Proposed Rules for the incorporation of nomina of higher-ranked zoological taxa in the International Code of Zoological Nomenclature. 1. Some general questions, concepts and terms of biological nomenclature
Alain DUBOIS Muséum national d’Histoire naturelle, Département Systématique et Évolution, USM 0602 Taxonomie-collections, Reptiles et Amphibiens, case postale 30, 25 rue Cuvier, F-75231 Paris cedex 05 (France) [email protected]
Dubois A. 2005. — Proposed Rules for the incorporation of nomina of higher-ranked zoo- logical taxa in the International Code of Zoological Nomenclature. 1. Some general ques- tions, concepts and terms of biological nomenclature. Zoosystema 27 (2) : 365-426.
ABSTRACT The scientific names (nomina) of higher-ranked taxa (above the superfamily) of animals are not regulated by the International Code of Zoological Nomenclature, but by “consensus” among workers. However, when there exists no real consensus, a frequent situation, some criteria must be used to establish which nomen should be considered valid for any given taxon. With the multiplication of taxa that follows the development of cladistic analyses, the implementation of such rules will become more and more necessary and important. To be acceptable by all zoologists worldwide, today and tomor- row, these rules should be independent from the philosophy of taxonomy adopted, but should allow unambiguous, automatic and universal allocation of a single nomen to each higher taxon, within the frame of any taxonomy, including “phylogenetic” ones. This first paper is devoted to the detailed dis- cussion of general theoretical and terminological problems related with this question. It is here argued that it is misleading and dangerous to try and make nomenclature artificially “simple”. The problems posed by the naming of mil- KEY WORDS lions of kinds of organisms, related through evolution and that have been Zoological nomenclature, studied for two and a half century under different approaches, are indeed definitions, higher-ranked taxa, complex: this complexity should be acknowledged, and the discipline in types, charge of this study should be recognized as a specific technical field, with its priority, own methods, concepts and terms. Among various proposals made in this usage, taxonomic category, paper, it is suggested to definitely abandon the misleading term “type” in tax- nomenclatural rank, onomy and nomenclature, objective categories for the “usage” of nomina are phylogenetic taxonomy, terminology, defined for the first time, and a distinction is made between taxonomic “cate- Rules. gories” and nomenclatural “ranks”.
ZOOSYSTEMA • 2005 • 27 (2) © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. www.zoosystema.com 365 Dubois A.
RÉSUMÉ Proposition de Règles pour l’incorporation des nomina de taxons de rangs supé- rieurs d’animaux dans le Code international de nomenclature zoologique. 1. Quelques questions générales, concepts et termes de la nomenclature biologique. Les noms scientifiques (nomina) des taxons de rangs supérieurs (au-dessus de la super-famille) d’animaux ne sont pas régis par le Code international de nomenclature zoologique, mais par un «consensus » entre chercheurs. Cepen- dant, lorsqu’il n’existe pas de vrai consensus, une situation fréquente, des cri- tères doivent être employés pour établir quel nomen doit être considéré valide pour un taxon donné. Avec la multiplication des taxons qui résulte du déve- loppement des analyses cladistiques, la mise en vigueur de règles va devenir de plus en plus nécessaire et importante. Pour être acceptables par tous les zoolo- gistes du monde, de nos jours et dans l’avenir, de telles règles devraient être indépendantes de la philosophie de la taxinomie adoptée, mais devraient per- mettre une attribution non-ambiguë, automatique et universelle d’un seul nomen à chaque taxon supérieur, dans le cadre de toute taxinomie, qu’elle soit «phylogénétique » ou non. Ce premier article est consacré à la discussion détaillée de divers problèmes théoriques et terminologiques relatifs à cette question. L’idée est défendue ici qu’il est trompeur et dangereux de tenter de rendre la nomenclature artificiellement «simple ». Les problèmes posés par la nomination de millions de sortes d’organismes, liés entre eux par l’évolution MOTS CLÉS et qui ont été étudiés depuis deux siècles et demi selon différentes approches, Nomenclature zoologique, sont indéniablement complexes : cette complexité doit être reconnue et assu- définitions, taxons de rangs supérieurs, mée et la discipline chargée de cette étude doit être reconnue comme un types, domaine technique spécifique, avec ses propres méthodes, concepts et termes. priorité, Parmi diverses propositions avancées dans cet article, il est suggéré d’aban- usage, catégorie taxinomique, donner définitivement le terme mensonger de «type » en taxinomie et en rang nomenclatural, nomenclature, des catégories objectives d’« usage » des nomina sont définies taxinomie phylogénétique, terminologie, pour la première fois et une distinction est faite entre «catégories » taxino- Règles. miques et « rangs » nomenclaturaux.
P RINTING CONVENTIONS on Zoological Nomenclature, “ BZN” to its official In the text below, species-series and genus-series organ, the Bulletin of Zoological Nomenclature, and nomina (see Dubois 2000b) are printed, as usual, the “ Code” to the fourth edition, currently in force, in lower case italics, whereas nomina of higher- of the International Code of Zoological ranked taxa are written in small capitals, with the Nomenclature (Anonymous 1999b). This latter following distinction: family-series nomina are in book is here quoted as “Anonymous” for reasons ITALICS, and class-series nomina in BOLD .The sec- explained in Dubois (2000b: 87), as it “does not ond paper of this series (Dubois in press) will give state the name(s) of [its] author(s)”, but is “attribu- precise definitions of most technical terms used ted to a body (e.g., a committee or a commission)”, below, especially all new ones. The abbreviation thus following the definition of the term “ anony- “ICZN” refers to the International Commission mous” in the Glossary of the Code itself (p. 100).
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PREAMBLE circles, thousands of pages have been devoted to this field over the last two centuries, as will be “When the golden rule of priority is once laid aside, exemplified by the publications quoted in the there seems to be no limit to these alterations; for, present work. In the pages below, criticism of the however appropriate a nomenclature may be, dabblers “ Phylocode” project will be developed. However, in science will always be found who will prefer terms of their own coinage to those which are already estab- there is one point for which all biologists interest- lished”. ed in nomenclatural matters can be grateful to Hugh Edward Strickland 1837: 128 the supporters of this project: it has rendered “There are only two ways of naming plants and ani- some of its letters patent of nobility to nomencla- mals, either to give them their oldest names or to give ture in scientific publications. Hundreds of pages them any names you please”. were published in the last decade, some in major David Starr Jordan in Frederick V. Coville 1895: 163 periodicals, dealing with the presentation, de- “As has been said, there are two ways of naming plants, according to the system of priority, or accord- fense and criticism of the “ Phylocode”. Never- ing to the personal system, which is no system at all”. theless, despite this “renewal”, the situation is not O. F. Cook 1895: 431 yet such as a free discussion develops in the international community about the “best system Among biological sciences, nomenclature is of nomenclature”. In our society for which every- rather disliked. It is often considered as a boring, thing that is (or appears) “new” is considered bet- almost useless, intellectually poor but technically ter than the past, it is much easier to develop a unnecessarily complex domain, that is not wor- new proposal, especially if it appears (slightly) thy to attract the attention of “great scientists” “revolutionary”, than defending an “ancient” sys- and should be left to “maniacs of names”: “a non- tem and simply stating that it works well: “there science that is the province of those biologists is no excitement in a story about retaining the with nothing better to do” (Knapp et al. 2004: ‘old’ way” (Nixon et al. 2003: 112). So if any- 612). For a zoologist, to devote time and energy thing dealing with the “ Phylocode”, especially in to this domain is clearly depreciating in the eyes support to this new system, is likely to be quickly of many, and is of little help in an academic published, the same is not true for papers placing career: rather, it can be used as evidence that a themselves within the frame of the current Code colleague is a mediocre archivist and not an active and “just” proposing changes meant to improve researcher. This social discrediting of the field is the latter (see Note 1). interesting and should attract the attention of If this difficulty to publish some kinds of ideas historians of science and epistemologists. can be understood from periodicals that are not However, in contrast with this prejudice, a specialised in taxonomy and nomenclature, it is serious examination of the matter shows that: less so for publications of the latter category, and 1) nomenclature is crucial for all biological especially for BZN, the official organ of ICZN. sciences, that cannot do without it: just like By the middle of the 20th century, this journal Molière’s Monsieur Jourdain who was making was largely open to contributions from all zoolo- prose without knowing it, all biologists use gists worldwide dealing with proposed changes to nomenclature, at least at a given time of their the Code or with specific nomenclatural problems. work, i.e. when they assign a name to the organ- This is well exemplified by the huge volume 15 isms they deal with; 2) the theory of nomencla- of this journal, published by Francis Hemming in ture or onymology (Dubois 2000b) is an 1957 and 1958, which only consists of free intellectually very complex and interesting field contributions to the discussion of various aspects whose mastering requires a broad culture, strong of the Code, before some changes were intro- methodological and conceptual rigour, and intel- duced in its “first” edition (Anonymous 1961), lectual adaptability; 3) despite the “official” when it replaced the last version of the previous neglect or contempt for this activity in academic Règles internationales de la nomenclature zoolo-
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gique (Hemming 1953). An interesting point of likely that any of these two “schools” will disap- this volume is that these discussions are not at all pear, as they correspond to basic philosophical obsolete today. Many of these papers deal with differences that go much beyond taxonomy and questions that had already been discussed long nomenclature. Therefore, as the tenants of both before, in fact since the beginning of scientific attitudes have to collaborate within a single classification, and that will no doubt be discussed nomenclatural system, both camps are bound to again in the future, such as the rôle of name-bear- make concessions and a compromise has to be ing types in nomenclature, priority versus usage, found between both attitudes. Carrying the joke or the relationships between nomenclature and too far might result in a secession within the taxonomy. This is easy to understand: whereas international community of zoologists, as was many of the hard discussions and debates that rightly pointed out by ICZN’s president Richard developed during the history of biology have Melville at a time when “conservationists” were finally come to an end because new data and/or dominant (1958: 1248): “Pursuing a defeated theories made the older theories obsolete, enemy to destruction may be good tactics, but it nothing of this kind is ever to be expected with is bad strategy when both parties have to work biological nomenclature, as nomenclature is not a together after the fight”. scientific discipline, but a technique, a tool, and Unfortunately, in the recent decades ICZN has there is nothing like “truth” to be expected in this not followed the sound advice of its former presi- field. No nomenclatural system is better than dent and has not repeated the experience of another for being “closer to the truth”, but differ- volume 15 of BZN. Most of the discussions ent approaches to nomenclatural questions preparatory to the last edition of the Code reflect different philosophical or practical (Anonymous 1999b) were not held in a public choices. Actually, if they take the time to read the and permanent medium such as BZN, where papers mentioned above, some of the recent they can be consulted again decades later, but on contradictors on these matters will certainly be an “internet forum”, i.e. a completely labile sup- surprised to realise that much of what they can port (see Dubois 2003b, 2004a), that further- write now had already been written, sometimes more at that time was accessible only to part of more clearly or persuasively, in these works of the the international community of zoologists. Only past. In particular, the opposition between two a small selection of the contributions of the inter- major “schools”, one in favour of “usage” and a national community to this discussion on the last posteriori “consensus”, and one in favour of uni- Code was published in BZN, although the total versal a priori stringent rules like priority, is as old volume of texts received by the ICZN secretariat as the discipline of biological nomenclature (e.g., (“more than 800 pages of documents from some De Candolle 1813; Strickland et al. 1843; 500 people or groups”, according to Minelli Blanchard 1889; Robinson 1895; Coville 1895; 2003: 650) was of an order of magnitude similar Ward 1895) and will no doubt continue to exist to that of volume 15 of BZN (1266 pages). A for decades. It is not justified, except on polemic number of contributions were never published, grounds, to call the first one “radical” and the and even then the published texts were authorita- second one “reactionary” (Stace 1991) – when in tively modified, mostly shortened, by the ICZN fact the former is more aptly designated as secretariat. Unfortunately, this is only one case “conservationist” (Melville 1958: 1248). It would among many, as in the recent decades a high be more exact to point out that this opposition is number of applications, dealing with problems a much more general one, that has existed for posed by specific cases of zoological nomencla- centuries between “ de jure” and “ de facto”, i.e. ture, were never published in BZN and these between Roman law, mostly based on a priori cases were never brought to the knowledge of the rules, and Anglo-Saxon law, relying largely on international community of zoologists, or had to jurisprudences established a posteriori. It is un- be so through other periodicals where they are
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more difficult to trace (see Note 2). Censorship either as a class (a group of organisms) or as an of zoologists by the ICZN secretariat is all the evolutionary individual (a clade), whereas a more problematic that, unlike in the case of a nomen is a label, by which a taxon can be unam- paper in any other scientific field, there exists no biguously designated within the frame of a given other periodical specialised in the field, where taxonomy, and which serves as a universal tool rejected papers are likely to be published. BZN for the storage and retrieval of the biological and being the only journal in the world devoted to non-biological information attached to this the discussion of theoretical and practical prob- taxon. Taxonomy is a scientific discipline, where- lems of zoological nomenclature, it is in a situa- as nomenclature is a technique. Both are partly tion of quasi- monopoly for papers in this field. It related but also partly independent. Nomina can should therefore be the responsibility of the edi- be given to “empty concepts”, i.e. created tors of this bulletin to make sure that every opin- without defining the taxa for which they were ion and problem presented by individual created (these are the “ nomina nuda” of nomen- zoologists be brought to the knowledge of the clature). On the contrary, taxa can be recognized, international community through this journal. defined or diagnosed but not named, and in the The only intervention of the ICZN secretariat past other systems have been proposed to desig- should be to check that authors of the applica- nate taxa, such as numbers (numericlature and tions have a good knowledge of the Code and that related systems: Little 1964; Rivas 1965; Hull the problems raised cannot be solved by a simple 1966; Johnson 1970; Heppell 1991), combina- and normal use of the provisions of this official tions of names, letters and numbers (Needham text. As long as this is not the case, zoologists who 1910) or artificial words resulting from a combi- wish to submit new suggestions regarding zoolog- nation of “coded letters” (Bergeret 1783; ical nomenclature to the international commu- Nicolson 1991; Knapp et al. 2004). The long- nity will have to use other journals, as is the case lasting use of a nomenclatural system for this here. purpose, however, is justified by the fact that in human language in general, and in scientific texts in particular, we usually communicate with INTRODUCTION words, not numbers, letters, abbreviations or other non-verbal systems. The current internation- Probably more than 10 million animal species al rules for the allocation of nomina to taxa and live on our planet, of which less than 2 million for the automatic establishment of the valid have been described so far by biologists nomen of any taxon within any taxonomy are (Hammond 1992; Heywood & Watson 1995). those of the current international codes of zoolog- In order to be able to communicate among us ical (Anonymous 1999b), botanical (Greuter et about these species, we need a taxonomic and al. 2000), bacteriological (Lapage & Sneath nomenclatural system, that will act as a “conve- 1992) and virological (Van Regenmortel et al. nient information storage and retrieval system” 2000) nomenclatures, which have force of law for about taxa, their characters, evolution and rela- all biologists and editors worldwide. The present tionships (Mayr 1981: 511). work is a continuation of my previous efforts at Although connected, taxonomy and nomenclature contributing to a clarification of some basic are different disciplines. Taxonomy recognizes concepts of biological nomenclature, which classificatory units or taxa, whereas nomenclature requires in some cases the establishment of a new attaches a given scientific name or nomen (see terminology, a process that is part of onymology, Dubois 2000b) to each of these units: of course, i.e. the study of the concepts and theory of biolog- “taxa must have nomina” (Dubois 1988b), but a ical nomenclature (Dubois 2000b). This paper taxon is not a nomen, and vice versa. A taxon is a is more specifically centered on zoological classificatory concept, which can be considered nomenclature, currently regulated by the zoological
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Code, although a number of discussions below taxon being designated by different authors have broader implications and also concern the under different nomina, a situation that is cer- nomenclature of other groups of organisms. tainly not satisfactory (Corliss 1983). In zoology, the current taxonomic system allows Recent amphibians are a good example of this to place all the known species unambiguously in situation, with the class being known either as a hierarchic arrangement of taxa (genus, family, A MPHIBIA or as B ATRACHIA, the subclass either as order, class, etc.), and the nomenclatural system B ATRACHIA or as L ISSAMPHIBIA, and the three allows us to designate unambiguously any of orders being designated by different authors these taxa by a single nomen, that should be under different nomina, the most common of agreed upon and used by all zoologists world- which are G YMNOPHIONA and A PODA for caeci- wide. Despite these general needs, however, the lians (worm-like amphibians), A NURA and current zoological Code only recognizes and S ALIENTIA for frogs (tailless amphibians), and regulates the nomina of three “nominal groups”, U RODELA and C AUDATA for salamanders (tailed renamed “nominal-series” by Dubois (2000b) in amphibians): whereas for the two former groups order to avoid possible semantic confusions: the the first cited nomen is more and more often “ family-series” (superfamily, family, subfamily, used by recent authors, uncertainty still strongly tribe, subtribe and additional intermediate taxa), persists for the third group (salamanders). The the “ genus-series” (genus, subgenus) and the “ spe- unclear status of these nomina is well exemplified cies-series” (species, subspecies, species-group, by the fact that in recent editions of the etc.). Nomina that may be given to forms or taxa Zoological Record up to volume 139, the orders below the subspecies (that can be called variety- were designated as follows: “ A NURA (= S ALIEN- series nomina) are ignored by the Code, as are TIA)”, “ C AUDATA (= U RODELA)” and “ G YMNO- nomina of taxa above the superfamily, i.e. class- PHIONA (= A PODA)”. It is also stressed by the fact series nomina (Dubois 2000b). that alternative nomina for the same taxon, e.g., With the development of cladistic studies dealing U RODELA and C AUDATA, or G YMNOPHIONA and with most groups of animals, new hypotheses of A PODA, are not rarely used either by different relationships between taxa are regularly proposed, authors in different chapters of the same book, and many of them are followed by new taxonom- or, quite often, by the same author in different ic proposals, that often include a multiplication publications, or even in the title of the same of higher-ranked or “higher” taxa (see Note 3), publication (for more details, see Dubois in especially within the frame of various systems of prep.). Such a situation may not seem very prob- “phylogenetic taxonomy” (De Queiroz & lematic for specialists of the zoological groups Gauthier 1992). The units which such systems concerned, who all know these different nomina, recognize are not taxa, in the traditional sense of but it may cause more confusion whenever non- the term, as they are defined by hypothesized cla- specialists (either zoologists working on other distic relationships, not necessarily by characters. animal groups or biologists or non-biologists They were called phylons by Dubois (1991a) and involved in disciplines far from systematics) need more aptly cladons by Mayr (1995: 425), and the to use the nomina of taxa to find information. system ordering them was designated as cladifica- Furthermore, with the growing computerisation tion (Mayr 1997) or cladonomy (a term coined of knowledge in databases, the risk is great that in and published independently in the same year by the future some references and pieces of informa- Brummitt 1997 and Dubois 1997b). tion may be lost or difficult to trace if the same The absence in the Code of rules for the nomina- taxon is designated by different nomina in differ- tion of taxa (or cladons) above the superfamily is ent works. a source of confusion and instability. Even when Here I will discuss some general problems of zoo- no new taxonomic proposals are involved, the logical nomenclature, in particular dealing with absence of rules sometimes results in the same higher taxa. In a second paper (Dubois in press),
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I will propose general rules for this nomenclature, classifications are of less significance to system- which may help to avoid the progressive develop- atics than is progress in analytical methods. It fol- ment of nomenclatural chaos for higher nomina. lows that the classification that serves as the best In a third paper (Dubois in prep.), as an example index to the analysis that produced it, and the of the use of the proposed rules I will present a easiest to use, is the wisest choice” (Ashlock detailed study of the status of the amphibian 1984: 44). nomina just mentioned. With these ideas in mind, Dubois (1991a: 63) Let us first address the following question: what stressed five properties that should be displayed properties should the taxonomic and nomencla- by any system of classification of biological taxa tural systems display to be most useful to taxono- (taxonomic system) to be theoretically satisfacto- mists, other biologists and non-biologists users of ry, acceptable by all biologists, workable and effi- nomina of taxa? cient. These properties, to which a sixth one, numbered (T4) below, can be added, are as fol- lows: SOME DESIRABLE PROPERTIES (T1) Unicity: this means that within any hierar- OF TAXONOMIC AND NOMENCLATURAL chical taxonomic system, there should be asingle SYSTEMS IN BIOLOGY taxonomic hierarchy for all living beings, not sev- eral. This applies in particular to taxa that show T AXONOMIC SYSTEM some evolutionary or biological peculiarities, Biological classifications may be of various kinds such as parthenogenetic or gynogenetic reproduc- according to the function we ask them to play tion, or hybridogenetic meiosis: they should also (Gilmour 1951). Two major functions of such be ascribed to the standard nomenclatural rank of classifications (see e.g., Mayr 1982, 1997) are a species, even if attention should be pointed to practical one (i.e. providing a universal system of their peculiarities in special ways, by referring storage and retrieval of information) and an them to particular taxonomic categories (Dubois explanatory one (i.e. providing an evolutionary & Günther 1982; Dubois 1991a). In this respect, interpretation and explanation of the diversity of the distinction between taxonomic category and organisms). Ignoring the first of these functions nomenclatural rank (Dubois 2005a) is relevant. to concentrate only on the second may seem On the other hand, this does not imply at all that appealing as a “purer” approach but is not doing a single taxonomic system should be imposed a service to science and its innumerable users in “from outside” on all taxonomists, as suggested all domains of human activity (Benton 2000). In by some (e.g., Godfray 2002, rebutted e.g., by contrast, emphasis on the first function is justi- Knapp et al. 2002). “Reaching consensus on spe- fied when classifications are understood as tools cies concepts (...) would indeed be desirable, but aiming at being useful to the whole scientific many serious obstacles would need to be over- community as well as to non-scientific users, not come before this could happen – not least of which designed for a “private usage” of taxonomists and is the failure of the species themselves to behave phylogeneticists: “Classifications merely store in a consistent and predictable way. Asking system- information; they do not tell us why or how the atists to resolve the problem of species concepts is information has the order it does” (Cracraft akin to asking ecologists to agree on which factors 1974: 79). “Classifications are not theories. control the distribution of organisms on the Several authors of quite diverse views (...) have landscape. That is to say, it is one of the central expressed the opinion that the best use of a for- issues of the discipline and is not likely to be mal classification is as an index to the analysis resolved definitively in the near future” (Smith from which it is derived and the organisms that it 2004). “Taxonomy is a complex and dynamic includes. Because formal classifications can only science, not a simple (albeit technically difficult) imperfectly express a full phylogenetic analysis, collation of facts like the nucleotide sequences of
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a genome survey. It is probably only at the most nothing surprising or shocking as any given taxon- complex levels of genetics, in dynamic proteo- omy is not only the result of previous works, but mics or developmental biology, that anything like also a working hypothesis, that may be confirmed the complexity of hypothesis-generation and test- or rejected by subsequent works: by itself, this ing involved in routine taxonomy and system- heuristic function of taxonomy has always acted atics is encountered” (Thiele & Yeates 2002). as a fruitful contribution to the development of Furthermore, the rôle of specific and subspecific our knowledge about organisms. It is much nomina in conservation biology poses particular better to assign a given, albeit provisional, place problems which require formal recognition of to most taxa in a taxonomy, rather than to have taxa for this mere purpose (e.g., Zink 2004; Mace ataxonomy with many “ incertae sedis” taxa: 2004). Emergence of a unified taxonomic system although there are certainly very few “certae sedis” could possibly occur in the future, through three taxa in our classifications, this in no way impedes possible processes: free adherence of most taxon- the latter from playing a useful rôle at every step omists to one of the current taxonomic schools, of the history of biology (Kottelat 1997; Dubois partial synthesis between these different concep- & Ohler 1999; Dubois 1999b). tions (Mayr 1981; Estabrook 1986; Rasnitsyn (T4) Hierarchy: in order to work as a system of 1996; Knox 1998) or emergence of a new storage and retrieval of information, a taxonomic conception (Crowson 1970; Dubois 1988c). The system must have a hierarchical structure. With- situation is clearly not ripe for any of these three out such a structure, all taxa having the same steps (Dubois 2003a), and for the time being the “level” or “rank”, scoring of millions of items existence of different conceptions must be respect- would be necessary before finding the proper ed, as all the history of science shows that author- one. This is just like the difference between an itarian imposition of an “official” theory or alphabetical index (all words being listed similar- opinion on some scientists by other scientists, or ly indented from the margin) and an alphabeti- by administrators or politicians, usually acts as a cal-thematic index (with successive indentations brake against the development of a scientific field from the margin showing subordination of rather than as a stimulating factor. concepts). To be efficient and coherent, a hierar- (T2) Exhaustiveness: the taxonomic system should chy must be unique, each taxon at a lower level be devised in such a way as to be able to accom- being referred to a single taxon at the next higher modate all living beings ever to be found in the level. This can be rigorously defined in mathe- real world, not only some of them. This need for matical terms (e.g., Woodger 1952: 11) and was a global coverage of all living beings results in a more simply called by Gould (2000: 66) “irrevo- need for taxonomic concepts to bear some deter- cable branching without subsequent amalgama- mined relationship to universally observable or tion”, “continuous branching with no subsequent demonstrable patterns and particularities of the joining of branches” or “successive (and excep- organisms of the real world, rather than being tionless) dichotomous branching”: “two species derived solely from some general theory, such as a in the same genus can’t reside in different fami- theory of evolutionary process, or a theory of bio- lies, and two orders in the same class can’t be logical classification. A taxonomic system that placed in different phyla”. More generally, this would for some reason exclude from its field of can be described as follows: “Taxa at the same competence some organisms (e.g., for being of rank are not included within one another; taxa at hybrid origin, or for being teratological) would higher ranks are not contained within taxa of not be acceptable. lower ranks” (Carpenter 2003: 84). Although (T3) Univocality: the taxonomic system should apparently obvious, such a rule was not always be univocal, i.e. any given living being should strictly followed by early taxonomists (see e.g., unambiguously be ascribed a single place in the the treatment of the genus Proteus in Laurenti system. This place may be provisional, which has 1768, discussed by Dubois 2004b) or even by
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some more recent ones (Sigal 1966). A good would be most inconvenient in use and poorly taxonomic hierarchy is “an organizational model informative. in which: 1) the highest level of organization (T6) Robustness: the taxonomic system should consists of a single entity; 2) an entity at a lower display at least rather important (but not com- level of organization is related to only one entity plete) stability, so that every new discovery or at the next higher level but can be related to more hypothesis should not be liable to modify it than one entity at the next lower level; and 3) partly or totally (but that some do). This stability entities at all lower levels are related by extension should concern both the classificatory pattern to the single entity at the highest level of organi- and the nomenclature. The major consequence zation” (Knox 1998: 7). of this requirement is that very rigid systems of (T5) Homogeneity: there should be some equiva- classification, in which any new fact, new taxon lence, by some criteria, between various taxa or new hypothesis of relationships (even when the ascribed to the same rank within the taxonomic change concerns only the phylogenetic position hierarchy. Although discussed by various authors of a single species) must be followed by a com- already (e.g., Hennig 1950, 1966; Van Valen plete change of the taxonomy (and consequently 1973; Schaefer 1976; Dupuis 1979; Dubois of the nomenclature), cannot be supported. A 1988a, c; Avise & Johns 1999; Minelli 2000), classificatory system should display at least some this difficult problem has not yet been satisfacto- inertia or flexibility, in order to incorporate at rily solved by the international community of least part of the new findings without taxonomic taxonomists. This means that taxa ascribed to a (and nomenclatural) changes. However, this does given rank in different clades may not be equiva- not mean that classification should remain the lent in various biological terms (numbers of sub- same forever: important new information must ordinate taxa; morphological, genetic or be acknowledged by a change in the taxonomic ecological diversity; speciation and extinction (and nomenclatural) patterns. As often in rates; genetic and evolutionary patterns, etc.; see biology (see Mayr 1997), the best solution to this e.g., the case of the taxa of birds as compared to problem is in a good equilibrium between two those of other vertebrates, discussed in detail in extreme and opposite points of view. The need Dubois 1988c). Consequently, comparisons that for relative stability of taxonomies would be most are sometimes made among lists or numbers of appropriately acknowledged by replacing the taxa in various works dealing with evolution, frequent request for stability by a request for stratigraphy, biogeography or ecology, may be robustness, which describes more adequately the irrelevant and spurious, and must be considered kind of “stability” we need in zoological taxon- very cautiously (Smith 1988; Sundberg & Pleijel omy and nomenclature (see also Dubois 1998). 1994; Minelli 2000; Pleijel & Rouse 2003). This Robustness could be defined as “stability + is not at all a good reason, however, to reject the change”. Whereas “taxonomic stability” alone hierarchical structure of the taxonomic system. would mean that taxonomy must be definitively Even if it could (and possibly will) be improved frozen and is not allowed to change, “taxonomic by making ranks more significant in biological robustness” means that taxonomy must be terms, this structure already allows taxonomy to flexible enough to be able to change, in order to play the practical rôle of an indexation system of take new information or ideas into account, but information, just like the hierarchical structure of that it cannot do so “easily”. keywords in an index or a database. In such sys- These six major criteria, to which a few others tems, keywords at a given level are only roughly could be added, could form the basis for a rela- equivalent (in terms of information content or tively simple and lax “code of zoological taxono- diversity, or of number of subordinate keywords) my” which could be accepted and adopted by all but are nevertheless quite useful to track down zoologists worldwide, independently from their the information. A taxonomy without ranks “taxonomic school”, their ideas on organic evolu-
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tion, and on the mutual relationships between with the request that all taxonomies be cladono- phylogenetic hypotheses and classification. In mies, these kinds of organisms should either be contrast, it is strongly to be feared that any more ignored or artificially included in the same taxa as stringent or “directive” taxonomic system would the organisms from which they originated, which be shared only by a part of the international zoo- is not acceptable. logical community and likely to be strongly Finally, in the future, completely new facts may modified later, as our ideas and knowledge about also have to be taken into account, such as the biological and evolutionary processes evolve. possibility of “lateral gene transfer” (LTG) be- As a matter of fact, it is quite likely that future tween organisms (Mourant 1971; Ruffié 1982: developments of biology will lead to further basic 213-215; Kidwell 1993; Doolittle 1999, and changes in our taxonomic thinking and practice references therein; Martin 1999; Nelson et al. (Jørgensen 2000). These may include the pro- 1999; Gould 2000; Kidwell & Lisch 2001; Oren gressive taking into account of new knowledge 2004), which might go much further than the concerning the evolution and functioning of series of symbiotic events that underpinned the genomes, in particular regarding various aspects evolution of eucaryotes (Stevens 2002): although of genetic regulation, which may ultimately lead no evidence currently exists for this phenomenon to the development of a new “experimental tax- in Metazoa, this cannot be theoretically ruled onomy” (Crowson 1970; Dubois 1988c). Our out, and actually phenomena of this kind might growing knowledge on the way the genome well account for some “strange” cases of shared works throws new light on long-known facts such characters between cladistically unrelated species as “character reversal” in some clades, that make occurring in the same region of the world, which sense if some parts of the genetic program, rather are well known to field naturalists, and usually than being lost, may be kept “sleeping” for a (although often “uncomfortably”) “explained” by while and “reactivated” later, as well exemplified “ad hoc” concepts such as “convergence” or by the frogs of the genus Gastrotheca (Del Pino & “mimetism”. As Doolittle (1999: 2124) put it: Escobar 1981; Wassersug & Duellman 1984). “If ‘chimerism’ or ‘lateral gene transfer’ cannot be The currently dominant idea that all taxa are dismissed as trivial in extent or limited to special created by cladogenesis has no generality, first of categories of genes, then no hierarchical universal all because viruses have no general cladistic rela- classification can be taken as natural. Molecular tionships between them (so they cannot be prop- phylogeneticists will have failed to find the ‘true erly “cladified”, except at a very low level, but tree’, not because their methods are inadequate or only classified), but also because evidence is now because they have chosen the wrong genes, but strong that, not only in the vegetal kingdom but because the history of life cannot properly be also in animals, reticulate phylogenetic relation- represented as a tree”. This would have drastic ships and speciation by hybridization (even be- consequences on our conceptions of biological tween species that are only distantly related, and classification, exemplified by this suggestion: at least not sister-species) are not uncommon: “While retaining useful names for recognized this applies to gynogenetic and hybridogenetic groups (Archaea and Bacteria), one could see kleptons, parthenogenetic klonons, and allopoly- these as taxonomic descriptors based directly on ploid species (e.g., Bogart & Wasserman 1972; shared genes, but only based indirectly and Dubois 1977, 1991a; Dubois & Günther 1982; unpredictably on shared ancestry” (Doolittle Dawley & Bogart 1989; Günther 1991). In par- 1999: 2128). ticular, hybridogenetic kleptons or zygokleptons All these possible future developments, and (Dubois 1991a) cannot be properly considered as others that cannot be thought of today, suggest clades, as hybridization occurs again at each that taxonomy is not a discipline of the past, but generation, so that such taxa do not result from a may show major changes in the coming century: single evolutionary event. In order to comply this implies that prudence is in order and that it
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would not be a good idea to “freeze” taxonomic source of complication of any nomenclatural sys- theory and practice according to our current tem is the existence of numerous exceptions to the thoughts. general rules. A good nomenclatural system must be in force in the vast majority of situations and N OMENCLATURAL SYSTEM must not include a forest of exceptions, for if the Similarly, it is possible to propose a set of proper- latter are too numerous they obscure the generali- ties that a formal and codified system of biologi- ty of the rules and their normative value. cal nomenclature should have in order to be (N4) Unicity: there should exist a single nomen- coherent, efficient and acceptable by all taxon- clatural system for all organisms. This aim is cur- omists worldwide, whatever their “school of rently not reached, as several distinct codes exist thought”. These, of course, include the same six for different kinds of organisms (animals, plants, properties outlined above for the taxonomic sys- bacteria, viruses, and even cultivated plants). It is tem (as both systems should be compatible and unlikely that these codes will ever be unified “parallel”), but they also include additional prop- under a single “ Biocode” (Hawksworth et al. erties, that can be requested from a set of rules 1994; Anonymous 1999b; Blackwell & Powell and could not be so in a free scientific discipline. 1999): as the botanical and zoological codes have Eleven “principles” can be outlined for any been in force for about one century, millions of biological nomenclatural system: nomina have been stabilized according to these (N1) Independence: the nomenclatural system respective codes, and homogenizing them would should be “theory-independent” (Lidén et al. necessarily entail major changes in one group of 1997: 738), i.e. devised in such a way as not to organisms or in both, which would go against the infringe upon the independence of taxonomic need of nomenclatural robustness that will be dis- thought and action. The function of a code of cussed below. It is however necessary to stop at nomenclature is simply, and should be restricted this point, and not to continue a process of sub- to, providing clear rules allowing one to know the division of the codes that was illustrated for valid nomen of any given taxon in any given taxo- example in zoology by the introduction of a par- nomy. It should be possible to apply the same ticular kind of “name-bearing types”, the hapan- rules to all taxonomies and taxonomic systems, totypes, for protistans (Anonymous 1999b). The without having to discuss the theoretical founda- risk would be to have a particular code for every tions and practical aspects of the latter. As nicely group of organisms, i.e. for every community of written by Knapp et al. (2004: 620): “Keep the “specialists”, which would be a threat against the science out of nomenclature!”. unity of biology. (N2) Exhaustiveness: like the taxonomic system, (N5) Universality: to be usable everywhere in the nomenclatural system should be devised in space and time, a nomenclatural code must be such a way as to be able to accomodate all living accepted by all biologists, whatever their country, beings ever to be found in the real world, not only language, etc. It must also be the same code for some of them. Here also, exclusion from its field all taxonomists, whatever their conception of of competence of some organisms (e.g., for being taxonomy. A universal code cannot be linked to a of hybrid origin, or for being teratological) would particular conception of taxonomy, as this would be unacceptable. not respect the first principle of independence (N3) Simplicity: the system must have a coher- between taxonomy and nomenclature presented ence, an internal logic and an elegance that make above. it easily assimilated by anyone who makes a real (N6) Univocality: in a good nomenclatural sys- effort to understand it. This means that useless tem, allocation of a nomen to a taxon must be complications should be avoided, but not neces- univocal, i.e. it should not allow ambiguity in sary complexities that are imposed by the particu- either sense: given a taxonomy, a taxon must re- larities of the matter themselves. An important ceive one single nomen, in all countries worlwide
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and by all taxonomists, whatever their opinions influence of their works, etc. If it were not the or tastes. Of course, this requirement does not case, one would no doubt assist very rapidly to concern distinct taxonomies of the same group: the leadership (not to say “dictatorship” or each taxonomy implies its own nomenclature. “tyranny”, to use words quoted below) of some (N7) Automaticity: the rules that command the biologists, e.g., of some countries, on those of the allocation of nomina to taxa must work in the rest of the world. This should be avoided by all most automatic possible way, without leaving means, not only for “ethical” reasons (although place to arbitrary or bureaucratic decisions. The such reasons have no doubt their place in the freedom of choice of taxonomists as concerns policy of scientific research), but also for a much nomenclature must be restricted to the indispens- more practical one: not doing so would no doubt able minimum, i.e. concretely to only two situa- imply a risk of secession within the international tions: at the creation of a nomen, and for every community of biologists. If several nomencla- nomenclatural act, i.e. every first-reviser action tures became stabilized in different countries or whenever such an action is necessary. In all other groups of countries, this would imply major cases, the decision should be automatic and problems of communication between biologists should, in all rigour, arouse no emotion from the worlwide, with risks of important consequences part of users. A good nomenclatural system in various fields such as conservation biology, should not only avoid arbitrary decisions, but medical and agricultural sciences, not to mention also limit to the minimum the recourse to basic biological research. In order to respect this bureaucratic action through committees, com- deontological criterion, a very important feature missions, courts or other boards, charged to take of any nomenclatural system is that the initial decisions whenever the rules leave some doubt in status of a nomen (i.e. its nomenclatural availabil- their application. The automaticity of the rules ity and its allocation to a biological taxon) be only should ideally be such that two taxonomists or at least primarily (in case of initial ambiguity) living on opposite sides of the planet should be determined by the original publication in which able, without communicating with each other, to this nomen was first proposed, and not by any reach the same nomenclatural conclusion when- subsequent, correct or incorrect, usage of this ever they adopt the same taxonomy. Any system nomen. requiring a frequent recourse to a bureaucratic (N9) Hierarchy: in order to allow taxonomy to authority would be too heavy and soon bound to play its role of system of storage and retrieval of become inefficient. Furthermore, the strong risk information, a nomenclatural system must have a exists that it could place the taxonomists of hierarchical structure. Nomina of different kinds different countries or having different opinions in should be given to taxa of different ranks. The an unbalanced situation, which leads us to the way such an indexation system through nomina next point. works is simple: even without being a specialist of (N8) Deontology: the basic rules must be such the taxonomic group concerned, any biologist or that all taxonomists worldwide can use them and non-biologist worldwide can find almost instant- contribute equally to biological nomenclature. ly, when reading nomina like “ Saga pedo (Insecta, These rules should not favour any group of Orthoptera, Tettigoniidae)”, “ Aubria masako taxonomists against the others. In case of prob- (Amphibia, Anura, Ranidae)” or “ Upupa epops lems, the criteria allowing decision should be (Aves, Coraciiformes, Upupidae)”, that the first independent from country, language, taxonomic one is a grasshopper, the second a frog and the school, opinion, date, support used to publish the third one a bird, and thus have immediate access data, and also from other more “material” para- to all the relevant literature concerning these meters such as the financial resources of researchers, taxa. This would not be possible, or much more laboratories or countries, their notoriety, their difficult, under a uninominal and non-hierarchi- “international importance”, the distribution or cal nomenclatural system.
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(N10) Homogeneity: in a good nomenclatural sys- only deals with nomina and their hierarchical tem, taxa of the same “nature”, i.e. in a hierarchi- arrangement). Quite independently from any cal taxonomic system, of same rank in the hierar- biological discussion on their genetic structure chy, should be treated in the same way, i.e. they and evolutionary particularities, kleptons, for should bear nomina formed in a similar way and example, must be named (Dubois 1988b) and following the same rules. This allows taxa of same given a place in the hierarchical system, in order rank to be recognized as such by simple examina- to be unambiguously designated in various kinds tion of the nomen. It is thus desirable that spe- of biological works or for more “trivial” purposes cific nomina cannot be mistaken for generic like placement in texts dealing with the manage- nomina, familial nomina for subfamilial nomina, ment of biodiversity (official “species lists” for etc. As discussed above, this does not mean that conservation biology, custom legislation, etc.). these taxa are necessarily “equivalent” in biologic- (N11) Robustness: finally, it is desirable that al terms, but that they have an equivalent place nomina of taxa do not change if taxa do not in the hierarchical taxonomic system. A good change, or if they change only “slightly”. Schuh example of this situation is provided by the spe- (2003: 62) aptly wrote in this respect: “by their cies rank. There are various concepts of the taxon- very nature, the codes are designed not so much omic category of species (see e.g., Mayden 1997; to create a stable nomenclature but possibly bet- Pleijel & Rouse 2000; Wheeler & Meier 2000). ter said, ‘an orderly one which minimizes changes Many taxonomists claim that their concept is in names as they are applied to taxonomic “the best” (Cracraft 2000) and should accommo- concepts’”. The problem is dual, as well outlined date all forms of living organisms. However, this by Thiele & Yeates (2002): “The heart of the results from a strong influence of the reductionist problem is the central duality and tension in mode of thinking on evolutionary biology. Given taxonomic names. On the one hand, names are the diversity of evolutionary processes, there is no convenient shorthand representations of scien- reason to think that all organisms should pertain tific hypotheses, and as such should be as volatile in units of the same “kind”. Systematics should as hypotheses in any other field – proposed, used, try and account for real evolutionary phenomena, modified and then perhaps discarded as evidence which requires fitting our taxonomic concepts to dictates. On the other hand, taxonomic names this reality, not the contrary. The correct attitude are keys to biological information, unique codes in this respect is probably to consider that, accord- that unlock the library of knowledge about those ing to the biological group and its evolutionary organisms, and for this purpose they must be history and particularities, different “kinds of non-volatile or the links will be lost. A solution units” should be recognized (Claridge et al. must adequately recognize these dual roles and 1997). Thus, so-called agamospecies or klonons, decouple the system that allows maximum free- or hybridogenetic or gynogenetic “species” or dom of hypothesis-generation from the system kleptons, have characteristics widely different that provides names for users”. Here also, like for from those of “bisexual Mendelian species”, but taxonomy above, the term robustness describes this does not entail that they should not be more adequately the kind of “stability” that we ascribed the nomenclatural rank of species in a need in nomenclature. Complete stability of hierarchical taxonomy, provided their particular nomina is indispensable only in some very spe- biological characteristics are pointed out, e.g., by cific and quite rare cases, i.e. when it concerns the use of special marks like interpolation of “kn.” nomina that are widely known and used in or “kl.” between the generic substantive and the various fields of human knowledge and commu- specific epithet (Dubois 1991a). Contrary to the nication outside the restricted domain of systematic statements of Minelli (2000), this stresses the research. This situation only concerns a very small independence between taxonomy (which refers proportion of all the millions of nomina available to biological concepts) and nomenclature (which in biological nomenclature. Beside these particu-
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lar cases, which of course must be duly con- stem from a debate around the relationships be- sidered, the request for “stability” of nomina tween reality and language. The long-discussed should not become a “religion”, in the shadow of question (see references and discussion in which the whole nomenclatural system can be Keller et al. 2003) whether taxa are classes (i.e. weakened and subsequently dismantled. It is true concepts) or individuals (i.e. real “things”) is that taxonomists need stability, but in reality what largely irrelevant, because they are both. they need is stability in the nomenclatural rules of Doubtless, “things” do exist (or have existed) in allocation of nomina to taxa, not stability of the the real world that, being the result of a real nomina themselves. Supporting simply “nomen- phylogenetic process, can qualify as “natural clatural stability” without further qualification clades” or “individuals” (this term being taken in would mean: 1) either that neither taxonomy nor an ontological sense). The phylogeny of any nomenclature are allowed to change; or 2) that group of organisms is a historical process that any change, even very minor, in taxonomy, must occurred only once and under a single pattern result in a nomenclatural change. In contrast, (real cladistic relationships). It is now widely “nomenclatural robustness” means that taxon- acknowledged that a major aim of evolutionary omy can evolve, but that a given change in taxon- systematics is to discover these cladistic relation- omy does not necessarily result in a nomencla- ships and to base biological taxonomy on them. tural change. More details on these points will be Whether this aim is ultimately within the given below. possible reach of science, using the (imperfect) Confrontation of the 11 “principles” above with tools that we have to reconstruct the past events the actual Code of Zoological Nomenclature cur- of the history of life on earth, is open to question. rently in force (Anonymous 1999b) shows that What is indisputable however is that, for the time the latter follows most of these principles. There being, and probably for many decades yet, this are a few exceptions however, and more so follow- aim is far from being reached. For the time being, ing some of the changes introduced in the last virtually any cladistic analysis of any living group two editions of the Code (Anonymous 1985, provides a cladogram that is slightly or largely 1999b), as compared to the previous ones different from the previous ones. Cladograms as (Anonymous 1961, 1964, 1974): this matter was we currently use them are working hypotheses, discussed in more detail elsewhere (Dubois 1985, liable to be refuted and replaced by better ones, 1999a, 2000a), and will need more discussion in they are not final results of science. They are the future. For the purpose of the present work, not facts or things of the real world, but however, the principles above will be taken as a scientific concepts. As Benton (2000: 634) put it: basic guideline for the reflection on new, univer- “phylogeny is real, classifications are not”. sal rules for the nomenclature of higher taxa in Actually, classifications also are real, as creations zoology. However, before going into the ques- of the human mind, but what is at stake is their tions posed by higher nomenclature, a few more relation to the actual history of life on earth, i.e. general theoretical questions concerning taxono- whether they reflect (or may, or will reflect) my and nomenclature must be discussed. exactly the reality of evolution, or not. Ghiselin (1966: 127) stated that: “Definitions only apply to words, not to the things to which SOME BASIC CONCEPTS the words correspond”. However, the term OF BIOLOGICAL TAXONOMY “words” is not the end here, as a word refers to a AND NOMENCLATURE concept, i.e. a tool used by man’s mind to inter- pret reality. Although one does not define reality, I NTENSION AND EXTENSION one does define concepts, that are aiming at deci- Many of the discussions that have developed pher and understand reality. Thus, the related around biological taxonomy and nomenclature statement of De Queiroz & Gauthier (1990:
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308) that “definitions are given to words, includ- phies of its members that are considered to estab- ing to names, not to the things that the names lish the fact that this taxon corresponds to a clade represent, that is, the taxa themselves”, would of its own); and 2) it may be based on known or apply to the (unknown) real clades of the real hypothesized relationships between organisms. world, but not to taxa, even when based on clado- Among such relationships are those expressed in grams: all taxa are working hypotheses that are the topology of a cladogram, i.e. hypothetic cla- defined by science, not discovered. Taxa defined distic relationships between organisms, that are on the basis of cladograms or cladons are concepts, used in the so-called phylogenetic definitions of they are not the “real clades of the real world”. taxa as advocated by De Queiroz & Gauthier Cladons are hypotheses about clades, they are not (1992) and others. However these are not the the clades: “taxonomic definitions (or diagnoses, only kinds of relationships used to define taxa, if you prefer) are hypotheses about entities in although this has not been much emphasized nature. Those hypotheses are subject to test and until now: another useful kind of information is rejection or revision” (Schuh 2003: 70). provided by criteria like hybridizability (crossabi- Although rather seldom mentioned in taxonomy, lity), allopatry/parapatry/sympatry or competi- the terms intension and extension, of traditional tive exclusion, which qualify as “relational use in philosophy, logics and didactics (Dupuis taxonomic criteria” (Dubois 1988c) or more 1988: 87; Rey 1993: 461, 767, 1038), are quite briefly relacters (Dubois 2004c). Ultimately adequate to describe the two ways of qualifying however, even under the second kind of defini- or characterizing taxa in biological classification tion by relationships, we need to have informa- (Buck & Hull 1966). Any concept can be quali- tion on characters to allocate organisms to taxa, fied in two distinct ways: either by its intension or except if one believes that this can be done simply comprehension, that is “the set of all the properties by way of faith or intuition. or attributes that characterize this concept” On the other hand, the extension or circumscrip- (Papavero et al. 2001: 6); or by its extension or tion is the content of the taxon, i.e. a list of its ele- circumscription, that is “the set of all objects that ments or members (individual organisms and/or satisfy the [intensional] definition of that taxa), or, which is strictly equivalent, statements concept” (Papavero et al. 2001: 6). These terms about its boundaries or limits, or even a list of its fully apply to all classificatory systems, including non-members (all organisms or taxa excluded taxonomies: the units or classes recognized can be from it). For more clarity in the discussion below, qualified and identified either by intension or by the first kind of qualification of a taxon, by a list extension. of its included members, will be called inclusive In any taxonomy, the intension or intensional extension, whereas the second kind, by a list of definition of a taxon is a set of statements about excluded elements (that are therefore referred to its characteristics, or its supposed differences one or several other taxa), will be designated as from other taxa, or its relationships with the lat- exclusive extension. ter. This definition may rely on different kinds of Of course, in any consistent and rigorous classifi- information according to the philosophy of taxon- catory system, both kinds of qualification are omy adopted: 1) it may be based on characters, strictly equivalent, as a given intension corre- the traditional practice in biological taxonomy, sponds to a given extension, and vice versa. being then either a description (list of all known However, concretely, in many cases, and especial- “major” and “relevant” characters of the taxon, ly in biological classification, they may give whether proper to this taxon or common with different results simply because some pieces of other taxa), a diagnosis (list of all known charac- information concerning some organisms may be ters that are supposed to distinguish this taxon (and often are) lacking or inexact: whatever the from all other taxa) or an apognosis (Dubois criteria used to build up a biological classification 1997b) (list of known or supposed synapomor- (phenetic, cladistic, “phyletic” sensu Rasnitsyn
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1996, or other), the result may be misleading as “defined”. At any rate, if “typification” is under- long as some taxa remain undiscovered or as stood as a “definition”, this can be only a defini- some characters remain unknown or inaccurately tion of the nomen, not of the taxon: whether or ascertained. This is the reason why quite fre- not a given organism should be given a certain quently some organisms or taxa are mistakenly nomen depends not on the definition of the referred to a taxon although their actual, but nomen but on that of the taxon it is referred to. unknown, characters, in fact exclude them from All of this was already briefly and strikingly stated the latter, and subsequent discovery of these in the nice words of Luella Weresub quoted by problems is a major reason why taxonomies often Nicolson (1977: 569) and Jørgensen (2000: have to change and cannot remain “stable” for 779): “Taxa have circumscriptions but no types ever. while names have types but no circumscriptions”. Taxa, which are concepts, may be fully qualified or defined either by intension or by extension: T HE ZOOLOGICAL C ODE, ATHREE- STOREY they have both an intensional definition, and a NOMENCLATURAL SYSTEM content and limits. On the other hand nomina, Biological nomenclature is not a system of classi- which are labels and not concepts, cannot be fully fication but of nomination of taxa. Its rôle is not defined either by intension or by extension, as to define taxa, be it by extension or by intension, will be discussed below. The discussion of De but to name them, i.e. to provide labels that will Queiroz & Gauthier (1990) about “character- allow their universal and unambiguous designa- based definitions” of nomina is irrelevant and tion. As argued above, the current Code is not stems from a typological concept of “types” using “character-based definitions of taxon (Lidén & Oxelman 1996) and a basic confusion names” (De Queiroz & Gauthier 1990: 308, between taxonomy and nomenclature (Benton 309). The Code indeed mentions the need to pro- 2000; Stuessy 2000, 2001): in recent taxonomy vide “character-based” definitions of taxa, but the and nomenclature, such “intensional definitions” status of nomina is not given by these definitions may apply to taxa but not to nomina, as accord- or by these characters. Characters as used in the ing to the Code nomina are not defined by inten- Code are in no way useful either for the allocation sion (characters or relationships) or extension, of nomina to taxa or for the validity of nomina: but attached to given members of the taxa (individ- their rôle is “simply”, but importantly, to allow uals or taxa) by ostension (Keller et al. 2003: 99). availability of nomina. For a better understand- Such so-called “ostensive definitions” of nomina ing of these questions, these three concepts must (De Queiroz & Cantino, 2001) are more aptly be clearly separated, which is not clearly done in designated as a process of “christening” or “label- the current Code and therefore deserves a special ling” rather than “defining” nomina (Stuessy discussion. 2001; Kojima 2003): “In the Linnaean system The current nomenclatural system for zoological the taxon names are not defined, but they are taxa is like a “three-storey” house, with three labels for taxon concepts that are defined or cir- independent steps leading to the establishment of cumscribed based on the observation of charac- the valid nomen of a taxon, availability-alloca- ters” (Kojima 2003: 55). Blackwelder (1967: tion-validity (Av-Al-Va): 1) rules for the nomen- 166) used another useful comparison, stating that clatural availability of nomina; 2) rules for the a type “provides an anchor for the name, indicat- taxonomic allocation of nomina; and 3) rules for ing the point in the diversity to which the name the nomenclatural validity of nomina. Each of is forever attached”. Although “labelling by these steps makes use of special technical tools to ostension” could be very formally designated as a reach its aim. Let us consider these three steps definition through “partial extension” of the separately. taxon, this is not a complete definition of the lat- 1) Nomenclatural availability (Av) of nomina is ter, as it provides no boundary for the taxon so determined by several criteria, the most notewor-
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thy of which are criteria of publication, criteria of greatest precautions, it is difficult to completely formation of nomina and statement of diagnostic avoid mistakes in this respect: the history of zoo- characters. This step is the only one in the logical nomenclature is full of examples of taxa nomenclatural process where characters are men- that were first established on the basis of incom- tioned. Why? Of course, mention of characters plete or inaccurate descriptions or definitions, plays a rôle in the taxonomic definition of the but this in no way affects the availability, alloca- taxon to which the new nomen is applied. But, in tion or validity of the nomina that were then terms of nomenclature, its rôle is merely to avoid given to them. This question was discussed at the introduction into biological nomenclature of more length elsewhere (Dubois & Ohler 1997). nomina based on hypothetical concepts rather than 2) According to the Code currently in force, taxon- on actual organisms, or of nomina based on uni- omic allocation (Al) of nomina does not depend in dentified material or undefined taxa ( nomina the least on the definitions, descriptions, diag- nuda). The existence of nomina nuda implies that noses or apognoses of the specimens or of the taxa descriptions be attached to nomina but not that to which the nomina are applied, but on the com- nomina have “intensional or extensional conno- bined use of three tools unique to biological no- tations” (Moore 1998: 562). What the Code asks menclature, those of so-called “nomenclatural is simply that a definition or diagnosis based on types”, of so-called “nominal taxa” and of coordi- characters observed on actual specimens be given, nate ranks. In this nomenclatural system, a nomen but not that this definition be exact or complete. is not attached to a taxon or to its definition, but to This is a very clever disposition of the Code one or several individual organism(s). These speci- indeed, because no definition or diagnosis of a mens, usually known as “types”, or better as ono- taxon can ever be complete, and, even with the matophores (see below), establish an objective link
W o r ld o f language (z ool o g i c al no m enc lat u r e ) N F 1 R eal w o r ld o f na t u r al p o p ula t i o n s NG2 NG1 Z o ne o f o bje c t i v e c o nnec t i o n b e t w een r eal w o r ld and w o r ld o f language N S 2 N S 1 N S 3 T a x o m i na (no m i nal t a x a )
O n y m o p h o r o n t O 1 O 3 O 2 O t h e r s p e c i m ens P 2
P 3 O n y m o t o p i c p o p ula t i o n P 1 O t h e r p o p ula t i o n P 5 P 4
F IG. 1. — The rôle of onomatophores as an objective connection between the real world of organismal populations and the world of language (biological nomenclature). Abbreviations: NF1, nominal family 1; NG1, NG2, nominal genera 1 and 2; NS1- NS3, nominal species 1 to 3; P1- P5, natural populations 1 to 5; O1- O3, onomatophores 1 to 3 (modified from Dubois & Ohler 1997).
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between the real world of organisms and the world teria (in some special but rare cases, i.e. to protect of language (nomenclature) (Fig. 1; Dubois & “well known” nomina, other criteria like “usage” Ohler 1997: 304). This permanent link between a may be called upon). real specimen (or a series of specimens) and a For reasons already discussed elsewhere (e.g., nomen is what the Code calls a “nominal taxon”, a Dubois & Ohler 1997; Dubois 2000a, b), the concept which was retermed taxomen (Dubois major advantages of such an onomatophore- 2000b) to stress the fact that a taxomen is not a based nomenclatural system are that: 1) it allows taxon, as it has no limits or defined contents. It is a clear relationship between the world of organ- not equivalent to a definition of the taxon, as it isms and the world of language; and 2) it is not does not refer to any character or relationships that linked to any given philosophy of taxonomy, would allow to establish whether a given organ- which would be the case of an intension-based ism should be referred to the taxon or not. The nomenclatural system. This independence be- definition, limits and contents of the taxon may tween taxonomy and nomenclature respects the change without being followed by a change in the basic philosophical foundation of the Code, nomen, as long as the onomatophore remains allo- which is expressed as follows in its preamble: cated to it. In a way, it may be argued that the “The objects of the Code are to promote stability taxomen is qualified by its extension, i.e. either by and universality in the scientific names of animals its original contents or by subsequently restricted and to ensure that the name of each taxon is contents (e.g., after designation of a lectotype unique and distinct. All its provisions and recom- among syntypes or of a type species among several mendations are subservient to those ends and originally included species), but this extension none restricts the freedom of taxonomic thought only qualifies the taxomen, not the taxon to which or actions” (Anonymous 1999b: 2). The need for it may apply. The allocation of a nomen to a given absolute freedom of taxonomic thought and taxon in a given taxonomy does not rely on any action was recognized long ago, as examplified by kind of definition, but on the onomatophore of this statement of Strickland et al. (1843: 259): this nomen being referred to this taxon in this taxon- “So long as naturalists differ in the views which omy: all these elements are liable to change as taxo- they are disposed to take of the natural affinities nomy evolves, whereas the taxomen remains a of animals there will always be diversities of clas- stable and permanent link between the onomato- sification, and the only way to arrive at the true phore and the nomen. Finally, as will be discussed system of nature is to allow perfect liberty to sys- in more detail below, within the frame of a given tematists in this respect”. So the principle of a taxonomy the hierarchical structure of the taxon- large (although not complete, which would be omic and nomenclatural system allows the un- impossible) independence between taxonomy ambiguous allocation of a nomen to a taxon of a and nomenclature should be followed in any given rank whenever the onomatophore of a nomenclatural system, not only because it is the nomen is included in several hierarchically coordi- only one that respects the necessary freedom of nate taxa. thought, writing and action of all scientists – a 3) Establishment of the validity (Va) of a nomen major need for a harmonious and fruitful devel- is the third and last step of the nomenclatural opment of any scientific field –, but also, and process briefly described here. In many cases, perhaps mostly, because no one knows what will several nomina are available for a given taxon in a be the future of taxonomy. The philosophical given taxonomy and might potentially be used to bases of zoological taxonomy have dramatically designate it, which would be a source of confu- changed during the last decades, with several peri- sion and instability. Among these competing ods where different paradigms were subsequently nomina termed synonyms, the rule of priority “dominant”, and there is no reason to think that allows to establish automatically the valid one, we have now reached the ultimate stage in this usually without consideration of any other cri- respect. As discussed above, taxonomy might
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show basic developments and changes in the (M4) stability of the nomina given to “kinds of coming century, and it would be extremely risky organisms”; (M5) stability of the nomina given to and presumptuous to tie nomenclature to a par- individual biological specimens; (M6) “metaphys- ticular conception of taxonomy. As noted by ical stability” of nomina as defined by Nixon & Moore (1998: 573), “the current system has sup- Carpenter (2000), i.e. stability of formal defini- ported artificial, natural, evolutionary, phenetic, tion of cladons, irrespective of their contents (i.e. and cladistic classifications” and it should not “be of the organisms referred to them). assumed that there will not be another theoretical Stability of the kind (M5) is of course totally shift some time in the future”. The evolution of impossible. Even if taxonomy, nomenclature and concepts and criteria in taxonomy should remain nomenclatural rules were absolutely clear and free and should not affect or be affected by stable, there would remain numerous cases of ini- nomenclatural rules. tially (or subsequently) wrongly identified speci- mens, i.e. specimens that were allocated to a U SAGE AND STABILITY wrong taxon according to the taxonomic stand- Two terms have had growing use in discussions ards then in force. Their nomina will have to about taxonomy and nomenclature in the recent change whenever their wrong identification is decades: those of “usage” and of “stability”. corrected, and this may be viewed by some biolog- However these terms have rarely been seriously ists of disciplines far from systematics as cases of defined and discussed. The basic idea here is as “nomenclatural instability”: a famous example in follows: nomina (scientific names) are used as frogs was the discovery that specimens used in universal references by all biologists and by non- several laboraties and in various studies under the biologists to designate living organisms; in order nomen Xenopus muelleri did in fact belong in the to play this rôle unambiguously and universally, quite distinct, although related, species Xenopus they should be stable and they should never borealis (Bisbee et al. 1977). change, or change only exceptionally. As a conse- Item (M4) is the kind of stability that most biolog- quence, some authors (e.g., Cornelius 1987; ists have in mind when they support “nomencla- Gould 1990; Savage 1990a, b, 1991; Bock 1994) tural stability”: it means that a given specimen have recently advocated the replacement of the correctly identified (according to the taxonomy basic principle, or “fundamental maxim” then prevalent) as Hyla arborea, member of the (Strickland et al. 1843: 262) of the Code, the family H YLIDAE and of the order A NURA in 1882 Rule of Priority, by a so-called “rule of usage”. will bear the same nomina in 1960 and in 2005. Others have been even farther, suggesting to However, whether this will be the case or not can “stop taxonomists” and to set up “international be due to different causes as it will depend on the standing commissions” excluding taxonomists to three items (M1) to (M3). decide on stable “lists of official names” (Barnett Concerning (M1), as was stressed repeatedly 1989, 1991). This results in various problems, (e.g., Hershkovitz 1958; Fosberg 1991; Dubois among which several can be stressed. 1998; Schuh 2003), the first and major cause of 1) The poorly defined term “stability of nomina” instability of nomina is not related to nomencla- covers at least six distinct meanings (and proba- ture but to the evolution of the scientific discipline bly more), that are confused by many authors: of taxonomy: new taxa are continuously being (M1) genuine “taxonomic stability”, i.e. stability recognized, and existing taxa are continuously of taxa; (M2) genuine “nomenclatural stability”, being re-analysed and re-defined, i.e. modified. i.e. stability of the nomina designating the taxa In the example above, discovery that two distinct (in the absence of changes in the latter); (M3) “sibling” species were hidden under the nomen stability in the nomenclatural rules, i.e. stability Hyla arborea results in all specimens from a part of the criteria allowing the identification of the of the original range now having to bear another valid nomen of a taxon in a given taxonomy; nomen. The only way to avoid this kind of
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“instability” would be to stop taxonomic research for conservation and rejection can mitigate some (Dubois 1998: 22), i.e. indeed to “stop taxon- of these negative effects, it rarely makes sense to omists” (Barnett 1989). Changes in taxonomy try to amend the Code to deal with highly specific include changes in the intensional definitions of situations. This does not mean that the Code can- taxa and changes in the contents of taxa: both not be modified or improved in certain areas, but may be linked, but they may also be independ- it needs to be done with great care” (McNeill ent, as a taxon may remain composed of the same 2000: 715). Examples of unexpected side-effects organisms but with a different intensional defini- brought by changes in subsequent editions of the tion, and on the contrary the discovery of new zoological Code were pointed out for example by organisms (new species, genera, etc.) may modify Van Valen (1963) and Dubois (1985). Changes in largely the contents of a taxon but without modi- the Rules are particularly pernicious as non-system- fying its intension: so the concept of “taxonomic atists have no way to understand them, and this instability” itself is a complex and multi-dimen- kind of changes is likely to have played a rôle in the sional one. negative appraisal of biological nomenclature by As for the second source of instability of nomina, many biologists outside the field of systematics. (M2), i.e. changes of nomina for purely nomen- Avoiding these cases of “instability” that are par- clatural reasons (e.g., discovery of a senior syno- ticularly perturbating for communication between nym or homonym of a nomen), despite a widely systematists and other biologists and non-biolog- held belief, serious studies of concrete situations ists is the kind of problems that depend on no- (e.g., Hoł ynski 1994; Dubois 1998) show that it menclatural rules and that must be addressed by plays a minor rôle at the scale of a zoological any nomenclatural code: “it is not reasonable to group, much less important than item (M1). Its think of nomenclature as stable in an absolute rôle is bound to decrease as serious, professional sense. It is much preferable to think of the meth- taxonomic and nomenclatural revisions of groups ods of applying names as being stable. This are conducted, and to increase if taxonomy and conclusion presupposes, of course, that names nomenclature become more and more “endan- cannot be perfectly stable if the taxonomy to gered” within biological sciences, and are aban- which they are attached is not stable” (Schuh doned by professional zoologists to “amateurs” 2003: 64). and poorly trained technicians. As for “metaphysical stability” of nomina (M6), Finally, item (M3) could be easily avoided if the it will be discussed below. Code was never modified by the introduction of 2) In zoological taxonomy and nomenclature, new rules having a retroactive strength. “universal usage” is quite rare. In the most fre- Unfortunately such changes have regularly been quent situation, different usages exist in parallel, introduced by ICZN into the rules, resulting in in different countries, among different taxonomic unnecessary nomenclatural changes. They can schools, or between different authors. Therefore, often have unexpected side-effects that may be in these cases, “usage” cannot be unambiguously more disturbating than the positive effects of the determined. As soon as different usages coexist, change in the rules. Botanists, who have experienced apart from the Rule of Priority there is virtually this more than zoologists, as changes are intro- no indisputable way to decide which usage duced in each edition of their code, i.e. every six should be followed. Usage may change over time years, emphasize this problem: “Fortunately it is (at different epochs) and space (in different coun- now almost universally realised that any change in tries). Quantitative criteria are not philosophical- the provisions of the Code, no matter how small, is ly acceptable here: in science, as soon as different certain to have some nomenclatural implications opinions are expressed and supported, each one is that will not necessarily have been thought entitled to respect and examination on its own through by the proposers or even detailed in the merits, irrespective of the number of persons who Rapporteurs’ comments. Whereas the provisions share it. In the history of science, new theories or
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unexpected discoveries are not always accepted by “Although stability is often considered an impor- the whole scientific community at once and their tant quality of classifications, I believe that it is supporters may remain a small minority for often a spurious and misleading indication of the decades: fortunately, this does not imply that attainment of phylogenetic ‘truth’. All of our no- their opinion should be silenced. Even if quanti- tions about phylogeny are hypotheses that could tative criteria were to be used to define “predomi- be wrong; they can never be proved correct. If a nant usage”, it would be difficult to choose classification is to have wide-ranging biologic use- between the following “numbers”: number of fulness, it must be susceptible to change”. Shortly authors having used a nomen over a given period after (Gaffney 1979a: 103), he commented again of time; number of books, periodicals, languages, on this point and added: “In fact, temporal stabili- countries, where this nomen was used; number of ty of classifications often reflects ignorance of rela- distributed copies of a publication using a tionships and lack of work. I hardly advocate nomen, etc. (see Note 4). Qualitative criteria are change for its own sake, but the maintenance of no more acceptable, as they would result in stating names for discarded concepts seems useless and that some scientists, or some publications, or misleading”. More recently, Bremer et al. (1990) some countries, are more relevant, important or wrote in forceful words: “Taxonomists should worthy of attention than others in science. pursue their scientific venture and stop worrying Commenting the projects of “lists of protected about instability in classification. Taxonomy is names in current use” in botany, Cronquist not a service function for labelling organisms, but (1991: 309) very aptly wrote: “how often must a a science of its own, dealing with variation, rela- name be used to be in current use? It may be tionships and phylogeny. Other biologists need to fifty, or even a hundred years after a perfectly keep themselves informed, and should realize that valid species is described and published, before removal of artificial groups and improvements in anyone has occasion to refer to it in print again. classification are desirable”. The same authors (...) Will there be a crew of botanists available to aptly pointed out that “The problem with taxon- search the revisionary and monographic literature omic instability is not one of science, but of in- for Names in Current Use? If not, are such works formation”, and that “we need international to be cast aside as meaningless? Any List of database systems carrying continuously updated Names in Current Use is likely to contain anom- taxonomic information and providing easy access alies. The same species may have different names to synonymy and recent literature”. Unlike any re- in current use in different countries. Which ligious or other dogma, science, by its very nature, country will prevail, and will the losers accept constantly modifies its (provisional) conclusions, their loss? Different names on the List may be and this is the basic reason for its permanent mutually incompatible because they reflect progress. Requiring a “freezing” of science would different taxonomic outlooks. Will not many mean killing it. botanists soon be tempted to ignore the Lists Actually, the question of “stability” of taxa and because the Lists are taxonomically obtuse? I nomina makes sense only for non-scientists. think so, and probably I would be among them. Therefore, discussions on stability should be Instead of fostering stability, such Lists would be strictly restricted to the very few taxa and nomina likely to foster confusion”. (probably much less than 1/10000 of the total) 3) “Stability” is not a scientific aim (Dubois which happen to be well known and widely used 1998): “the goal of absolute nomenclatural sta- by non-taxonomists. Thus, any statement that bility is illusory and misguided” (Schuh 2003: “protection” of a nomen because of “usage” and 60). As Gaffney (1979a: 103), Dominguez & against priority should be considered is potentially Wheeler (1997) and Benton (2000) put it, in tax- acceptable only if based on evidence coming from onomy, “stability is ignorance”. More than outside the field of specialized publications deal- 25 years ago, Gaffney (1977: 23) already wrote: ing with taxonomy and related domains.
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4) Naming a taxon is a three-dimensional process by new nomina. Such criteria could be quantita- that involves the nomen of the taxon, the extension tive or qualitative. Quantitative criteria could be, of the taxon (its contents and limits) and the inten- for example, that whenever a taxon is split into sion of the taxon (its definition by characters or rela- two new taxa, its nomen remains attached to the tionships). Thus, “nomenclatural stability” taxon that contains the highest number of species, depends on the kind of link that exists between or that whenever two taxa are lumped together these three elements: it is a problem of relationships they take the nomen of the most speciose one. A between taxonomy and nomenclature. Two major very short reflection shows that any system of this kinds of relationships can exist in this respect: kind would very soon result in problems of inter- a) There is an absolute and definitive connection pretation and decision, and would create a great between nomen, extension and intension of the risk of progressive drift of any nomen from the taxon: this means that every time either the taxon for which it had been originally proposed to contents or the intensional definition of the other ones that in the end may have nothing to do taxon change(s), the nomen must be abandoned with this original concept. The solution to this and replaced by a new one. This kind of relation- problem is to use not quantitative but qualitative ship between taxonomy and nomenclature may criteria. Those used in the Code are simple, univer- seem more “logical” and is praised by some non- sal and unambiguous. They are the concept of biologists, in particular by some data processing onomatophore, the concept of taxomen, the and artificial intelligence specialists, as it allows concept of priority and the rule of coordination. unambiguous and definitive qualification of the The Code uses these rules only for the nomina of nomen. But this stability is in fact a source of the three nominal-series it recognizes. How such considerable instability, as taxonomy is always rules could be applied to the nomina of higher changing, and would require therefore a per- taxa is more problematic, and will need a special manent increase in the number of nomina. This discussion (Dubois in press). would not be only “cumbersome”, it would also The discussion above shows that a seemingly obscure the historical understanding of the simple formula like “taxonomic stability” or “no- taxonomy of any given taxon as given by its syno- menclatural stability” is a complex matter that nymy. This is the case today in the nomenclature needs careful study before peremptory statements of higher taxa, where no continuity exists between can be formulated. Whereas it may be fashionable many nomina that are introduced in the litera- and easy to publish that “Taxonomic instability ture as new taxa are recognized. continues to irritate” (Crisp & Fogg 1988), few b) The other solution is to have a certain dissocia- journals would agree to publish a paper stating tion between nomen, extension and intension of that the current neglect (not to say quasi-destruc- the taxon. There are two possibilities then: either tion) of taxonomy by the international scientific the nomen is linked to a certain content of the community is more than “irritating”, but a major taxon, so that the intension may change but the error in the scientific policy of our times (Wheeler nomen may remain the same; or the nomen is 2004). At any rate, “irritation” is not a solution to connected with a certain intension of the taxon, the many problems mentioned above, which need so that the content may change but the nomen to be solved by taxonomists, not by arrogant outsid- may remain the same. The first situation is that ers of the field. in the rules of the present Code, whereas the second one is that advocated e.g., by the support- O PENING THE P ANDORA’ SBOX: ers of the “Phylocode” (see below). THE RÔLE OF ICZN The need for nomenclatural robustness, as defined The consequences of the recent movement in above, requires to have criteria allowing nomina to favor of “usage” vs priority in zoological nomen- follow in a certain way the evolution of taxonomic clature are on the whole largely negative, mostly concepts without having constantly to be replaced because this “philosophy” results in strongly
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undermining the legislative value of the Code in the basis of priority alone, without interfering the eyes of all zoologists. with other subjective criteria like the opinions or As a matter of fact, some authors recently sug- tastes of authors, or judgements on “respectabili- gested a more or less complete abandonment of the ty” of usages or authors. Another consequence Rule of Priority as the major structuring principle has been that the editors of many periodicals of zoological nomenclature, stating that this rule (including “high-ranked” ones) and books now amounts to a “tyranny of the past” (Savage accept to publish papers containing major 1990a, b, 1991). They apparently did not realise nomenclatural mistakes (see list of examples in that their struggle against international Rules Dubois 2003a), thus continuing to undermine may eventually result in another, much more the power of the Code, with strong negative long- real, “tyranny”, that of “bureaucracy” (Fosberg term consequences regarding the unity of zoology 1991), or, even worse, of lobbies and partisan as a science (Dubois 1999b). groups having important financial resources, i.e. in the end, a tyranny of money on science. Thus Q UEIRAUTHIAN NOMENCLATURE doing, they opened a Pandora’s box and played a Rather than improving the current Code (in par- major rôle in the recent challenging of the legisla- ticular in extending its use to higher nomina) and tive function of the Code and in the development caring for its being more seriously applied in zoo- of proposals of alternative nomenclatural systems. logical publications, some recent authors, follow- Unfortunately, their analysis was partly shared by ing De Queiroz & Gauthier (1990, 1994), ICZN, which in the recent decades often adopt- suggested to abandon it and replace it by a brand ed a lax attitude and repeatedly accepted to su- new “ Phylocode” (see Note 6). I propose to call spend the Rules in order to “conserve” so-called such a nomenclatural system “Queirauthian”, a “usage”. It did so even in cases where this “usage” term combining parts of the patronyms of both concerned only a few “confidential” taxonomic its original proponents. Even if in the future the publications. In so validating gross recent “ Phylocode” turns out to be quite different from nomenclatural errors, especially when committed their original proposal, this is quite similar to the by “prominent zoologists”, ICZN has failed to situation regarding the so-called “Linnaean” play properly its rôle of “Keeper of the Law” and nomenclature, which is now very different from has contributed to spread among the internation- that originally advocated by Linnaeus (Moore al community of zoologists the feeling that the 2003). Strictly speaking, the Code currently in Code is not a system of clear and stringent rules, force is no less “Linnaean” than the “ Phylocode”, but an impressionist set of “recommendations” as most of its Rules were implemented much and “suggestions” that zoologists are more or less after the works of Linnaeus, whereas its basic free to follow or not (for more details on these logic is Aristotelian (see McNeill in Brummitt points, see e.g., Kiriakoff 1962; Dubois et al. 1997: 723). It is particularly misleading to state 1988; Ng 1991; Hoł ynski 1994; Dubois & e.g., that “the absence of a system of priority for Ohler 1997; Dubois 1999a, b, 2000a, b; and suprafamiliar [ sic, for suprafamilial] taxa” is a fea- Note 5). This recent tendency of ICZN to ignore ture of “Linnaean taxonomy” (Laurin 1998: 8), some basic rules of the Code, besides adding for two reasons at least: 1) no current taxonomy confusion and difficulties in the daily work of can be considered to be “Linnaean”, as, whatever taxonomists, has resulted in clearly weakening the philosophy of taxonomy adopted, not one the legislative value and strength of this text. In current author worldwide still relies on Linnaeus’ the eyes of many zoologists, the Code appears philosophy in this respect; and 2) the Code is not now as an obscure and obsolete text of little a system of taxonomy but a system of nomencla- consequence, rather than a set of stringent Rules ture, which is quite different (see Note 7). allowing automatic allocation of a nomen to a As a matter of fact, the Queirauthian proposals taxon within the frame of a given taxonomy, on are based in part on misunderstandings of the
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current Code and above all on a basic confusion term “ nominotypes” for such multiple onomato- between nomenclature and taxonomy (Dubois phores, but this proposal was not adopted by their 1999b, 2000a; Benton 2000; Stuessy 2000, fellow workers. In the proposed “ Phylocode”, the 2001). Although it is presented as a code of status of nomina in biological taxonomy would be nomenclature, the “ Phylocode” aims at playing in determined not by specimens but by “phylogene- fact a double rôle, that of a code of taxonomy and tic definitions” using “ specifiers”. Although speci- that of a code of nomenclature formally linked to fiers may be specimens or taxa, they may also be the former by rigorous rules. This led Stuessy characters (apomorphies, i.e. concepts), a reversal (2001) to rename it “ Phylosystem” instead of to the early days of taxonomy when “the method “ Phylocode” (see Note 8). The basic philosophical of concepts” or “conceptual types” was used in- foundation of this system is fully different from stead of “the method of types” or “nomenclatural that of the current Code. The many negative types” (Cook 1895, 1898a, b, 1900; Moore 1998: comments on this proposal in the recent years 564). Thus the “ Phylocode” is not a consistent (for references to several works that cover most of onomatophore-based nomenclatural system. In the question, see Wheeler 2004: 577) and the accepting that nomina can be qualified by charac- replies to these criticisms (e.g., Bryant & Cantino ters, this system is, at least partly, a purely circular 2002) largely remind a dialogue of the deaf, as system inside language, dealing only with abstrac- the same terms (e.g., “definition” or “reality”) are tions (intensions), whereas the major strength of often used with different meanings by the contra- the current Code is its connection with the world dictors. The purpose of the present discussion is of real animals through onomatophores. Any in- not to review all the problems posed by this proj- tension-based nomenclatural system is bound to ect, but some of its aspects only, that are directly be linked to a philosophy of taxonomy, which is related to the matter of higher nomenclature. not the case in an onomatophore-based system, in Four theoretical points will be tackled here: 1) which the reference for the allocation of nomina Querauthian nomenclature is not a consistent to taxa is not a conception of the taxa, but simply onomatophore-based nomenclatural system; 2) it the inclusion of one or several specimens or taxa in proposes “metaphysical stability” of nomina, but the latter. This is the case in the present Code for not genuine nomenclatural robustness; 3) it family-, genus- and species-series nomina, and I rejects the use of ranks for taxa; and 4) it does not propose to generalize this basic rule to class-series care for the original taxonomic allocation of nomina (see Note 9). Lidén & Oxelman (1996: nomina, with their original authorship and date, 183) were correct when they wrote: “Misunder- and rejects priority as the basis of validity of standing of the type method is probably behind nomina. Besides, and even perhaps more impor- the enthusiam for ‘phylogenetic taxonomy’ as tantly, this project poses considerable practical reflected in recent discussions on the world wide problems, that also need to be dully considered. web”. Abandonment of the onomatophore principle in Theoretical problems the Querauthian system requires to link nomen- 1) Contrary to the original statements of De clatural rules with a philosophy of taxonomy, Queiroz & Gauthier (1990, 1994), nomina recog- namely “phylogenetic taxonomy” or cladonomy, nized by the current Code are not based on inten- which infringes upon taxonomic freedom (Forey sional definitions of taxa, but on the (complete or 2002). It results in nomina defined within this sys- partial) original contents of the latter, and more tem having no existence within any other possible precisely on their original onomatophores, through system, so it will tend to act as a powerful tool to the use of the taxomen. Some supporters of the impede evolution of the discipline of taxonomy, by Querauthian system are in favour of using several trying to impose a “definitive taxonomic system” onomatophores to define nomina in this system. whatever the future discoveries and evolutions of Schander & Thollesson (1995: 266) proposed the concepts in biological sciences. Actually, this
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“freezing of ideas” may be, perhaps inconsciously the Code, this is simply not true: in both systems, in the minds of some, its major aim. nomina are unambiguously applied to a given 2) Proponents of the Querauthian system taxon in a given taxonomy, but any taxonomic contend that it allows a better stability of change entails a change in the nomina given to nomina, but this is due to confusion between the some organisms. In both systems, specimens of various senses of the term “stability of nomina” the genera Agama and Chamaeleo belong in two mentioned above. Discussing this requires first to distinct cladons (or taxa) under the taxonomy of remind a seemingly trivial fact, i.e. that the pur- Figure 2A and in the same cladon (or taxon) pose of biological nomenclature is to name organ- under the taxonomy of Figure 2B: as a result, in isms in order to be able to designate them in both cases a new cladogram may results in new scientific and non-scientific language. It is not to contents for the taxa designated by the same nomina, name concepts, be them hypothesized clades (cla- so that the same nomen ( A GAMIDAE or C HA- dons sensu Mayr 1995) or “germ lines” (see in MAELEONIDAE) applies to different individuals. In this respect: Echelle 1990a, b; Dubois 1991a: 70, Figure 2E, the nomen C HAMAELEONIDAE applies note 3). The Querauthian system provides stabi- in part to specimens to which it does not apply in lity in the nomenclatural designation of concepts Figure 2A, but the exactly same situation occurs (“clades”, a clade being defined as an ancestor in Figure 2F regarding the nomen A GAMIDAE! The and all of its descendants), a kind of stability only difference in this respect between the two aptly designated by Nixon & Carpenter (2000) systems is the way in which nomina are attached as “metaphysical stability”, but not stability in to taxa, either through allocation of onomato- the nomenclature of organisms, i.e. stability of the phores to taxa or through their inclusion in the (M4) kind above. The nomenclatural stability cladon defined as stemming from a most common provided by the Querauthian system is a priori ancestor, but in both cases there is no ambiguity stability by definitions, but such a system is not in the nomination of given individuals within a falsifiable: “Predictivity is not tested. Particular given taxonomy. Nixon & Carpenter (2000) dis- clades, once names (created), are always true: cussed other examples of similar situations. these clades henceforth always exist. This is trivi- According to the Code, a nomen potentially al: every higher level taxon name that has ever applies to all taxa including its onomatophore, been proposed is verified in this sense” (Nixon & and among several such nomina, the valid one is Carpenter 2000: 315). determined by priority: thus, the definition of the Let us consider the example of the nomina A GA- nomen is unambiguous, but it does not tell us MIDAE and C HAMAELEONIDAE discussed by De which organisms should bear this nomen, as this Queiroz & Gauthier (1990, 1994) (Fig. 2). is a matter of taxonomy. Similarly, in Under the Code, allocation of these nomina to Querauthian nomenclature, a nomen applies familial taxa is established by the simple taxon- unambiguously to a defined cladon, but this does omic allocation of the “type genera” of these not tell us which organisms are members of this nomina to the taxa, irrespective of the criteria cladon: this is to be established through a cladis- used to define these taxa, and irrespective of the tic analysis resulting in a cladistic hypothesis. In place of these genera in a cladogram. In contrast, both cases, the “definition” of the nomen in the Querauthian system, nomina are associated remains the same whatever the taxonomy adopt- with “phylogenetic definitions”, so that a nomen ed, but any change in cladistic hypotheses may will designate the clade including all taxa result in taxonomic and therefore nomenclatural stemming from the most common recent ancestor changes. In the Querauthian system, just like in of all taxa included in the original taxon designated the Code, total nomenclatural stability would by this nomen. Despite the claim of De Queiroz exist only if taxonomy was final and stable, but as & Gauthier (1990, 1994) that their system long as taxonomies will evolve, nomina given to provides a better “nomenclatural stability” than individual organisms will have to change.
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A B A gamidae C h amae l e onidae A gamidae C h amae l e onidae
C D L e i ole p ididae = A gamidae = C h amae l e onidae L e i ole p ididae A gamidae C h amae l e onidae
E F C h amae l e onidae A gamidae C h amae l e onidae
F IG. 2. — The “classical” example of the nomina A GAMIDAE and C HAMAELEONIDAE discussed by De Queiroz & Gauthier (1990, 1994). These nomina were first proposed for two families, i.e. implicitly two clades, the former including two groups or subfamilies ( A GAMINAE and L EIOLEPIDINAE) ( A ). Under a revised hypothesis of relationships, the A GAMIDAE appear as paraphyletic relative to the C HAMAELEONIDAE ( B ). In any phylogenetic taxonomic system, paraphyletic groups cannot be recognized as taxa, which requires a new taxonomy, either with three taxa ( C ) or with a single taxon ( D ). According to the Code, there is no ambiguity for naming the taxa under both taxonomies, using the taxonomic allocation of the “type genera” of the familial nomina, the taxonomic hierarchy and pri- ority among nomina: if three families are recognized, they will be called A GAMIDAE, C HAMAELEONIDAE and L EIOLEPIDIDAE ( C ), but if the three groups are placed in a single family, these three nomina are synonymous ( D ), and the nomen C HAMAELEONIDAE has priority ( E ). In the Querauthian system, nomina are associated with “phylogenetic definitions”, and the nomen A GAMIDAE will designate the clade including all taxa stemming from the most common recent ancestor of all taxa included in the original A GAMIDAE ( A ), so that it will apply to the group including both the latter and the original C HAMAELEONIDAE ( F ).
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This is in fact fully recognized by supporters of the An additional problem regarding stability is posed Querauthian system, such as Bryant (1996: 174): by the fact that “specifiers” need not only be mem- “The clade referred to by the phylogenetic defini- bers of the taxon designated by the nomen they tion of a taxon name depends on 1) the reference qualify (as is the case for onomatophores), but phylogeny, 2) the meanings of taxon names refer- have also to be hypothesized ancestors of the cladon red to in the definition, and 3) whether the name defined by the nomen: therefore their own ances- is defined as a node-, stem-, or apomorphy-based tors or sister-taxa cannot be included in the taxon. taxon”. Therefore the meaning of a nomen in this If the specifier is not well chosen, the cladon so de- system depends on a reference phylogeny, i.e. the fined may exclude some taxa that have always be same nomen has different meanings under differ- included in the taxon designated by this nomen, as ent taxonomic arrangements. This is exactly simi- remarked by Lidén et al. (1997: 737): “If by mis- lar to the Code system, except that the latter is take or ignorance (and we are ignorant, that is why more flexible as it does not impose a kind of defini- we pursue our research) an unhappy typification is tion for taxa, which will make it compatible with made, we might – as fuller knowledge is gained – any future taxonomic systems that may develop. end up with having Aves (birds, traditionally a ‘hi- De Queiroz (1997: 139) acknowledged this when gher’ taxon) subordinate to Struthioniformes (os- he wrote: “neither phylogenetic nor Linnaean sys- triches, traditionally a ‘lower’ taxon)”. Given the tems guarantee clarity, universality and stability in high lability of many of our current cladistic hyp- terms of hypotheses about the relationships and otheses in the whole animal kingdom (illustrated composition of taxa”. by the permanent publication of new phylogenetic To put the same things differently, the interpretations in many zoological groups), there is Querauthian system defines concepts, but does not no need to be a soothsayer to predict that this provide means to apply these concepts to real or- situation is likely to appear very frequently. Of ganisms. To be sure, the concept “clade composed course, it will then be possible to emend the of an ancestral species A and all its descendants” is definition of ill-defined nomina (Bryant 1996: fully unambiguous, but whether any given speci- 186), but this will be a heavy procedure and there men belongs or not in this clade is and will remain is no advantage in terms of stability of this new ambiguous. The traditional way of attaching system over the traditional one. nomina to taxa (through inclusion of onomato- 3) Supporters of the Queirauthian system claim phores in taxa) has more generality and practicabi- that taxonomic ranks should be abandoned as they lity than the new proposed criterion (through are subjective and arbitrary, and do not warrant ancestor-descendant relationships). All organisms comparisons between taxa of same rank in differ- can be allocated to taxa. In some cases this can be ent groups. The latter is indeed true (families and done through cladistic analysis, but in many orders of birds are not equivalent by any criterion others this is impossible, either for practical rea- to families and orders of amphibians: see Dubois sons, e.g., in the many groups whose majority of 1988a, c), but the same applies to many other species are still undescribed and that first need scientific concepts and tools, which does not im- basic descriptive and monographic revision works pede them from being useful. The hierarchy ex- (Berry 2002; Janovec et al. 2003), or for more pressed by taxonomic-nomenclatural ranks basic reasons, e.g., in cases of reticulate evolution certainly does not “exist in nature”. “But what or of “sexual parasitism” mentioned above. In the does it mean to ask if hierarchies really exist in na- latter cases, individual organisms cannot be refer- ture? A hierarchy is not a natural object. It is an or- red to cladons. To attach nomina to taxa, the cri- ganizational model, and as such, it is a human terion of inclusion of onomatophore in taxon has conceptual construct. This organizational model therefore more generality than that of membership of relationship, like others (e.g., networks, causal in a hypothetic clade and is to be preferred on that chains, vicious cycles, etc.) is an abstraction, but account. that does not make it any more imaginary than
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numbers and mathematical relationships” (Knox most actual taxonomies, ranks afforded to taxa 1998: 3). “Hierarchies are organizational models are not given automatically by any criterion but that are useful for keeping track of inter-level rela- are chosen through a combination of criteria, tionships among entities” (Knox 1998: 42). The including robustness of cladistic hypotheses (as fact that nomenclatural rank has been used by long as a strong uncertainty remains regarding some biologists to make unwarranted compar- the relationships between clades, it is better to isons as if taxa at the same hierarchical rank were maintain the latter in the same taxon, in order to somehow equivalent is not a good reason for re- avoid frequent taxonomic and nomenclatural jecting ranks: “Even if some biologists misuse ranks, back-and-forth changes), degree of divergence, I do not think that the elimination of ranks is war- and taxonomic homogeneity within a given ranted. Just because some doctors misprescribe a higher taxon. This is certainly not satisfactory particular drug, should other doctors be banned from a purely theoretical point of view, but it from prescribing it? Certainly not” (Moore 2003: allows taxonomy to play its major rôle of an 13). Just like in the case, discussed below, of the indexation system for information, a rôle that a misunderstandings surrounding the use of the non-hierarchical system cannot play: “The term “type” in biological nomenclature, the prob- context in which biological classifications must lem at stake here is mostly related to the current function is a hierarchical one. In this sense the low level of training and knowledge of most biolog- Linnaean system has served biology effectively for ists regarding the basic concepts, methods and nearly 250 years. Although some workers may achievements of taxonomy and nomenclature, a have misconstrued the use of the ‘family’ category problem that should seriously be addressed but to mean that placement of a taxon at that level which has little to do with the function of ranks in conveys some particular (essential) properties, taxonomy. most investigators using the system in this day As reminded above, the major function of taxon- and age are well aware that ‘family’ and all other omic ranks is not to provide biological or histori- categorical ranks are used to denote a scheme of cal information on taxa, but to provide a useful nested hierarchical relationships, not to convey organisation of the information offered by the information on some additional aspect of reality taxonomy, according to the basic criteria used over and above the characters by which the taxa to build the latter. Taxonomic-nomenclatural are diagnosed” (Schuh 2003: 76). From a practi- hierarchies are “just” organizational models of cal point of view, this hierarchy allows biologists taxonomic information, not “theories of relation- of various disciplines to make useful compari- ships” or other theories. Thus, in purely phenetic sons, provided certain precautions are taken: taxonomies, the information provided by ranks is ranks provide “a hierarchical series of partitions mostly related to the “degree of divergence” be- of biodiversity, however imperfect that might be. tween taxa, whereas in phylogenetic taxonomies, An ecologist can work with the generic and fami- hierarchical ranks provide information on the ly diversity of insects in a field site, and compare structure of the cladogram, and in evolutionary this with other field sites. He or she knows that or phyletic taxonomies it provides information genera or family distinctions are arbitrary, but both on cladistic relationships and on “subse- this matters less in comparative studies using the quent modification” (Knox 1998). But, more same taxa. (...) Were taxonomists to retreat from importantly, in both cases, the existence of this ranked taxa, they would be re-invented by hierarchy, that acts as an indented index, facili- ecologists, naturalists and horticulturalist, and we tates the research of the information relative to believe that it would be better done by taxon- any taxon in the system, which would be much omists” (Godfray & Knapp 2004: 561). more difficult and long if the nomina of all taxa As long as cladograms will evolve, no final taxon- were given as a mere list, just like in an non- omy, and therefore no stable nomenclature, will indented, non-hierarchic alphabetical index. In exist, and the stability in the definition of nomina
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provided by the Querauthian system will not tem. This drastic difference should be made clear result in stability in the nomina given to individ- to all zoologists and other users through use of ual organisms. Even if a final “tree of life” was different “kinds” of nomina, e.g., including spe- ever to be agreed upon by all biologists, a final cial characteristics making them easily recognis- taxonomic and nomenclatural stability would able, as is the case in some other recently require that all dichotomies in this tree be taxon- proposed nomenclatural systems (Kluge 2000; omically and nomenclaturally recognized. It is Papavero et al. 2001). true that, with this “final cladogram” and with Furthermore, the tendency to consider author- “phylogenetic definitions” of nomina, ranks ship of a nomen as a kind of “ownership” and as would not be necessary to have an unambiguous more important than contributing to the know- allocation of nomina to individual organisms, but ledge of organisms, once aptly called mihilism such a system using a “cornucopia of categories” (Bruun 1950), has never completely disappeared (Colless 1977) would be so cumbersome and from zootaxonomy, but fight for the “ Phylocode” unpalatable that it would be as uninformative as may give a new vigour to this attitude. A possible mere chaos, as up to n -1 supraspecific taxa might way of reducing its importance would be to be required to express taxonomically the cladistic remove the names of the authors of nomina, but relationships of all n species that ever existed not their dates, from the nominal-complexes that (Szalay 1977: 363; Dupuis 1979: 45). This is designate zoological taxa (for detailed proposals why the “ Phylocode” (Note 2.1.2) expressly states in this respect, see Dubois 2000b: 62-65). As the that “it is not necessary that all clades be named”. “ Phylocode” is a new system, its proponents could Dupuis (1979: 46) already wrote in this respect: well have decided not to cite the authors of nomina “Rien n’interdit en taxinomie cladiste ou autre de in the nominal-complexes and to adopt such an ne nommer, par convention, que les taxa opéra- “anonymous” form (but including date) for tionnellement utiles”. How can these “operation- nomina of cladons, but they did not. According ally useful taxa” be recognized remains a mystery. to the “ Phylocode” supporters, the new meanings Lee (1996: 188) worded the same idea differently: given to the same nomina will be acknowledged “Phylogenetic taxonomists are free to name only by crediting these nomina with new authors and those clades they feel are important and robust dates, corresponding to the first publications enough to warrant formal recognition”. But then, where these nomina were used with Querauthian if it depends on the “feelings” of Querauthian definitions. Such a practice is strikingly shocking cladonomists, the decision regarding which for any honest taxonomist. Papers are now being clades should be named is as arbitrary as in all published in which it is seriously discussed, e.g., other taxonomic and nomenclatural systems whether the nomen T ETRAPODA should be cred- (Colless 1977; Forey 2001: 90). ited to Gaffney (1979b) or to Gauthier et al. In conclusion, the system of ranks as used in the (1989) (Laurin 1998, 2002; Lee 1999; Anderson Code provides a convenient and workable system 2002; Laurin & Anderson 2004), although this of indexation of taxonomic information and it nomen was first used in zoological nomenclature would simply be destructive and “masochist” for by Fischer (1808)! Such an attitude certainly does zoologists to abandon it. not qualify as “courtesy” (Corliss 1972). For 4) In the project of “ Phylocode”, “the names of those who have a certain experience of study of clades may be established through conversion of early taxonomic works, this very much looks like preexisting names or introduction of new names” a return to “prehistorical” times of zoological (Rule 9.1). This heterogeneous rule is liable to nomenclature: any author who would propose a cause considerable confusion in the future. The new definition or content for a taxon would then basic philosophy of Querauthian nomenclature is feel entitled to use for it an existing nomen, but incompatible with that of the Code, and cannot to credit himself with authorship of the nomen be considered as a modification of the latter sys- (in a new, “emended”, sense), and to write “ mihi”
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or “ nobis” after the nomen, instead of the name for “protection” on the basis of “usage”. If of the real author of that nomen. However, a Querauthian nomina are not clearly distin- nomen is not a taxon, and it is a common (and guished from nomina ruled by the Code, problems fully valid) practice in zoological nomenclature to of homonymy between nomina following either change either the definition or the content of a rules will rapidly become a considerable source of taxon, but not its nomen and the author of the confusion and error (Cantino 2000: 87; latter (see below for more details). Once available Blackwell 2002: 152). As Stevens (2002: 22) under the current Code, a nomen keeps its author remarked: “If different groups of people apply and date whatever the vicissitudes of the taxon- the same name to different groups of organisms, omic concept designated by this nomen, and the or different names to the same group of organ- rule of priority relies on the date of the original isms, it will be decidedly unsetting for society and publication of the nomen, not of any “redefini- perhaps damaging for our discipline”. Greuter tion” of the nomen. The status of the nomen (2004: 24) put the same idea even more bluntly: regarding its allocation to a taxon within the “the PhyloCode operation might still be judged frame of any given taxonomy is established once dispassionately if it had the decency to leave the and for all either at the creation of the nomen or current system of nomenclature alone; if it opted through a first-reviser action whenever necessary, for using its own, independent set of names, and cannot be changed by subsequent authors. perhaps in English if not in Esperanto, but any- This is just to render unto Caesar the things that way clearly distinct from the current scientific are Caesar’s: nomenclature is not taxonomy, and names of plants, animals and bacteria. The the author of a nomen is not necessarily the disturbing fact is that the PhyloCode chose to author of the concept of the taxon designated parasitise the extant system of names. In conse- today by this nomen. The idea of “borrowing” quence, its adoption would inevitably lead to the nomina from earlier authors and “converting” or dual use of the same name for distinct and often “redefining” them is not acceptable. In the fu- different concepts, thus undermining the princi- ture, clarification will be necessary: although pal function of biological nomenclature: to give “freedom of taxonomic thought or actions” no access to biological information with a minimum doubt allows any taxonomist to follow a of ambiguity. If, by misfortune, phylogenetic Querauthian philosophy of taxonomy and nomenclature should prove as successful as its nomenclature, editors of scientific periodicals promoters hope, this would result in the eventual and books should not accept to publish works in collapse of the whole edifice of organismic which nomina established under the Code would nomenclature – without anything available to be used under the “ Phylocode” rules. For use replace it”. An excellent example of the kind of under the latter rules, new nomina should be “parasitism” mentioned in this quotation is pro- created, and the fact that these nomina are vided by the recent paper by Hillis & Wilcox defined in a Querauthian way should be made (2005). It is therefore urgent that the internation- clear to any reader by special marks. al community of taxonomists, through the edito- Without such a provision, “converting” ancient rial rules and the action of the editorial boards of nomina to the new system, may be a “strategic” scientific periodicals and books, impose on way to introduce progressively and “surrepti- Querauthian taxonomists the obligation to use tiously” Querauthian nomenclature in zoological special nomina that cannot be mistaken for scien- publications: the new nomenclature looking simi- tific nomina as recognized by the Code. If this is lar to the traditional one, many zoologists may viewed by some as a restriction of freedom of not realise that such a drastic change has been taxonomic thought and action, this should be brought in the way nomina are defined and used, compared with the following statement by and these nomina may become progressively Greuter & Nicolson (1993: 927): “For Taxon we “entrenched in zoological literature”, thus eligible will now certainly not accept any more papers
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contrary to these principles, but we will instead could follow directly the nomen, e.g., urge authors to submit proposals as appropriate”.
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status of nomina cannot be based on their histo- bility of nomenclature”. It should no doubt be ry, on subsequent “redefinitions” or on “prevail- improved, but this system has shown consider- ing” or “dominant” usage: “The current usage of able flexibility and it can be used with any kind the term T ETRAPODA (or the vernacular term of taxonomic systems, including cladistic or tetrapod) is sufficiently variable to leave the ques- “phylogenetic” ones. But, of course, for this, like tion of the prevailing usage of this name unre- for any human tool, its good use requires a good solved, especially if we consider both intended knowledge and understanding of its characteris- and literal usage” (Laurin in Laurin & Anderson tics. Since its adoption by the international com- 2004: 77). The status of nomina must be based munity of zoologists, the Code has had a major only on information provided in the original impact on publications in zoological systematics. publication where they were coined, irrespective Serious, consistent and permanent enforcement of subsequent avatars of the use of these nomina. of the principles of onomatophore and of priority This is the case already in zoology for all nomina by thousands of zoologists working on all animal regulated by the Code, and my proposals (Dubois groups has allowed to establish the proper status in press) are meant at extending this practice also of millions of nomina proposed since 1758 for to class-series nomina. Once its status is estab- animal taxa. The result has been a progressive sta- lished through such non-ambiguous rules, alloca- bilization of nomenclature of the vast majority of tion of a nomen to a taxon in the frame of a given animal groups. The proportion of nomina that taxonomy is an automatic procedure which leaves remain unallocated is very small indeed com- no place to personal feelings, preferences, majori- pared to the number of nomina the status of which ty, power or authoritarism. This is the kind of is now clarified and stabilized. The Principle of nomenclatural rules zoological taxonomy needs. Priority has proved in fact to be a very powerful, The four points mentioned above are not the honest and automatic rule for the allocation of only ones that raise theoretical problems in the nomina to taxa and the Code based on this rule “ Phylocode” project, but may be the most impor- has proved so efficient that one may wonder tant theoretical flaws of this new system, especial- “how to improve the excellent” (Hoł ynski 1994). ly regarding higher nomenclature. However, As Benton (2000: 647) ironically wrote, “if it beside these basic philosophical problems, proper ain’t broke, don’t fix it”. Its replacement by com- caution should be paid to more pragmatic, but pletely new rules would no doubt be followed by not trivial, problems. considerable confusion and loss of information, and cause many more new problems than those it Practical problems would be supposed to solve (see e.g., Dubois The current rules were not born at once, but are 2000a, b). As mentioned in the preamble of the the results of a process that spread over several Code, the latter is not meant to interfere with hundred years (Melville 1995) and which in- taxonomic theories, concepts and practices, but volved thousands of zoologists worldwide. This simply to give automatic rules for the unambig- Code is not an abstract set of “ideal” rules that uous allocation of a single nomen to a taxon would have resulted from a purely “theoretical” within the frame of a given taxonomy. This is all approach to zoological nomenclature, but the and this should remain the only function of any result of a concrete confrontation, through a nomenclatural code. “trial-and-error process”, of such possible “ideal” A practical but crucial question must be addressed rules with the daily work of taxonomists. The here. We are not living in an abstract world of Code is of course not perfect (is any juridical text concepts and theories, but in a material and social perfect?), but its bases are quite sound (Ng 1991; world which imposes a certain number of Hoł ynski 1994; Dubois & Ohler 1997; Dubois contraints upon us. One of them is the highly in- 2000a, b) and it is indeed an excellent tool to sufficient number of taxonomists worldwide and achieve its aim, namely “the universality and sta- the financial and other limitations for their work
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over the whole planet. The taxonomic impediment ther contribute to the taxonomic impediment by is so strong, both quantitatively and qualitatively, diverting funds and energies from the crucial work that it is unlikely to be significantly reduced dur- of taxonomic inventory of the planet. ing the coming century. Before being able to Furthermore, “in this urgent climate of seeking to build a reasonably reliable “Tree of Life”, we need inventory the world’s biota (...), and requesting first a much more complete “Catalog of Life” on funds from the rest of society to do so, it would be our planet than is currently available (Berry highly counterproductive to simultaneously re- 2002). This would only be “sad and disappoint- commend whole-scale change of names of organ- ing” if “we had time” and if we could think that isms for any reason” (Stuessy 1997: 117). The this impediment will be progressively reduced in consequences of such a move could be deadly for the next centuries, but the biodiversity crisis that is the whole field of taxonomic research: “The whole- currently developing on earth, and that certainly sale abandonment of the Linnaean naming system qualifies our times as the century of extinctions and its replacement by a new and untried method (Dubois 2003a), throws a different light on these would destroy the support base for the field and questions (Wilson 1985, 2000). The work of ex- imperil its survival” (Godfray & Knapp 2004: ploration of the biodiversity of our planet that will 562) (see Note 10). not have been done in the coming decades will be By and large, the international scientific commu- impossible to carry out later, because many species nity has not yet recognized the urgency and pri- will have disappeared. This particular situation ority of accelerating collection and inventory of imposes on all taxonomists worldwide the need to biodiversity before it is too late, and few voices are realize that we are not living in the same world as loud enough to draw efficiently attention on the one century ago and that strong acceleration of the current dramatic situation: despite nice words taxonomic exploration of our planet is an urgent (e.g., Wheeler et al. 2004), few concrete decisions need and a scientific priority that requires a major (in terms of large-scale research programs, alloca- re-examination of our research programs (May tion of budgets and working positions) are yet to 2004; Wheeler 2004). This was forcefully stated be seen in most countries. But at the time where by a few biologists in the recent years, e.g.: “There thousands of species are getting extinct under our is really no excuse for perpetuating our level of eyes, it would be completely foolish and irrespon- ignorance about nematodes, fungi or mites and at sible for the community of taxonomists, or a signi- the same time [having] no hope, in my opinion, of ficant part of it, to decide to devote a huge completing world catalogues before many of them amount of effort, working time and funds to rede- disappear” (Raven 2004: 730). “Funds nominally fining the nomina of thousands of taxa in order to allocated to taxonomy go largely to reconstruct have a “better” nomenclatural system rather than molecular phylogenies, while thousands of species working on the discovery and study of the mil- ate threatened by imminent extinction. (...) In the lions of other taxa that we still never collected. As face of the biodiversity crisis, the need for urgency argued above, the current paradigm of “phyloge- could be no greater. (...) Our generation is the first netic taxonomy” is unlikely to be the “last word to fully comprehend the threat of the biodiversity forever” in this respect: for example if the LGT crisis and the last with the opportunity to explore problem mentioned above came out as a major and document the species diversity of our planet. problem, taxonomists might have to develop a Time is rapidly running out. (...) The grand biolog- new paradigm of biological classification. As for ical challenge of our age is to create a legacy of today, even if it were true that Querauthian knowledge for a planet that is soon to be biologic- nomenclature is “better” (a statement that many ally decimated. To meet it, 21st-century taxon- taxonomists are not ready to endorse), “con- omists and museums must have the right tools” verting” the information stored in the existing (Wheeler et al. 2004). Clearly, implementing the nomenclatural system into the new system would “ Phylocode” cannot be one of these tools, but ra- be a much less urgent task than studying the still
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unknown organisms of our planet. In an abstract should not be supported by zoologists, despite the world with unlimited human and financial “favorable fanfare in the popular media” resources, it might be considered “theoretically (Carpenter 2003: 79) that has been organized better” to replace the nomenclatural system by a around this project: 1) transferring all the taxon- “better one” at each step of the history of compara- omic information currently stored under the no- tive biological sciences, i.e. every second or third menclatural system of the Code to another system decade, but this is not true in the real world. would not only be very time-, energy- and re- According to Wilson (2004), there are at present sources-consuming, at the expense of other more about 6000 working taxonomists worldwide, i.e. useful taxonomic works, but would also undoubt- “a tiny slice of the biological community as a edly cause losses of information or introduce mis- whole”, and their discipline is “one of the weakest takes and confusions: the current nomenclatural and most underfounded”. In the last decade, the system is not perfect, which is impossible, but it many discussions, meetings and papers devoted to has been working for two and a half centuries, and the conception and promotion of Querauthian continuity of this system should be maintained (of taxonomy, as well as to its rebuttal, have already course, this system should be improved whenever diverted thousands of working hours (i.e. collec- needed and possible); 2) the “ Phylocode” is not a tively, months or years) of competent taxonomists, consistent onomatophore-based nomenclatural who would have been better inspired to use this system, i.e. it is a system that works partly only precious time (provided by their salaries) and their within language; in contrast, through onomat- energy to collect, study and describe the biodiver- ophores, the current Code maintains a permanent sity of our planet that is getting extinct at an accel- material connection between the world of lan- erated rate in our indifference. It is more than guage and the real world of specimens, and is time to stop this “pure folly” (Carpenter 2003): highly to be preferred on that account; and 3) the “The sooner the Phylo Code is buried the better “ Phylocode” is both a code of nomenclature and a for biology” (Greuter 2004: 24). For this to hap- code of taxonomy, and as such it infringes upon pen however, it may not be enough to witness taxonomic freedom, which is a major weakness of “indifference from biological systematists in gen- this system as compared to the current Code (see eral” to these proposals (Lidén et al. 1997) and to Note 11). For all these reasons, the proposals “boycott meetings” (Wheeler 2004: 577) but “we made below regarding the nomenclature of higher need community involvement” against “a small taxa are not based on a Querauthian philosophy of group of people with very significant resources, nomenclature, but on the same theoretical prem- and thus influence”: “If the community does not isses as those of the current Code for the nomina of become actively involved in fighting the lower taxa. PhyloCode, they will succeed, and in so doing, demolish much of the hard work that our prede- T HE CONCEPT OF “ AMBIGUITY” AND THE NEED cessors have built” (Nixon et al. 2003: 119). It FOR STABILITY OF NOMINA DURING would be irresponsible for the majority of taxon- THE EVOLUTION OF TAXONOMY omists to decide to embark on the “ Phylocode” One of the reasons given by supporters of the project in the coming years, and future genera- Querauthian nomenclature for their proposed tions, if they happen to be more clever than ours, changes is that the new system would allow may pass a severe judgement on such a choice if it unambiguous and definitive allocation of a nomen was to be done (May 1988, 2004; Dubois 2003a). to a taxon, whatever the changes in taxonomies. The question may therefore be asked whether it is Conclusion acceptable to keep the same nomen for a taxon To summarize the statements made above, three when the definition and content of the taxon major problems are posed by the project of have changed, which is the case under the current “ Phylocode”, which suggests that this project Code. Under a very rigid conception of nomen-
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clature, this would be unacceptable and any Linnaeus (1758): although indeed different, this change in the taxon’s definition and content concept is connected to the original one through would require introduction of a new nomen. the history of the nomen, and through the per- Such a request may sound appealing in particular manence of the (real or virtual) onomatophore to people having an approach to zoological on which this nomen was based. nomenclature through computerized data bases More attention must be given to the meaning of and data processing: a given nomen would cor- the term “ambiguity”. This term may be defined respond only to a fixed definition and content of as “expression capable of more than one mean- a taxon, and, as soon as one or both of these char- ing” (Fowler & Fowler 1929: 35), i.e. as liability acteristics change, the nomen should change. to be understood in more than one way. However, this is not what biologists and laymen However, there are two kinds of unambiguity, need. Biological nomenclature is not a “perfect that can be designated as absolute and relative system” designed only for taxonomists, but has unambiguity. In any classificatory system, i.e. a the function to be useful to all biologists and even system that sorts elements or units into groups or to society as a whole, far beyond biologists. The classes, a fully unambiguous system would be one usefulness of biological nomenclature relies on its that would define a class so that not only its being a simple and handy system, not only for intension but also its extension is definitive and naming the taxa (an abbreviated way of referring not liable to be modified: in other words, it to them, rather than by full description of all the would be a classificatory system based on closed data we have obtained about them), but also for definitions. In taxonomy, the discipline whose tracing the taxonomic and other biological infor- aim is to sort all living organisms into classes mation about these taxa that may have accumu- called taxa, it would be a system in which any lated through the history of biology. What is taxon would be defined as including a closed and important is to be able to find rapidly, through definitive list of organisms, any addition or the nomen, or through a list of nomina and refer- subtraction of any single organism resulting in ences in a logonymy (see Dubois 2000b), all or the definition of a new, distinct taxon. But this is most of the references to a given taxon and its not the case in any real taxonomy, whatever the close relatives, and thus to obtain, for those who philosophy of taxonomy adopted. The definition wish to, not only information on what is current- of the taxon may be unambiguous, but the ly understood under this nomen, but also on the content of the taxon as recognized by taxon- history of the taxon’s concept. Even if today what omists will never be so until we have reached is understood under the nomen Rana esculenta is the End of Research: “The meaning of a word has quite different from what was understood under nothing whatever to do with the practical prob- the same nomen in 1758, 1897 or 1960 (see lem of identifying the things which may happen Dubois 1998), the continuity of use of the latter to fit the definition” (Ghiselin 1966: 127). nomen through centuries allows any interested Let us take an example outside taxonomy to person, even if completely unaware of the history make this clearer. Let’s consider a human society of the taxonomy of this group of frogs, to find in which the law would state that any child must rapidly (e.g., in a logonymy or by use of a data- bear the patronym of his/her biological father. base like the Zoological Record) all major references This is a fully unambiguous definition of classes, to understand this history and the current mean- according to which all humans bearing the same ing of the nomen Rana esculenta. The same patronym share a common male ancestor. would not be true if any new conception of the However, when one comes to reality, it is inesca- taxonomy of this group of frogs had resulted in pable that in any such society some children bear abandonment of this nomen or of its followers, a patronym although their biological father was on account of the fact that today’s concept of not the “official” one. Although methods would Rana esculenta is very different from that of exist (through genetic analysis) that would
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potentially allow clarification of this point, they (a very frequent process in taxonomy), or every are unlikely ever to be applied: a man who has time a taxon, or even a single specimen, is added to travelled much around the world may potentially the taxon or removed from it, this would require have “left” children in many countries, so that it the establishment of a new nomen to account for will be virtually impossible ever to establish the this “new taxon”: such a system would result in a complete list of his descendants. So, a fully permanent and endless increase in the number of unambiguous definition of a class may not nomina at the scale of all zoology, each nomen ap- correspond to a fully unambiguous list of mem- pearing in only a single or a few publications, bers of the class: the practical tool used in such a without providing any system allowing us to trace case may be that all children legally recognized the taxonomic history of a taxon through conti- by a man and bearing his patronym may be nuity of its nomen. Nomina could therefore not considered his descendants, but this is only an be used to establish historical connections be- approximation of the “real” class (which no tween pieces of information added by subsequent doubt exists in the real world but which may authors, and zoological nomenclature would be- remain unknown forever). come a kind of game, a useless discipline, unable All real taxonomies are approximations of the to play a rôle of information storage and retrieval. same kind, whatever the philosophy of classifica- Actually, it is just such a system which is currently tion adopted: they all rely on relative unam- at work as concerns the nomenclature of higher biguity, i.e. on open definitions. In any taxonomy, taxa. The untold “rule” that prevails among many taxa are defined by characters or properties that taxonomists is that, as nomina of higher taxa are may be fully unambiguous, but the characters or not governed by the Code, zoologists are free to use properties of some organisms may be unknown any nomen for such taxa. They then often tend to or wrong, so that their taxonomic allocation may consider that such nomina are unambiguously change. In this respect, although the Quer- and definitively associated with the original authian proposal is to use unambiguous “phylo- content and/or definition of the taxa to which genetic definitions” of taxa, the taxonomic they were first applied, so that every time the allocation of organisms to taxa remains in part concepts on taxonomy or phylogeny change, ambiguous in this system, quite as much as in the nomina have to change also. In the long run, such “traditional” approach to taxonomy. a tendency may not only unnecessary and dramat- To play its rôle of information storage and retriev- ically increase the number of available higher al system usefully, nomenclature must rely both nomina, but also, by suppressing all continuity, on strict and stable Rules for the allocation of nomina through the on-going use of the same nomina de- to taxa and for the establishment of the valid spite progressive changes in taxonomy, between nomen of each taxon, but also on a rather high subsequent taxonomies of a group, lead to a pro- flexibility in the relationship between taxon and gressive chaos, where storage and retrieval of taxon- nomen: the taxon must be allowed to change, omic information would become increasingly both in its definition and content, without change difficult. in the nomen, which remains characterized by its Class-series nomina should therefore follow rules permanent connection with the onomatophore similar to those of the other nomina, that are through the taxomen. Under these rules, the ques- regulated by the Code: after establishment of a tion of the allocation of a nomen to a taxon cannot higher taxon, the latter should be liable to be be considered “in general” or in an abstract way, modified in its definition and content but not in but must be addressed within the context of a given its nomen, as long as the onomatophore remains taxonomy. If a nomen was unambiguously and de- included in the taxon. This is indeed what occur- finitively linked to a biological definition or a closed red very often in the past (see below for details list of included individuals or taxa, every time the and examples), and this should continue in the definition of the taxon changes, even slightly future: change (through addition of taxa) in the
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content of a higher taxon should not require relevant characters of the latter: any other speci- modification of its nomen, that remains in all men showing “the same” characters could then cases attached to its onomatophore. The rules be considered as a “double” or a “duplicate” of devised for higher nomenclature should allow the type(s), belonging in the same taxon, whereas unambiguous allocation of nomina to taxa within any specimen showing “different” characters the frame of a given taxonomy, but not “in gener- would have to be allocated to a distinct taxon. In al”, i.e. independently from the taxonomy adopt- such a typological approach to taxonomy, the ed: in other words, nomenclature should be at types were mostly understood as specimens the service of taxonomy, not the reverse. “bearing the characters” that allow recognition of the taxa in which they belong. However, with the development of population thinking and of the NEW CONCEPTUAL AND “new systematics”, the idea that species were TERMINOLOGICAL CLARIFICATIONS composed of identical (or almost so) specimens was replaced by a concept of species showing A PROPOSED FAREWELL TO THETERM “ TYPE” IN (sometimes important) internal polymorphism. BIOLOGICAL TAXONOMY AND NOMENCLATURE This was the end of the typological or essential- Time may have come for seriously considering a istic concept of species and its replacement by an progressive, but ultimately complete, abandon- evolutionary or populational species concept ment of the term “type” traditionally used in bio- (Mayr et al. 1953; Mayr 1969, 1981, 1982, logical taxonomy and nomenclature. As a matter 1995, 1997; Mayr & Ashlock 1991). of fact, this term is both ambiguous (as it desig- However, in parallel with the progressive nates both a taxonomic and a nomenclatural consciousness that no specimen can be consid- tool) and misleading (as it suggests that biological ered as “typical” or fully representative of a spe- taxonomy is still typological). It is here argued cies, taxonomists also discovered that, beside that this term should be abandoned, and replaced their function as semaphoronts, “types” had a by unambiguous and well defined terms. second and independent function, that of being a material, objective reference for the allocation of Hypodigm and onomatophore nomina to taxa. The idea was already clearly The term “type” is highly ambiguous as it has expressed at the end of the 19th century: “It is been used with various meanings in many fields still protested by the surviving idealists that no of human activities. Even within evolutionary single specimen can give an adequate idea of the biology, it may designate either an evolutionary, species, and nobody claims that it can, but the a taxonomic or a nomenclatural concept. As an desirability of a single definite nexus between evolutionary concept, a type or “ground-plan” nature and science is rapidly becoming patent to (see e.g., Danser 1950) “is a class whose members all” (Cook 1898a: 187). This led to the persis- share a certain set of defining traits”, and gaps are tence of “types” in biological taxonomy, although thought to exist between types (Levinton 1988: mostly as a nomenclatural, instead of taxonomic, 17). In most of these acceptations, the term tool, i.e. as a “name-bearer” (Blackwelder 1967: “type” designates a concept, i.e. an abstraction (see 166). Unfortunately, the perpetuation of the use e.g., Bloch 1956: 68-69). In contrast, in biolog- of the term “type” to designate this tool resulted ical systematics, “types” have long been used as in ambiguities. As well outlined, among others, reference specimens both for taxonomy and by Rickett (1959: 24), “The word type is unfor- nomenclature. These were the specimens avail- tunate”. Its continued use is confusing for some able to the author(s) who proposed a new specific taxonomists (e.g., Oldroyd 1966), but mostly for or subpecific taxon and its nomen. In the early non-taxonomists, who sometimes consider that, days of taxonomy, these specimens were believed because they are still using “types”, taxonomists to be “typical” of the taxon, i.e. to show all the are still relying on a typological concept of species
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(e.g., Ruffié 1982: 405-408). This is still a mological tool of onomatophore and a pre-evolu- widespread belief, as illustrated, among many tionary, typological thinking in taxonomy. The others, by the following statement under the pen time should now come for progressively consider- of an editorialist of the journal Science: “Under ing going further in the direction indicated by the traditional system, a taxonomist (...) selects the authors cited above. Of course, the specimens the most representative species to be the ‘type’ for which are traditionally called “types” or “type each genus, then the most representative genus to specimens” play a central rôle in taxonomy and be the type of the family and so forth. (...) as new nomenclature, not because they would be “typi- specimens with similar characteristics are found, cal” in the essentialistic sense of the term, but they are deemed part of a known species, or even because they constitute an objective, material a new genus based on how closely they resemble reference to what the author who created a taxon the type specimen” (Pennisi 2001: 2304). and a nomen had in hand. This double rôle, both Disinformation or simple ignorance are wide- taxonomic and nomenclatural, of these speci- spread in this domain, as also shown by the for- mens, should be accounted for by the use of two mula “Linnaean type concept” used by some distinct terms, especially when dealing with (Hansen 1961: 11), although Linnaeus himself microtaxonomy (Mayr & Ashlock 1991), i.e. did not refer to types. taxonomy at the level of species and other lower To avoid definitively such misunderstanding, and taxa (subspecies, superspecies, etc.). The term to fully distinguish between the double (nomen- hypodigm was introduced by Simpson (1940, clatural and taxonomic) function of “types”, these 1961) to designate the set of specimens used by two functions should be designated by different an author to establish and describe a new species- terms. A first clarification in this respect was due series taxon: this refers to a concept useful in taxon- to Dennler (1939), followed by Simpson (1940, omy. On the other hand, the term onomatophore 1961), who both proposed the replacement of the has only a nomenclatural usefulness, as it desig- term type, respectively by the terms testimonium or nates only the specimen(s) on which the nomen more briefly test (which is ambiguous), and ono- given to the taxon objectively relies. Hypodigm matophore (which is clearly more appropriate). and onomatophore are not equivalent: in many That this change was really needed is stressed by cases, especially in recent taxonomy, the ono- the fact that, a few years later and independently, matophore consists only of a small part of the Schopf (1960) made a similar proposal, intro- hypodigm, often being restricted to one of its ducing the new term nomenifer, which is a strict specimens (the holotype or lectotype), or even synonym of test and onomatophore. In contrast, being a specimen that was not part of the original Hennig (1950: 9; 1966: 6) introduced the term hypodigm (the neotype). Rather than adopting semaphoront, which means “bearing the signs, the the well defined term of onomatophore, which characters” to designate asecond and independent clearly refers to a technical tool specific to biolog- function of specimens, including “type speci- ical nomenclature, the Code employs the unpal- mens”. All these terminological proposals show atable formula “name-bearing type”, which that the idea that “types” are “typical” and have furthermore maintains the use of the term “type” the function to give the characters of a taxon is in taxonomy and nomenclature. I suggest that in wrong and misleading, and should be extirpated the future not only this formula be abandoned from taxonomy and nomenclature. and replaced by onomatophore, but also that the The continued use, despite these judicious pro- technical terms designating the various kinds of posals, of the term “type” in the Code, probably onomatophores be changed. played a non-negligible rôle in the fact that many authors, both within and also outside taxonomy Two major kinds of onomatophores (see e.g., Dubois 2000a), have had difficulties As will be exemplified below in this paper, it understanding the difference between the ony- would be useful to dispose of a simple, unique
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term to designate any of the various kinds of ono- types” by Farber (1976) and Moore (1998), are matophores that exist in biological nomenclature. taxomina (type species, type genus), that are the This would avoid using unpalatable plurinominal reference for the status of nomina of higher ranks designations such as “onomatophoric specimen”, (genus, family, etc.). Because of the hierarchical “onomatophoric genus” or “onomatophoric local- structure of the system, all taxomina refer ulti- ity”. Combination of various terms with the root mately to type-specimens, so that in this system “onomatophore” or “nomenifer” would also pro- all nomina are connected to the real world of duce long, unpalatable terms which would have living populations through the latter (Fig. 1). So, little chance to be adopted by the community of in order to distinguish these two major kinds of taxonomists. For the terms designating different onomatophores, I propose: 1) for all type speci- kinds of onomatophores, I therefore propose to mens, the term onymophoront (from the Greek use shorter and simpler roots. To avoid the for- terms onymos, “name”, - phoros,“bearing”, and on mation of heterogeneous terms, I propose to use ontos, “being, individual”); and 2) for all ono- two distinct roots to point to the function of matophores which are taxomina, the term nucleo- onomatophore, i.e. to indicate that these are men (plural nucleomina; from the Latin terms technical tools having the function of bearing a nucleus and nomen). In both these categories of nomen. In some cases I simply propose to use the onomatophores, subcategories can be distinguished. root - phore (from the Greek - phoros,“bearing”, from the verb phero, “I bear”), but associated, just Onymophoronts like in the term semaphoront, with the root on For the four kinds of type specimens (onymo- ontos, “being, individual”, derived from the verb phoronts) recognized by the Code, I propose the eimi, “I am”: this means that such onomato- following terms: holophoront (instead of holotype; phores are real organisms that exist in the world from the Greek holos, “whole, entire”), symphoront independently of science. In other cases I propose (instead of syntype; from the Greek sun, “with, to use another, short, root, that of nucleus, together”), lectophoront (instead of lectotype; from “nucleus, core, stone” (from the Latin nux, the Greek lektos, “chosen”, from the verb lego, “I “nut”). This term is used in various domains choose”) and neophoront (instead of neotype; from (astronomy, cell biology, sociology, history, etc.) the Greek neos, “new”). After designation of a to designate a “central part or thing round which lectophoront among symphoronts, the remaining others are collected, (…) beginning meant to symphoronts become exonymophoronts (for para- receive additions” (Fowler & Fowler 1929: 776), lectotypes) and lose their status of onomatophores which quite well corresponds to the status of (see e.g., Dubois & Ohler 1997: 309, fig. 2). I do onomatophores. In biology, a cell usually cannot not propose replacement terms for the terms live without its nucleus, and in biological nomen- paratype, allotype, cotype and topotype, which clear- clature a nomen has no status (allocation to a ly do not designate onomatophores and should, taxon) without its onomatophore. This does not in my opinion, not be regulated by any code of mean that the onomatophore is “typical”, but that nomenclature, as they are in fact taxonomic, not it is “vital”, indispensable and irreplaceable. nomenclatural, tools: such specimens are in fact I suggest to use these two different roots because, just particular members of the hypodigm (for the as discussed in more detail elsewhere (Dubois first three terms) or “simple specimens” collected 2000b), two major kinds of onomatophores must from the same locality as the hypodigm or part of first be distinguished: 1) some, called “collection the latter (for the last term). The suggestion of types” by Farber (1976) and Moore (1998), are Fricke (1985) to use modified spellings in - oid specimens (holotype, syntypes, lectotype or neo- (e.g., paratypoid, allotypoid) might be followed for type), that are the objective reference by which the those who find it useful to have special terms for status of nomina of taxa of rank species or below these special members of the hypodigm: this pro- is established; and 2) others, called “classification posal has the merit to point that such specimens
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are not “real types”, but the weakness to still use In the species-series of nomina, a taxomen can be the root typ- in their designation. I also refrain created with several joint onomatophores, the from proposing a replacement term for the terms symphoronts, and subsequent selection among hapantotype of the zoological Code, or epitype of them of a lectophoront restricts the onomato- the botanical code, as I consider the concepts phore function to the latter. Similarly, in the designated by these terms as belonging to typolog- genus-series of nomina, a taxomen can be created ical thinking: onomatophores are not sema- with several joint onomatophores, among which phoronts and do not have the function to give a subsequent choice allows to restrict the onomato- the characters of the taxon. In my opinion, all phore function to one of them. I propose to call nomina, including those of protists, should rely the initial joint onomatophores in such cases pre- on holophoronts, symphoronts, lectophoronts or nucleomina (from the Latin prae, in the sense of neophoronts, and in case the latter do not allow “before”), and more precisely prenucleospecies for unambiguous allocation of a nomen to a taxon, the originally included nominal species of a new they should be replaced by adequate neophoronts, nominal genus or subgenus established with sev- a process for which the Code should provide clear eral included species, before subsequent designa- rules (this matter has not yet been properly tion among them of a single nucleospecies. In the addressed regarding the zoological Code). family-series, the situation is simpler: although An additional proposal meant to eradicate the use one could formally consider that all originally of the term “type” in biological taxonomy and included nominal genera of a new family-series nomenclature is the term onymotope (from the taxomen established with several included genera Greek onymos, “name”, and topos, “place”), to are its prenucleogenera, this almost never occurs designate the place of collection of the onymo- concretely, as to be available a family-series phoront(s), called type-locality in the Code, or nomen must be based on an available genus-series sometimes also terra typica or locus typicus by nomen, which is therefore its nucleogenus by some zoologists. Any specimen collected from the “implicit etymological designation” (Dubois same locality, whether or not an original member 1984b) from the very moment of its creation; of the hypodigm and/or of the onomatophore, however, as explained and discussed in more can be designated as onymotopic (for topotypic in detail in Dubois (in press), the category of prenu- the Code). A “topotype” may thus be designated clogenera is useful in a special situation, namely as an “onymotopic specimen”. to establish the onomatophore of an unavailable family-series nomen not being based on an avail- Nucleomina able genus-series nomen. For taxa of nominal-series above the species- For reasons that will be explained in detail in series, onomatophores are not specimens (onymo- Dubois (in press), in the class-series it is here pro- phoronts) but taxomina (nucleomina). posed that the onomatophore of a classomen may Nucleomina are of two kinds only: 1) a “ type spe- be either a single nucleogenus or a permanent, cies” is a speciomen (i.e. a taxomen of rank spe- indissoluble set of two or several nucleogenera, cies or below; see Note 12) that is the among which no subsequent restriction or desig- onomatophore of a nomen of the genus-series nation can be made. In the latter case these can (i.e. of rank genus or subgenus); and 2) and a be termed conucleogenera (from the Latin cum, “ type genus” is a generomen (i.e. a taxomen of “with”), which means “nucleogenera together”. rank genus) that is the onomatophore of a nomen of a rank above genus (i.e. of the family- or of the Additional proposals class-series). To avoid the formation of composite Also in order to avoid using the root “type”, I pro- Latin-Greek terms, I propose the replacement pose the new term monophory (from the Greek terms nucleospecies for “type species” and nucleo- monos, “single, unique”, and - phoros, “bearing”) to genus for “type genus”. designate the concept called monotypy in the Code.
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Whenever a new taxomen was established based few terminological novelties concerning the desig- on more than one onomatophore (symphoronts nation of various taxonomic and nomenclatural for a new species, prenucleospecies for a new situations usually known under the general term genus, conucleogenera for a new class, or even of “synonymy”. These proposals are complement- rarely prenucleogenera for a new family), I propose ary to those I already gave elsewhere (Dubois to designate this situation under the general term 2000b). For the time being, the term synonymy of symphory (from the Greek sun, “with, to- designates either a taxonomic or a nomenclatural gether”). In most cases, symphory is a transitional situation, or both together. In order to distin- situation, as a lectotype often is designated among guish these two situations, I propose the new symphoronts or a nucleospecies among prenu- term synotaxy (from the Greek terms syn, “with, cleospecies, but in the class-series, for reasons ex- together”, and taxis, “action to put in order”) to plained in Dubois (in press), conucleogenera designate all cases of different taxa having a remain the onomatophore together. Finally, for partially or totally identical or common content, the situation where a new taxomen was established and to restrict the use of the term synonymy (from without designated onomatophore, I propose the the Greek terms syn, “with, together”, and onymos, term aphory (from the Greek a- or an-, “without”): “name”) to the designation of all cases of different a taxon created with original aphory may later be nomina applying to the same taxon. These two provided an onomatophore in another publica- kinds of relations between taxa and nomina are tion (“subsequent type designation” under the ter- not necessarily superposed: two taxa can have the minology of the Code). same content and the same nomen, the same Finally, I propose to replace the term nominotypi- content and different nomina, the same nomen cal, used in the Code to designate taxa hierarchi- and different contents, or different contents and cally connected by the rule of coordination and nomina. having the same onomatophore, by the terms of The basic distinction between the onymological coordinate (general) and of subordinate (hierarchi- tools of nomen and taxomen (“nominal taxon” in cally lower) and superordinate (hierarchically the Code) on one hand, and the taxonomic higher). concept of taxon on the other, must be kept in The terminological proposals made here will be mind (see Dubois 2000b). Concerning nomina, summarised in Dubois (in press). The complete several distinctions can also be made (Dubois eradication of the term “type” in biological tax- 2000b). Any nomen can exist under several mor- onomy and nomenclature would clarify the mat- phonyms, i.e. several spellings or ranks. For each ters in two respects: 1) it would completely nomen, it is possible to distinguish the protonym extirpate reference to essentialistic or typological (original morphonym of the nomen) from its apo- terminology from biological taxonomy; and 2) it nyms (any subsequent, modified, morphonyms), would suppress the ambiguity between the taxon- which together constitute a group of paronyms. omic function of the hypodigm and the onymo- Similarly, during the history of the taxonomy of a logical function of onomatophores. To be fully group, definitions and contents of taxa change efficient, these terminological clarifications frequently, so that any taxon can exist under should first be adopted by individual taxonomists, several morphotaxa (from the Greek morphos, and later incorporated into the existing codes of “form”). nomenclature. It may be useful and interesting to know which taxon was initially in the mind of the author of a S YNOTAXY AND TOPOTAXY new taxomen. I here propose to distinguish the Synotaxy; morphotaxy; protaxon and apotaxon; protaxon (from the Greek pro, in the sense of ergotaxon “first”, “primitive”, “original”; see Note 13), i.e. In order to facilitate the discussion that will fol- the taxon with its original definition and content, low (Dubois in press), I am also led to propose a from its apotaxa (from the Greek apo, “coming
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from”), i.e. any subsequent, modified, definition Topotaxy and/or content of the taxon. The original taxono- Several kinds of synonymies and of synotaxies my may be known as a protaxonomy, and the can be distinguished. Let us first consider the derived taxonomies as apotaxonomies. All the dif- situation of synonyms, as here redefined. Two taxa ferent morphotaxa considered by subsequent can bear the same nomen for two different kinds authors under the same nomen (i.e. referring to of reasons (Dubois 2000b). Nomenclatural or the same taxomen), i.e. the protaxon and all its objective synonyms, more shortly isonyms (Dubois apotaxa, constitute a “family” of related parotaxa 2000b), are nomina that are based on the same (from the Greek para, “close to”; see Note 14) onomatophore, whereas taxonomic or subjective referring to the same taxomen. The different synonyms, or doxisonyms (Dubois 2000b), have morphotaxa usually differ both by their intension different onomatophores. Isonyms will always and extension, except if some mistake was made remain so, whereas the relation between doxiso- by an author when defining a taxon or when list- nyms is liable to change as taxonomy evolves: two ing its members or its non-members. nomina that were once considered subjective syno- Among the apotaxa bearing a given nomen that nyms may later be considered to denote distinct may have been recognized in various taxonomies, taxa, which can be more shortly stated by saying some may today no longer be considered valid that they are then no longer considered syno- whereas others are currently recognized by some nyms but have become xenonyms (from the Greek or all taxonomists. I propose to call ergotaxon xenos, “foreigner”). Doxisonyms and xenonyms (from the Greek ergon, “work, action”) any taxon are the two different kinds of anisonyms (from the recognized as valid by a given author or in a given Greek a- or an-, “without”, and isos, “equal”), i.e. taxonomy. An ergotaxon may be either a protaxon nomina based on different onomatophores. or an apotaxon. Any taxonomy used by a given au- Similarly, there exist several relationships between thor, whether well stabilized or highly provisional taxa regarding their contents (Fig. 3): I propose to (“working taxonomy” as defined by Dubois call these relationships topotaxy (from the Greek 1999b) can be qualified as an ergotaxonomy. topos, “place”). One can call xenotaxa two taxa As will be shown below, apotaxa always share with whose contents are completely distinct, without their protaxa a minimal common extension, i.e. any overlapping (i.e. in mathematical terms, two the original onomatophore, as exclusion of part or sets whose intersection is empty), and synotaxa all the onomatophore results in fact in the creation two taxa whose contents are partially overlapping of a new taxon. Therefore, all apotaxa are by or are identical (i.e. in mathematical terms, two definition obtained by addition into the taxon of sets whose intersection is not empty). An example new included specimens or taxa to the ono- of xenotaxy is that of two taxa X1 and X2 com- matophore. In this respect there are two major posed respectively of the individuals (or taxa) A, B, kinds of apotaxa. Some are obtained by mere addi- C, D and E, and F, G, H, I and J. As for synotaxy, tion of new specimens or taxa of low ranks, i.e. three major situations can be distinguished: 1) iso- they do not result in a modification of the inten- taxy (from the Greek isos, “equal”), i.e. absolute sion of the taxon. In contrast, some are obtained identity of contents of both isotaxa: e.g., both I1 by addition of new immediately subordinate high- and I2 and composed of A, B, C, D and E; 2) all er taxa (ametoendotaxa; see below) and in a modi- other cases belong in the general category of aniso- fied definition or diagnosis of the taxon. For those taxy (from the Greek a- or an-, “without”, and isos, of the first kind I propose the term menapotaxon “equal”), with two subcategories: a) peritaxy (from (from the Greek meno, “I remain”), and for the the Greek peri, “around”), i.e. complete inclusion second kind the term auxapotaxon (from the of one taxon ( endotaxon, from the Greek endon, Greek auxo, “I increase”). Nomina used for such “inside of”) within the other taxon ( angiotaxon, taxa can be either protonyms or aponyms, and from the Greek aggeion, “hull, capsule”): e.g., the may be designated as menonyms and auxonyms. angiotaxon A1 is composed of A, B, C, D and E,
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and the endotaxon E1 is composed of B, C and D; X enotax y and b) gephyrotaxy (from the Greek gephyra, “bridge”), i.e. partial overlap of the contents of both X enotax on X 1 X enotax on X 2 gephyrotaxa: e.g., the gephyrotaxon G1 is com- posed of A, B, C, D and E, and the gephyrotaxon A B F G G2 of C, D, E, F and G. These different kinds of relationships between taxa are diagrammatically C H shown in Figure 3. The following substantives and D E I J adjectives can be used to designate the relations between two taxa: isotaxy/xic, anisotaxy/xic, peri- taxy/xic, angiotaxy/xic, endotaxy/xic and gephyro- taxy/xic. Thus, the convenient formula “two I s otax y isotaxic nomina” is a shortened way of stating that I s otax on I 1 I s otax on I 2 these two nomina refer to exactly identical taxa or A B isotaxa (see Note 15). All these situations can be C considered among the protaxa of any given pro- taxonomy as defined above. This can be designat- D E ed as the relationships of protopotaxy: it is thus possible to distinguish xenoprotaxa from synopro- taxa, and among the latter category to distinguish A n g i otax on A 1 isoprotaxa, periprotaxa, angioprotaxa, endoprotaxa P e r i tax y and gephyroprotaxa. Similarly, relationships of A ergotopotaxy exist between the ergotaxa of any er- gotaxonomy: it is possible to distinguish xenoergo- B C D taxa from synoergotaxa, and among the latter category to distinguish isoergotaxa, periergotaxa, E angioergotaxa, endoergotaxa and gephyroergotaxa. E n d otax on E 1 In any hierarchical taxonomic system, all taxa G e p h y r otax y occupy a given hierarchical rank. Within each of the three nominal-series recognized by the Code, G e p h y r otax on G 1 G e p h y r otax on G 2 a hierarchy of taxa includes: 1) peritaxic (coordi- nate) taxa, i.e. endotaxic (subordinate) and angio- A F taxic (superordinate) taxa, that are hierarchically C D E related, and that may share the same nomen and B G onomatophore (“nominotypical” taxa) or not (“non-nominotypical” taxa); and 2) xenotaxic taxa, that always exclude mutually the onomato- F IG. 3. — Diagrammatic illustration of the different kinds of rela- tionships between taxa (A1, E1, G1, G2, I1, I2, X1, X2) concern- phore of each other. ing their contents or extensions. A to J are either individuals Whenever a taxonomist proposes a new taxon, (specimens) in the case of species-series taxa, or taxa in the case of taxa of all other nominal-series. See text for explanation. the latter occupies a given position in the hierar- chy, and several different situations can be distin- guished. Relative to any other taxon of this the various situations at stake may be designated hierarchy, the new protaxon may be xenoprotaxic by different terms. In some rare cases, a new or periprotaxic. In the latter case, relative to the taxon when first introduced is considered to be immediately superordinate taxon, the new taxon completely isolated, not closely related to any may either be alone or share its rank with other already recognized taxon, and it is the only taxon taxa. For more clarity in the following discussion, of a given rank subordinate to the taxon of
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immediate higher rank: it then qualifies as an taxa of the same rank that are immediately subor- eremoprotaxon (from the Greek eremos, “desert, dinate to the same ametoangiotaxon can be desig- isolated”). In all other cases, the new taxon shares nated as ametoxenotaxy, with two particular cases, its rank with other xenotaxa immediately subordi- ametoxenoprotaxy and ametoxenoergotaxy. In nate to the same higher taxon: it then qualifies as contrast, relationship between teletaxa can be their ametoxenoprotaxon (from the Greek ametos, designated as teletaxy ( teleprotaxy and telergotaxy). “immediate”). Two different situations may then All these terms that designate categories of proto- be distinguished. In some cases, the new taxon is potaxy and ergotopotaxy will be useful to designate simply added, beside other already recognized in a concise and precise way the kinds of relation- taxa of the same rank, as a new subdivision of a ships that exist between taxa in any protaxonomy taxon of immediately higher rank; this often or ergotaxonomy. requires some changes in the intensions and A final clarification is needed concerning the rela- extensions of these already existing taxa, but not tionship between endotaxy and isotaxy. Because their creations. These taxa, as well as the newly of the hierarchical structure of taxonomies, two created taxon, are ametoxenoprotaxa that were taxa may be isotaxic at one taxonomic level but not created together. They can be designated as endotaxic at another level. A taxon of a given hier- adelphoprotaxa (from the Greek adelphos, “bro- archical rank having several levels of hierarchically ther”), and their nomina as adelphonyms. In the subordinate taxa may be isotaxic with another second situation, a previously existing taxon is taxon of same rank including representatives divided by an author into two or more new taxa: (members) of all its ametoendotaxa (taxa of the the latter are created together, in the process of immediately subordinate rank) but angiotaxic splitting and redefining an earlier taxon, and relative to this taxon at lower levels if the latter together they cover the whole extension of the includes only some of its lower subordinate taxa, latter as redefined. They can be called didymopro- at some ranks of the hierarchy only. In the family- taxa (from the Greek didumos, “double, twin”), series, stating that a family is endotaxic to another and their nomina as didymonyms. means that it contains only a part of the tribes or In contrast to ametoxenotaxa, teletaxa (singular genera of the latter (and no genus excluded from teletaxon; from the Greek tele, “far from”) are taxa it). In the class-series, more possibilities are of- that are more remotely related, i.e. taxa, whether fered as a class is endotaxic to another if it contains of the same rank or not, that are not members of only part of its families, tribes or genera, even if it the same immediately superordinate higher is isotaxic to it at order level. Therefore it is pos- taxon. sible to distinguish between panisotaxic taxa (from To put the same ideas more generally, in any the Greek pas, “all, each”), that are isotaxic at all hierarchical taxonomy, peritaxy exists at different ranks, and merisotaxic taxa (from the Greek meros, levels: a class includes several orders, but also “part”) that are isotaxic only at the higher levels several families, several genera, etc. Of particular but peritaxic at lower hierarchical ranks. interest are the taxa of the immediately higher The relations of xenotaxy and synotaxy here (superordinate) and immediately lower (subordi- applied to nomina of the family- and class-series nate) ranks. This kind of relationship can be desig- of nomina are also liable to be applied to nomina nated as ametoperitaxy (from the Greek ametos, of the genus-, species- and variety-series, but this “immediate”): it includes ametoangiotaxy and question won’t be explored further in this paper. ametoendotaxy. These categories can be recog- On the other hand, all the terms just introduced nized both in any protaxonomy ( ametoperiprotaxy, will be useful to describe the taxonomic and ametoangioprotaxy, ametoendoprotaxy) and in any nomenclatural situations that are met with when ergotaxonomy ( ametoperiergotaxy, ametoangio- one examines the evolution of higher zoological ergotaxy, ametoendoergotaxy). Furthermore, in any classifications, as will be exemplified for the taxonomic hierarchy, the relationship between amphibians (Dubois in press, in prep.).
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C ATEGORIES OF USAGE foundations of our zoological nomenclature. It is Some additional comments on the concept of usage still more regrettable that the Conservation in zoological nomenclature are also necessary. In Principle should be applied to names ‘in general recent decades, a number of zoologists have sup- current use’, for we today are the very zoologists ported the idea that the basic principle of zoologi- who have the least excuse for errors in nomencla- cal nomenclature, that of priority, should be ture having had the International Rules in force moderated by a “subsidiary principle”, that of for a longer period than any of our predecessors. “usage”. As well analyzed by Hoł ynski (1994), the (...) Is the New Code designed to amplify the transition from a simple system with a single existing Rules to cover as many cases as possible, to structuring principle to a new system with two make the rules clearer, to facilitate their applica- such principles is a source of terrible confusion tion and so hasten law and order in nomenclature? and instability, and is certainly not to be viewed as Or is it designed to please as many conflicting per- progress (see also Dubois 2000a): “The co-exis- sonal preferences as possible and thus prolong dif- tence in one code of diametrically opposed articles ferences of usage?”. cannot possibly contribute to stability and univer- Besides, basic problems exist regarding the real sality of nomenclature” (Melville 1958: 1249). meaning of the term usage. Several recent authors This is by far not a new question, as it was very have insisted that “usage” should be “protected” clearly analysed more than one century ago by the in many cases, even when it is not correct accord- most acute theoreticians of biological nomencla- ing to the rules of the Code, because nomina are ture, whose writings are still worth reading, such not meant for the needs of taxonomists only but as those of O. F. Cook, who wrote for example: are also employed by many “non-systematists” “Usage has never produced any general or per- (such as scientists working in various fields of manent uniformity in manners, government, litera- biology, medicine, agronomy, etc., or even non- ture or science, and no reasons are apparent for scientists such as members of administrations, supposing that it ever will. There could scarcely be governments, customs, conservation organiza- a uniform logical system founded upon usage. tions, etc.). This is no doubt true, and agreed The idea involves a contradiction in terms, and a upon by most taxonomists. However, in a num- plea for usage is, in effect, a plea for anarchy” ber of cases, the same authors argued for the (Cook 1898a: 189). Sixty years later, Tottenham conservation of an incorrect usage although the (1958: 1080-1081) also rightly remarked: “While latter had only existed among systematists, i.e. fully in agreement with the Principle of taxonomists or phylogeneticists (see Note 16), Conservation in itself, I deeply regret its introduc- which is either incoherent or dishonest. As a mat- tion into the Code as a Rule. It is often directly ter of fact, there is a real intellectual dishonesty in contradictory to the Principle of Priority and there- both stating that nomenclatural stability is neces- fore its introduction means a fundamental change sary for non-systematists, users of taxonomies, of Rule. If the Rules themselves are subject to but then to provide evidence for a “need of pro- such fundamental change and are themselves un- tecting usage” based on purely taxonomic or stable, how can the nomenclature which is based phylogenetic publications. In such a context, the upon them be stable? Furthermore, its introduc- formula “widely used” should be understood as tion as a Rule would mean that a large amount of meaning “widely used outside systematic litera- zoological nomenclature would be based upon the ture”, as was already pointed out: “When a name errors of present day zoologists; this is wholly un- has been used only or almost only in taxonomic desirable; accuracy is a hall-mark of good scientific publications, the only biologists that are really work. It is sad to think that this system of basing acquainted with this name are taxonomists. If much nomenclature on errors should be intro- discovery of a senior ‘overlooked’ synonym duced into the Code this year when we are cele- occurs, all taxonomists can understand the reason brating the bicentenary of Linnaeus who laid the why the junior name must be replaced by the
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senior one, and no real disturbance of their work line, the Commission, according to the suggested follows this change” (Dubois 1997a: 319). It procedure, must reject the senior synonym and might be added that, if some systematists are place the junior synonym on the appropriate unable to understand this, then they should Official List of Valid Names”. These words read undertake some refresher courses or decide to particularly prophetic in the light of a number of change to another activity. cases and decisions taken by ICZN in the recent Another important comment regarding “usage” is years. Hershkovitz (1958: 913) went as far as that, if “universal usage” (by non-systematists) is concluding his paper by the following a valid reason for asking for the protection of a “Recommendation”: “Any principle, rule or law nomen, the same is not true for a so-called “pre- that specifically provides for annulment or cir- dominant usage”. In all cultural domains includ- cumvention of the Law of Priority shall not be in- ing science, as soon as two different opinions corporated into the International Rules of exist, both have their philosophical justification Zoological Nomenclature”. and are entitled to consideration, irrespective of All these clarifications allow the proposal of more their numbers of supporters, of the institutional precise definitions than those currently available “importance” of the latter, of their financial situa- for the different categories of usage of nomina. tion or other similar criteria: no quantitative cri- Clarity of the zoological Code would certainly terion is useful to decide which opinion is “more benefit from adopting such well defined catego- important” or “more significant” and therefore ries for the usage of nomina, instead of the vague should be followed (actually, the history of sci- concept of “predominant usage” introduced in ence is full of examples where the majority was the last edition of this book, which takes into later shown to have been completely wrong). account only nomina treated as valid but not Finally, consideration of “usage” as a reason for available, so that a nomen duly listed as an inval- protecting a nomen should be restricted to publi- id synonym in hundreds of publications since cations that appeared previous to the public discus- 1899 may now be considered a “nomen oblitum”. sion of the case, because otherwise risks are great In what follows, “usage” of a nomen designates that “‘current usage’ be deliberately rigged or ma- two different situations: 1) genuine usage of a nipulated to support one’s interpretation” (Dubois nomen as the valid nomen for a taxon; and 2) 1997a: 319). More generally, the risk of manipu- explicit mention of this nomen (e.g., in a synony- lation of usage (a risk that does not exist with pri- my) as an available but invalid junior synonym or ority) was identified long ago, e.g., by Hershkovitz homonym of another nomen considered the valid (1958: 912) who very aptly wrote, in a public dis- one for the taxon; this category includes the cases cussion on the “Principle of Conservation” (an of due consideration given to this nomen as a earlier avatar of the “rule of predominant usage”): senior homonym of a junior nomen, which is “According to the procedure suggested for use of therefore considered invalid. In the case 2, the Principle of Conservation, a proposal for the although the nomen is not used in a given taxon- rejection of a senior synonym may be made by one omy, it is considered available (and therefore can- person. The proposal may be based, consciously or not qualify as a “forgotten” nomen) but unconsciously, on prejudice. The pertinent “inactive”, a situation that may change whenever bibliographic data may be deliberately or inadver- the taxonomy of the group changes (this corre- tently one-sided and/or misleading. The proposer sponds to the rather common situation of “resur- is not required to present personal credentials, his rection” of a doxisonym during the revision of a motives may not be openly questioned, his com- group); furthermore, in the special case of a petence as a zoologist, if he is one, is not weighted. nomen treated as a senior homonym making a In the absence of a protest to a proposal because of junior homonym invalid, this nomen, although ignorance, indifference, or inability on the part of invalid, is fully “active” in zoological nomencla- an objecting specialist to meet the two-year dead- ture.
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After their creation, nomina can have four kinds December 1899, whereas a class-series nomen of fate. 1) Some nomina are adopted universally that has never been used as a valid nomen (or an as valid by all subsequent authors, either for the available senior homonym) by any author after original taxon for which they had been coined, or that date must be considered an aphonym (if for slightly or widely different taxa. Such nomina, mentioned as available but invalid) or an eneo- used harmoniously by all authors, can be called nym (if never mentioned); all other nomina, that symphonyms (from the Greek symphonos, “harmon- have been used as valid (or as available senior ious”). 2) Some nomina are adopted as valid by homonyms) after that date by only some of the some of the subsequent authors, while some authors and in only some of the publications other authors use another nomen or other (whatever the respective numbers or proportions nomina for the same taxon or for related taxa. of these different uses) are to be treated as di- Independent of the respective numbers of uses of aphonyms. Therefore, any class-series nomen the competing nomina (see below), these can be created after 31 December 1899 cannot be an called diaphonyms (from the Greek diaphonos, aphonym or an eneonym, but is either a sympho- “discordant”). 3) Some nomina are regularly nym or a diaphonym. listed as available but invalid (for being junior As tackled above, the idea that a nomen should be synonyms or homonyms of valid nomina). These protected because it has had a “common” (al- can be called aphonyms (from the Greek aphonos, though not universal) usage should be qualified “silent”). This category includes the nomina that further. If some diaphonyms are to be protected in are considered invalid but “active” as available such cases because they are “well known to non- senior homonyms that make their junior homo- specialists”, then the evidence for this statement nyms invalid. 4) Finally, some nomina are com- should rely on “non-systematic” publications pletely forgotten by all authors and never used as only. Some nomina, although known by specia- available nomina for a long time. Such nomina lists (taxonomists or phylogeneticists), are never, can be called eneonyms (from the Greek eneos, or very seldom, used in non-systematic literature, “dumb”). The category of “ nomina oblita” as and the need of their “protection” cannot be sup- recognized by the Code is a strongly heterog- ported for this reason, whereas this argument can eneous one, that mixes two widely different situa- be used to protect nomina that have been widely tions, that of aphonyms and that of eneonyms. used outside systematics: the case of the nomina These four categories of nomina have general B ATRACHIA and L ISSAMPHIBIA, presented in detail value for all zoological nomina, but here special in Dubois (2004b, in prep.), exemplifies this dis- attention will be given to their status and use in tinction well. In other cases, different nomina are class-series nomenclature. The meaning and use used alternatively but almost as frequently by dif- of these categories in the three other nominal- ferent authors (or even by the same author) to de- series, and in particular relative to the definition signate the same taxon, as exemplified by the case of “usage” and “predominant usage” as recog- of the nomina U RODELA and C AUDATA. To clarify nized in the Code for these nomina, is a distinct these distinctions, I recognize several subcatego- matter that will not be addressed here. ries within the general category of diaphonym. I Following some of the Rules of the Code for the propose the term stenodiaphonym (from the Greek nomina of other nominal series (e.g., Art. stenos, “narrow”) for any nomen that has been 11.7.2), I suggest that the deadline of 31 used until now only, or almost only, within the December 1899 be used to establish the status of narrow frame of systematic (i.e. taxonomic and/or class-series nomina relative to these four catego- phylogenetic) literature after 31 December 1899, ries: to be considered a symphonym, a nomen as a valid nomen or as a senior homonym making must have been recognized as valid for the taxon it a junior homonym invalid; therefore, such a denotes or for similar, synotaxic taxa (see Note 17) nomen has not been significantly used as valid in in all publications and by all authors after 31 non-systematic literature after 31 December
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1899. In contrast, the term eurydiaphonym (from that may be known but are not universally adopt- the Greek eurus, “wide”) designates a nomen that ed, can be grouped in a general category of dis- has been significantly used as valid in non-systema- tagmonyms (from the Greek distagmos, “doubt, tic literature after 31 December 1899 for a given uncertainty”). taxon. To qualify the term “significantly” further, A last clarification should be added on this ques- I propose below some practical rules, which are of- tion of usage. As explained by Hoł ynski (1994: fered for discussion. A further distinction can be 14-16) in his excellent rebuttal of the ill-inspired made between two kinds of eurydiaphonyms: a paper of Gould (1990), the fact that a nomen has paneurydiaphonym (from the Greek pas, “all, had widespread use and should be treated as a each”) is a nomen that is the only eurydiaphonym to sozonym according to the proposals above, have been used as valid for the taxon it designates implies by no means a judgement on the “value” in numerous non-systematic publications, where- of this nomen or on the “quality” of its original as a schizeurydiaphonym (from the Greek skhizo, publication. It only depends on the subsequent “I split, I separate”) designates an eurydiaphonym history of this nomen and of other nomina in the used as valid but alternative to another eurydi- nomenclature of the zoological group concerned. aphonym for the same taxon in numerous non- In fact, in many cases a nomen has gained univer- systematic publications. Of course, allocation of a sal or general acceptance although it has been nomen to one of the categories above is not neces- poorly introduced for a poorly defined or heterog- sarily definitive, as a stenodiaphonym can become eneous taxon, as well examplified by the nomen an eurydiaphonym if it becomes regularly used in A MPHIBIA (Dubois2004b). Protection of such a general literature. nomen is only justified by the need of nomencla- Altogether, symphonyms and paneurydiapho- tural stability for well known taxa, irrespective nyms form a special category of nomina that have of the quality of the publications where their been applied by all or almost all authors in many nomina were originally proposed. works, including non-systematic ones, to the same taxon or to closely related ones, whereas no T AXONOMIC CATEGORIES AND RANKS other nomen has been used significantly for the In zoological taxonomy and nomenclature, the same taxon. Such nomina are therefore unambig- terms “category” and “rank” are often used as uous and are well known not only to system- synonyms. For example, in the Glossary of the Code, atists, but also to other biologists and to laymen. the term rank is defined as “The level, for nomen- In the class-series especially, where the only rule clatural purposes, of a taxon in a taxonomic hier- in force until now for the validity of nomina has archy”, whereas the phrase taxonomic hierarchy is been “consensus”, such nomina no doubt merit defined as “A system of classification based on a se- protection and conservation, even if they had not quence of taxonomic categories ranked by their been originally coined for the taxon that they increasing levels of inclusiveness”. As for the term now refer to in the eyes of many. I propose the category, no definition is provided in the English term of sozonym (from the Greek sozo, “to keep, Glossary, but in the French Glossaire of the Code to protect”) to designate any such nomen. This is the following definition is provided: “Classe des a Greek-derived term equivalent to the Latin- taxons d’un certain rang, p. ex. l’espèce, la sous-fa- derived expression nomen conservandum that is mille etc.”, which could be translated into: “Class sometimes found in taxonomic literature, and of taxa of a certain rank, e.g., species, subfamily, that was introduced in the last edition of the etc.”. Mayr & Ashlock (1991: 431) defined tax- Code under the form of nomen protectum onomic category as follows: “Rank in a hierarchy (Anonymous 1999b: Article 23.9.2). of levels to which taxa are assigned, such as sub- In contrast, schizeurydiaphonyms, stenodiapho- species, species, and genus. A class whose members nyms, aphonyms and eneonyms, which cannot are all taxa assigned a given rank”. The same au- be considered well known to non-systematists, or thors provided the following definition of ranking:
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“The placement of a taxon in the appropriate cate- Second, the category of plesion (Patterson & gory in the hierarchy of categories” (Mayr & Rosen 1977: 160), used by some authors to desig- Ashlock 1991: 427). Many other authors who nate fossil taxa that cannot properly be placed in provided similar definitions could be mentioned. the taxonomic hierarchy, is an unranked taxon- It is here proposed to make a distinction between omic category that has no nomenclatural rank the two terms, by considering category as designat- counterpart. The same applies to the common ing a taxonomic concept and rank as designating use in all taxonomic literature of the phrase incer- anomenclatural concept. Whereas all ranks cor- tae sedis to designate taxa that cannot be properly respond to a category occupying the same position ascribed a place in the taxonomic hierarchy of in the hierarchy ( ranked categories), a few catego- their group. ries do not correspond to hierarchical ranks and In most cases in zoological taxonomy, rank and could be called unranked categories. These are of category correspond strictly, thus being a single two kinds. taxonomic-nomenclatural category-rank, but in First, they include the categories termed by the few cases just mentioned the two are dissociat- Bernardi (1980) “taxonomic categories of evolu- ed: the taxonomic category does not directly cor- tionary systematics”. These are a number of cate- respond to a nomenclatural rank. For many gories recognized either between the rank genus purposes, including non-systematic usage (e.g., in and the rank species, or below the rank species, to legislation, customs, trade, etc.), taxonomic cate- point either to some particular cases of species (so- gory is not an important information but nomen- called “sibling species” or dualspecies; allopatric clatural rank is indispensable: to take just one species or prospecies; incipient species; all surveyed example, in order to build up a strategy of conserv- by Bernardi 1980), or of subspecies ( grex, quasispe- ation biology for a locality, country or conti- cies, vicespecies and exerge, all surveyed by nent, it is necessary to have lists of species-rank Bernardi 1980; morph of Edwards 1954; strong taxa, whatever the biological characteristics of subspecies and weak subspecies of Deuve 1994, these “species” (i.e. whether dualspecies, prospe- 2004), or to taxa that do not have normal meiosis cies or kleptons). Thus the distinction between and reproduction, referable to categories such as nomenclatural rank (of general use) and taxon- “parthenogenetic species” ( klonon), “gynogenetic omic category (of more specialised use in some species” (gynoklepton) or “hybridogenetic species” cases) is relevant, and should be spread in all zoo- ( zygoklepton) (Dubois 1991a). These are properly logical taxonomy. recognized as taxonomic categories, but they are not occupying a particular rank in the taxon- omic-nomenclatural hierarchy. All these categories CONCLUSION are referable to the species or subspecies rank. These “evolutionary categories” are useful to pro- The present proposals are the result of a 20-year vide biological information on the “kind of taxa” reflection, already published in part (Dubois that they designate, but no information on rank- 1984b, 2000b). I do not expect immediate ing. Thus, in the species-series, and also in the understanding and agreement from all colleagues variety-series, such unranked categories can be to these proposals: rather, I have no illusion used in parallel with the ranked categories that about the fact that introduction of all these new correspond to the nomenclatural ranks. At this terms will probably entail a strong resistance, if level (species-subspecies-variety), a third, distinct, not aggressive comments, in the community of item that should be distinguished, is the recogni- taxonomists, as nothing is more difficult than tion of special evolutionary or populational changing an “old tradition” in science, and espe- concepts, such as deme (Gilmour & Gregor cially in an old discipline. It may be easier to 1939) or cline (Huxley 1939), that are neither introduce completely new rules and terms, under taxonomic categories nor nomenclatural ranks. a new philosophy, than trying to improve an
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existing system. However, the latter attitude seems International Trust for Zoological Nomenclature, more justified in view of the fact that the current London, xviii + 176 p. system of zoological taxonomy and nomencla- A NONYMOUS [International Commission on Zoological Nomenclature] 1964. — International ture, although not perfect, has a number of merits, Code of Zoological Nomenclature. 2nd edition. has been in existence for more than 250 years and International Trust for Zoological Nomenclature, that all the information stored in this system London, xx + 176 p. should be saved rather than thrown away; this A NONYMOUS [International Commission on Zoological Nomenclature] 1974. — Amendments does not mean that this information should to the International Code of Zoological Nomenclature remained unchanged, but in order for it to evolve adopted since the XVIth international congress of and improve, it should remain available. Even if zoology, Washington, 1963. Bulletin of Zoological the process is slow, I do hope that in the long run Nomenclature 31: 77-101. A NONYMOUS [International Commission on the merits of completely abandoning the term Zoological Nomenclature] 1985. — International “type” in biological taxonomy and nomenclature, Code of Zoological Nomenclature. 3rd edition. and of using the other terms proposed above, will International Trust for zoological Nomenclature: be progressively recognized. The next papers of London, xx + 338 p. A NONYMOUS [International Commission on this series (Dubois in press, in prep.) are meant to Zoological Nomenclature] 1990. — Opinion 1604. illustrate how the use of these terms can help in Ichthyophiidae Taylor, 1968 (Amphibia, clarifying some complex nomenclatural situations Gymnophiona): conserved. Bulletin of Zoological that arise concerning the nomina of higher taxa Nomenclature 47 (2): 166-167. because of the long absence of rules for these A NONYMOUS [International Commission on Zoological Nomenclature] 1993. — Opinion 1749. nomina, and, particularly, of the absence of a rule Epicrium Wagler, 1828 and Ichthyophiidae Taylor, of coordination among them. 1968 (Amphibia, Gymnophiona): conserved, and Epicriidae Berlese, 1885 (Arachnida, Acari): con- served by the emendation of Epicriidae Fitzinger, 1843 (Amphibia, Gymnophiona) to Epicriumidae. Acknowledgements Bulletin of Zoological Nomenclature 50 (3): 261-263. For comments on previous drafts of this paper, A NONYMOUS [International Commission on Zoological Nomenclature] 1997. — Opinion 1873. discussions on some onymological concepts and Hemidactyliini Hallowell, 1856 (Amphibia, their uses, and for providing me with bibliogra- Caudata): conserved. Bulletin of Zoological phical references, I am grateful to Philippe Nomenclature 54: 140-141. Bouchet, Roger Bour, Thierry Bourgoin, A NONYMOUS [International Commission on Zoological Nomenclature] 1999a. — Opinion Frédéric Chérot, Patrick David, Thierry Deuve, 1921. Petropedetinae Noble, 1931, Cacosterninae Claude Dupuis, Jacques Le Renard, Valéry Noble, 1931 and Phrynobatrachinae Laurent, 1941 Malécot, Alessandro Minelli, André Nel, Fredrik (Amphibia, Anura): given precedence over Pleijel, Odile Poncy, Roger Roy, and particularly Hemimantidae Hoffmann, 1878, and Phryno- batrachinae: not given precedence over Annemarie Ohler, who also prepared the figures Petropedetinae. Bulletin of Zoological Nomenclature of this paper. I thank Myrianne Brival and 56 (1): 96-100. Victoire Koyamba for their help in bibliographic A NONYMOUS [International Commission on research. I sincerely thank two anonymous review- Zoological Nomenclature] 1999b. — International Code of Zoological Nomenclature. 4th edition. ers for their careful reading and their constructive International Trust for Zoological Nomenclature, comments on this paper. London, xxix + 306 p. A NDERSON J. S. 2002. — Use of well-known names in phylogenetic nomenclature: a reply to Laurin. Systematic Biology 51: 822-827. REFERENCES A SHLOCK P. D. 1984. — Monophyly: its meaning and importance, in D UNCAN T. & S TUESSY T. F. (eds), A NONYMOUS [International Commission on Cladistics: Perspectives on the Reconstruction of Zoological Nomenclature] 1961. — International Evolutionary History. Columbia University Press, Code of Zoological Nomenclature. 1st edition. New York: 39-46.
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Should internet sites Mémoires du Muséum national d’Histoire naturelle be mentioned in the bibliographies of scientific (A) 140: 1-123. publications? Alytes 21 (1-2): 1-2. D UBOIS A. 1989a. —Leptobrachium parvum D UBOIS A. 2004a. — Editorial. Alytes, a different jour- Boulenger, 1893 (Amphibia, Anura): proposed con- nal. Alytes 21 (3-4): 105-110. servation. Alytes 7 (3): 97-100. D UBOIS A. 2004b. — The higher nomenclature of D UBOIS A. 1989b. — Hyla reinwardtii Schleger, recent amphibians. Alytes 22 (1-2): 1-14. 1840(?) (Amphibia, Anura): proposed conservation. D UBOIS A. 2004c. — Developmental pathway, specia- Alytes 7 (3): 101-104. tion and supraspecific taxonomy in amphibians. 2. D UBOIS A. 1991a. — Nomenclature of partheno- Developmental pathway, hybridizability and gener- genetic, gynogenetic and “hybridogenetic” verte- ic taxonomy. Alytes 22 (1-2): 38-52. brate taxons: new proposals. Alytes 8 (3-4): 61-74. D UBOIS A. in press. — Proposed Rules for the incor- D UBOIS A. 1991b. — Comments on the suppression poration of nomina of higher-ranked zoological of Epicrium Wagler, 1828 and Epicriidae Fitzinger, taxa in the International Code of Zoological 1843 (Amphibia, Gymnophiona). Bulletin of Nomenclature. 2. The proposed Rules and their Zoological Nomenclature 48 (2): 152-154. rationale. Zoosystema. D UBOIS A. 1994a. — Case 2362. Phrynobatrachinae D UBOIS A. in prep. — Proposed Rules for the incorpor- Laurent, 1941 (Amphibia, Anura): proposed con- ation of nomina of higher-ranked zoological taxa in servation. Bulletin of Zoological Nomenclature the International Code of Zoological Nomenclature. 3. 51 (3): 240-246. A detailed example: the nomina of higher-ranked D UBOIS A. 1994b. — Comment on the proposed con- taxa of recent Amphibia. servation of Hemidactyliini Hallowell, 1856 D UBOIS A. unpublished-a. — Comments on the sup- (Amphibia, Caudata). Bulletin of Zoological pression of Epicrium Wagler, 1828 and Epicriidae Nomenclature 51 (3): 264-265. Fitzinger, 1843 (Amphibia, Gymnophiona). D UBOIS A. 1995a. — Comments on the proposed Original manuscript (3 pages) submitted to Philip conservation of Hemidactyliini Hallowell, 1856 Tubbs, Secretary of the International Commission
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Bulletin of Zoological Nomenclature tion of the names Epicrium Wagler, 1828 and 48 (4): 335-336. Ichthyophiidae Taylor, 1968 (Amphibia,
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Gymnophiona), and on the conservation of NOTES Epicriidae Berlese, 1885 (Arachnida, Acari). Bulletin of Zoological Nomenclature 49 (2): 153-154. 1. — One of my recent papers (Dubois 2000b), that V AN R EGENMORTEL M. H. V., FAUQUET C. M., BISHOP contains a number of points worthy, in my opinion, D. H. L., C ARSTENS E. B., E STES M. K., L EMON S. to be discussed by zoologists, submitted in 1998 to a M., M ANILOFF J., M AYO M. A., MC G EOCH D. J., “major” journal, was rejected with the following P RINGLE C. R. & W ICKNER R. B. 2000. — Virus comment: “I would hate to see nomenclature be- Taxonomy: the Classification and Nomenclature of coming even more complicated with lots of new Viruses. Academic Press, San Diego, 1167 p. technical terms, however logical. We should try to V AN V ALEN L. 1963. — Some results of the 1961 simplify the rules so that practising zoologists will Code of Nomenclature. Systematic Zoology 12 (1): follow them, and I think that this manuscript points 39-40. in the wrong direction”. In other words, it was re- V AN V ALEN L. 1973. — Are categories in different jected because the managing editor whom it had been phyla comparable? Taxon 22: 333-373. sent to had an opinion different from mine regarding W ARD L. F. 1895.— The nomenclature question. the way zoological nomenclature should evolve, and Bulletin of the Torrey Botanical Club 22 (7): 308- regarding the meaning of terms like “simple” and 329, 2 tab. “complicated” (as in my opinion terminological clar- W ASSERSUG R. J. & D UELLMAN W. E. 1984. — Oral ity simplify matters that are obscure with imprecise structures and their development in egg-brooding terms). This is not acceptable, as widely different hylid frog embryos and larvae: evolutionary and eco- opinions regarding this controversial field have been logical implications. Journal of Morphology 182: 1-37. defended, sometimes very bitterly, by different au- W EBB R. G., DUNDEE H. A., GARCIA P ARIS M., C OX thors over the last two centuries: rejecting a paper in M. J., T HOMAS R. A., HILLIS D. M., C OOK F. R., this field on account of differences in opinion (i.e. O TA H. & C HIPPINDALE P. 1994. — Comments on not of demonstrable flaws in the reasoning or of ig- the proposed conservation of Hemidactyliini norance or misunderstanding of some nomencla- Hallowell, 1856 (Amphibia, Caudata). Bulletin of tural rules) merits only one qualification: this is Zoological Nomenclature 51: 153-156. censorship. Silencing colleagues who disagree with W HEELER Q. D. 2004. — Taxonomic triage and the one’s opinions or ideas is certainly less difficult than poverty of phylogeny. Philosophical Transactions of having to refute their point of view and, above all, to the Royal Society of London (B) 359: 571-583. convince other colleagues on the basis of arguments. W HEELER Q. D. & M EIER R. (eds) 2000. — Species Another possible explanation of this rejection could Concepts and Phylogenetic Theory. Columbia be the mere length of the manuscript, dealing with University Press, New York, xii + 230 p. several questions at once, but this does not hold, as W HEELER Q. D., RAVEN P. H. & W ILSON E. O. other much shorter papers, dealing with a single 2004. — Taxonomy: impediment or expedient? problem, were also rejected as not presenting some of Science 303: 285. the “fashionable” ideas that have today the favor of W ILKINSON M. & N USSBAUM R. A. 1988. — Case the “popular medias” (see e.g., list of references in 2616. Ichthyophiidae Taylor, 1968 (Amphibia, Carpenter 2003: 79). This difficulty to simply make Gymnophiona): proposed conservation. Bulletin of some ideas widely available is one of the reasons why Zoological Nomenclature 45 (3): 207-209. the present work presents detailed approaches to sev- W ILSON E. O. 1985. — The global biodiversity crisis: eral problems altogether. I am not prepared to spend a challenge to science. Issues in Science & Technology months submitting manuscripts to journals before 2: 20-29 [not seen]. receiving negative replies simply because the referees W ILSON E. O. 2000. — A global biodiversity map. have “opinions” different from mine regarding Science 289: 2279. nomenclature. W ILSON E. O. 2004. — Taxonomy as a fundamental 2. — Hoł ynski’s (1994) in my view very important discipline. Philosophical Transactions of the Royal contribution to the discussion on the last version of Society of London (B) 359: 739. the Code had to be published in a Hungarian jour- W OODGER J. H. 1952. —From biology to mathemat- nal because it had been refused by BZN. The same ics. British Journal for the Philosophy of Science 3 (9): is true of 14 of my own texts submitted to the 1-21. ICZN secretariat for publication in BZN: six of Z INK R. M. 2004. — The role of subspecies in obscur- them were finally published elsewhere (Dubois ing avian biological diversity and misleading conser- 1987c-e, 1989a, b, 1997a) and eight of them vation policy. Philosophical Transactions of the Royal (Dubois unpublished-a-f; Bour & Dubois unpub- Society of London (B) 359: 561-564. lished-a, b) are still unpublished. Interestingly, one of them (Dubois unpublished-a) was used by the ICZN secretary to write part of a paper published Submitted on 23 October 2003; under his name (Tubbs 1992) and my text was accepted on 27 January 2005. never brought to the knowledge of the members of
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ICZN before vote on this case (Anonymous 1993). 5. — In the recent years, in order to support its action Similarly, ICZN members voted on the “famous” against a strict respect of the Rules, the ICZN P ETROPEDETINAE-PHRYNOBATRACHINAE case Secretariat has deliberately biased the international (Anonymous 1999a) in the ignorance of my last public discussion which should normally have manuscript on this case (Dubois unpublished-f) developed freely in its Bulletin: it has done so both where I had shown that the previous published text through active support to some interpretations of about it (Frost & Savage 1995) contained gross fac- the cases and repeated censorship of adverse opin- tual mistakes in the presentation of the facts and ions submitted to BZN (see Note 2 above). A recent demonstrably incomplete bibliographical coverage practice, quite foreign to the intellectual tradition of of the case. These examples are not given for “grum- science but closer to political, economic or other bling” but for pointing out a real problem of cen- social activities, has been the support brought by sorship of opinions and texts by ICZN, a problem ICZN to clear “lobby” actions undertaken by some that should attract the attention of zoologists world- researchers in order for various colleagues to speak wide. I know of several other cases of censorship for “their” nomenclature, despite its clear invalidity that touched several other colleagues, who either according to the Rules (e.g., Jennings et al. 1994; decided to forget it (and not to care any more for Webb et al. 1994). That such suddenly numerous ICZN when dealing with nomenclatural problems) comments cannot have been “spontaneous” but or to discuss these problems “privately” with ICZN were duly organized is made clear by the usual com- officers in order not to “create problems”, but in my plete or almost complete absence of comments in opinion it is not acceptable for zoologists to face similar cases in the same zoological group and at the such an attitude from officers that should serve the same period, many votes of ICZN having been community of zoologists. Furthermore, most col- made in the absence of even a single published com- leagues who ever published a text in BZN know ment in BZN. how their text can be authoritatively shortened or 6. — Future historians of science will perhaps be inter- modified before publication, not rarely in a sense ested to realize that, at the time of giving this to quite different from that of the originally submitted print (March 2005), I am unable to provide a bibli- manuscript. Indisputably, in the recent decades ographic reference to a paper-published complete BZN has worked as the “private journal” of a small version of the proposed “ Phylocode”. A version number of persons, not as the official journal of an dated 17 June 2004 and signed P. D. Cantino & K. international body supposedly at the service of zool- De Queiroz is available on-line (http://www.ohiou. ogy and zoologists worldwide. edu/phylocode), but as internet sites have no long- 3. — Strictly speaking, taxa of ranks above superfamily term permanency (see Dubois 2003b, 2004a), the should be designated as “high-level taxa”, “taxa of text as it is today on this site may be unavailable to higher ranks” or “higher-ranked taxa”, but for sake future readers. This will appear all the more strange of brevity, because of the frequent use of this con- that a number of papers have recently been pub- cept in this paper and followers, the shorter formula lished commenting on this “phantom” text. “higher taxa”, frequently found in the literature, Discussing this unpublished text in publications is including in titles of papers (e.g., Chitwood 1958; similar to discussing phantom unpublished manu- Levine 1958; Ghiselin 1977; Lee 1999; Minelli scripts, quoting them as “personal communication”, 2000), will be employed here. “in press”, “submitted” or “in preparation” (Dubois 4. — With such a criterion, nomina like those used in 1999b). the book of Fei et al. (1991), which was distributed 7. — It is even more misleading to write that “there is in thousands of schools all over China, would cer- much more consensus at the familiar [ sic, for fami- tainly qualify much more for “protection” than lial], generic and specific level [ sic, for levels]” nomina used in any book of which only a few hun- (Laurin 1998: 8). The notion of “consensus”, here, dred copies were printed and distributed only to is relevant only as concerns taxonomy, but not specialists of the academic world in western coun- nomenclature. For nomina regulated by the Code, tries. The words of Melville (1958: 1249) are also their validity is not fixed by “consensus” but by worth remembering here: “It is to my mind impos- Rules, that should be considered stringent, not sible to define numerically the amount of usage that optional, by all zoologists. Needless to say, the a junior synonym must have enjoyed to qualify for recent tendency of ICZN to “validate” a so-called conservation, or the amount of neglect that a senior (although incomplete) “consensus”, as opposed to synonym must have suffered to necessitate its rejec- the Rules, has certainly contributed to spread this tion in a way that will cover both the household idea that the nomina of taxa could be fixed by “con- words of applied zoology (in the medical, agricul- sensus”. This example is not the only one where an tural, veterinary and geological fields) and the usage active supporter of the Querauthian nomenclatural of highly restricted specialist fields; five usages in system and opponent of the current Code showed the latter may be as important to those concerned as ignorance of the latter. Here is another example: five hundred in the former”. “the standard suffixes for the various Linnean ranks
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do not exist in traditional zoological nomenclature. “fashionable” field, especially using a costly In zoological nomenclature, only taxon names tra- methodology (see Dubois 2003a), e.g., molecular ditionally assigned to the family level have a stand- phylogeny, will usually prove more fruitful than ard ending (-idae), and even here there are many describing hundreds of new species of obscure trop- exceptions” (Lee 1996: 189). The same author ical mites or worms. This is why the major problem mentioned elsewhere the “rare instances where at stake here is not one of individual tastes or orien- names have the same root but different endings tations but one of political choices by decisions- (e.g., Iguanidae, Iguaninae, Iguana)” (Lee 1999: makers regarding the scientific priorities of the 365), without apparently realizing that such “rare coming century. instances” are in fact very common, because of the 11. — Bryant & Cantino (2002: 52) stated that Rule of Coordination, a rule seldom, if ever, men- “Taxonomic freedom is also a principle of phyloge- tioned by Querauthian taxonomists and that netic nomenclature”, which is of course a mislead- imposes the existence of a nominotypical subordi- ing statement, as “taxonomic freedom” means that nate taxon in any taxon having subordinate taxa. taxonomists are free to adopt, or not, principles of The title of the last paper (on “stability of (…) taxa “phylogenetic taxonomy”, not that, in the frame of in (…) nomenclature”) also well illustrates the basic such principles, they are granted “freedom of taxon- confusion between taxa and nomina, i.e. between omic opinion with regard to hypotheses about taxonomy and nomenclature, that characterizes relationships”! many Querauthian militants: as far as nomenclature 12. — To designate a “taxomen of the species-series”, is concerned, one can only speak of stability of Dubois (2000b) had proposed the term nomina, not of taxa, a matter that does not belong “speciesomen”. The root used to coin this term was in nomenclature. That such gross factual mistakes the complete term “species”, although a correct ety- can be quietly published in a well known refereed mological formation, based on the stem of the term international journal of zoology gives support to the “species”, would have led to the term “speciomen”. statement that “low quality taxonomic works, or I had felt that the latter term would have been liable works containing severe methodological taxonomic to cause confusion with the term “specimen”. mistakes, can be and are regularly published in However, several colleagues pointed to me that the highly-ranked periodicals” (Dubois 2003a: S14). term “speciesomen” was particularly unpalatable 8. — This confusion is plainly acknowledged by some and suggested use of the correct spelling “spe- Querauthian taxonomists, for example when they ciomen”, a suggestion followed here. In the plural support “an alternative system, phylogenetic taxon- form, there is no risk of confusion between “speci- omy (or phylogenetic nomenclature)” (Pleijel & mens” and “speciomina”, but in the singular atten- Rouse 2000: 627). tion should be paid to this possible risk of misprint 9. — In the future, this rule should also be generalized or confusion. to variety-series nomina, in order to include them 13. — I do not advocate the use of the root proto here, also in the Code: as a matter of fact, the exclusion for reasons of euphonism (it would entail a heavy from the zoological Code of nomina for taxa below repetition of the letter t ) and of length of term the subspecies is an unacceptable case of “restriction (especially in composed terms derived from protax- of the freedom of taxonomic thought and action”. on). The root pro, from Greek and Latin languages, Unlike in zoology, the botanical code (Greuter et al. may take several meanings, including the one used 2000) provides nomenclatural rules for the nomina here, which is not the most frequent but exists in of lower taxa like varieties or forms. several other scientific terms (e.g., pronephros). 10. — Johnson (1970: 205) provided the following 14. — The new taxonomic term parotaxon, similar in interesting comment: “In general, whether good or aspect to its nomenclatural counterpart paronym, is bad, theoretically disposed or otherwise, taxono- distinct from the term parataxon which has the same mists themselves, like other scientists and scholars, etymology. The latter term was introduced by Moore find their own activities self-justifying because they & Sylvester-Bradley (1957) to designate classifica- bring some degree of intellectual satisfaction. This is tory units that do not apply to living organisms but doubtless the chief motivation for most of us, what- to parts of organisms or to traces left by living organ- ever rationalizations we offer about usefulness when isms. However, strictly speaking, the term taxon is called upon to show why society would support us. only relevant to designate a classificatory unit for Nevertheless, society does support us, niggardly organisms, so that, for the classificatory units that do though we claim this support to be”. Such a com- not apply to complete living organisms, I suggest ment would be only in part true today. “Intellectual abandonment of the term of parataxon (which statisfaction” may not be the only motivation for would be appropriate to designate a “kind” of taxon) chosing a field of research rather than another, even and to replace it by the new term pseudotaxon (from within a general disciplinary field like systematics. If the Greek pseudes, “liar, deceiver”). we speak in terms of career, notoriety and income, 15. — Strictly speaking however, it should be clear it is hardly questionable that choosing a “modern”, that the taxa may be isotaxic, but not their nomina,
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as nomina are only labels and have no contents, a so-called general usage outside taxonomy, which extension or intension. simply did not exist. In fact, “nomenclatural stabili- 16. — Very enlightening examples in this respect were ty” may not be the real motivation of some of the provided in the recent years in amphibians, in the militants for “usage” in the cases above, as suggested “campaigns” developed by adversaries of the recently by the fact that one of them took the exact- Principle of Priority for the conservation of the ly opposite attitude when he applied (Savage 2003) nomina I CHTHYOPHIIDAE versus E PICRIIDAE (see e.g., for “suppression” by ICZN of a widely known book Dubois 1984a, 1985, 1991b, unpublished-a; (de la Cepède 1788a, b) which contains a number Wilkinson & Nussbaum 1988; Smith 1989, 1991; of nomina used without discontinuity in zoology Anonymous 1990, 1993; Tubbs 1991, 1992), since their creation. As shown by Bour & Dubois P ETROPEDETINAE versus H EMIMANTINAE (Dubois (1984), if such a rigid application of the Rules was 1981, 1982, 1984b, 1985, 1987a, b, 1994a, 1995b, to be followed in this case, it should be so in the unpublished-f; Poynton 1995; Frost & Savage case of the classical works of Schlosser (1768) and 1995; Clarke 1995; Anonymous 1999a) or, even Boddaert (1770a, b, 1772a, b) as well. If ICZN more caricatural, H EMIDACTYLIINI versus decided the suppression of de la Cepède’s works, M YCETOGLOSSINI (Dubois 1984a, 1985, 1994b, simple consistency requires that it also suppresses 1995a; Smith & Wake 1993, 1994, 1996; Webb et the seemingly classical book of Laurenti (1768), al. 1994; Anonymous 1997), although these nomina that contains several trinominal nomina for species, had never or very rarely appeared in non-systematic as well a striking non-respect of the Linnaean hier- publications. In such cases, these “campaigns” archical structure of taxonomy in the case of the should certainly be interpreted mostly as a way of genus Proteus (see Dubois 2004b). “testing” the willingness of the majority of ICZN to 17. — This is an important qualification: of course, an replace at least partly the Principle of Priority by a author who does not recognize a taxon will not use “principle of usage”, rather than as a way to preserve its nomen as valid.
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