Skip to main content Skip to main navigation menu Skip to site footer
Type: Articles
Published: 2007-07-02
Page range: 27–68
Abstract views: 31
PDF downloaded: 2

Phylogeny, taxonomy and nomenclature:the problem of taxonomic categories and of nomenclatural ranks

Vertébrés: Reptiles & Amphibiens, USM 0602 Taxonomie & Collections, Département de Systématique & Evolution, Muséum national d’Histoire naturelle, 25 rue Cuvier, 75005 Paris, France
General Taxonomic impediment Biodiversity crisis Nomenclatural ranks Taxonomic categories Equivalence between taxa Nomenclatural redundancy Rule of Coordination Nomenclatural parsimony Polysemy Monosemy Google taxonomy Onomatophores Code Phy

Abstract

The use of ranks and nominal-series in zoological nomenclature has recently been challenged by some authors who support unranked systems of nomenclature. It is here shown that this criticism is based on a double misunderstanding: (1) the confusion between nomenclatural ranks and taxonomic categories; (2) the request for a monosemic nomenclatural system, not for scientific reasons, but to please non-taxonomists, especially customers of the web. It is here argued that nomenclatural ranks and taxonomic categories should be clearly distinguished and designated by different terms, and that the Code should be modified in order to make this distinction clear. Whereas taxonomic categories have biological definitions, nomenclatural ranks do not, as they express only a position in a taxonomic hierarchy. If used consistently (which is not always the case), the system of nomenclatural ranks is very useful for the storage and retrieval of taxonomic and phylogenetic information. Taxa referred to a given rank in different groups cannot therefore be considered equivalent by any criterion, so that using ranks for comparisons between taxa (e.g., for biodiversity richness assessment) is irrelevant and misleading. Although the current Code needs to be improved in several respects, the superiority of this nomenclatural system, which is theory-free regarding taxonomy as it relies on ostensional allocation of nomina to taxa rather than on intensional definitions of nomina, is again stressed. It is suggested that all taxonomists should follow the Code for the allocation and validity of nomina, whatever taxonomic theory they favour, and in particular whatever kinds of definitions or diagnoses they wish to use for taxa. This would avoid the considerable loss of manpower, time and energy that would be required by the implementation of a new nomenclatural system (e.g., in order to require “phylogenetic definitions” for nomina, or to make nomenclature fully monosemic), and the confusion that would result for most users of nomina. The new paradigm imposed to biology by the combination of the taxonomic impediment and of the biodiversity crisis requires from taxonomists, who are already considerably much less numerous than required by this new situation, to concentrate on what should be their priority at the beginning of the century of extinctions, namely the inventory of the living species of our planet before they get extinct.

References

  1. Anonymous [Systematics Agenda 2000] (1994) Charting the biosphere: a global initiative to discover, describe and classify the world’s species. Technical report. American Museum of Natural History, American Society of Plant Taxonomy, Society of Systematic Biologists and the Willi Hennig Society, NewYork, 34 pp.

    Anonymous [International Commission on Zoological Nomenclature] (1999) International code of zoological nomenclature. Fourth edition. International Trust for zoological Nomenclature, London, 335 pp.

    Anonymous (2005) Discussion. In: Pleijel, F., Le projet de “Phylocode” et la nomenclature des taxons supérieurs, in: Dubois, A., Poncy, O., Malécot, V. & Léger, N. (Eds.), Comment nommer les taxons de rang supérieur en zoologie et en botanique?, Biosystema, 23, 67–71.

    Ashlock, P.D. (1971) Monophyly and associated terms. Systematic Zoology, 20, 63–69.

    Ashlock, P.D. (1984) Monophyly: its meaning and importance. In: Duncan, T. & Stuessy, T. F. (Eds.), Cladistics: perspectives on the reconstruction of evolutionary history, Columbia University Press, New York, pp. 39–46.

    Avise, J.C. & Johns, G.C. (1999) Proposal for a standardized temporal scheme of biological classification for extant species. Proceeding of the national Academy of Sciences of the USA, 96, 7358–7363.

    Avise, J.C. & Mitchell, D. (2007) Time to standardize taxonomies. Systematic Biology, 56, 130–133.

    Bănărescu, P. (1973) Principiile şi metodele zoologiei sistematice. Editura Academiei Republicii Socialiste România, Bucureşti, 220 pp.

    Barigozzi, C. (Ed.) (1982) Mechanisms of speciation. Liss, New York, 560 pp.

    Benton, M.J. (2000) Stems, nodes, crown clades, and rank-free lists: is Linnaeus dead? Biological Reviews, 75, 633–648.

    Bernardi, G. (1956) Contribution à l’étude des catégories taxonomiques. I. Avant-propos, nomenclature et définitions. Bulletin de la Société entomologique de France, 61, 194–200.

    Bernardi, G. (1957) Contribution à l’étude des catégories taxonomiques. II. Les Règles internationales de la nomenclature zoologique et la notation des catégories taxonomiques. Bulletin de la Société entomologique de France, 62, 224–250.

    Bernardi, G. (1980) Les catégories taxonomiques de la systématique évolutive. In: Bocquet, C., Génermont, J. & Lamotte, M. (Eds.), Les problèmes de l’espèce dans le règne animal, Vol. 3, Mémoires de la Société zoologique de France, 40, 373–425.

    Blackwelder, R.E. (1967) Taxonomy: a text and reference book. John Wiley & Sons, New York, 713 pp.

    Brummitt, R.K. (1997) Taxonomy versus cladonomy, a fundamental controversy in biological systematics. Taxon, 46, 723–734.

    Bryant, H.N. (1996) Explicitness, stability, and universality in the phylogenetic definition and usage of taxon names: a case study of the phylogenetic taxonomy of the Carnivora (Mammalia). Systematic Biology, 45, 174–189.

    Cantino, P.D. (2000) Phylogenetic nomenclature: addressing some concerns. Taxon, 49, 85–93.

    Cantino, P.D., Olmstead, R.G. & Wagstaff, S.J. (1997) A comparison of phylogenetic nomenclature with the current system: a botanical case study. Systematic Biology, 46, 313–331.

    Carpenter, J.M. (2003) Critique of pure folly. The botanical Review, 69, 79–92.

    Claridge, M.F., Dawah, H.A. & Wilson, M.R. (Eds.) (1997a) Species, the units of biodiversity. Chapman & Hall, London, 455 pp.

    Claridge, M.F., Dawah, H.A. & Wilson, M.R. (1997b) Practical approaches to species concepts for living organisms. In: Claridge et al. (1997a), pp. 1–15.

    Cracraft, J. (1974) Phylogenetic models and classification. Systematic Zoology, 23, 71–90.

    Cracraft, J. (2000) Species concepts in theoretical and applied biology: a systematic debate with consequences. In: Wheeler & Meier (2000), pp. 3–14.

    Crowson, R.A. (1970) Classification and biology. Heinemann, London, 359 pp.

    Cuénot, L. & Tétry, A. (1951) L’évolution biologique. Les faits, les incertitudes. Masson, Paris, 597 pp.

    De Carvalho, M.R., Bockmann, F.A., Amorim, D.S., de Vivo, M., de Toledo-Piza, M., Menezes, N.A., de Figueiredo, J.L., Castro, R.M.C., Gill, A.C., McEachran, J.D., Compagno, L.J.V., Schelly, R.C., Britz, R., Lundberg, J.G., Vari, R.P. & Nelson, G. (2005) Revisiting the taxonomic impediment. Science, 307, 353.

    De Queiroz, K. (2006) The PhyloCode and the distinction between taxonomy and nomenclature. Systematic Biology, 55, 160–162.

    De Queiroz, K. & Cantino, P.D. (2001) Taxon names, not taxa, are defined. Taxon, 50, 821–826.

    De Queiroz, K. & Gauthier, J. (1990) Phylogeny as a central principle in taxonomy: phylogenetic definitions of taxon names. Systematic Zoology, 39, 307–322.

    De Queiroz, K. & Gauthier, J. (1992) Phylogenetic taxonomy. Annual Review of Ecology & Systematics, 23, 449–480.

    De Queiroz, K. & Gauthier, J. (1994) Toward a phylogenetic system of biological nomenclature. Trends in Ecology & Evolution, 9, 27–31.

    Delorme, M., Dubois, A., Grosjean, S. & Ohler, A. (2006) Une nouvelle ergotaxinomie des Megophryidae (Amphibia, Anura). Alytes, 24, 6–21.

    Delorme, M., Dubois, A., Kosuch, J. & Vences, M. (2004) Molecular phylogenetic relationships of Lankanectes corrugatus from Sri Lanka: endemism of South Asian frogs and the concept of monophyly in phylogenetic studies. Alytes, 22, 53–64.

    Dominguez, E. & Wheeler, Q.D. (1997) Taxonomic stability is ignorance. Cladistics, 13, 367–372.

    Doolittle, W.F. (1999) Phylogenetic classification and the universal tree. Science, 284, 2124–2128.

    Dubois, A. (1981) Hybridation interspécifique et notion de genre en zoologie. Comptes rendus de l’Académie des Sciences, (3), 292 (A), 201–203.

    Dubois, A. (1986) A propos de l’emploi controversé du terme “monophylétique”: nouvelles propositions. Bulletin mensuel de la Société linnéenne de Lyon, 55, 248–254.

    Dubois, A. (1988) The genus in zoology: a contribution to the theory of evolutionary systematics. Mémoires du Muséum national d’Histoire naturelle, (A), 140, 1–123.

    Dubois, A. (1991) Nomenclature of parthenogenetic, gynogenetic and “hybridogenetic” vertebrate taxons: new proposals. Alytes, 8, 61–74.

    Dubois, A. (1997) An evolutionary biologist’s view on the science of biology. Alytes, 15, 133–136.

    Dubois, A. (1998) List of European species of amphibians and reptiles: will we soon be reaching “stability”? Amphibia-Reptilia, 19, 1–28.

    Dubois, A. (2000) Synonymies and related lists in zoology: general proposals, with examples in herpetology. Dumerilia, 4, 33–98.

    Dubois, A. (2003) The relationships between taxonomy and conservation biology in the century of extinctions. Comptes rendus Biologies, 326 (suppl. 1), S9–S21.

    Dubois, A. (2004a) Developmental pathway, speciation and supraspecific taxonomy in amphibians. 1. Why are there so many frog species in Sri Lanka? Alytes, 22, 19–37.

    Dubois, A. (2004b) Developmental pathway, speciation and supraspecific taxonomy in amphibians. 2. Developmental pathway, hybridizability and generic taxonomy. Alytes, 22, 38–52.

    Dubois, A. (2005a) 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. Zoosystema, 27, 365–426.

    Dubois, A. (2005b) Proposals for the incorporation of nomina of higher-ranked taxa into the Code. Bulletin of zoological Nomenclature, 62, 200–209.

    Dubois, A. (2006a) Proposed Rules for the incorporation of nomina of higher-ranked zoological taxa in the International Code of Zoological Nomenclature. 2. The proposed Rules and their rationale. Zoosystema, 28, 165–258.

    Dubois, A. (2006b) New proposals for naming lower-ranked taxa within the frame of the International Code of Zoological Nomenclature. Comptes rendus Biologies, 329, 823–840.

    Dubois, A. (2006c) Incorporation of nomina of higher-ranked taxa into the International Code of Zoological Nomenclature: some basic questions. Zootaxa, 1337, 1–37.

    Dubois, A. (2006d) Naming taxa from cladograms: a cautionary tale. Molecular Phylogenetics & Evolution, 42, 317–330.

    Dubois, A. (2007) Naming taxa from cladograms: some confusions, misleading statements, and necessary clarifications. Cladistics, in press [doi: 10.1111/j.1096-0031.2007.00151.x].

    Dubois, A. & Günther, R. (1982) Klepton and synklepton: two new evolutionary systematics categories in zoology. Zoologischer Jahrbuch Systematik, 109, 290–305.

    Dubois, A. & Ohler, A. (1997) Early scientific names of Amphibia Anura. I. Introduction. Bulletin du Muséum national d’Histoire naturelle, (4), 18, 297–320.

    Dupuis, C. (1979) Permanence et actualité de la systématique: la “systématique phylogénétique” de W. Hennig (historique, discussion, choix de références). Cahiers des Naturalistes, Bulletin des Naturalistes parisiens, (n.s.), “1978”, 34 (1), 1–69.

    Eck, S. (1978) Die Entwicklung superspezifischer Begriffe in der zoologischen Taxonomie seit der Jahrhundertwende. Biologische Rundschau, 15, 98–103.

    Frost, D.R., Grant, T., Faivovich, J., Bazin, R.H., Haas, A., Haddad, C.F.B., de Sá, R.O., Channing, A., Wilkinson, M., Donnellan, S.C., Raxworthy, C.J., Campbell, J.A., Blotto, B.L., Moler, P., Drewes, R.C., Nussbaum, R.A., Lynch, J.D., Green, D.M. & Wheeler, W.C. (2006) The amphibian tree of life. Bulletin of the American Museum of Natural History, 297, 1–370.

    Gaffney, E.S. (1977) The side-necked turtle family Chelidae: a theory of relationships using shared derived characters. American Museum Novitates, 2620, 1–28.

    Gaffney, E.S. (1979) An introduction to the logic of phylogeny reconstruction. In: Cracraft, J. & Eldredge, N. (Eds.), Phylogenetic analysis and paleontology, Columbia University Press, New York, pp. 79–111.

    Génermont, J. & Lamotte, M (1980) Le concept biologique de l’espèce dans la zoologie contemporaine. In: Bocquet, C., Génermont, J. & Lamotte, M. (Eds.), Les problèmes de l’espèce dans le règne animal, Vol. 3, Mémoires de la Société zoologique de France, 40, 427–452.

    Ghiselin, M.T. (1966) An application of the theory of definitions to systematic principles. Systematic Zoology, 15, 127–130.

    Godfray, H.C.J. & Knapp, S., (eds.) (2004a) Taxonomy for the twenty-first century. Philosophical Transactions of the royal Society of London, (B), 359, 559–739.

    Godfray, H.C.J. & Knapp, S. (2004) Introduction. In: Godfray & Knapp (2004b), pp. 559–569.

    Haeckel, E. (1866) Generelle Morphologie der Organismen. Zweiter Band. Allgemeine Entwickelunggsgeschichte der Organismen. Georg Kramer, Berlin, 630 pp.

    Haffer, J. (1986) Superspecies and species limits in vertebrates. Zeitschrift für zoologische Systematik- und Evolutionsforschung, 24, 169–190.

    Hennig, W. (1936) Beziehungen zwischen geographischer Verbreitung und systematischer Gliederung bei einigen Dipterenfamilien: ein Beitrag zum Problem der Gliederung systematischer Kategorien höherer Ordnung. Zoologischer Anzeiger, 116, 161–175.

    Hennig, W. (1950) Grundzüge einer Theorie der phylogenetischen Systematik. Deutscher Zentralverlag, Berlin, 370 pp.

    Hennig, W. (1966) Phylogenetic systematics. Univ. Illinois Press, Urbana, Chicago & London, 270 pp.

    Hillis, D.M. (2006) Constraints in naming parts of the tree of life. Molecular Phylogenetics & Evolution, 42, 331–338.

    Hillis, D.M., Chamberlain, D.A., Wilcox, T.P. & Chippindale, P.T. (2001) A new species of subterranean blind salamander (Plethodontidae: Hemidactyliini: Eurycea: Typhlomolge) from Austin, Texas, and a systematic revision of central Texas paedomorphic salamanders. Herpetologica, 57, 266–280.

    Hillis, D.M. & Wilcox, T.P. (2005) Phylogeny of the New World true frogs (Rana). Molecular Phylogenetics and Evolution, 34, 299–314.

    Hubbell, S.P. (2001) The unified neutral theory of biodiversity and biogeography. Princeton University Press, Princeton, 458 pp.

    Jørgensen, P.M. (2000) Names are defined, but not as taxa. Taxon, 49, 779.

    Joyce, W.G., Parham, J.F. & Gauthier, J.A. (2004) Developing a protocol for the conversion of rank-based taxon names to phylogenetically defined clade names, as exemplified by turtles. Journal of Paleontology, 78, 989–1013.

    Kiriakoff, S.G. (1954) Chorologie et systématique phylogénétique. Bulletin et Annales de la Société royale d’Entologie de Belgique, 90, 185–198.

    Kiriakoff, S.G. (1965) La vicariance géographique et la taxonomie. Comptes rendus de la Société de Biogéographie, 41, 103–115.

    Kluge, A.G. (2005) Taxonomy in theory and practice, with arguments for a new phylogenetic system of taxonomy. In: Donnelly, M.H., Crother, B.I., Guyer, C., Wake, M.H. & White, M.E. (Eds.), Ecology and evolution in the tropics: a herpetological perspective, University of Chicago Press, Chicago, pp. 39–46.

    Knox, E. (1998) The use of hierarchies as organizational models in systematics. Biological Journal of the Linnean Society, 63, 1–49.

    Köhler, J., Vieites, D.R., Bonett, R.M., Hita García, F., Glaw, F., Steinke, D. & Vences, M. (2005) New amphibians and global conservation: a boost in species discoveries in a highly endangered vertebrate group. BioScience, 55, 693–696.

    Kuntner, M. & Agnarsson, I. (2006) Are the Linnean and phylogenetic nomenclatural systems combinable? Recommendations for biological nomenclature. Systematic Biology, 55, 774–784.

    Laurin, M. (2005) The advantages of phylogenetic nomenclature over Linnean nomenclature. In: Minelli, A., Ortalli, G. & Sanga, G. (Eds.), Animal names, Istituto Veneto di Scienze, Lettere ed Arti, Venezia, pp. 67–97.

    Laurin, M. & Cantino, P.D. (2004) First International Phylogenetic Nomenclature Meeting: a report. Zoologica scripta, 33, 475–479.

    Laurin, M. & Cantino, P.D. (2006) Second Meeting of the International Society for Phylogenetic Nomenclature: a report. Zoologica scripta, 36, 109–117.

    Laurin, M., de Queiroz, K. & Cantino, P.D. (2006) Sense and stability in taxon names. Zoologica scripta, 35, 113–114.

    Lecointre, G., Philippe, H., Lê, H.L.V. & Le Guyader, H. (1993) Species sampling has a major impact on phylogenetic inference. Molecular Phylogenetics & Evolution, 2, 205–224.

    Lherminier, P. & Solignac, M. (2000) L’espèce: définitions d’auteurs. Comptes rendus de l’Académie des Sciences de Paris, Sciences de la Vie, 323, 153–165.

    Lidén, M., Oxelman, B., Backlund, A., Andersson, L., Bremer, B., Eriksson, R., Moberg, R., Nordal, I., Persson, K., Thulin, M. & Zimmer, B. (1997) Charlie is our darling. Taxon, 46, 735–738.

    Linnaeus, C. (1758) Systema Naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio decima, reformata. Tomus I. Laurentii Salvii, Holmiae, 828 pp.

    Lotsy, J.P. (1916) Evolution by means of hybridization. Martinus Nijhoff, The Hague, 176 pp.

    Lotsy, J.P. (1918) La quintessence de la théorie du croisement. Archives néerlandaises des Sciences exactes et naturelles, (3B), 3, 351–353.

    Mayden, R.L. (1997) A hierarchy of species concepts: the denouement of the saga of the species problem. In: Claridge et al. (1997), pp. 381–424.

    Mayr, E. (1982) The growth of biological thought. Belknap Press, Cambridge, Mass. & London, 987 pp.

    Mayr, E. (1995) Systems of ordering data. Biology & Philosophy, 10, 419–434.

    Mayr, E. (1997) This is biology. The science of the living world. Belknap Press, Cambridge, Mass. & London, 344 pp.

    Mayr, E. & Ashlock, P.D. (1991) Principles of systematic zoology. Second edition. McGraw-Hill, New York, 495 pp.

    McNeill, J., Barrie, F.R., Burdet, H.M., Demoulin, V., Hawksworth, D.L., Marhold, K., Nicolson, D.H., Prado, J., Silva, P.C., Skog, J.E., Wiersema, J.H. & Turland, N.J., (Eds.) (2006) International code of botanical nomenclature (Vienna Code) adopted by the Seventeenth International Botanical Congress, Vienna, Austria, July 2005. Koeltz, Königstein, 586 pp.

    Minelli, A. (1991) Biological systematics: the state of the art. Chapman & Hall, London, 403 pp.

    Minelli, A. (2000) The ranks and the names of species and higher taxa, or a dangerous inertia of the language of natural history. In: Ghiselin, M. T. & Leviton, A. E. (Eds.), Cultures and institutions of natural history: essays in the history and philosophy of sciences, California Academy of Sciences, San Francisco, pp. 339–351.

    Moore, G. (1998) A comparison of traditional and phylogenetic nomenclature. Taxon, 47, 561–579.

    Moore, G. (2003) Should taxon names be explicitly defined? The botanical Review, 69, 2–21.

    Ohler, A. (2005) Le mode de fonctionnement des systèmes nomenclaturaux en zoologie et en botanique. In: Dubois, A., Poncy, O., Malécot, V. & Léger, N. (Eds.), Comment nommer les taxons de rang supérieur en zoologie et en botanique?, Biosystema, 23, 11–16.

    Padial J.M. & De la Riva, I. (2006) Taxonomic inflation and the stability of species lists: the perils of ostrich’s behavior. Systematic Biology, 55, 859–867.

    Papavero, N., Llorente-Bousquets, J. & Abe, J.M. (2001) Proposal of a new system of nomenclature for phylogenetic systematics. Arquivos de Zoologia, São Paulo, 36, 1–145.

    Patterson, C. & Rosen, D.E. (1977) Review of ichthyodectiform and other Mesozoic teleost fishes and the theory and practice of classifying fossils. Bulletin of the american Museum of natural History, 158, 81–172.

    Pennisi, E. (2001) Linnaeus’s last stand? Science, 291, 2304–2305, 2307.

    Pickett, K.M. (2005) Is the PhyloCode now roughly analogous to the actual codes? A reply to Laurin et al. Cladistics, 21, 608–610.

    Pleijel, F. & Rouse, G.W. (2000) Least-inclusive taxonomic unit: a new taxonomic concept for biology. Proceedings of the royal Society of London, (B), 267, 627–630.

    Pleijel, F. & Rouse, G.W. (2003) Ceci n’est pas une pipe: names, clades and phylogenetic nomenclature. Journal of systematic and evolutionary Research, 41, 162–174.

    Polaszek, A. & Wilson, E.O. (2005) Sense and stability in animal names. Trends in Ecology and Evolution, 20, 421–422.

    Raikow, R.J. (1985) Problems in avian classification. In: Johnston, R. F. (Ed.), Current Ornithology, Vol. 2, Plenum, New York, pp. 187–212.

    Reif, W.-E. (2003) Problematic issues of cladistics. 1. Ancestor recognition and phylogenetic classification. Neues Jahrbuch für Geologie und Paläontologie Abhandlungen, 230, 97–143.

    Rieppel, O. (2006) The PhyloCode: a critical discussion of its theoretical foundation. Cladistics, 22, 186–197.

    Schaefer, W.W. (1976) The reality of the higher taxonomic categories. Zeitschrift für zoologischer Systematik und Evolutions-Forschung, 14, 1–10.

    Schuh, R.T. (2003) The Linnaean system and its 250-year persistence. The botanical Review, 69, 59–78.

    Sibley, C.G. & Ahlquist, J.E. (1982) The relationships of the yellow-breasted chat (Icteria virens) and the alleged slowdown in the rate of macromolecular evolution in birds. Postilla, 187, 1-19.

    Sibley, C.G. & Ahlquist, J.E. (1990) Phylogeny and classification of birds. A study in molecular evolution. Yale University Press, New Haven & London, 999 pp.

    Simpson, G.G. (1940) Types in modern taxonomy. American Journal of Science, 238, 413–431.

    Simpson, G.G. (1961) Principles of animal taxonomy. Columbia Univ. Press, New York, 259 pp.

    Sluys, R., Martens, K. & Schram, F.R. (2004) The PhyloCode: naming of biodiversity at a crossroads. Trends in Ecology and Evolution, 19, 280–281.

    Smith, A.B. (1988) Patterns of diversification and extinction in early Palaeozoic echinoderms. Palaeontology, 31, 799–828.

    Stuart, S.N., Chanson, J.S., Cox, N.A., Young, B.E., Rodrigues, A.S.L., Fischman, D.L. & Waller, R.W. (2004) Status and trends of amphibian declines and extinctions worldwide. Science, 306, 1783–1786.

    Stuessy, T.F. (2000) Taxon names are not defined. Taxon, 49, 231–233.

    Stuessy, T.F. (2001) Taxon names are still not defined. Taxon, 50, 185–186.

    Sundberg, P. & Pleijel, F. (1994) Phylogenetic classification and the definition of taxon names. Zoologica scripta, 23, 19–25.

    Turesson, G. (1922) The genotypical response of the plant species to the habitat. Hereditas, 3, 211–350.

    Van Valen, L. (1973) Are categories in different phyla comparable? Taxon, 22, 333–373.

    Vuilleumier, F. (1976) La notion d’espèce en ornithologie. In: Bocquet, C., Génermont, J. & Lamotte M. (Eds.), Les problèmes de l’espèce dans le règne animal, Vol. 1, Mémoires de la Société zoologique de France, 38, 29-65.

    Wheeler, Q.D. & Meier, R. (Eds.) (2000) Species concepts and phylogenetic theory. Columbia University Press, New York, 243 pp.

    Wheeler, Q.D., Raven, P.H. & Wilson, E.O. (2004) Taxonomy: impediment or expedient? Science, 303, 285.

    White, M.J.D. (1978) Modes of speciation. Freeman, San Francisco, 463 pp.

    Wiley, E.O. (1981) Phylogenetics. The theory and practice of phylogenetic systematics. Wiley, New York, 454 pp.

    Wilson, E.O. (1985) The global biodiversity crisis:a challenge to science. Issues in Science & Technology, 2, 20–29.

    Wilson, E.O. (2004) Taxonomy as a fundamental discipline. In: Godfray & Knapp (2004a), p. 739.