Abstract
Taxon names are at the core of communication related to biodiversity. Thus, systems that regulate such names should prevent unnecessary changes. Unfortunately, the current regulatory codes have some articles that produce confusion or, even, generate name instability. To promote nomenclatural stability, while maintaining a system that effectively conveys information about biological diversity and its evolution, we argue in favor of emendations to the current International Code of Zoological Nomenclature (ICZN). We propose that taxon names should be created within an evolutionary framework; ideally, this would mean including a phylogenetic tree or, at the very least, formulating a hypothesis in words based on explicit arguments (homologous traits) for the evolutionary relatedness of taxa, avoiding subjective perceptions. Additionally, we consider that some relatively minor changes of the ICZN would greatly increase name stability; the most relevant modifications that would facilitate long-term stability are: (i) immutability of the species epithet of species names, (ii) intercalation of unranked taxa between formal Linnaean ranks, (iii) elimination of homonyms in taxon names at the genus-group level, and (iv) inclusion of flexible phylogenetic definitions when supraspecific taxa are named.
References
Artois, T. (2001) Phylogenetic nomenclature: the end of binominal nomenclature? Belgian Journal of Zoology, 131, 87–89.
Avise, J.C. (2008) Three ambitious (and rather unorthodox) assignments for the field of biodiversity genetics. Proceedings of the National Academy of Sciences USA, 105, 11564–11570.
Bartolous, A. (2008) Error Cascades in the Biological Sciences: The Unwanted Consequences of Using Bad Taxonomy in Ecology. AMBIO: A Journal of the Human Environment, 37, 114–118.
Baum, D.A., Alverson, W.S., Nyffeler, R. (1998) A durian by any other name: taxonomy and nomenclature of the core Malvales. Harvard Papers in Botany, 3, 313–330.
Benton, M.J. (2000) Stems, nodes, crown clades, and rank-free lists: is Linnaeus dead? Biological Reviews, 75, 633–648.
Berry, P.E. (2002) Biological inventories and the PhyloCode. Taxon, 51, 27–29.
Bertrand, Y. & Härlin, M. (2006) Stability and universality in the application of taxon names in phylogenetic nomenclature. Systematic Biology, 55, 848–858.
Bryant, H.N. (1994) Comments on the phylogenetic definition of taxon names and conventions regarding naming of crown clades. Systematic Biology, 43, 124–130.
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. & de Queiroz, K. (2007) PhyloCode: International Code of Phylogenetic Nomenclature. Version 4b. Available from: http://www.ohiou.edu/phylocode/toc.html (accessed 13 July 2020)
Cantino, P.D. & de Queiroz, K. (2010) PhyloCode: International Code of Phylogenetic Nomenclature. Version 4c. Available from: http://www.ohio.edu/phylocode/PhyloCode4c.pdf (accessed 13 July 2020)
Cantino, P.D., Bryant, H.N., de Queiroz, K., Donoghue, M.J., Eriksson, T., Hillis, D. & Lee, M.S.Y. (1999) Species names in phylogenetic nomenclature. Systematic Biology, 48, 790–807.
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.
Dayrat, B., Cantino, P.D., Clarke, J.A. & de Queiroz, K. (2008) Species names in the PhyloCode: The approach adopted by the international society for phylogenetic nomenclature. Systematic Biology, 57, 507–514.
de Queiroz, K. (1995) Phylogenetic approaches to classification and nomenclature, and the history of taxonomy (an alternative interpretation). Herpetological Review, 26, 79–81.
de Queiroz, K. (1997a) Misunderstandings about the phylogenetic approach to biological nomenclature: a reply to Lid´en and Oxelman. Zoologica Scripta, 26, 67–70.
de Queiroz, K. (1997b) The Linnean hierarchy and the evolutionization of taxonomy, with emphasis on the problem of nomenclature. Aliso, 15, 115–144.
de Queiroz, K. (2000) The definitions of taxon names: a reply to Stuessy. Taxon, 49, 533–537.
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) Phylogenetic nomenclature and the PhyloCode. Bulletin of Zoological Nomenclature, 58, 254–271.
de Queiroz, K. & Donoghue, M. (2001) Taxing debate for taxonomists. Science, 292, 2249–2250.
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 and Systematics, 23, 449–480.
de Queiroz, K. & Gauthier, J. (1994) Toward a phylogenetic system of biological nomenclature. Trends in Ecology and Evolution, 9, 27–31.
Darwin, C.R. (1859) On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life. 1st Edition. John Murray, London, x + 502 pp.
Dubois, A. (2007a) Naming taxa from cladograms: a cautionary tale. Molecular Phylogenetics and Evolution, 42, 317–330.
Dubois, A. (2007b) Naming taxa from cladograms: some confusions, misleading statements, and necessary clarifications. Cladistics, 23, 390–402.
Dubois, A. (2008) Phylogenetic hypotheses, taxa and nomina in zoology. Zootaxa, 1950, 51–86.
Dubois, A. (2001) The International Code of Zoological Nomenclature must be drastically improved before it is too late. Bionomina, 2, 1–104.
Dubois, A., Bauer, A.M., Ceríaco, L.M.P., Dusoulier, F., Frétey, T., Lobl, I., Lorvelec, O., Ohler, A., Stopiglia, R. & Aescht, E. (2019) The Linz Zoocode project: a set of new proposals regarding the terminology, the Principles and Rules of zoological nomenclature. First report of activities (2014‒2019). Bionomina, 111, 1–111.
Forey, P.L. (2001) The PhyloCode: description and commentary. Bulletin of Zoological Nomenclature, 58, 81–96.
Forey, P.L. (2002) PhyloCode: pain, no gain. Taxon, 51, 43–54.
Frost, D.R., Grant, T., Faivovich, J., Bain, 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.E., 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
Hillis, D. (2006) Constrains in naming parts of the tree of life. Molecular Phylogenetics & Evolution, 42, 331–338.
Guayasamin, J.M., Castroviejo-Fisher, S., Trueb, L., Ayarzagüena, J., Rada, M. & Vilà, C. (2009) Phylogenetic systematics of glassfrogs (Amphibia: Centrolenidae) and their sister taxon Allophryne ruthveni. Zootaxa, 2100, 1–97.
Greuter, W., Hawksworth, D.L., McNeill, J., Mayo, M.A., Minelli, A., Sneath, P.H.A., Tindall, B.J., Trehane, P. & Tubbs, P. (1998) Draft BioCode (1997): the prospective international rules for the scientific names of organisms. Taxon, 47, 127–149.
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. University Illinois Press, Urbana, Chicago & London, III + 263 S.
Hillis, D.M. (2007) Constraints in naming parts of the tree of life. Molecular Phylogenetics and Evolution, 42, 331–338.
Hillis, D.M. & Wilcox, T.P. (2005) Phylogeny of the New World true frogs (Rana). Molecular Phylogenetics and Evolution, 34, 299–314.
International Code of Nomenclature of Bacteria (1992) American Society for Microbiology.
International Commission on Zoological Nomenclature. (1999) International Code of Zoological Nomenclature. 4th Edition. International Trust for Zoological Nomenclature, London, 306 pp.
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 Paleontolology, 78, 989–1013.
Kaiser, H., Crother, B.I., Kelly, C.M.R., Luiselli, L., O’shea, M., Hidetoshi, O., Passos, P., Schleip, W.D. & Wüster, W. (2013) Best practices: In the 21st century, taxonomic decisions in herpetology are acceptable only when supported by a body of evidence and published via peer review. Herpetological Review, 44, 8–23.
Kojima, J. (2003) Apomorphy-based definition also pinpoints a node, and PhyloCode names prevent effective communication. Botanical Review, 69, 44–58.
Kraus, O. (2004) Phylogeny, classification and nomenclature: a reply to F. Pleijel and G. W. Rouse. Journal of Zoological Systematics and Evolutionary Research, 42, 159–161.
Kuntner, M. & Agnarsson, I. (2006) Are the Linnean and phylogenetic nomenclatural systems combinable? Recommendations for biological nomenclature. Systematic Biology, 55, 774–784.
Laurin, M. & Cantino, P.D. (2007) Second Meeting of the International Society for Phylogenetic Nomenclature: a Report. Zoologica Scripta, 36, 109–117.
Laurin, M., de Queiroz, K. & Cantino, P. (2006) Sense and stability of taxon names. Zoologica Scripta, 35, 113–114.
Laurin, M., de Queiroz, K., Cantino, P., Cellinese, N. & Olmstead, R. (2005) The PhyloCode, types, ranks and monophyly: a response to Pickett. Cladistics, 21, 605–607.
Lee, M.S.Y. (1999) Reference taxa and phylogenetic nomenclature. Taxon, 48, 31–34.
Lee, M.S.Y. (2001) On recent arguments for phylogenetic nomenclature. Taxon, 50, 175–180.
Lee, M.S.Y. (2005) Choosing reference taxa in phylogenetic nomenclature. Zoologica Scripta, 34, 329–331.
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, Homiae, 823 pp.
Moore, G. (1998) A comparison of traditional and phylogenetic nomenclature. Taxon, 47, 561–579.
Nixon, K.C., Carpenter, J.M. & Stevenson, D.W. (2003) The PhyloCode is fatally flawed, and the “Linnean” system can easily be fixed. Botanical Review, 69, 111–120.
Pennisi, E. (2001) Linnaeus’s Last Stand? Science, 291, 2304–2307.
Pickett, K.M. (2005) The new and improved PhyloCode, now with types, ranks, and even polyphyly: a conference report from the First International Phylogenetic Nomenclature Meeting. Cladistics, 21, 79–82.
Stuessy, T.F. (2000) Taxon names are not defined. Taxon, 49, 231–233.
Taylor, E.H. (1951) Two new genera and a new family of tropical American frogs. Proceedings of the Biological Society of Washington, 64, 33–40.
Thomson, S.A., Pyle, R.L., Ahyong, S.T., Alonso-Zarazaga, M., Ammirati, J., Araya, J.F., Ascher, J.S., Audisio, T.L., Azevedo-Santos, V.M., Bailly, N., Baker, W.J., Balke, M., Barclay, M.V.L., Barrett, R.L., Benine, R.C., Bickerstaff, J.R.M., Bouchard, P., Bour, R., Bourgoin, T. & Zhou, H.-Z. (2018) Taxonomy based on science is necessary for global conservation. PLOS Biology, 16 (3), e2005075.
https://doi.org/10.1371/journal.pbio.2005075
Vences, M., Guayasamin, J.M., Miralles, A. & de La Riva, I. (2013) To name or not to name: criteria to promote economy of change in Linnaean classification schemes. Zootaxa, 3636, 201–244.
Withgott, J. (2000) Is it “So Long, Linnaeus?” Bioscience, 50, 646–651.
Wolsan, M. (2007) Naming species in phylogenetic nomenclature. Systematic Biology, 56, 1011–1021.