Content

[Introduction] [About the name Lingula] [About the name anatina]
[About the usage of the term linguliform ...] [Bibliographic references] [Figures] and ... [Tables]

On the history of the names Lingula, anatina, and
on the confusion of the forms assigned them among the Brachiopoda

Christian C. Emig

BrachNet, 20, Rue Chaix, F-13007 Marseille (France)

Manuscript online since December 10, 2008

Abstract

The first descriptions of Lingula were made from then extant specimens by three famous French scientists: Bruguière, Cuvier, and Lamarck. The genus Lingula was created in 1791 (not 1797) by Bruguière and in 1801 Lamarck named the first species L. anatina, which was then studied by Cuvier (1802). In 1812 the first fossil lingulids were discovered in the Mesozoic and Palaeozoic strata of the U.K. and were referred to Lingula on the basis of similarity in the form of the shell. In the 1840's other linguliform brachiopods from the Palaeozoic were described. The similarity of the shell form of the extant Lingula and these fossils led Darwin in 1859 to create the description "living fossil" in his book "On the Origin of Species". Thereafter, this Darwinian concept became traditional in that Lingula was considered to lack morphological evolutionary changes. Although denounced as scientifically incorrect for more than two decades, the concept still remains in many books, publications and Web sites, perhaps a witness to palaeontological conservatism.

Key Words

Lingula; brachiopod; anatina; Darwin; living fossil.

Citation

Emig C.C. (2008).- On the history of the names Lingula, anatina, and on the confusion of the forms assigned them among the Brachiopoda.- Carnets de Géologie / Notebooks on Geology, Brest, Article 2008/08 (CG2008_A08)

Résumé

De l'origine historique des noms lingule, Lingula, anatina, et de la confusion des formes chez les Brachiopodes.- Les premiers auteurs à publier sur les lingules – Lingula - furent français : Bruguière créa le genre Lingula en 1791 [et non en 1797], cette date acceptée par tous les auteurs jusqu'à la fin du XIX^ème siècle et confirmée dans ce travail. Elle fut remise en cause à partir du début du XX^ème siècle et remplacée officiellement par 1797 par la Commission Internationale de Nomenclature zoologique en 1982, puis confirmée en 1985. La signification de lingule ou lingula (ou ligula) est communément considérée comme un diminutif (signifiant languette ou élément en forme de langue) du latin lingua "langue". D'autres possibilités peuvent être suggérées, comme un diminutif savant du latin lingua + le suffixe –ula [en français –ule] ou encore une autre traduction du latin ligula ou lingula à savoir cuillère.
Lamarck (1801) puis Cuvier (1802) décrivirent la première espèce du genre sous Lingula anatina. L'origine du nom d'espèce est inconnue, mais, en latin, ce nom évoque une ressemblance avec le "bec de cane" cité par Cuvier (1798). Auparavant Seba (1758) indiquait qu'il s'agit "d'une espèce particulière de conque anatifère", anatina signifiant en latin du canard ou appartenant au canard.
J. Sowerby (1812) se basa sur la ressemblance de forme de la coquille pour décrire les premières lingules fossiles dans le Jurassique de Grande-Bretagne. À partir des années 1840, notamment avec la description des "Lingula-flags" dans le Paléozoïque inférieur du Pays de Galles, il y a amalgame, devenu classique, de la forme – linguliforme – entre les espèces actuelles de Lingula et les lingulides fossiles paléozoïques et mésozoïques.
L'extension géologique des Lingula à travers tout le Phanérozoïque a influencé Darwin (1859) lors de la création du terme "fossile vivant". Dans son livre "Sur l'origine des espèces", il fait plusieurs fois référence aux lingules.
Depuis cette époque, les Lingula ont été décrites, dans la plupart des traités de paléontologie, "avec une forme pratiquement inchangée depuis leur origine au Cambrien, il y a environ 550 Millions d'années". La conséquence en est que tout brachiopode fossile linguliforme a été décrit, et l'est parfois encore de nos jours, comme appartenant au genre Lingula sur ce seul critère.
Pourtant, le traditionnel concept introduit par Darwin que les Lingula sont des "fossiles vivants" a été rejeté depuis plus de 20 ans, et auparavant par Cuvier (1798). La persistance de cette hérésie scientifique ne peut s'expliquer que par le conservatisme de la communauté internationale des paléontologues avec des modèles difficilement remis en question.

Mots-Clefs

Lingula ; brachiopode ; anatina ; Darwin ; fossile-vivant.

Introduction

Offered by the Web (Internet) the facility to examine old scientific work in facsimile makes it possible to review and to clarify the origin of the name Lingula and that of its type species L. anatina. The Web also makes it possible to show that (Darwin, 1859) who originated the concept of living fossil for Lingula, was wrong because his conclusion was not based on systematics, and has been denounced for more than 20 years. Nevertheless, it continues to be postulated by some palaeontologists in their publications, books and Web sites.

Until the end of the XIXth century the brachiopods were classified as molluscs. Duméril (1806) proposed the creation of a 5th order of molluscs under the name Brachiopoda. However, he noted (1806, p. 154) that nine years earlier Cuvier had already suggested this subdivision. Indeed, Cuvier (1798), then Lamarck (1801), subdivided Molluscs, inserting in the Acephal Molluscs the four brachiopod genera known at that time: Lingula, Orbicula, Crania and Terebratula. Bosc (1802) included these groups in the "Coquilles inaequivalves" and all the lamellibranchs in the "équivalves". This classification has been largely overlooked as Muir-Wood (1955) has pointed out. Menke (1828) divided the class Brachiopoda into three families, one being the family Lingulaceae (now the superfamily Linguloidea Menke, 1828 and the family Lingulidae Menke, 1828) with Lingula.

At the end of the XIXth century, Hatschek (1888, p. 40) separated the Brachiopods from the Molluscs. He included Brachiopoda, Bryozoa (Ectoprocta) and Phoronida in his Tentaculata (= Molluscoidea) (see also Muir-Wood, 1955).

About the name Lingula

The first authors who published on Lingula were French, respectively Bruguière, Lamarck¹ and Cuvier¹ (Table 1 ). Before the creation of the genus Lingula by Bruguière (1791), Rumphius (1705) called it a gastropod which was later named Scrutus Montfort, 1810. Linnaeus (1758, 1767) placed these animals as gastropod molluscs under the name Patella unguis (specimen from Ambon island in the Moluccas, Indonesia), but as stated by Cuvier (1798, 1802) (Figs. 1 - 2 - 3 ) he described only one detached valve without a pedicle. Along the coast of this island three species of Lingula have been reported: L. anatina by various authors of the XVIIIth and XIXth centuries, and recently L. reevei and L. rostrum by Emig and Cals (1979). No specimen of Lingula has been found in Linnaeus's collections. Some authors like Gmelin (1789) confirmed the name Patella unguis. According to Cuvier and Latreille (1817) "(...) Solander [1786] et Chemnitz [1786] qui surent qu'elle avait deux valves, lui donnèrent l'un le nom de mytilus lingua, l'autre celui de pinna unguis" (see also Davidson, 1888; Fig. 4 ).

Mytilus lingua is catalogued by Lightfoot (1786) on the annotated list (which is referred to him) for the auction of the collection of Lady Margaret Cavendish Bentinck, duchess of Portland, after her death in 1785. For some taxa he used the names given them by Daniel Carl Solander (1733-1782), a pupil of Karl von Linné and curator of the collection of the duchess. He indicated these taxa by an "S" following the name. Solander's manuscripts remain unpublished and are preserved at the Natural History Museum of London. Until 1965, most of the quotations of the "Portland Catalogue" were attributed to Solander (see Davidson, 1888; Fig. 4 ). Then, Kay (1965) assigned all the names in the catalogue to Lightfoot (1786), and this revision was adopted by many malacologists. Several authors, such as Cuvier & Latreille (1817) and Davidson (1888), assigned Mytilus lingua to Solander (Fig. 4 ). I was not able to consult Solander's manuscripts Lightfoot's list (1786), or the work of Kay (1965).

Moreover oddly enough Mytilus lingua was considered (see Rowell, 1964; I.C.Z.N., 1982) as a possibility for the type species of the genus Lingula. But in the end L. anatina Lamarck, 1801 was accepted and Mytilus lingua removed (I.C.Z.N., 1985), owing to a doubtful identification and the absence of acceptance (see below).

Bruguière (1791 [1789]) created the genus Lingula (Fig. 5 ) in Volume 1 of the Tableau encyclopédique et méthodique des trois règnes de la nature : vers, coquilles, mollusques et polypes divers, written by Bruguière as the sole author. Three volumes were published by Charles Joseph Panckoucke (Paris and Liège) and in several editions.

Volume 1 in which Lingula is figured is dated 1789, but was published in 1791 (Fig. 5 ), as confirmed in the "Trésor de la Langue Française informatisé" (A.T.I.L.F., 2007): "1791 lingule (JG Bruguière, Tableau encyclopédique et méthodique des trois règnes de la nature, I, 151a, Pl. 250 dans Compte rendu de R. Arveiller sur les datations de Quem. DDL t. 20, à paraître dans R. Ling. rom. t. 47)" (see Arveiller 1982; Quemada 1982). The etymology of lingule (French), and lingula or ligula (Latin) is probably a diminutive meaning tongue or tongue-shaped, from the Latin lingua "tongue". Another possibility proposed in the "Trésor de la Langue Française informatisé" (A.T.I.L.F., 2007) is that it is a scholarly diminutive of the Latin lingua+suffix-ule (= little, small): this suffix is associated with a name from the Latin: -ulus, -ula, -ulum.This usage is found mainly in scientific works, particularly those of the life sciences. Another etymology: lingula may be derived from the Latin ligula, sometimes lingula, "spoon", with reference to the spoon-shape of the animal formed by the shell and the pedicle (Pers. Comm. R. Gourvennec).

Some vernacular names for Lingula: "moule-à-queue" in New Caledonia; "bec de cane" along some coasts in the Indian Ocean (Cuvier, 1798); "shamisen-gai" in Japan. The name is related to its similarity to the shamisen, a Japanese lute of Chinese origin.

Cuvier (1798) (Fig. 1 ) assigned the genus Lingula to Bruguière (1791) (Fig. 3 ). Throughout the XIXth century Davidson (1880) and several other authors indicated the year 1789 (Fig. 4 ), as follows: Lingula anatina, Bruguière, Hist. des vers. Encycl. Méth. Pl. CCl, fig. 1 a, b, c, 1789. Others like d'Orbigny (1852) list 1791. I could not find this reference 1789, but found two under Bruguière (1791) with plates 172-314 and Bruguière (ca. 1793) with plates 199-488.

Starting in the XXth century, various authors indicate Bruguière, 1797 as the date of the creation of the genus Lingula. Rowell (1964) in accordance with the publication dates of the Encyclopédie Méthodique as printed by Sherborn and Woodward (1906, p. 581), obtained in 1966 from the International Commission on Zoological Nomenclature (I.C.Z.N., 1982, 1985) a change of year for Lingula from 1791 to 1797. Recently, the data published by Evenhuis (2003) and Evenhuis and Petit (2003) confirm Rowell's proposal. The volumes of plates for the Histoire Naturelle des Vers volumes were published in the form of a Tableau encyclopédique et méthodique. However, their review of the publication of the Encyclopédie méthodique and of the Tableau, has several gaps in their compilation of the editions that appeared before 1797, as well as some incomplete references. For example Bruguière's Tome 1, edited in 1792 should be quoted as: Encyclopédie Méthodique, ou par ordre de matières: par une société de gens de lettres, de savants et d'artistes : précédée d'un vocabulaire universel, servant de table pour tout l'ouvrage, ornée des portraits de MM. Diderot et d'Alembert, premiers éditeurs de l'Encyclopédie [26, a], 1 Histoire Naturelle des Vers, tome 1, p. 1-757. Panckoucke, Paris.

So the date 1797 for the first description of Lingula is wrong and we must return to the original date of 1791, which is the correct one. Many successive editions under various titles, as well as gaps in the bibliographic searches may explain this error. After the death of Charles-Joseph Panckoucke in 1798, his daughter Thérèse-Charlotte Agasse, wife of Henri Agasse, the associate of Panckoucke, assured the publication. Such errors in the dates of the first publication of old books are common. Another example concerns the first editions of Seba's volume 3: its year and pagination range between 1758, the correct one, 1759 (p. [1-24], 1-108, Pl. 1-116; or 212 p., 116 plates; or 511 p., 116 Pls.) and 1761 (511 p., 116 Pls.). Several other editions have been published by scientific editors, e.g., Cuvier, Saint-Hilaire and Lesson under Seba et alii (1828), and recently Müsch et alii (2005).

About the name anatina

Lingula Bruguière, 1791 as a genus was named 10 years before the description of the first species L. anatina Lamarck, 1801. In Plate 16, fig. 4, Seba (1758) figured for the first time two lingulide² specimens stored in his own collection (Fig. 6 ). After Albert Seba's death in 1736, many of the specimens in his collection were auctioned off in 1752 and dispersed all over Europe. Among them the lingulids² and other specimens figured in the plates were acquired by the Muséum National d'Histoire Naturelle de Paris. These lingulide specimens were used by Lamarck (1801) to describe the new species Lingula anatina (Fig. 7 ). Appointed in 1793 at the Muséum de Paris as a professor on insects and worms, Lamarck was not as skilled as Cuvier in making detailed analyses, so he gave Cuvier easy access to his specimens, in particular those from the Seba collection. They were described again by Cuvier (1802) in the Mémoire sur l'animal de la Lingule. This memoir was reissued in 1817, the date under which this memoir is usually quoted. In 1802 (republished in 1824 and 1836), Bosc too described Lingula anatina in his "Histoire naturelle des coquilles" (Fig. 8 ).

The origin of the species name is unknown, but in Latin the adjective anatinus, a, um, means "duck or belonging to the duck" perhaps because a resemblance to the "bec de cane" (female duck bill) as indicated by Cuvier (1798; Fig. 1 ) for the Lingula shell? Seba (1758) had defined it in a few words "d'une espèce particulière de conque anatifère" (quoted by Cuvier, 1802, p. 2; Fig. 2 ). The French word anatifère is composed of the Latin radical anas, anatis (meaning "duck") and the suffix -fère (from the Latin -fer "which carries") - see "Trésor de la Langue Française informatisé" (A.T.I.L.F., 2007). Note that Goose barnacle is Anatife in French (see Lamarck, 1801; and Fig. 7 ).

Lingula anatina was recognized as the type species of the genus Lingula only late in 1892 (see Rowell, 1964; I.C.Z.N., 1982, 1985). Yet G.B. Sowerby (1847) had already attributed the genus Lingula to Bruguière and the type species L. anatina to Lamarck. In addition, he listed the current extant species: Lingula anatina Lamarck, 1801; L. hians Swains, 1823, [now L. rostrum (Shaw, 1798)]; L. audebarti Broderip, 1835 [now Glottidia audebarti]; L. semen Broderip, 1835 [now Glottidia semen (Broderip, 1835) ]; L. tumidula Reeve, 1841; L. ovalis Reeve, 1841 [now L. reevii Davidson, 1880]; L. albida Hinds, 1841 [now Glottidia albida].

Davidson (1888) introducing the family Lingulidae wrote: "The recent species belonging to this family are representatives of the genus Lingula, Bruguière (1789), and to the genus or subgenus Glottidia, Dall (1870)". And he attributed Lingula anatina to Lamarck, 1819 on the list of Lingula species, but to Bruguière in the species description (Fig. 4 ).

Goulven (2000) in an analysis of Lamarck's text (1819) wrote: "En ce qui concerne les Lingules, Lamarck ne retient que la lingule actuelle, Lingula anatina, qui habite l'Océan des Moluques" (p. 258). But he added: "l'animal de la térébratule est fort rapproché de celui de la lingule par ses rapports" (p. 244). Cuvier (1802) also compared Lingula with Terebratula and on p. 9 in discussing the craniid now named Novocrania anomala (Müller, 1776) he wrote: "Il suffit de jeter les yeux sur la figure que Müller a donnée de l'animal de son Patella anomala, pour voir qu'il ressemble à la lingule par des bras ciliés et en spirale".

About the usage of the term linguliform and its consequences

I could not find the origin of the adjective linguliform. However, the earliest definition is in King (1859, p. 260): linguliform, tongue shaped. This descriptive term is employed in a specific scientific sense in several groups of invertebrates (crustaceans, brachiopods), and in botany and medicine. As concerns brachiopods it appeared in the 1880's.

Linguliform means having the form of a tongue, tongue-shaped. It is derived from the Latin linguliformis. Cuvier et alii (1834) used it for Mytilus describing "the linguliform of the appendage of the foot". More interesting is that this adjective was not used in the description of Lingula in the same book (p. 131) in which Lingula anatina is attributed to Cuvier.

At the beginning of his memoir of 1802 Cuvier was the first to discuss the shape of the shell but these sentences have not been heeded:
"Il n'est pas de genres de testacés qui prouve mieux que ne fait celui des Lingules, la nécessité de connoître [= connaître] l'animal, et ne pas se borner à la coquille, pour ranger convenablement ces mollusques dans une méthode naturelle.
En effet les coquilles des Lingules, quoique de forme assez particulière, ne pouvoient [= pouvaient] faire soupçonner les grandes différences qui séparent leur animal des autres genres de sa classe ; et tant qu'on n'a connu qu'elles, on les a ballottées arbitrairement de genre en genre."

From the Jurassic of Great-Britain came the first descriptions of fossil lingulides, i.e., Lingula mytilloides, L. ovalis, L. tenuis. They were made by J. Sowerby (1812; Fig. 9 ) based on Linnaeus's (1758) and Cuvier's (1802) studies and on similarity in the shape of the shell with those of the living species of Lingula. Lingula mytilloides at least is assignable to Lingularia Biernat & Emig, 1993. Broderip (1835) was probably the first who affirmed the similarity between fossil and recent forms (Fig. 10 ). However the two Lingula species he described were later referred to Glottidia.

In the 1840's the similarity of the linguliform shell in extant and fossils forms was emphasized during the studies of the "Lingula flags" in the lower Palaeozoic of North Wales, particularly of those from the Cambrian (see Murchison, 1847). In 1845, Lyell wrote about the occurrence of Lingula in the "Potsdam sandstone" (New York): "(...) it is highly interesting that one of its commonest organic remains should belong to a living genus (Lingula), and that its form should come very near to species now existing." (p. 132).

Many works and books published between 1845 and 1865 discuss or describe this genus. Consequently the number of species of Palaeozoic Lingula grew rapidly. In various French and English publications during the second half of the XIXth century the geological range of Lingula: "Palaeozoic to Present" is found. (Boule, 1910, p. 29): "Dans les couches les plus inférieure des terrains primaires, on trouve des lingules, tout à fait semblables aux lingules des mers actuelles (fig. 27 et 28); c'est là un premier exemple de longévité extraordinaire de certains types d'animaux". Consequently, based solely on the shape of the shell any fossil linguliform brachiopod was assigned this genus! However, Emig (1982, 2002), Biernat and Emig (1993) have demonstrated that this shape has no taxonomic value. The linguliform shape also occurs in several other inarticulated brachiopod families, i.e., the Pseudolingulidae, Obolidae, and Eoobolidae, of which many species were originally referred to the genus Lingula.

The broad geological range and the similarity in the shape of the lingulide shell throughout the Phanerozoic led Darwin (1859) to create the term living fossil for his book "On the Origin of Species" (Table 2 ), in which several sentences refer to Lingula.

Since Darwin's times Lingula is considered as in an almost unchanged form since first appearing in the Cambrian period around 550 MA ago. Once found in more widespread environments, today's Lingula are confined to brackish intertidal habitats where they live in burrows. Such a statement concerning persistence is known to be wrong (see Emig, 1997). The taxa of the superfamily Linguloidea show morphological evolutionary changes despite the panchronic characteristics of this group among the Recent Brachiopods. Consequently, Darwin's statement that Lingula is a "living fossil" must be rejected (see Emig, 2003).

Today this anachronism, condemned for more than two decades and blatantly erroneous for those who work on evolution, can still be read on Web sites and in publications. Some recent examples among others:

Williams et alii (1996) created a new subphylum of Brachiopoda named "Linguliformea" which includes all the former Inarticulata Brachiopoda, except the former Craniida which become the subphylum Craniiformea. Linguliformea appears to be inappropriate as the name for a group of which the majority of taxa do not have a linguliform shell.

Milsom and Rigby (2003) write: "Lingula. Linguliform brachiopod. Ordovician-Recent. A small (about 2 cm from the beak to the anterior edge), smooth, phosphatic brachiopod known as a "living fossil" as its morphology has not changed significantly since the Ordovician. Fully infaunal, it lives in burrows with its anterior edge close to the sediment-water interface. The pedicle anchors the brachiopod to the mud whilst the valves rotate and grind through the sediment. Modern Lingula mainly exploit marginal habitats, but fossil Lingula are known from shelf and basin environments." NB: Lingulide species may reach 7-8 cm, are unable to live in a muddy substrate, but live in true marine conditions and habitats (Emig, 1986, 1997)

J. Moore (2006): "Living fossils: this is a potentially confusing description of animals that have changed remarkably little over long periods of time. Examples include brachiopod Lingula, found in Cambrian fossils and persisting today." So the confusion is maintained.

As a result of this blind consensus even today fossil linguliform brachiopods are assigned the genus Lingula based only on their linguliform appearance. Nevertheless, this scientific anachronism has been impugned for over two decades by Emig (1982, 1997, 2002), Biernat and Emig (1993). These authors have demonstrated as pointed out two centuries ago by Cuvier (1802) that the shape of the lingulide shell is not a taxonomically valid character. The concept introduced by Darwin that Lingula is a "living fossil" must be rejected (see Emig, 2003).

The persistence of this scientific heresy may be related to the conservatism of the palaeontological community. A partial explanation is contained in the answer of an American specialist in Mesozoic brachiopods, who in 2003 wrote me in an email: "(...) people who study faunal lists and databases instead of anatomy and taxonomy, and thus perpetuate older nomenclature. As the ecology and distribution of lingulides does not seem to have changed dramatically since their origin, the name "Lingula" has a tremendous amount of inertia."

An anonymous referee, an Anglophone palaeontologist (according to the editor of the Journal of Experimental Marine Biology and Ecology) in 1982 wrote this short sentence as comment on my submittal to that journal: "All what is written in this manuscript is opposite of what can be read on Lingula in any treatise of palaeontology. To be rejected". My work was published in Marine Biology (Emig, 1983).

Evidence of an approach to the acceptance of the possibility that linguliform brachiopods are not all assignable to a single unique genus is found in the successive editions of the Treatise of Invertebrate Paleontology. In the first (R.C. Moore, 1965) the stratigraphic distribution of Lingula was: "? Ord., Sil.-- Rec., Cosmopolitan." The 2nd edition (Holmer and Popov, 2000) states: "?Cretaceous, Tertiary-Holocene;? Cosmopolitan (exact stratigraphic and geographic distribution of fossil forms is very uncertain)". The study of these distributions has advanced considerably (Schmid et alii, 2001; Emig, 2003; Emig and Bitner, 2005; Bitner and Dulai, 2008) but no update regarding them was made in the revision of Holmer and Popov's contribution in the Treatise on Invertebrate Paleontology: Volume 6 (2007) part H, Brachiopoda.

Finally, in the future may authors stop using the so-easy copy and paste, and instead read recent papers on the matter.

Acknowledgments

Many of the works quoted here are available on the Web, e.g., http://darwin-online.org.uk/ - http://www.lamarck.cnrs.fr/ - http://www.buffon.cnrs.fr/ - http://www.literature.org/ - http://books.google.com/ - Universal Bibliophiles Association: http://abu.cnam.fr/ (more at http://paleopolis.rediris.es/BrachNet/ANNONCES/JOURNAL/e-library.html). I would like thank Martin Liebetruth of Niedersächsische Staats- und Universitätsbibliothek Göttingen (http://www.sub.uni-goettingen.de/ - http://www.animalbase.uni-goettingen.de/zooweb/servlet/AnimalBase/search) and of the Zentrum Göttinger Digitalisierung (GDZ: http://gdz.sub.uni-goettingen.de/) for permission to reproduce some of facsimiles made available on these sites. I am most grateful to Remy Gourvennec and to two anonymous referees for comments on the earlier version, and to Nestor Sander for improvements in the English of this version.

Footnotes:

¹ Lamarck and Cuvier are the founders of the science of palaeontology, both vertebrate and invertebrate (see Goulven, 2000). Lamarck also created the word "biology" for the science of living beings.

² In conventional terminology "lingulid" designates taxa or specimens of the Order Lingulida; preferably this word should now be restricted to indicate members of the superfamily Linguloidea. And the term, "lingulide" is used to indicate taxa and specimens of the family Lingulida.

Bibliographic references

A.T.I.L.F. - Analyse et Traitement Informatique de la Langue Française (2007).- Trésor de la Langue Française informatisé (Le).- C.N.R.S., http://atilf.atilf.fr/

Arveiller R. (1982).- Compte rendu sur les datations de Bernard Quemada, Datations et documents lexicographiques.- Revue de Linguistique romane, Paris, vol. 46, p. 453-461. [Lingula p. 460]

Biernat G. & Emig C.C. (1993).- Anatomical distinctions of the Mesozoic lingulide brachiopods.- Acta Palaeontologica Polonica, Warszawa, vol. 38, n° 1, p. 1-20.

Bitner M.A. & Dulai A. (2008).- Eocene micromorphic brachiopods from north-western Hungary.- Geologica Carpathica, Bratislava, vol. 59, n° 1, p. 31-43.

Bosc L.A.G. (1802 = an X).- Histoire naturelle des coquilles, contenant leur description, les mœurs des animaux qui les habitent et leurs usages.- Déterville, Paris, vol. 2. 330 p.

Bosc L.A.G. (1836). Histoire naturelle des coquilles contenant leur description, les mœurs des animaux qui les habitent et leurs usages [Lingula: p. 169-172 ; Pls. 45.].- Librairie encyclopédique de Roret, Paris, 3e éd., vol. 1, 539 p.

Boule M. (1910).- Conférences de géologie.- Masson, Paris, 160 p. [p. 29]

Broderip W.J. (1835).- Descriptions of some new species of Cuvier's family of Brachiopoda.- Transactions of the zoological Society of London, vol. 1, p. 141-144.

Bruguière J.G. (1791 [1789]).- Tableau Encyclopédique et Méthodique des trois Règnes de la Nature : vers, coquilles, mollusques et polypes divers.- Panckoucke, Paris, T. 1, Pls. 172-344. [Lingula: Pl. 250, fig. 1a-c]

Bruguière J.G. (ca 1793).- Histoire naturelle des vers.- Panckoucke, Paris, Tableau [2.] Planches 199/488.

Bruguière J.G. (1797).- Tableau Encyclopédique et Méthodique des trois Règnes de la Nature : vers, coquilles, mollusques et polypes divers.- Panckoucke, Paris. T. 2, Pls. 190-286. [Lingula: Pl. 250, fig. 1a-c]

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