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Les Planches inédites de Foraminifères d’Alcide d'Orbigny

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Marie-Thérèse Vénec-Peyré

Introduction

Texte intégral

1The Planches inédites were drawn by d’Orbigny to illustrate the Tableau méthodique de la classe des Céphalopodes, more usually called the Tableau méthodique. In this work of 1826, he raised to ordinal rank a group of microscopic animals which, according to him, differed from the other cephalopods, as understood at this time, in its lack of siphon and the presence of one or several apertures (foramens) allowing communication between chambers. He presented the first classification of this order that he called “Foraminifères”, including in this nomenclatural framework the names of 544 species, 335 of which were new to science, but no description was given. Unfortunately, a lack of time — since the authorities of the Muséum national d’Histoire naturelle (France), impressed by his first work, named him as the naturalist explorer for an expedition to South America (1826-1834) — and perhaps also financial difficulties, prevented him from publishing the 73 plates illustrating most of the species listed in the Tableau méthodique. Only 31 species were figured on 8 plates which accompanied the text of 1826 and were different from the original plates that remain unpublished.

2On his return from South America, d’Orbigny illustrated in his further works on foraminifera some of the species’ names enumerated in 1826. However, it was not until after his death that a large number of the other names became available when other authors published d’Orbigny’s own drawings or illustration of the “Models” of foraminifera that he had sculpted and to which he referred in his publication of 1826.

3One hundred and two of the species described by d’Orbigny between 1826 and 1852 are designated as type-species of genera (cf. Loeblich & Tappan 1987). Among them, twenty-two are figured in the Planches inédites reproduced in the present volume. The purpose of this present work is to pay homage to this first comprehensive work of Alcide d’Orbigny, to retrace the nomenclatural history of certain of the species figured in these plates, to clarify a number of nomenclatural points and explain certain other unclear aspects, to give information on the type material preserved in the collections, and finally to place the species in the current scientific context.

The Planches inédites: their content

4The Planches inédites form a set of 57 booklets, each one representing a genus. The name of the family and the heading “Céphalopodes, Foraminifères” appear on a brown-paper cover. Each booklet contains plates of various types corresponding to different stages of completion:

  • 1) Finished plates (Plates 1-45, reproduced here), grouping the different species belonging to a single genus inside a frame. The generic name is handwritten at the top of the plates, and the captions at the bottom, outside the frame. Most of the figures are drawn in pencil and wash, delicately tinted. The remainder are in ink. The names of the species, written at the bottom of the plates, are surrounded by two numbers. The preceding one corresponds to the figure number on the plate, the one following indicates the serial number of the species in the Tableau méthodique, but several errors in this second numbering can be observed. Each species is illustrated by one or more enlargements and, in most cases, also a full-sized figure. In view of the size of the specimens, this may be represented by no more than a point or a small circle, or, for the largest species, by a miniature of the enlarged figure.

  • 2) Unfinished plates with a frame. In some of them the drawings are finished but they do not fill the full plate, in others the drawings are only sketched.

  • 3) Plates with a frame but in a different style: the outlines of species and ornamentations are drawn or inked over (Indian ink or sepia drawing); these drawings, numbered but without reference to a caption, are rarely coloured. No species’ name is given, but the origin is indicated beneath the figures, inside the frame. The headings “free shells” or “fixed shells” are written outside the frame.

  • 4) Plates without a frame, representing the first stages of completion of the drawings (Illustration 1). The majority are sketches made in pencil and afterwards inked over, on small, odd scraps of paper. For each genus, they are grouped together, without particular care, on a same plate.

  • 5) Plates containing figures cut from works published by d’Orbigny after 1826.

5This set contains approximately 300 plates illustrating more than 500 species. Beyond their scientific interest, it is touching to be able to witness the successive stages of completion of the drawings which led the young author, and perhaps the child, from the first sketches to the definitive plates. We can have no doubt that the drawings are his own. A letter addressed by Charles-Marie d’Orbigny to Louis Benjamin Fleuriau de Bellevue (Heron-Allen 1917: 7) explains that as his eyesight weakened with age, it was the young Alcide who observed the small organisms under the microscope and drew them.

6Of a high quality, the Planches inédites testify not only to their author’s keen sense of observation, but also to his artistic talent. Alcide d’Orbigny’s drawings are all the more admirable considering the optical material he used, which was obviously less efficient than modern equipment.

7As stated by Lys (1948a: 11), “no date was given for the Planches inédites […] If we want to be exact, their creation extended over a period running from his youth to his death, but the greater part was complete in 1825, for the presentation at the Academy. [...] We can therefore give as an approximate date the Tableau méthodique, that is, 1826. I consider as a decisive argument in favour of this the heading ‘ Céphalopodes Foraminifères’, which was a term he abandoned on his return from America after Dujardin’s discoveries in 1835”.

8In this set of plates, only 45 are finished; they correspond to the type (1) described previously and represent 198 species. I have chosen to present only these plates in this book, since, having been finished by d’Orbigny, they can be published without a risk of false interpretation.

The foraminiferal collections of Alcide d’Orbigny: an indivisible set, an inestimable patrimimony

9The Planches inédites, the Tableau méthodique, the Models and the collections of foraminifera of Alcide d’Orbigny constitute an indivisible set and an inestimable patrimony which is stored in the Museum national d’Histoire naturelle (MNHN) in Paris (France).

10The Museum purchased the Planches inédites when the family auctioned the library of Alcide d’Orbigny on January 6, 1858 (Lys & Sigal 1947). His huge collections were registered in June 1858 in the catalogue of the Paleontology Laboratory of this Institution (Lauriat-Rage 2002: 620, fig. 1). Olry Terquem was the first to realize the importance of the Planches inédites and of d’Orbigny’s collections. We are indebted to him for having understood that these Plates “referred to the Tableau méthodique and reproduced exactly the order and the numbers” (Terquem 1878: 2). It is also to Terquem that we owe the bringing to light of the collections, “a second stroke of chance, equally delightful, brought the discovery of a cardboard series containing shells numbered and determined according to figures. It was then easy to see that the Tableau, the plates and the shells constituted a set which could be used in the study and the determination of shells, despite the absence of text” (Terquem 1878: 2).

11The Planches inédites were certainly intended to illustrate the Tableau méthodique published in 1826 in Les Annales de Sciences Naturelles, but d’Orbigny had also conceived another method of illustration, as he explains in this important work (1826: 248), “With a view to giving greater publicity to the work I had undertaken, and with the object of making it available to everybody, without entailing the necessity of observing the numerous genera of foraminifera under the microscope, I have had the idea of sculpting a shell of each genus and each sub-genus of this order about 1 1/2 inches in size; from these I have made matrices by means of which we can have many specimens of each shell”. A set of 100 three-dimensional Models, the diameters of which have been enlarged 40 to 200 times, are sculptured in a very fine limestone (Illustration 2). Like the drawings of the Planches inédites, they testify to the artistic talent of their author and, once again, merit our admiration. In the Tableau méthodique, d’Orbigny often refers to the Models, but he does not illustrate them. These sculptures were replicated in plaster. Some letters between Alcide and Charles-Marie d’Orbigny and scientists of the Museum, recently published (Béraud & Vénec-Peyré 2003), provide information on this operation. The replicas were sold in the form of instalments as indicated in the label, a translation of which has been published by Parker et al. (1865: 2), “There are four fasciculi, each comprising twenty five models; besides, for the first sixty subscribers and, in addition, three or four shells. The great rarity of the originals does not allow any more to be promised. (The specimens are in glass boxes, which must be opened with great care). [...] The four fasciculi will be issued in the course of the first six months of the year 1823”. This particular care of associating real shells to the reproductions of the Models is evidence of Alcide d’Orbigny’s attention to detail, and of his constant wish to educate; as we shall see later, it has compensated for the loss of certain types in the original collection. Drawings of Models have been published by Deshayes (1828a) and Parker et al. (1865), as well as a photograph of a set of plaster replicas produced by Basset (1885).

12In addition to this set, we also have important topotypic material composed of sand stored in small bottles (1100, according to Terquem 1882: 11), coming from various parts of the world and from which d’Orbigny obtained the species that he described or illustrated throughout his career. These sediments were partially collected by himself or supplied by colleagues, traveller-naturalists, doctors and naval officers. In acknowledgement, d’Orbigny (1826: 249) dedicated certain taxa to these donators, as shown by the specific names deriving from the names of their patronyms. These bottles of sand have proved to be extremely valuable in the revision of d’Orbigny’s species, since they allow us to find topotypes to perform detailed morphological studies, and designate neotypes when required for the stability of the Nomenclature.

The fate of the Planches inédites: incomplete projects of publications

13Certain drawings of the Planches inédites were published when d’Orbigny was alive. He was obliged to considerably limit his illustration of the Tableau méthodique to the main genera, and extracted from the Planches inédites the drawings of 31 species that he grouped together in 8 plates published in 1826. He also gave some of his drawings (representing 22 species) to Guérin-Méneville, who used them to illustrate Cuvier’s “Règne Animal” (1829-1843).

14Admired by specialists who have had the privilege of consulting them or insuring their conservation, the Planches inédites have inspired several projects of publication. Terquem was probably the first to become aware of their value and the necessity to publish them. However, the extent of such a task and a lack of financial means forced him to abandon the project. He described and illustrated 41 species of the Tableau méthodique, but did not reproduce d’Orbigny’s original drawings. He wrote: “We hope that, appreciating the importance of d’Orbigny’s work, someone will complete the work that we have begun, and publish all the plates; this task, which today exceeds our capabilities, could be undertaken in series, which would include a certain number of genera; this work would fill numerous gaps and would serve as a guide for the study, not only of living, but also fossil Foraminifera” (Terquem 1878: 3). Later, with the same enthusiasm, Georges Berthelin carefully traced the outlines of many of d’Orbigny’s original sketches, and bequeathed these copies to Carlo Fornasini. This was related by Fornasini (1898: 11) in a footnote reporting a testamentary letter of Berthelin: “among my books there is an exercise book, large 4 °, entitled d’Orbigny, tableau méthodique; it is handwritten and it includes plates of tracings related to unpublished species of d’Orbigny. I wish that it will be sent under my name to Monsieur Carlo Fornasini, 24 delle Lame, Bologna, Italy. I hope that it will interest him”. Fornasini completed this task by publishing these copies with conscientiousness and patience between 1897 and 1908. In turn, Edward Heron-Allen also wished to publish the Planches inédites, as he declared on two occasions: “We may say in this place that it is our hope and intention in due course to reorganise the d’Orbigny type specimens in Paris [...] and to publish with the co-operation of Prof. Marcelin Boule, Director of the Musée Paléontologique, the whole of the Planches inédites of d’Orbigny” (Heron-Hallen & Earland 1915: 545), and then, “I have made it my duty to make a minute and categorical examination of these Planches in Paris and compare them with the Tableau méthodique, and it is my intention to bring about the publication of the whole of them….” (Heron-Hallen 1917: 34). Unfortunately, the first world war and its consequences would once again prevent the completion of this project.

15The tremendous impulse that was given to the study of foraminifera by the development of oil research gave rise to new hopes, and Maurice Lys, who was at the time an Engineer at the French Petroleum Institute, attempted to save this inestimable patrimony and bring it to the attention of the micropaleontological community. Thus, in the footnote of an article (Lys & Sigal 1947: 16) he indicates, “the publication of the whole of the Planches inédites is envisaged very shortly, accompanied by personal interpretations”. In the same text, however, a certain amount of anxiety can be perceived concerning the outcome of this project when he writes (p. 8), “it must be hoped that purely material means will not stop this project now”. Later, in 1948, he added, “I pray that this project, which inspired such a renowned master, will come up against no more obstacles, and that it will put an end to the uncertainties and inaccuracies which cloud the validity of the names of species given by d’Orbigny during his fruitful and prodigious career”. This new attempt was also motivated by the creation, in parallel, of micropaleontological files, called “authors’ files”, undertaken by the Institute of Petroleum’s Sedimentological Laboratory (Lys & Sigal 1947). Lys was responsible for the creation of d’Orbigny’s file: “The Planches inédites are, as a precious relic, stored in their original state, and admired and consulted by French and foreign specialists. I attempted to go one further than my predecessors, and began to systematically revise the whole of d’Orbigny’s collection” (1948: 7). This file is dedicated to the revision of the 1077 species described or represented by his illustrious predecessor. He carefully examined the foraminiferal mounts, the sand and the Planches inédites. Each species is presented as an index card, including the original diagnosis and drawings (those of the Planches inédites for the species of 1826), the synonymy, and the stratigraphical and geographical characteristics. An optical microscope photograph of the type, together with Lys’s personal observations are also reported. The reproduction of these files “was envisaged in the form of either microfilms or photographs in the desired format”. For details of the constitution and the description of this file, the reader is invited to refer to Lys & Sigal (1947) and Lys (1948a). Like the Planches inédites, this file is also unpublished. It is, however, always available for consultation in the Departement Histoire de la Terre, MNHN. The admiration this file arouses in the researchers who use it does honour to Maurice Lys as being one of the researchers who has most contributed to the significance of d’Orbigny’s micropaleontological collections.

Illustration 2 Les modèles de Foraminifères sculptés par d’Orbigny. (cliché: P. Lafaite, MNHN)
Models of foraminifera sculpted by d’Orbigny. (photo: P. Lafaite, MNHN)

16All these large-scale attempts have failed up to now. Nevertheless, some of the Planches inédites have been published by various authors in black and white or in coulours. This is the case of the plates concerning the following genera: Rotalia (Heron-Allen 1917: pl. 9; Norling 1997: 5); Rosalina (Heron-Allen 1917: pl. 8; Vénec-Peyré 1981: pl. 2; Vénec-Peyré 2002: fig. 3); Triloculina (Lys 1948a: pl. 1) and of one of the plates entitled “free shells” (Lys 1948a: pl. 3). Two plates of sketches, unframed, were published by Lys (1948a: pl. 2) and Hottinger (1960: 104, fig. 55).

Significance of the collections of Alcide d’Orbigny

17The foraminiferal collection of d’Orbigny has undergone several damages, the importance of which had been related by Heron-Allen & Earland (1915) and later referred to by Lys (1947), Y. Le Calvez (1974) and Poignant (1998) who revised d’Orbigny’s collection. A glance at the synthesis table presented at the end of this book shows that the group of Nodosaria is the most damaged. The first among the incriminated causes is repeated curatorial absence over many years. Thus, between the work of Maurice Lys and the nomination of Yolande Le Calvez as curator, the collection obviously suffered due to a lack of vigilance, as suggested by the lack of some specimens, however illustrated by photographs in Lys’s file. Edward Heron-Allen evoked the damage caused by the rising of the Seine River in 1912 [effectively 1910] to d’Orbigny’s collection which was at the time stored in the cellars of the Museum. He also observed that many specimens were invaded or destroyed by mycelium and fungus. Y. Le Calvez (1974) again accused “the action of the glue, that of the salt which filled the shell of the specimens not washed in fresh water”. The destructive power of bacteria, possibly originating from using a brush that was moistened by saliva to manipulate specimens, has equally been brought to question. To this we must add the consequences of indelicate manipulations, and the rashness or lack of attention of Olry Terquem, mentioned by Lys who regrets (1948a: 5), “that he did not work methodically and that he did not always distinguish d’Orbigny’s material from his own; he also neglected to properly store the samples”. It is thus possible that some of d’Orbigny’s types were put in other mounts by Terquem and stored in his own collection, which is also housed in the MNHN.

18I feel it necessary in this section to stress the important role played by certain micropaleontologists who have enhanced the scientific value of the foraminiferal patrimony of Alcide d’Orbigny.

19Whatever the reproach aimed at Olry Terquem (debatable interpretation or carelessness), we are also indebted to him for having partially preserved the specimens selected by d’Orbigny by placing them in small glass tubes, work that, unfortunately, he could not complete (Lys in Lys & Sigal 1947: 189). We must also praise Charles Schlumberger’s attentiveness, as recalled by Heron-Allen and Earland (1915), “We grieve to record that the original tubes of d’Orbigny’s material, which were kept in the cellars of the Musée de Paléontologie, were flooded in the rising of the Seine in the year 1912 and many of the labels are lost. Fortunately, however, when Prof. Schlumberger went over the collection during his Directorate of the Musée, he put labels inside many of the bottles (including the Madagascar samples), and we owe it to him that the loss to science was not greater than it is”.

20The work of Maurice Lys, mentioned above, is exemplary in the significance of d’Orbigny’s patrimony. Although it remains unpublished, the file which he produced has been consulted and used by the specialists who have worked on d’Orbigny’s collections, as I did for this work.

21More recently, lectotypes were selected from d’Orbigny’s collection by Loeblich & Tappan (1964), and by Banner & Clarke (the latter authors having also designated two neotypes; their list was published in 1962). The lectotypes designated by Alfred Loeblich & Helen Tappan were separated from the tubes containing syntypes and transferred by the authors to more modern foraminiferal slides. Hansen (1967) found in the collections of the Mineralogisk Museum of Copenhagen 19 specimens of recent foraminifera accompanying d’Orbigny’s collection of Models. According to the author, the date of reception of these specimens is between 1825 and 1829. This discovery allowed him to designate 7 lectotypes of species described in 1826 (types that are missing from the Parisian collection): Textularia gibbosa, Fasciolites oblonga (= Alveolina oblonga), Gyroidina orbicularis, Pararotalia bisaculeata, Asterigerina rosacea, Anomalina elegans, Nummulina discoidalis (since syntypes of Anomalina elegans have been found in the original collection of the MNHN by Stephan Revets in 1994).

22Since the 1970s, the development of the scanning electron microscope has given an impetus to new studies on the systematics of foraminifera, and has led to the re-examination of numerous species. The species described by Alcide d’Orbigny (some of which are figured in the Planches inédites) have themselves been revised. Among the most important, we must first mention the revisions of the collections of the Canary Islands and Cuba by Y. Le Calvez (1974, 1977a, b), who, following Maurice Lys, greatly contributed to the preservation of this patrimony as Curator of the MNHN foraminiferal collections between 1967 and 1978. Certain authors have revised parts of d’Orbigny’s collection relating to stratigraphic units, such as the Tertiary of the Vienna Basin by Papp and Schmid (1985) (this work was performed on the Austrian collection, see below), and the Eocene-Miocene of Aquitaine by Poignant (1998). Other researchers have carried out taxonomic revisions on certain genera or families (Revets 1990, 1993, 1996, 1997).

The classifications of foraminifera from d’Orbigny to Nowadays

23With the work he completed in his youth, Alcide d’Orbigny made known a large number of foraminiferal species, as he wrote in his introduction (1826), “Until now, a hundred species at most were well known; we have greatly increased this number, and present six to seven hundred different species”. The Tableau méthodique enumerated 544 species; some of them were previously described by Defrance, Fichtel & Moll, Lamarck, Linné, Denys de Monfort, Montagu, Plancus and Soldani, 335 were new.

24His great achievement was to have also multiplied comparisons between taxa, and to have attempted, for the first time, to put order into the nomenclature of these microorganisms which was in a chaotic state. Although the English school was severe with d’Orbigny as recently recalled by Lipps (2002), Parker et al. (1871: 145) stated “with all its faults, and they are neither few nor small, the’Tableau méthodique’by Alcide Dessalines d’Orbigny must be regarded as the alphabet of the nomenclature of the fora minifera”.

25Although in 1826, d’Orbigny, as the other scientists, still considered this group as belonging to the Cephalopods, he was the first to elevate them to ordinal rank, the order of “Foraminifères”. He based this distinction on the presence of foramens (openings between the successive chambers), as opposed to cephalopods which have a siphon, that he called Siphonifera. It was only some years later (1835) that Dujardin discovered the real nature of these unicellular microorganisms and judged that they should form a new class, that of the Rhizopods, a conclusion wholeheartedly approved by d’Orbigny. He explained the evolution of his ideas in the introduction to his work on Cuba (1839a), in which he recapitulated the history of the work dedicated to this group. Among the four periods which he considered to be decisive in the understanding of these microscopic animals, he mentions: “The third period was when, after having carried out more detailed observations, we not only realised that the small shells should not be grouped in the same families as the large Cephalopods, but also that they should be completely separated from them and form adifferent group, which we named Foraminifères [...] The fourth and final change is that which took place in 1825 [sic, it was actually 1835], when Dujardin was the first to publish observations on the animals of these shells; observations which, as we had also aknowledged, compel to remove them not only from the order of the Cephalopods, but also from the class of Molluscs, and to relegate them to the lower classes of animalisation where, as we are inclined to agree, they should henceforth stay”.

26D’Orbigny’s classification (1826) included 52 genera grouped into 5 families defined on the single character of plan of growth or chamber arrangement:

  • The “Stichostègues”: “chambers piled end to end along a single axis, protuding or not and with a more or less important lateral overlap; no spiral”.

  • The “Enallostègues”: “chambers completely or partly linked alternately, or variously piled along two or three distinct axes but without regular or clear spiral”.

  • The “Hélicostègues”: “chambers linked together along one or two distinct axes but making a regular, clear, “turriculate” or discoidal spiral”.

  • The “Agathistègues”: “chambers variously clustered along a common axes, each coiled along the total length of the test; thence the aperture, always with an appendage, is alternatively situated at an extremity or the other”.

  • The “Entomostègues”: “chambers divided into several cavities by walls or small tubes”.

27Later, in his works published in 1839, d’Orbigny made some changes in the taxonomical hierarchy, he classified foraminifera as a class, his families of 1826 as orders, and introduced a new concept of family the name of which was formed by “adding a uniform and euphonic [idae] suffix to the name of the dominant genus showing the most obvious character” (1850a: xlv). At this time he also defined the order “Monostègues”.

28The criteria used in 1826 by d’Orbigny to define the families were severely criticized. The first controversy came from Deshayes (1828a) who reproduced the outlines of 21 species based on the Models, again to illustrate Cuvier’s “Règne Animal”, and relations between the two authors, which were at first cordial, deteriorated in their writtings. In the papers he published in the Encyclopédie Méthodique (1830-1832), Gérard Paul Deshayes severely attacked the Tableau méthodique, using the most unfriendly terms, and modified d’Orbigny’s classification. D’Orbigny’s response was late coming (d’Orbigny 1839a: xxv), more elegant, but just as merciless, “... we regret that Mr Deshayes did not better understand the models of our genera that we had given him, and that he could not distance himself from the conchyliological forms of the Molluscs with which he is familiar; perhaps he would not have tried to upset in one day the result of six years’work upon animals quite different to any he knew, and for which, today, after nineteen years of research, we believe that nothing should be changed in the main cuts that we have primitively established and which was generally adopted in Europe, whereas his classification has not been followed”. Then as mentioned above, the English school severely criticized d’Orbigny’s classification. William Parker, Rupert Jones, Henry Brady and William Carpenter reproached him for splitting the species and regarded most of them as varieties. Opinion quite excessive explained by Cifelli (1990: 13-14) “The English took an extremely broad view of variation and their concern with it nearly reached the point of obsession. […] they refused to admit the existence of such [species] limits”.

29Since d’Orbigny, a large number of taxa has been discovered and former classifications have given way to new ones. Most were analysed by Richard Cifelli, in a text carefully commented in the light of 20th century advances in the understanding of this group. These notes were published 6 years after his death, thanks to the tenacity and generosity of Suzan Richardson (Cifelli 1990, Richardson 1990). They are particularly important to the understanding of the philosophy of these classifications and the historic context in which they emerged. The classifications which marked the second half of the 20 th century are the work of two eminent authorities in micropaleontology, Alfred Richard Loeblich and Helen Tappan. They published two treatises in 1964 and 1987, followed in both cases by several modifications of their classification (1989). Although it represented a monumental piece of work aiming to classify 296 families and 2446 genera, the classification of 1987 suffers from imperfections which were analysed by Haynes (1990).

30All these classifications are the result of reflection based on the state of knowledge advancement at a given time. As Lecointre & Le Guyader (2001:10), formulate “We must lose the reflex of thinking that classifications are unchanging”. It is thus not surprising that the classification of d’Orbigny did not resist the progress of knowledge. Likewise, it seems logical to envisage the emergence of new classifications in the future, in line with the development of studies on living foraminifera, and more particularly of analyses of molecular phylogeny.

31None of the classifications proposed until now corresponds to a natural classification, because they only take into account the characters of the test. As stated by J. Le Calvez (1953: 210) following his work on the biology of foraminifera, “An ideal classification would thus be one based not only on phylogeny, enlightened by compared morphology and stratigraphy, on species ontogeny deduced from the development of microspherical tests, but which would also use biologic information provided by living animals: life-cycles, reproductive elements, cytoplasm and nucleus characters. The small number of species whose life-cycle and cytoplasm are well-known, prevents such a classification at the present time”. D’Orbigny’s classification (1826) is based on the chamber arrangement. Since d’Orbigny, further classifications have used other criteria, such as the nature of the test (organic, agglutinated, calcareous), phylogenetic relationships, or oral characters, but none have taken into account biological criteria. It must be said, in favour of the authors, that biologists have shown little interest in the study of foraminifera. Thus, the reproduction cycle of only 25 species is known in its totality, and that of about 40 partially (Lee et al. 1991), which is a small number out of the 5 000 species living at present. Sexual and asexual phases may alternate during the cycle and create a polymorphism of the test. The first analyses of molecular biology carried out on foraminifera were not undertaken until after 1990 and concern only very few species for the moment. Fossil foraminifera, on the contrary, have been at the centre of a large number of geological studies. For more than 150 years they have been used as stratigraphical indicators, for 70 years as paleoenvironmental indicators, and for about 40 years as indispensable proxies in paleoclimatic and paleoceanographic reconstructions. Today, we estimate around 50 000 foraminiferal species (living and fossil). This brief account explains the discrepancies which exist between the amount of knowledge concerning fossil species and the little information available on the biology of this group, in spite of the steady efforts of certain successful but rare teams throughout the world. It also explains why the comments of Jean Le Calvez still apply today, half a century later, “We must be content with a mostly analytical taxonomy, more paleontological than zoological, based exclusively on the test” (1953: 210).

32For the moment, it is difficult to obtain a consensus on the taxonomic position of Foraminifera which can appear either as an order, a class or a phylum.

The present book

Conception of this volume

33The present work does not aim to be a taxonomic revision of taxa, a great number of species described by d’Orbigny have yet been revised by different specialists. Its objective is to outline the nomenclatural history of the different species figured in some of the finished plates of the Planches inédites and to make the status on the related collection. It also aims to unravel, as much as possible, the errors, the misinterpretations and the confusion which have been responsible for the erroneous interpretation of the concept of certain species by some authors.

34It goes without saying that the content of the 45 Plates illustrated in this volume is the sole work of Alcide d’Orbigny, and that in this present work I have the role of editor and not of author of these plates.

35For the sake of convenience, the taxa are presented in this book according to their order in plate captions which follows the classification established by d’Orbigny in the Tableau méthodique, now obsolete. For the same reason, the family names (“Stichostègues”, etc.) are used in the text, although they are invalid (International Code of Zoological Nomenclature: article 29). I have attempted to give the current name of species. This was not, however, always possible to do, because most of the genera have been split since d’Orbigny’s work. The present day generic assignation is difficult to apply to those species whose original drawings are not sufficiently accurate, and whose reference specimens are not to be found in the collections, as is the case for the Nodosaria (sensu d’Orbigny). For this reason, I have proposed some generic names with a question mark and some species are left in open nomenclature. A synthesis table, at the end of the book, presents the species illustrated in the Plates with the name given by d’Orbigny, their current name, the cross-reference of each to the page, figure and plate numbers, and the collection number of the specimens.

36In writing this book, I referred to a large bibliography. Maurice Lys’s file (Lys & Sigal 1947) and his observations served as a guide to find works published prior to 1948. It helped me to go back to the original sources and occasionally this allowed me to complete the information. I then updated the information we have on the different taxa which had been revised after the completion of Lys’s file. The important works by Ellis and Messina (1940) and Sherborn (1955) provide also a valuable help.

37The list of synonymy presented here is far from exhaustive. I have deliberately limited it to:

  • references of the papers in which species cited by d’Orbigny were described or figured by previous authors such as Lamarck, Soldani, Denys de Montfort, Fichtel & Moll;

  • the references of the Tableau méthodique;

  • subsequent references which made available species’names for the nomina nuda of 1826;

  • other important references in the history of taxa, such as those of taxonomical revisions. In a general way, I have only cited the appropriate synonymies; I have made an exception for erroneous synonymies subsequently applied to drawings of Planches inédites and Models.

  • one or two references giving a recent illustration of species, such as scanning electron microscope photographs when possible.

38For each species, I have given the references of the different foraminiferal mounts labelled to this species’ name in d’Orbigny’s collection. As I have already mentioned, this collection has undergone certain damage, resulting in the poor state of conservation and the temporary or permanent loss of certain specimens (some may simply be mislaid. I therefore prefer to use the term “not found”, as several previous authors have done).

39For the sake of convenience, the references to plates and figures related to the Planches inédites are capitalized (Pl., Fig.) in the following text, those related to other papers use lower case letters (pl., fig.).

40In the present work, dedicated to the analysis of the Plates finished by d’Orbigny and to the related collection, I designate 14 lectotypes and 1 neotype.

Validity of nomenclatural acts related to the species illustrated in the Planches inédites

41Despite the prodigious work that he accomplished, d’Orbigny bequeathed to his successors a number of nomenclatural problems by leaving unpublished a great part of his original drawings; these problems relate to the authorship and date of availability of the species’names that he had chosen and used in the Tableau méthodique as well as the status of specimens in his collections. Moreover, considering 1) the great number of researchers who have studied d’Orbigny’s work over the last 150 years, 2) the explosion in known species due to the considerable development in foraminiferal research driven by oil prospection, and 3) the lack of precise nomenclatural rules until the establishment of the International Rules of Zoological Nomenclature in 1905, the various works inevitably contained a certain number of imperfections or inaccuracies; they emphasize the need to update and clarify certain nomenclatural points.

Treatment of names

42In the Tableau méthodique brief diagnoses are published for higher rank taxa (orders, families, genera and sub-genera), but none for species which are only presented with a reference-list and information on “habitat”. Although d’Orbigny first used the term “foraminifères” in 1826, the author of the scientific name Foraminiferida is C. E. von Eichwald who latinized the vernacular name in 1830. D’Orbigny did not used latinized names for families in his works of 1826.

43He applied the Linnean nomenclature; nevertheless some of the species names used in the Tableau méthodique and in the captions of the Planches inédites raise some nomenclatural problems. It is important to discuss them in this introduction especially because they are related to the pioneer work of foraminiferal studies. Nevertheless, we have to acknowledge, in all fairness, that d’Orbigny’s works and those of the first revisers were produced before the institution of the Règles internationales de la Nomenclature zoologique in 1905, followed by the International Code of Zoological Nomenclature (ICZN; first edition in 1961, latest edition in 1999).

44When d’Orbigny produced his first work, the unique rule in force at that time recommended the use of binominal nomenclature introduced by Carl von Linné. By deontology and as was the custom at that time, d’Orbigny complied with the use of priority, giving precedence to the oldest species names and cited also the patronym of the authors responsible for these names. Nevertheless, when changing generic combinations for species described by earlier workers, he assumed the authorship. To explain this practice, which is inconsistent with the present ICNZ, he wrote in the Prodrome, 2pages summarized as followed (1850a: liv): “whenever one changes the generic name of a species, one must place his patronym after it, with the date of publication of this change […] We have adopted this nomenclature which we consider to be the best”. Indeed, he thought it unjust to “give the authorship to an author who did not know the genus and to deprive the reformer of the result of his work”. Moreover, he added: “by placing behind the designated name, the synonymy of the first describer, one does him justice and considerably simplifies the research” (1850a: lviii).

45The names used by d’Orbigny to designate the specimens illustrated in the Planches inédites, were cited in the Tableau méthodique. Some of them satisfy the provisions of articles 10 to 20 of the ICZN. They correspond to the available names established by Defrance, Fichtel & Moll, Lamarck, Linné, Montagu and Denys de Montfort, as well as to the 31 new names illustrated by d’Orbigny in the 8 plates of the Tableau méthodique. Other new names refer to the figures published by Ambrogio Soldani but d’Orbigny also published numerous names without illustration or description, these nomina nuda (between quotation marks herein) were made available by d’Orbigny in his later works or by other authors. These last cases created problems which are discussed below.

  • The species referring to Soldani’s work. As often mentioned, Ambrogio Soldani illustrated between 1789 and 1798 a great number of foraminiferal specimens using names (mainly groups of descriptive adjectives) which were not consistently binominal. In the Tableau méthodique d’Orbigny established new species names on the basis of Soldani’s drawings. Since he applied the Linnean nomenclature, d’Orbigny is credited with authorship. Because they fail to conform to the provisions of the Code and are not governed by its rules the names used by Ambrogio Soldani are between quotation marks and printed as in Soldani (fide Parker et al. 1871) in the following text.

  • The nomina nuda of 1826. D’Orbigny made available numerous nomina nuda in his works of 1839a and b, 1846, 1850 and 1852, when he described and illustrated these species only cited in 1826.

46Several authors made available some of the nomina nuda of 1826 in publishing drawings of the Planches inédites. Guérin-Méneville (1829-1843) used 22 of them to illustrate Cuvier’s “Règne Animal”. According to Lys (1948a: 11), “d’Orbigny himself indicated those species which could be more easily reproduced (by a cross which can still be seen on the Plates)”. Between 1887 and 1908, Carlo Fornasini validated a large number of names by reproducing the tracings of the Planches inédites, bequeathed to him by Berthelin. Since the original drawings are from d’Orbigny’s hand, the authorship is credited to d’Orbigny; nevertheless, the date of the publication is that of Guérin-Méneville or Fornasini’s papers; then, the authorship can be expressed “d’Orbigny in Guérin-Méneville” or “d’Orbigny in Fornasini”.

47The case related to the authorship of species names documented by the Models is different. Deshayes (1828a) was the first to publish the outlines of 21 Models to illustrate Cuvier’s “Règne Animal”; then Parker et al. (1865), revised the species represented by the 100 Models (without reference to Deshayes’s work); as the Models are the result of the observations made by d’Orbigny they credited the authorship to him. Unfortunately, these sculptures do not provide an indication required by article 12 of the Code; therefore, these names must be credited to the authors who drew the Models’outlines and were the first to publish them. Moreover, Deshayes (1830, 1832) is also responsible for several names cited in the Tableau méthodique and in the Planches inédites because he was the first to describe these taxa. Nevertheless taking into account that he used the d’Orbigny documents, he attributed the name to d’Orbigny, as many micropaleontologists after him.

48Olry Terquem also made available some of the nomina nuda of 1826. Nevertheless, the work of Terquem, apothecary by trade, is not exempt from criticism. As reported by Lys (1948a: 5), “he liberally uses the precious booklets in his determinations; he uses references to them in his synonymy of the species of Rhodes and of the Eocene of the Paris Basin; but his interpretation of the species is very debatable, if not false, and it would have been preferable to us all for him not to have created these confusions”. Despite this, through the Principle of Priority, Terquem is the author responsible for species names “created” by d’Orbigny for species that he understood differently. He is also the author of several junior homonyms of names established by d’Orbigny in 1850-1852.

Typification

49D’Orbigny was well aware of the need of references or standards which he often alluded to in his works. Nevertheless he did not designate type-specimens due to the lack of typification rules at this time. He recognized the high variability of foraminiferal species despite the contrary opinion of the English school. If there is nothing on this topic in the Tableau méthodique, in his later works d’Orbigny insisted upon “species limits” and the morphological variations which “could be related to habitat influence […] sexual influence […] variations related to age […] growth of species” (1852: xlix). For this reason, he always (or almost) grouped together in the same slide several specimens that he thought belonged to the same species. To better understand some of the difficulties researchers had when dealing with the d’Orbigny collection, a few points need to be cleared up.

50For the species based on Soldani’s drawings, the specimens of this author as well as the species of the d’Orbigny collection belong to the type-serie (ICZN: article 72: 72.4.1, 72.4.1.1, 72.5.6), however the Soldani collection seems to be lost. In the Planches inédites the illustrations are not replicates of Soldani’s drawings. Louis Benjamin Fleuriau de Bellevue obtained sand samples collected by Soldani and gave them to d’Orbigny (1826: 249). The young author examined these sands and produced new drawings.

51D’Orbigny established several species illustrated in the Planches inédites (and cited in the Tableau méthodique) in his work on the Foraminifera of the Vienna Basin (1846) based on material given by Joseph von Hauer. In their revision of the material of Vienna, Papp & Schmid (1985) stated: “For quite some time one was of the opinion that the original material of d’Orbigny (1846) was deposited in Paris. The discovery of a document finally disclosed that this material was located at the ‘Geologische Bundesanstalt’ (formerly the ‘K. K. Geologische Reichsanstalt’) in Vienna”. This collection is composed of vials containing one ore more specimens (sometimes several hundreds), labelled with a number on the cork corresponding to the serial order of each species in d’Orbigny’s memoir (1846). There is also a collection related to this monograph at the MNHN in Paris. In each mount, labelled with the species name and locality, one to several specimens are glued on a thin glass plate placed inside a small tube. Two indications in the Prodrome confirm that this collection was performed by d’Orbigny: 1) he said in volume 1 (1850a: iv) “the asterisk, that we place before the serial number of a species, indicates that we possess it in our collection”, 2) the species described in the Vienna Basin listed in volume 3 (1852) are indeed preceded by this mark. He made its own collection and sent back the other part of the material to Joseph von Hauer, which is now in Vienna. Lectotypes have been designated in the Austrian collection by Adolph Papp & Manfred Schmid and in the Parisian collection by other authors. In such case with syntypes in several collections, the Code recommends to choose lectotypes in the collection “preferably of the institution containing the largest number of syntypes of the nominal species-group taxon, or containing the collection upon which the author of the nominal species-group taxon worked, or containing the majority of that author’s types” (Recommendation 74D). With regard to the species described in the Vienna Basin memoir, the Parisian collection perfectly complies with the two last criteria (it is not the case of the collection housed in Austria), and in several case with the first one for some species.

52Among the subsequent acts of typification some errors have to be mentionned. Sometimes the terms “holotype” and “paratypes” were used inappropriately. When one specimen is present in a foraminiferal slide, nothing allows to infer that d’Orbigny had based his species concept on this single specimen because his collection has undergone many manipulations and damages. Thereby, the “holotypes” subsequently designated have the status of “lectotypes” (ICZN: article 74.6) and the specimens labelled “paratypes” are “paralectotypes”. Banner & Clarke (1962) wrote that they have selected 11 lectotypes and 2 neotypes among the species described in 1826. As they stated “… specimens of the following species have been selected by us as types and are now clearly marked on the original mountings….”, they probably correspond to the specimens encircled by a red line in d’Orbigny’s collection. They added “We propose, in conjunction with other workers, to validate these selections in strict accordance with the International Code of Zoological Nomenclature”. Since this validation was not made (fide Loeblich & Tappan 1987), I formally designate these specimens as lectotypes in the following paragraphs dealing with the relevant species.

Difficulties related to the non completion of the Planches inédites

53Beyond this nomenclatural aspect, some errors made by d’Orbigny have added to the difficulties of revisers. Most of them would probably have been corrected if d’Orbigny could have completed this work. They will be noted for each species concerned. The most obvious are related to the genus Robulina and Uvigerina and are responsible for systematic imbroglios that I attempt to explain. The Planches inédites were annotated and corrected by d’Orbigny himself. For example, the corrections in the title of Plates 15 and 16 (this volume), where the generic name Polymorphina s. g. Polymorphina replaces Anomalina, were certainly made by d’Orbigny, because he cannot have mixed up the two genera. Two points testify to this: 1) two plates, with a frame but unfinished, are entitled Polymorphina s. g. Polymorphina with a calligraphy identical to the correction of Plates 15 and 16; 2) the abbreviation “Poly.” followed by the species name appears in the caption of certain sketches. It is possible that d’Orbigny took advantage of an empty frame that was already drawn and intended for Anomalina, to insert his drawings on polymorphinids. It is also possible that he himself corrected certain captions, however other amendments, such as certain corrections in the figures indicating the species’ serial number in the Tableau méthodique, or the addition of the species’ origin in the caption, were probably not made by d’Orbigny.

54Other difficulties relate to the fact that he used adjectives characterizing morphological traits as specific names (costata, elongata etc.). This led to secondary homonymies when they were placed subsequently in combination with other generic names. This risk, evaluated later, perhaps led d’Orbigny to state in the introduction of the Prodrome: “although we do not attach importance to the name, which is, for us, a convention intended to designate an animal, we prefer those which do not recall the morphology” (1850a: l).

55One cannot close this critical aspect related to the nomenclatural problems left by the first foraminiferal works of d’Orbigny without saying again that it was the work of a youth, produced in a period before the Règles internationales de la Nomenclature zoologique. It is only fair to recall his care for the notion of standards or references and his concern for order and method in order to put an end to the “chaos” (a word often mentioned in his works) prevailing in systematic and stratigraphy. These qualities are more apparent in his later works on foraminifera and also in his books, the Paléontologie française (1840-1860), the Prodrome (1850-1852), and the Cours élémentaire de Paléontologie (1849-1852). In these huge works he reevaluated 40 000 species names of invertebrates and foraminifera and finally only kept 18 000 valid names. He arranged the valid species in stratigraphical order, and defined the stratigraphic scale including the concept of reference localities (today’s stratotypes). In the introduction of the Prodrome d’Orbigny emphasized the importance of nomenclature and the necessity to “improve nomenclature which could turn the wheels of Science” (1850a: liv).

Table des illustrations

Légende Illustration 2 Les modèles de Foraminifères sculptés par d’Orbigny. (cliché: P. Lafaite, MNHN)Models of foraminifera sculpted by d’Orbigny. (photo: P. Lafaite, MNHN)
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