# Lectotype designation for Alveolina boscii and Fabularia discolites (Foraminifera) from the rediscovered part of the Defrance collection

Jérôme Tréguier, François Le Coze, Roch-Alexandre Benoit

► To cite this version:

Jérôme Tréguier, François Le Coze, Roch-Alexandre Benoit. Lectotype designation for Alveolina boscii and Fabularia discolites (Foraminifera) from the rediscovered part of the Defrance collection. Revue de Micropaléontologie, 2019, 63, pp.37-43. 10.1016/j.revmic.2019.03.001. insu-02159252

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1 Lectotype designation for *Alveolina boscii* and *Fabularia discolites*

2 (Foraminifera) from the rediscovered part of the Defrance collection

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J. TRÉGUIER¹, F. LE COZE² & R.-A. BENOIT¹

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¹ Musée des Sciences, Place de Hercé, 53000 Laval, France

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(jerome.treguier@laval.fr)

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² World Foraminifera Database, François Le Coze, 80 rue Jean Parot, 42100 Saint-Etienne, France

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(francois.lecoze@9online.fr)

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10 Abstract

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12 Five lots of Foraminifera, belonging to the Marin Defrance (1758-1850) collection, were discovered in the collection of
13 the *Musée des Sciences* in Laval (Mayenne, France). The original material used by Defrance to describe *Alveolina
14 boscii* (Defrance in Bronn, 1825) and *Fabularia discolites* Defrance in Bronn, 1825 was identified among them,
15 allowing the designation of a lectotype and paralectotypes for those two species. The systematic description of *A. boscii*
16 and *F. discolites* is reviewed. The presence of these specimens in the museum collections is still puzzling, since the
17 complete Defrance's collection was thought to have been lost during World War II.

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19 Résumé

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21 Cinq lots de foraminifères appartenant à la collection Marin Defrance (1758-1850) ont été retrouvés au Musée des
22 Sciences de Laval (Mayenne, France). Le matériel original utilisé par Defrance pour décrire *Alveolina boscii* (Defrance
23 *in* Bronn, 1825) et *Fabularia discolites* Defrance *in* Bronn, 1825 a été identifié au sein de ces échantillons, permettant la
24 désignation d'un lectotype et de paralectotypes pour ces deux espèces. La systématique d'A. boscii et F. discolites est
25 révisée. La présence de ces spécimens dans les collections de ce musée est toujours énigmatique, puisque la totalité de
26 la collection Defrance semblait avoir été perdue pendant la Seconde Guerre mondiale.

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28 Keywords

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30 Foraminifera, Eocene, Defrance's collection, Systematic revision, Musée des Sciences in Laval

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32 Mots clés

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34 Foraminifères, Éocène, Collection Defrance, Systématique, Musée des Sciences de Laval

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## 1. Introduction

During the inventory of the palaeontological collections held in the *Musée des Sciences* in Laval (Mayenne, France), seventeen lots were found, all part of the collection of famous French naturalist Jacques-Louis Marin Defrance (1758-1850) (Tréguier *et al.*, 2017; Pacaud *et al.*, 2017). Five of these are constituted by Foraminifera, and include specimens of *Alveolina boscii* and *Fabularia discolites*.

Bosc (1802) first described the species as "Alvéolite grain de fétuque (Fetusca seed)" and "Alvéolite grain de millet (Millet seed)", Alvéolite being the name given by Lamarck (1801) to what he thought was a genus of coral. Defrance (1820) referred to Bosc "Alvéolites" to describe two new genera and two new species "Oryzaire-Bosc" and "Fabulaire discolithe", Latinised by Bronn in 1825.

The original material used by Defrance to describe these two species was identified among the five lots cited herein, allowing the designation of a lectotype and paralectotype. The systematic description of *A. boscii* and *F. discolites* is reviewed.

## 2. History of the collection

### 2.1 Formation

The Defrance collection of malacological fossils was probably one of the largest in Europe during the late XVIII<sup>th</sup> and the early XIX<sup>th</sup> centuries (von Humbolt, 1825, p. 205). During field work, he collected in Grignon (now Thiverval-Grignon, Yvelines, France) around 500 species of fossil shells, the vast majority of which had not been described in any natural history book before (Lamarck, 1802, p. 305). Lamarck described and published a large number of these new species between 1803
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and 1811. The *Muséum National d'Histoire Naturelle* in Paris commissioned the illustration of half of the specimens collected by Defrance in Grignon to enhance its vellum collection. During his career, Defrance described, among others, around 40 new Foraminifera taxa, including 7 new genera, such as *Frondicularia*, *Pyrgo* and *Textularia* to name the best known only, essentially in the "Dictionnaire des Sciences Naturelles" volumes, from 1816 to 1829.

## 2.2 Loss and partial rediscovery

After he passed away, Defrance's family bequeathed his collection to the *Musée d'Histoire Naturelle* in Caen (Calvados, France), where it was deposited in 1869 after some deplorable misfortune (Tréguier *et al.*, 2017). Unfortunately, this institution, along with all its collections, was destroyed on the 7th of July 1944, during an Allied bombing. Therefore, these collections were thought to be lost, once and for all.

Somehow, several specimens of the Defrance collection found their way to the *Musée des Sciences* in Laval (5 lots of Foraminifera, 3 of Brachiopoda, 7 of Gastropoda and 5 of Bivalvia). They have been identified by the presence of typical labels used by Defrance (Bigot, 1906), and also thanks to the inventories compiled by Bigot (1905, 1906, 1907) and Bigot and Matte (1903, 1904) in which are cited precisely which specimens were found in the *Musée des Sciences* in Laval, as well as a manuscript note from Maurice Cossmann found associated with specimens of Bivalvia. Until now, it is still unclear how they ended up in this museum, but Daniel Œhlert (1849-1920), who was curator at the time, might have played a part, as he acquired a lot of specimens with different taxa and/or periods.

## 3. Material and Methods
109 Both *Alveolina* and *Fabularia* belong to the Larger Benthic Foraminifera (LBF). LBF, and especially the alveolinids, are studied through various oriented sections (mainly axial and equatorial). The standard procedure for studying alveolinids is detailed in Hottinger (1974, pp. 19 to 26).

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114 Almost all the specimens of the Defrance collection are glued onto blue card which are kept in glass tubes; these are then glued onto beige card. Defrance made several axial sections for *A. boscii* and *F. discolites* from which it was possible to choose lectotypes. We assessed the historical and museum value of the material in the collection. We did not cut equatorial sections from free specimens.

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120 In the past ten years, the routine use of X-ray microtomography (microCT) has allowed to study the complex inner architecture of LBF tests (Speijer et al. (2008), Briguglio and Benedetti (2012), Briguglio et al. (2013)). This safe and non-destructive method of analysis is useful to study museum collections without altering historical material.

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125 For the rediscovered samples holding species which were not described by Defrance, we undertook our identifications based on the locality where they were collected. We tried to compare the shape of the specimens with known species described from the same localities. Without further investigations, our identifications remain indicative especially for *Alveolina* species.

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130 Furthermore, the two main samples of *A. boscii* and *F. discolites* in the Defrance collection were found in Grignon, locality from which some additional material was recently collected, allowing to compare complete tests of the former with the latter.

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The determination of Grignon as the type locality for *A. boscii* and for *F. discolites* allows also to fix the type level as Middle Lutetian for both species. Lower Lutetian is absent from Grignon and the poorly visible Upper Lutetian is too indurated to extract similar specimens to those collected by Defrance.

Measurements were made with a stereomicroscope (Leica MZ95) with an integrated camera (Leica DC 300), using the imaging software Leica IM50 at the University of Rennes 1. The specimens are stored in the collections of the *Musée des Sciences* in Laval (Mayenne, France).

### 3.1 *Alveolina* specimens

Three glass tubes containing *Alveolina* specimens.

Lot n°ML-PAL-22432: This lot contains eight complete specimens and five specimens cut longitudinally, registered as ML-PAL-22432-1 to ML-PAL-22432-13 (Pl. 1, fig. 1-4) (Tréguier *et al.*, 2017, p. 23, figs. 4A-E). They were identified as *Oryzaria bosci* (Def.) by Defrance and were collected from the Lutetian sands of Thiverval-Grignon (Yvelines), their type locality. The length of the axis of coiling is comprised between 2.97 mm and 4.46 mm, the equatorial diameter ranges from 0.95 mm to 1.31 mm, and the elongation index from 2.36 to 3.88. However, the last whorls are missing on all the specimens. A complete description of the morphological features is given in the lectotype designation paragraph. This glass tube is identified as containing the original material used by Defrance to describe *A. boscii* on the account of the label glued on it, which indicates « Types Dict. Sc. nat. t XVI. 1820 p. 104. coll. Defrance » referring to the original publication of Oryzaire-Bosc.

Lot n°ML-PAL-22330: This lot contains a single specimen, complete and identified as both
160 Oryzaire ovoides (Def.) and *Oryzaria bosci* (Def.) (fig. 1 in text) (Tréguier *et al.*, 2017, p. 23, figs. 4F-H). It was collected in Vendémies (Aude, France), which gave its name to the Vendémies Formation from the Lower Ypresian (Ford *et al.*, 2016). The length of the axis of coiling is 3.83 mm, the equatorial diameter is 1.89 mm, giving an elongation index of 2.03. This specimen is neither *A. ovoidea* d'Orbigny, 1826 nor *A. boscii* as Defrance's identifications could have suggested.
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165 The origin of the specimen and its dimensions indicate that it possibly belongs to the group of *A. subpyrenaica* Leymerie, 1846 but based on a unique complete specimen the identification remains tentative.
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171 Lot n°ML-PAL-22350: This lot contains seven complete specimens and three cut longitudinally (Tréguier et al., 2017, p. 23, figs. 4I-L) (fig. 2 in text), collected from Chaumont-en-Vexin (Oise).
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173 The length of the axis of coiling ranges from 2.30 mm to 3.42 mm, the equatorial diameter from 0.95 mm to 1.44 mm and the elongation index from 1.97 to 2.56. The origin of the specimens and their dimensions indicate that they possibly belong to *Alveolina oblonga* d'Orbigny, 1826. These specimens have been compared with specimens collected and identified as *Alveolina oblonga* by Abrard from the same locality (Cuisian level) and housed in the *Muséum National d'Histoire Naturelle* in Paris (MNHN-GG-GG2004-76822). They are very similar in shape and share the same iron oxide redish coloration. The cut sections left by Defrance are covered with glue and difficult to interpret.
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184 **3.2 Fabularia specimens**

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186 Two glass tubes containing *Fabularia* specimens.
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Lot n°ML-PAL-24962: This lot contains thirteen complete specimens and four specimens cut longitudinally, registered as ML-PAL-24962-1 to ML-PAL-24962-17 (Pl. 1, fig. 5-8) (Tréguier *et al.*, 2017, p. 25, figs. 6A-E). Seven of them are still glued to blue card. At the top of this card is written "Vélin 48 f.16 alvéolite grain de Millet Bosc. B des Sc. n°61 f.4", which refers to both the vellum illustration and the original description by Bosc (1802). They are labelled under the vernacular name "Fabulaire discolithe (Def.)" and were collected from the Lutetian sands of Thiverval-Grignon (Yvelines), the type-locality of *F. discolites* but also of *F. ovata* its senior synonym. The length of these specimens ranges between 3.9 mm and 6.48 mm, with a width between 3.00 mm and 4.92 mm. All the specimens are identified as *F. discolites*. The identification is fairly easy because there is no other *Fabularia* species than *F. discolites* in the Paris Basin and the specimens are very characteristic.

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Lot n°ML-PAL-29059: This lot contains eight complete specimens (Tréguier *et al.*, 2017, p. 25, figs. 6F-I) (fig. 3 in text), collected from the Lutetian of Hautteville-Bocage (Manche, France). The label by Defrance mentions: "Fabularia discolithes var.". They are ranging from 5.7 mm to 8.6 mm in length, and from 4.1 mm to 5.9 mm in width. They have been identified as *F. compressa* d'Orbigny, 1850, which is regarded as a junior synonym of *F. discolites* by various authors (Nicolas (1945), Le Calvez (1947, 1970). The larger dimensions correspond to the description given by d'Orbigny (1850) for specimens of *F. compressa* collected around Valognes (Manche, France).
According to Nicolas (1945, p. 538), Defrance considered specimens from Valognes to be a variety of *F. discolites* from Grignon which is in turn confirmed by the label.

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212 **4. Systematic description and lectotypes designation**

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214 Phylum FORAMINIFERA d'Orbigny, 1826

215 Class TUBOTHALAMEA Pawlowski, Holzman & Tyszka, 2013

216 Order MILIOLIDA Delage & Hérouard, 1896

217 Suborder MILIOLINA Delage & Hérouard, 1896

218 Superfamily ALVEOLINOIDEA Ehrenberg, 1839

219 Family ALVEOLINIDAE Ehrenberg, 1839

220 Genus *Alveolina* d'Orbigny, 1826

221 Type species: *Oryzaria boscii* Defrance in Bronn, 1825 (subsequent designation, ICZN Opinion

222 1501 (1988))

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224 *Alveolina boscii* (Defrance in Bronn, 1825)

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226 1802 Alvéolite grain de fétuque Bosc. p. 99, pl. 5, fig. 3 A-C

227 1808 *Miliolites sabulosus* Montfort. Vol. 1, p. 175

228 1820 Oryzaire-Bosc Defrance. p. 104

229 1825 *Oryzaria boscii* Defrance in Bronn. p. 44

230 1826 *Alveolina Boscii* [sic] d'Orbigny. p. 306 n° 5, Modèles, n° 50, deuxième livraison

231 1960 *Alveolina boscii*. Hottinger, pp. 149-151, text. figs 78-80, pl. 10, figs 19-22

232 1978 *Fasciolites (Microfasciolites) boscii*. Gaemers p. 107 pl. 1, figs 19-24

233

234 The long nomenclatural story of the genus *Alveolina* and its type species has been exhaustively

235 documented by Loeblich and Tappan when they submitted Case 2356 to the International

236 Commission on Zoological Nomenclature (ICZN) in 1987.
237 Answering the Case, the ICZN in its Opinion 1501 in 1988, selected *O. boscii* as the type species for the genus *Alveolina*. The genera *Fasciolites* Parkinson, 1811, *Oryzaria* Defrance in Bronn, 238 1825, *Microfasciolites* Gaemers, 1978 and the binomen *Miliolites sabulosus* Montfort, 1808 were 239 rejected. 240

241 The ICZN also invited its members to designate the figured specimen of "alvéolites grain de festuque" [sic] by Bosc (1802, p. 99, pl. 5 fig. 3 A-C) as the lectotype of *O. boscii* but this part of 242 the vote was withdrawn from the final decision. 243

244 Lectotype designation 245

246 A specimen from lot n°ML-PAL-22432 is supposedly illustrated on the vellum from the *Muséum National d'Histoire Naturelle* in Paris (vellum n° 48, fig. 17, a, b) (Fig. 4 in text), but the drawings 247 are not accurate enough to identify which specimen of this batch is actually illustrated. 248

249 Therefore specimen ML-PAL-22432-4 (Pl. 1, fig. 2, axial section) is designated as lectotype, based 250 on good preservation, and presence of diagnostic features of *A. boscii* on this specimen. 251

251 Description: Megalospheric form, 12 visible whorls, last whorls are missing, axial diameter: 4.28 252 mm; equatorial diameter: 1.31 mm; elongation index: 3.27; megalosphere diameter: 115 µm. 253

253 Specimen ML-PAL-22432-7 (Pl. 1, fig. 3, axial section) is not as well preserved as the latter, but 254 still good enough to bring further information concerning *A. boscii* (microspheric form). It is 255 designated as paralectotype. 256

256 Description: Microspheric form, 12 visible whorls, last whorls are missing, axial diameter: 3.65 257 mm; equatorial diameter: 1.17 mm; elongation index: 3.12. 258

258 The others specimens showing axial section (ML-PAL-22432-10, ML-PAL-22432-11, ML- 259 PAL22432-12) are too damaged and/or incomplete to be designated as paralectotypes. 260

260 Pending further studies, the complete specimens [ML-PAL-22432-1, ML-PAL-22432-2, ML- 261 PAL22432-3 (Pl. 1, fig. 4), ML-PAL-22432-5, ML-PAL-22432-6, ML-PAL-22432-8, ML-PAL- 262 22432-9, ML-PAL-22432-13] are identified as belonging to the same species. 263
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266 Diagnosis

267 Fusiform test, elongated along the coiling axis with pointed poles in adult stages, small size with
268 very tight equatorial spiral and numerous, very small cavities in the shell, no supplementary
269 passages in the basal layers (Hottinger, 1974). Bosc (1802) already compared his species to a "grain
270 de fétuque", an elongated grass seed with pointed endings. Hottinger defined A. boscii as a "small
271 sized Alveolina with an extremly fine structure" (1960, p. 150) and the whole of his description
272 (1960, pp. 149-151) remains the most detailed. He mentioned that the forms from the Paris Basin
273 are easily collected but their last whorls are often missing. He described well preserved specimens
274 from Grignon with an axial diameter of 4.5-5.5 mm, an elongation index for the adult forms of
275 2.43.2 and 3.0-3.8 in the first 12 whorls. Hottinger (1999, p. 391) mentioned also an elongation
276 index for A. boscii from the lower Middle Eocene of the Paris Basin "reaching 4 at most". Hottinger
277 added that the diameter of the megalosphere never exceeded 125 microns and that the morphology
278 of microspheric and megalospheric forms are similar except for the tighter spire in the first whorls
279 of the megalospheric forms.

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281 Stratigraphy

282 Hottinger mentionned A. boscii in the Lower and Middle Lutetian (1960, p. 149). This is in
283 agreement with the SBZ (Shallow Benthic Zones) 13 and part of SBZ 14 as in Papazzoni et al.
284 (2017, p. 3).

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286 Occurrence

287 A. boscii is found in all the Lutetian of the Paris Basin and it is abundant in Grignon, its type
288 locality (Hottinger, 1960 and our own observations). Hottinger mentionned it in Cahurt
289 (SainteMarie-de-Gosse, Bas-Adour, France), Biron (Bas-Adour, France) and in the basal layers of
Colombres from the coast of Cantabria (San Vincente de la Barquera, Spain). Gaemers (1978) found *A. boscii* in the Tremp Basin (Tremp section, Puente de Montañana, Beniure) South-Central Pyrenees (Spain). It is also found in the Belgian Basin. As noticed by Sztrákos *et al*. (2010, p. 28), there is a similarity of the Alveolina fauna from the Lower Lutetian of the Aquitanian Basin and the Paris-Belgian Basin which is characterised by the "odd pair" of elongated Alveolina: *A. boscii* - *A. stipes* (Hottinger, 1999). Drobne *et al*. (2011, table 3, p. 729) mentioned also *A. boscii* in the Paleogene Adriatic Platform (Osp, Rakitovec) and in the East-Tethyan (Libya).

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298 Remarks

Hottinger (1960) listed *A. boscii* in the fusiformis *Alveolina* category but did not include it in any of his new erected groups. Hottinger considered *A. boscii* as a late intermediary form between *Glomalveolina* and *Alveolina stricto sensu* (1960, pp. 149, 150).

Jámbor-Kness (1981) tentatively proposed the new non phylogenetic *Alveolina boscii* group to comprise species not included in any group by Hottinger (1960) and Drobne (1977). This has not been followed by any other authors and it must not be mistaken with the *Alveolina boscii* group of Hottinger (1974). In his group, Hottinger separated specimens from the Aquitaine Basin, he included in *A. boscii* in 1960, which have a greater elongation index and "may represent a stratigraphic follower of *A. boscii*" (1974, p. 63). Drobne (1977, p. 53) described for the first time forms closely related to *Alveolina* (*Alveolina* aff. *boscii* from the Cuisian (Late Ypresian) of Slavec (Slovenia). More recently Vecchio *et al*. (2007, p. 35) highlighted the marginal status of the Paris Basin compared with the Mediterranean Neotethys and the Aquitaine Basin; a low faunal diversity giving "few but exceptionally variable species". In their observations *A. boscii* "represents a whole phyletic lineage ranging at least from the Ypresian to the Upper Lutetian" and specimens of "similar size and shape relationship" need to be reviewed "as soon as enough material" is collected for study.

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316 Family FABULARIIDAE Ehrenberg, 1839

317 Genus *Fabularia* Defrance, 1820

318 Type species: *Fabularia discolites* Defrance in Bronn, 1825 (subsequent designation by monotypy,

319 Defrance in Bronn, 1825, p. 43) = *Nummulites ovata* Roissy, 1805

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321 *Fabularia ovata* (Roissy, 1805)

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323 1802 Alvéolite grain de millet Bosc. p. 99 fig. 4 A-C

324 1802 Discolithes n° 9 Fortis. p. 109-110, pl. II fig. Z, 1, 2

325 1805 *Nummulites ovata* Roissy. p. 59

326 1820 Fabulaire discolithe Defrance. p. 103 (vernacular, not available)

327 1825 *Fabularia discolites* Defrance in Bronn. p. 43

328 1826 *Fabularia discolithes* [sic]. D'Orbigny, p. 307 n° 1, pl. 17 figs 14-17, Modèles, n° 100,
329 quatrième livraison

330 1850 *Fabularia compressa* d'Orbigny. p. 409

331 1861 *Fabularia ovata* = *F. discolithes* [sic]. Parker and Jones, p. 162

332 1905 *Fabularia discolithes* [sic]. Schlumberger, pp. 129-131, text-figs 22-24, pl. 3 figs. 37-38

333 1970 *Fabularia discolithes* [sic]. Le Calvez, p. 56, pl. 46, fig. 9

334

335 Roissy (1805) gave a description of this species on the basis of Fortis drawings (1802) naming it

336 *Nummulites ovata*. Defrance (1820) described Bosc's "alvéolite grain de millet" as *F. discolithe*,

337 later Latinised as *F. discolites* in Bronn (1825), the unique species quoted under his new genus

338 *Fabularia*. Parker and Jones (1861) were the first to conclude that *F. ovata* was a senior synonym

339 of *F. discolites*. Unlike English-speaking authors, several French-speaking authors continued to use

340 *F. discolithes* [sic], following d'Orbigny, instead of *F. ovata*. Nicolas (1945) compared topotypes of

341 *F.*
342 *discolites* from Grignon and specimens from the Schlumberger's collection from Valognes (Manche), the type locality of *F. compressa*, and concluded that the two species are synonyms. Le Calvez (1947, p. 34) also considered the synonymy between *F. discolites* and *F. compressa* which she confirmed after studying the material left by d'Orbigny (1970, p. 56).

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347 Lectotype designation

348 A specimen from lot n° ML-PAL-24962, glued onto a card, is supposed to be illustrated on a vellum from the *Muséum National d'Histoire Naturelle* in Paris (vellum n° 48, fig. 16a-b) (Fig. 4 in texte), but the drawings are not accurate enough to confirm which specimen of this batch is actually represented on it.

352 Specimen ML-PAL-24962-1 (Pl. 1, fig. 6, axial section) is designated as lectotype, based on good preservation, and the presence of the diagnostic features of *F. discolites*. Axial diameter: 5.31 mm; equatorial diameter: 3.96 mm.

355 Specimen ML-PAL-24962-4 (Pl. 1, fig. 7, axial section) is less well preserved than the previous, but still good enough to present additional information concerning *F. discolites*. It is designated as paralectotype. Axial diameter: 4.00 mm; equatorial diameter: 2.97 mm.

358 The others specimens showing axial section (ML-PAL-24962-3, ML-PAL-24962-5) are too damaged and/or incomplete to be designated as paralectotypes.

360 Pending further studies, the complete specimens [ML-PAL-24962-2, ML-PAL-24962-6, ML-PAL-24962-7, ML-PAL-24962-8, ML-PAL-24962-9, ML-PAL-24962-10, ML-PAL-24962-11, ML-PAL-24962-12, ML-PAL-24962-13, ML-PAL-24962-14, ML-PAL-24962-15, ML-PAL-24962-16 (Pl. 1, fig. 8), ML-PAL-24962-17] are identified as belonging to the same species.

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365 Diagnosis

366 Ovate test, flattened, quinqueloculine in early stage, bilocular in appearance due to the chambers arrangement in later stages, cribrate aperture (trematophore) often missing or incomplete in free
368 specimens. Well preserved specimens are smooth. The others appear striated when they show the internal canalicular structure. Chambers have thick walls and are subdivided by subepidermal partitions, forming a series of chambers that are connected by pre- and post- septal passages.

371 Munier-Chalmas and Schlumberger (1883) were the first to describe micro- (B) and megalospheric (A) forms of very distinct sizes. Their analysis of the internal structure and the drawings they have left, are still valid. The recovered Defrance collection contains only microspherical forms which are much larger and more abundant than the megalospheric forms in all the deposits.

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376 Stratigraphy

377 Le Calvez (1970) mentionned it in the Middle Lutetian (zone IVa) and in the Upper Lutetian outside the Paris Basin. It is mentionned in SBZ 13 pro parte in Drobne (2002), currently SBZ 13 and 14 pro parte as in Papazzoni *et al.* (2017, p. 3).

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381 Occurrence

382 Le Calvez (1970) found it in about fifteen deposits in the Paris Basin. It is abundant in Grignon, the type locality of *F. ovata*. It is a very characteristic species of the Paris Basin. Its presence in Belgium and Aquitaine is not demonstrated. It is found in the eastern and western Channel area in the Upper Lutetian (Bignot and Neumann, 1991), demonstrating the extension of an equatorial belt of shallow, warm waters (Bignot and Pomerol, 2010).

387
388 Remarks

389 Another species is worth mentioning; Kaasschieter (1961) described *F. bella* from the sands of Lede in Belgium, which he regarded to be of Late Eocene age. The Lede Formation is now known to be Middle Lutetian (National Commission for Stratigraphy Belgium). Therefore, the affinities between *F. ovata* (= *F. discolites*) and *F. bella*, two coeval species from two adjacent basins, should be studied again.
394

395 **5. Conclusion**

396

397 We herein designate a lectotype for *Alveolina boscii* (Defrance *in* Bronn, 1825), the type species of

398 *Alveolina* d'Orbigny, 1826 and *Fabularia discolites* Defrance *in* Bronn, 1825, the type species of

399 *Fabularia* Defrance, 1820. We herein also designate paralectotypes for those two species. The

400 lectotypes and paralectotypes are housed in the micropalaeontological collections of the *Musée des*

401 *Sciences* in Laval.

402 The discovery, in the collections of this institution, of some samples from the Defrance's collection

403 corroborates the value of inventoring systematically museum and university collections, which has

404 been undertaken for some years now, and should be carried on.

405

406 **Acknowledgments**

407

408 The authors wish to thank Jean-Michel Pacaud, from the National Museum of Natural History of

409 Paris, for his bibliographical research and the material he found in Chaumont-en-Vexin; Didier

410 Néraudeau and Vincent Perrichot from the University of Rennes 1 for the material support

411 concerning the pictures and samples' measurements. They also thank Didier Debailleux, Hervé

412 Dineur and Jean-François Lhomme for the samples from Grignon and the Paris Basin; Karoly

413 Sztrákos and Daniel Vachard for the information and improvement of the manuscript; and Antonino

414 Briguglio for his major contribution in the construction of this article. They finally thank Yves

415 Candela from the National Museums Scotland for his great improvement of the English quality of

416 this manuscript.

417

418

419
420

421 **References**

422

423 Bignot, G., Neumann, M., 1991. Les « grands » foraminifères du Crétacé terminal et du Paléogène du Nord-Ouest européen. Recensement et extensions chronologiques. *Bulletin d'information des géologues du Bassin de Paris* 28 (2), 13-30.

426 Bignot, G., Pomerol, M., 2010. Extension de la ceinture équatoriale d'eaux littorales chaudes au cours de l'Éocène moyen, fondée sur la distribution latitudinale des grands Foraminifères en Europe occidentale et en Afrique australe. *In*: Whittaker, J.H., Hart, M.B. (Eds.), *Micropalaeontology, Sedimentary Environments and Stratigraphy: a Tribute to Dennis Curry* (1912-2001). The Micropalaeontological Society, Special publication, 91-98.

431 Bigot, M.A., 1905. Catalogue critique de la Collection Defrance, conservée au Musée d'histoire naturelle de Caen - Deuxième partie : Céphalopodes, Arthropodes et Vers. *Bulletin de la Société Linnéenne de Normandie*, ser. 5, 8, 251-273.

434 Bigot, M.A., 1906. Catalogue critique de la Collection Defrance, conservée au Musée d'histoire naturelle de Caen - Troisième partie : Pélécypodes décrits et figurés par Lamarck d'après des échantillons de cette collection. *Bulletin de la Société Linnéenne de Normandie*, ser. 5, 9, 3-40.

437 Bigot, M.A., 1907. Catalogue critique de la Collection Defrance, conservée au Musée d'histoire naturelle de Caen - Quatrième partie : gastéropodes décrits et figurés par Lamarck d'après des échantillons de cette collection. *Bulletin de la Société Linnéenne de Normandie*, ser. 5, 10, 81135.

441 Bigot, M.A., Matte, H., 1903. Catalogue critique de la Collection Defrance, conservée au Musée d'histoire naturelle de Caen. *Bulletin de la Société Linnéenne de Normandie*, ser. 5, 6, 152-185.

443 Bigot, M.A., Matte, H., 1904. Catalogue critique de la Collection Defrance, conservée au Musée d'histoire naturelle de Caen - Première partie : Pélécypodes (suite). *Bulletin de la Société Linnéenne de Normandie*, ser. 5, 7, 243-268.
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497 l'arrangement établi dans les galeries du Muséum d'Histoire Naturelle, parmi leurs dépouilles
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502 dans la collection des vélins du Muséum. *Annales du Muséum national d'Histoire naturelle par
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508 Loeblich, A.R., Tappan, H., 1987. Case 2356: *Alveolina* d'Orbigny, 1826 (Foraminiferida):
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510

511 Montfort, P.D.de, 1808. *Conchyliologie systématique et classification méthodique des coquilles*.
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541
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546 Sztrákos, K., Blondeau, A., Hottinger, L., 2010. Lithostratigraphie et biostratigraphie des formations marines paléocènes et éocènes nord-aquitaines (bassins de Contis et Parentis, seuil et plate-forme nord-aquitains). *Géologie de la France* 2, 3-52.

549 Tréguier, J., Pacaud, J.-M., Prugneaux, V., 2017. Des types porte-noms de la collection Defrance conservés au Musée des Sciences de Laval. *Fossiles* 32, 21-34.

551 Vecchio, E., Barattolo, F., Hottinger, L., 2007. Alveolina horizons in the Trentinara Formation (Southern Apennines, Italy): Stratigraphic and paleogeographic implications. *Rivista Italiana di Paleontologia e Stratigrafia* 113 (1), 21-42.

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577

Captions for figures in text

578

579

Figure 1. Complete specimen of *Alveolina subpyrenaica* Leymerie, 1846 ?, lot ML-PAL-22330, Vendémies (Aude, France), scale bar: 1 mm.

580

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582

Figure 2. Axial section of *Alveolina oblonga* d'Orbigny, 1826 ? from lot ML-PAL-22350-7, Chaumont-en-Vexin (Oise, France), scale bar: 1 mm.

583

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Figure 3. Complete specimen of *Fabularia compressa* d'Orbigny, 1850 ?, lot ML-PAL-29059-3, Hautteville-Bocage (Manche, France), scale bar: 1 mm.

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588

Figure 4. Reproduction of p. 60 of Palmer (1977) which illustrates, among others, Lamarck's vellum, stored in the *Musée Naitonal d'Histoire Naturelle* in Paris, n°48, fig. 16 a-b (*Fabularia discolites*) and fig. 17 a-b (*Alveolina boscii*).

589

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592

Plate 1

593

(Scale bar: 1 mm)

594

595

1. Card with thirteen specimens of *Alveolina boscii* (Defrance in Bronn, 1825), lot ML-PAL22432, Thiverval-Grignon (Yveline, France), lectotype (n°4) and paralectotype (n°7).

596
597 2. Axial section of *Alveolina boscii* (Defrance in Bronn, 1825), ML-PAL-22432-4, Thiverval-Grignon (Yvelines, France), lectotype.

599 3. Axial section of *Alveolina boscii* (Defrance in Bronn, 1825), ML-PAL-22432-7, Thiverval-Grignon (Yvelines, France), paralectotype.

601 4. Complete specimen of *Alveolina boscii* (Defrance in Bronn, 1825), ML-PAL-22432-3, Thiverval-Grignon (Yvelines, France).

603 5. Card with seven out of seventeen specimens of *Fabularia discolites* Defrance in Bronn, 1825 from lot ML-PAL-24962, Thiverval-Grignon (Yveline, France), lectotype (n°1) and paralectotype (n°4).

606 6. Axial section of *Fabularia discolites* Defrance in Bronn, 1825, ML-PAL-24962-1, Thiverval-Grignon (Yveline, France), lectotype.
569 Figure 7. Axial section of *Fabularia discolites* Defrance *in* Bronn, 1825, ML-PAL-24962-4, 570
Thiverval-Grignon (Yveline, France), paralectotype.

571

572 Figure 8. Complete specimen of *Fabularia discolites* Defrance *in* Bronn, 1825, A. lateral view, B.

573 aperture view, C. lateral view, ML-PAL-24962-16, Thiverval-Grignon (Yveline, France).
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