 8 pars

1991

Marinella lugeoni.- Schlagintweit, p. 47, Pl. 16, figs. 6-7

1993

Marinella lugeoni.- Leinfelder & Werner, p. 107-116, Pl. 1, figs. 1-4 pars, 5, 6-7 pars & 8-10; Pl. 2, figs. 1-12; Pl. 3, figs. 1-2 pars & 3-9

1994

Marinella lugeoni.- Bucur, p. 162, Pl. XIX, figs. 9-12

non

2007

Lithothamnion angolense.- Tomás et al., p. 91, Fig. 8

2008

Marinella lugeoni.- Granier et al., p. 175, Pl. 2, fig. 1

2015

Marinella lugeoni.- Granier et al., p. 527, Pl. 9

Fig. 5
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Figure 5: A-B) Lithothamnion angolense Romanes, 1916; A) "Longitudinal section of 290" (excerpt from Romanes, 1916, Pl., fig. 2); B) "Transverse section of 290, showing lines produced by compression, due to growth" (excerpt from Romanes, 1916, Pl., fig. 4). It is the lectotype defined here; C) transverse section of Marinella lugeoni Pfender, 1939, for comparison. Thin section UPAFSE 149, locality SE 6, Sergipe, Brazil; Albian, Riachuelo Formation (enlarged view of Pl. 9, up right figure in Granier, 2015). [A) scale bar 500 µm; B-C) scale bar 250 µm]

Fig. 6
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Figure 6: Marinella lugeoni in thin section USNM no. D992 a843 of the Jesse Harlan Johnson Collection (Johnson, 1961, Pl. 31, fig. 1). [Scale bar 500 µm]

Fig. 7
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Figure 7: Marinella lugeoni in thin section USNM no. D992 a844 of the Jesse Harlan Johnson Collection (Johnson, 1961, Pl. 31, fig. 2). [Scale bar 100 µm]

Fig. 8
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Figure 8: Marinella lugeoni in thin section USNM no. 42606 of the Jesse Harlan Johnson Collection as Lithophyllum ? sp. (Johnson, 1968, Pl. 6, fig. 1). [Scale bar 500 µm]

Taxonomic note: Considering the International Code of Nomenclature for algae, fungi, and plants (McNeill et al., 2012), Lithothamnion angolense Romanes, 1916, or Lithothamnium ? regonis Howe in Maury, 1937, have priority [cf. Art. 11 of the ICBN] over Marinella lugeoni Pfender, 1939. However, the use of the last name is by far the most common and it is permanently attached (the nomenclatural type) to the genus Marinella Pfender, 1939, the validity of which is established beyond doubt; on the other hand, the first two names were correctly used only once, in their original descriptions (remark: the Lithothamnion angolense quoted by Tomás et al., 2007, refers to a discrete species, i.e., a genuine Lithothamnion species, but not the Angolan form). For these reasons and for stability of the nomenclature, as in Granier et al. (1991) earlier, the first author (B.G.) proposes to conserve the name Marinella lugeoni Pfender, 1939.

Description: Except for an additional remark on the occurrence of cross partitions within the filaments, there is no need to alter the original description (Pfender, 1939, p. 215-216):

"Il s'agit probablement d'une algue filamenteuse, (...) zonée, genre 'bouffée de pipe' de H. Derville (…). Le thalle est formé par des filaments juxtaposés, sans cloisons*, mais avec des constrictions ; ils changent de direction fréquemment, tout en restant rayonnants, jamais pelotonnés. Le tissu forme un chevelu plutôt qu'un réseau. L'ensemble constitue un thalle digité, cranté, en éventail, d'aspect sombre, car le diamètre des filaments tubuleux est très petit, 6 à 9 µ généralement (pl. II, fig. 1). Les constrictions, dans les filaments, sont souvent à un même niveau et forment des lignes concentriques sur le thalle (pl. II, fig. 2). La section transversale des tubes, juxtaposés, est polygonale ou arrondie et semble de dimension un peu variable ; 9 µ de diamètre généralement. Ces thalles sont aisément reconnaissables, dans les préparations, où ils paraissent plus sombres, étant formés d'un tissu plus serré, plus régulier, et non orientés concentriquement comme les filaments des Girvanelles."

["It is probably a filamentous alga, (...) zoned with the H. Derville's 'puff of pipe smoke' type (…). The thallus consists of juxtaposed filaments, without cross partitions [*], but with constrictions; they frequently vary in direction, while remaining radiating, never curled up. The fabric looks hairy rather than forming a lattice. The whole thing forms a finger thallus, notched, fan-shaped, dark, because the diameter of the tube-like filaments tubuleux is very small, usually 6 to 9 µ (Pl. II, fig. 1). The constrictions within the filaments are often at the same level and form concentric lines on the thallus (Pl. II, fig. 2). Transverse sections of juxtaposed tubes are either polygonal or rounded and apparently may slightly vary in size; with a diameter of 9 µ in average. These thalli are readily identifiable in the thin sections where they appear darker, being formed of a denser, more regular fabric, which is not concentrically oriented like the Girvanella filaments."]

Finally, as pointed out by our predecessors (e.g., Johnson, 1965; Johnson & Kaska, 1965; Barattolo & del Re, 1984, inter alia), one should add that there is "no evidence of reproductive structures".

Age of the stratum typicum

According to Lugeon (in Pfender, 1939, p. 213), the stratum typicum should correspond to the lower Liassic (lowermost Jurassic). However that was not the opinion of Bataller (1943, p. 9) who stated: "Recientemente M.lle Pfender (...) ha descrito unas algas nuevas procedentes de Navajas (Castellón de la Plana) y que según Lugeon y A. Marin, que las recogíeron, pertenecen al Liasico inferior sin mas precisión de nivel: por el aspecto de la roca simplemente, pues no hemos explorado nunca esta zona, hubiéramos creído tratarse del Jurásico superior, pues el Liasico tiene aquí poco desarrollo en comparación con el que se encuentra más al Norte, especialmente en la provincia de Teruel" [Recently Miss Pfender (...) described some new algae from Navajas (Castellón de la Plana) and that according to Lugeon and A. Marin, who collected them, belong to lower Liassic strata without further precision on the level: simply on the basis of the aspect of the rock, because we have never explored this area, we would have believed it to be Upper Jurassic, because the Liassic in this region is very little developed compared with that which is found farther northwards, especially in the Province of Teruel]. In the notice of the Mapa geologico de España for Jerica (Ortí Cabo, 1977), the strata bearing Marinella, together with Alveosepta jaccardi (Schrodt, 1894), are given a Middle-Late Kimmeridgian age. The latest foraminifer, which is a marker for the Upper Oxfordian - Kimmeridgian interval, is present in our set of thin sections (Fig. 9.A-E & G-H ). Pfender (1939) probably misidentified it as "très petites Choffatella Schlumberger" ["very small Choffatella Schlumberger"].

Fig. 9
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Figure 9: A-E & G-H) Alveosepta jaccardi (Schrodt, 1894); F) Nautiloculina oolithica Mohler, 1938. Thin sections: A) Navajas 12, B) Navajas 11, C-E) Navajas 7, F) Navajas 3, G-H) Navajas 5, Juliette Pfender Collection. [All photos with 250 µm scale bar]

The Brazilian morphotypes

In some thin sections from the Albian limestones of the Riachuelo Formation (Sergipe basin, Brazil), we identified two type of grains or structures with Marinella: 1) Marinella is a main contributor to some biolithites (framestones) and rhodolithes, commonly asymmetrical, 2) Marinella is also found in the form of small lumps, rarely exceeding 1 mm in diameter. The characteristic feature of these Marinella lumps is to have a hollow structure (filled by a late drusy calcitic cement), subcircular (Pl. 2 , figs. B-F & I-K; Granier, 2015, Pl. 9, down left figure) to ovoid (Pl. 2 , fig. G) in section, and eventually elongated and rounded on one end and open on the opposite end (Pl. 2 , figs. A & H). This hole is not a sporangia nor a microboring (see discussion on "empty sporangial complexes versus trace fossils" in Woelkerling et al., 2014). It is located in the core of the lump and at the base of the Marinella filaments. Such cylindrical holes (with a maximum diameter of 300 µm) probably represent moulds of parts of larger organisms that did not fossilize, such as the stipes of non-calcifying brown, green or red algae or of seagrasses, upon which Marinella was growing. When these organisms died, lumps were freed like the pearls of a broken necklace.

Conclusions

This study of Marinella specimens, which include type-materials from Juliette Pfender and Jesse Harlan Johnson collections, helped us to sort out some of the details pertaining to the alga systematics, its stratigraphy and its paleoenvironments:

Acknowledgments

This publication follows the "Revision of the Juliette Pfender Collection - Part 1" (Granier & Dias-Brito, 2016) and the "Revision of the Jesse Harlan Johnson Collection - Part 1" (Granier et al., 2013). In 2015, the first author (B.G.) was the laureate of a Franco-Brazilian professorship at the UNESPetro in Rio Claro (São Paulo State, Brazil) that enabled him to work on Brazilian material. Earlier, in 2012, he also benefited from a Smithsonian Fellowship allowing him to investigate the J. Harlan Johnson Collection stored in the premises of the Smithsonian Institution. He would like to thank the staff of the Department of Paleobiology at the Smithsonian National Museum of Natural History and particularly William A. DiMichele and Jonathan G. Wingerath for their hospitality and having facilitated his work there. He is also grateful for the support provided by the successive curators of paleontological collections at the Université Pierre et Marie Curie, first Jean-Pierre Bellier, then Isabelle Rouget. Special thanks to Christine Appia, Mitsuru Arai, Michael P.A. Howe, and Hermès Dias-Brito who helped with documentation. Phil Salvador helped polishing the English text. This research is associated with the "Carbonatos do Brasil Project" linked to the Brazilian Sedimentology/Stratigraphy Net sponsored by Petrobras; it was also partly sponsored by the Foundation "Carnets de Géologie".

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Appendix (list of species)

Plates

Pl. 01
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Plate 1: Two rock samples representing the original material used to make the thin sections. Figs. A-B) label 29261 / Algues / Lias inf. / Navajas (Espagne) / Province de Valence / M. Lugeon et Marin 1928 / M. 2379. Figs. C-D) label 29261 / Algues / Lias inf. / Navajas / Province de Valence / Espagne / M. Lugeon 1928 / M. 2379 ; fig. C) see Pfender, 1939, Pl. I, fig. 1. [All photos with 1 cm scale bar]

Pl. 02
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Plate 2: Marinella lumps. Fig. A) thin section UPAFSE 117; fig. B) thin section UPAFSE 99; fig. C) thin section UPAFSE 118; fig. D) thin section UPAFSE 94; fig. E) thin section UPAFSE 117; fig. F) thin section UPAFSE 121; fig. G) thin section UPAFSE 99; fig. H) thin section UPAFSE 99; fig. I) thin section UPAFSE 108; fig. J) thin section UPAFSE 118b; fig. K) thin section UPAFSE 118. Fig. D, locality SE1, and figs. A-C and E-K, locality SE 3, Sergipe, Brazil; Albian, Riachuelo Formation. [All photos with 250 µm scale bar]


X

Actually, Barattolo and del Re (1984: Fig. 6.c-d; Pl. II, figs. 2-3; Pl. VIII, figs. 1-2) and later Leinfelder and Werner (1993: Pl. 2, figs. 2-4) and (Bucur, 1994, Pl. XIX, figs. 10-12) did document the occurrence of such partitions (rather rare).