Australian Systematic Botany Australian Systematic Botany Society
Taxonomy, biogeography and evolution of plants
L. A. S. JOHNSON REVIEW

An end to all things? — plants and their names

Peter F. Stevens

Missouri Botanical Garden, PO Box 299, St Louis, MO 63166-0299, USA. Email: peter.stevens@mobot.org

Australian Systematic Botany 19(2) 115-133 https://doi.org/10.1071/SB05011
Submitted: 6 May 2005  Accepted: 3 October 2005   Published: 28 April 2006

Abstract

Great advances in our understanding of phylogenetic relationships have been made over the last decade and a half. Major clades in many groups, including flowering plants, now show substantial stability both in terms of content and relationships. This makes possible the development of a system in which only monophyletic ( = holophyletic) entities are named, entities that represent all and only the descendants of a common ancestor. However, some argue that use of Linnaean ranked names is inappropriate in such circumstances; this argument is bolstered by appeals to history and philosophy. Those who doubt the wisdom and / or very possibility of naming only monophyletic groups also argue that their position follows from history, or that ancestors cannot be incorporated into a Linnaean-type classification and that ancestors are an integral part of monophyletic groups. However, I argue that most of the apparently more cosmic issues brought up in this debate are based on a combination of a misunderstanding of the nature and purpose of language, fallacious reasoning and dubious—and largely irrelevant—interpretations of history. A flagged hierarchy helps memory and communication. Binomials in particular simply represent the noun–adjective combinations of ordinary language in a Latinised form, and are too valuable a communication device to be discarded because rank has been demonised. However, hierarchies can be misinterpreted and cannot be made complex enough to cope with the much more detailed phylogenies being produced. Thinking of naming systems as conventions may help clarify what we should be doing, if we are not to squander both the time and the reputation of systematics. Time is in short supply and our reputation not what it might be; solving the less cosmic issues may involve a self-discipline that also seems in short supply in the systematic community.


Acknowledgments

I thank two reviewers for their comments, and Brent Mishler and Marc Sosef for raising some interesting issues. I am particularly grateful to Andrew Doust, Micah Dunthorn, Lucia Lohmann, Beto Vicentini and Felipe Zapata for discussion and help with the figures, and to participants at the 2003 annual conference of the Australian Systematic Botany Society, Oregon State University, the Systematics Discussion Group at the Missouri Botanical Garden, and the BioLunch at the University of Missouri, Saint Louis, for helping me clarify several points.


References


Aarsleff, H (1976). Wilkins, John. In ‘Dictionary of scientific biography, vol. XIV, Verrill-Zwelfer’. pp. 361–381. (Charles Scribner’s Sons: New York)

Anderberg AA, Rydin C, Kallersjö M (2002) Phylogenetic relationships in the order Ericales s.l.: analyses of molecular data from five genes from the plastid and mitochondrial genomes. American Journal of Botany 89, 677–687. open url image1

Angiosperm Phylogeny Group Bremer K, Chase MW, Stevens PF (1998) An ordinal classification for the families of flowering plants. Annals of the Missouri Botanical Garden 85, 531–553. open url image1

Angiosperm Phylogeny Group Bremer B, Bremer K, Chase MW, Reveal JL, Soltis DE, Soltis PS, Stevens PF (2003) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Botanical Journal of the Linnean Society 141, 399–436.
CrossRef | open url image1

Ashlock PD (1971) Monophyly and associated terms. Systematic Zoology 20, 63–69. open url image1

Backlund A, Bremer K (1998) To be or not to be—principles of classification and monotypic plant families. Taxon 47, 391–401. open url image1

Barkley TM, DePriest P, Funk V, Kiger RW, Kress WJ, Moore G (2004a) Linnaean nomenclature in the 21st century: a report from a workshop on integrating traditional nomenclature and phylogenetic classification. Taxon 53, 153–158. open url image1

Barkley TM, DePriest P, Funk V, Kiger RW, Kress WJ, McNeill J, Moore G, Nicolson DH, Stevenson DW, Wheeler QD (2004b) A review of the International Code of Botanical Nomenclature with respect to its compatibility with phylogenetic classification. Taxon 53, 159–161. open url image1

Baum, DA (2004). Reticulate genealogy and its bearing on the PhyloCode. In ‘First international phylogenetic nomenclature meeting’. [Abstracts]. pp. 1.

Baum DA, Alverson WS, Nyffeler R (1998) A durian by any other name: taxonomy and nomenclature of core Malvales. Harvard Papers in Botany 3, 315–330. open url image1

Baum DA, Smith SW, Yen A, Alverson WS, Nyffeler R, Whitlock BA, Oldham RA (2004) Phylogenetic relationships of Malvatheca (Bombacoideae and Malvoideae; Malvaceae sensu lato) as inferred from plastid DNA sequences. American Journal of Botany 91, 1863–1871. open url image1

Bentham G (1875) On the recent progress and present state of systematic botany. Report of the British Association for the Advancement of Science 1874, 27–54. open url image1

Benton MJ (2000) Stems, nodes, crown clades, and rank-free lists: is Linnaeus dead? Biological Review 75, 633–648. open url image1

Bergthorsson U, Richardson AO, Young GJ, Goertzen LR, Palmer JD (2004) Massive horizontal transfer of mitochondrial genes from diverse land plant donors to the basal angiosperm Amborella. Proceedings of the National Academy of Sciences USA 101, 17747–17752.
CrossRef | open url image1

Berlin, B (1992). ‘Ethnobiological classification: principles of categorization of plants and animals in traditional societies.’ (Princeton University Press: Princeton)

Bonde, N ,  and  Westergaard, B (2004). New non-Linnaean, neo-cladistic nomenclature and classification conventions exemplified by recent and fossil hominids. In ‘First international phylogenetic nomenclature meeting’. [Abstracts]. pp. 34.

Boyd, R (1999). Homeostasis, species and higher taxa. In ‘Species: new interdisciplinary essays’. pp. 141–185. (MIT Press: Cambridge, MA)

Briggs BG (1996) L.A.S. Johnson—a botanical career. Telopea 6, 511–520. open url image1

Brown, R (1818). Observations, systematical and geographical, on Professor Christian Smith’s collection of plants from the vicinity of the River Congo. In ‘Narrative of an expedition to explore the River Zaire,...’. pp. 420–488. (John Murray: London)

Brummitt RK (1997) Taxonomy versus cladonomy, a fundamental controversy in biological systematics. Taxon 46, 723–734. open url image1

Brummitt RK (2002) How to chop up a tree. Taxon 51, 31–41. open url image1

Brummitt RK (2003) Further dogged defense of paraphyletic taxa. Taxon 52, 803–804. open url image1

Brummitt RK, Sosef MSM (1998) Paraphyletic taxa are inherent in Linnaean classification—a reply to Freudenstein. Taxon 47, 411–412. open url image1

Bryant HN (1994) Comments on the phylogenetic definition of taxon names and conventions regarding the naming of crown clades. Systematic Biology 43, 124–130. open url image1

Bryant HN (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. open url image1

Bryant HN, Cantino PD (2002) A review of criticisms of phylogenetic nomenclature: is taxonomic freedom the fundamental issue? Biological Reviews 77, 39–55. open url image1

Buck RC, Hull DL (1969) Reply to Gregg. Systematic Zoology 18, 354–357. open url image1

Cain AJ (1958) Logic and memory in Linnaeus’s system of taxonomy. Proceedings of the Linnean Society of London 169, 144–163. open url image1

Cain AJ (1959a) Deductive and inductive methods in post-Linnaean taxonomy. Proceedings of the Linnean Society of London 170, 185–217. open url image1

Cain AJ (1959b) The post-Linnaean development of taxonomy. Proceedings of the Linnean Society of London 170, 233–244. open url image1

de Candolle, A-P (1813). ‘Théorie élémentaire de la botanique.’ (Déterville: Paris)

de Candolle, A-P (1844). ‘Théorie élémentaire de la botanique’. 3rd edn . (Roret: Paris)

Cantino PD (1998) Binomials, hyphenated uninomials, and phylogenetic nomenclature. Taxon 47, 425–429. open url image1

Cantino PD (2000) Phylogenetic nomenclature: addressing some concerns. Taxon 49, 85–93. open url image1

Cantino PD (2004) Classifying species versus naming clades. Taxon 53, 795–798. open url image1

Cantino, PD ,  and  de Queiroz, K (2005). ‘PhyloCode: a phylogenetic code of biological nomenclature.’ [ consulted 6 Jan 2005].

Cantino PD, Olmstead RG, Wagstaff SJ (1997) A comparison of phylogenetic nomenclature with the current system: a botanical case study. Systematic Biology 46, 313–331. open url image1

Cantino PD, Bryant HN, De Queiroz K, Donoghue MJ, Eriksson T, Hillis DM, Lee MSY (1999) Species names in phylogenetic nomenclature. Systematic Biology 48, 790–807.
CrossRef | open url image1

Carpenter JM (2003) Critique of pure folly. Botanical Review 69, 79–92. open url image1

Cellinese, N ,  and  Donoghue, MJ (2004). The demise of the ranked system in the arrangement of herbaria: utopia or reality? In ‘First international phylogenetic nomenclature meeting’. [Abstracts]. pp. 2.

Chung C (2003) On the origin of the typological / population distinction in Ernst Mayr’s changing views of species, 1942–1959. Studies in History and Philosophy of Biology and Biomedical Sciences 34C, 277–296.
CrossRef | open url image1

Coyne, JA ,  and  Orr, HA (2004). ‘Speciation.’ (Sinauer: Sunderland, MA)

Crisp MJ, Chandler GT (1996) Paraphyletic species. Telopea 6, 813–844. open url image1

Cronquist, A (1981). ‘An integrated system of classification of flowering plants.’ (Columbia University Press: New York)

Cronquist A (1987) A botanical critique of cladism. Botanical Review 53, 1–52. open url image1

Davis CC, Chase MW (2004) Elatinaceae are sister to Malpighiaceae: Peridiscaceae belong to Saxifragales. American Journal of Botany 91, 262–273. open url image1

Davis CC, Wurdack KJ (2004) Host-to-parasite gene transfer: evidence from Malpighiales. Science 305, 676–678.
CrossRef | open url image1

Davis, PD ,  and  Heywood, VH (1963). ‘Principles of angiosperm taxonomy.’ (Oliver & Boyd: Edinburgh)

Darwin, CD (1859). ‘On the origin of species...’ (John Murray: London)

Dayrat, B (2004). Selecting a form of species name in the PhyloCode. In ‘First international phylogenetic nomenclature meeting’. [Abstracts]. pp. 3.

Dayrat B, Schander C, Angielczyk KD (2004) Suggestions for a new species nomenclature. Taxon 53, 485–491. open url image1

Dobzhansky, T (1951). ‘Genetics and the origin of species.’ 3rd edn . (Columbia University Press: New York)

Dominguez E, Wheeler QD (1997) Taxonomic stability is ignorance. Cladistics 13, 367–372. open url image1

Donoghue MJ, Cantino PD (1988) Paraphyly, ancestors and the goals of taxonomy: a botanical defence of cladism. Botanical Review 54, 107–128. open url image1

Donoghue, MJ ,  and  Doyle, JA (1989). Phylogenetic analysis of angiosperms and relationships of Hamamelidae. In ‘Evolution, systematics and fossil history of the Hamamelidae, vol. 1. Introduction and ‘lower’ Hamamelidae’. pp. 17–45. (Clarendon Press: Oxford)

Eigen E (1997) Overcoming first impressions: Georges Cuvier’s types. Journal of the History of Biology 30, 179–209.
CrossRef | open url image1

Ereshefsky, M (2000). ‘The poverty of the Linnaean hierarchy: a philosophical study of biological taxonomy.’ (Cambridge University Press: Cambridge)

Evans RC, Campbell CS (2002) The origin of the apple subfamily (Maloideae; Rosaceae) is clarified by DNA sequence data from duplicated GBSSI genes. American Journal of Botany 89, 1478–1484. open url image1

Farber PL (1976) The type concept in zoology in the first half of the nineteenth century. Journal of the History of Biology 9, 93–119.
CrossRef | open url image1

Fisher, KM ,  and  Mishler, BD (2004). Monography and the PhyloCode: a practical example from the moss clade Leucophanella. In ‘First international phylogenetic nomenclature meeting’. [Abstracts]. pp. 12.

Forey PL (2002) Phylocode—no pain, no gain. Taxon 51, 43–54. open url image1

Forey PL (2005) Naming the world: is there anything left of Linnaeus? Proceedings of the California Academy of Sciences 56, suppl. 1, 182–195. open url image1

Freudenstein JV (1998) Paraphyly, ancestors and classification—a response to Sosef and Brummitt. Taxon 47, 95–104. open url image1

Futuyuma, D (1998). ‘Evolutionary biology.’ 3rd edn . (Sinauer: Sunderland, MA)

Gauthier, JA , de Queiroz, K , Joyce, WG , Parham, JF , Rowe, T ,  and  Calrake, J (2004). A phylogenetic nomenclature for the major clades of Amniota Haeckel 1866, with emphasis on non-avian Reptilia Laurentus 1768. In ‘ First international phylogenetic nomenclature meeting’. [Abstracts]. pp. 14.

Gelman, SA (2003). ‘The essential child.’ (Oxford University Press: Oxford)

Ghiselin MT (1974) A radical solution to the species problem. Systematic Zoology 23, 536–544. open url image1

Ghiselin MT (2005) Taxonomy as the organization of knowledge. Proceedings of the California Academy of Sciences 56, suppl. 1, 161–169. open url image1

Gilmour, JSL (1940). Taxonomy and philosophy. In ‘The new systematics’. pp. 461–474. (Clarendon Press: Oxford)

Govaerts R (2003) How many species of seed plants are there? — a response. Taxon 52, 583–584. open url image1

Grant V (2003) Incongruence between cladistic and taxonomic systems. American Journal of Botany 90, 1263–1270. open url image1

Grass Phylogeny Working Group (2001) Phylogeny and subfamilial classification of the grasses (Poaceae). Annals of the Missouri Botanical Garden 88, 373–457. open url image1

Gregg JR (1950) Taxonomy, language and reality. American Zoologist 84, 419–435. open url image1

Gregg JR (1968) Buck and Hull: a critical rejoinder. Systematic Zoology 17, 342–345. open url image1

Greuter W (2005) Linnean nomenclature and our nomenclatural codes: how many do we need? Symbolae Botanicae Upsalienses 33(3), 119–125. open url image1

Greuter, W , McNeill, J , Barrie, FR , Burdet, HM , Demoulin, V , Filguieras, TS , Nicholson, DH , Sliva, PC , Skog, JE , Trehane, P , Turland, NJ ,  and  Hawksworth, DL (2000). ‘The international code of botanical nomenclature.’ (Koeltz: Königstein)

Griffiths GCD (1973) Some fundamental problems in biological classification. Systematic Zoology 22, 338–343. open url image1

Griffiths CGD (1974) On the foundations of biological systematics. Acta Biotheoretica 23, 85–131.
CrossRef | open url image1

Griffiths GCD (1976) The future of Linnaean nomenclature. Systematic Zoology 25, 168–173. open url image1

Hennig, W (1966). ‘Phylogenetic systematics’. (University of Illinois Press: Urbana)

Heywood V (2001) Floristics and monography—an uncertain future? Taxon 50, 361–380. open url image1

Hibbett DS, Donoghue MJ (1998) Intergrating phylogenetic analysis and classification in fungi. Mycologia 90, 347–356. open url image1

Hibbett DS, Nilsson RH, Snyder M, Fonseca M, Costanzo J, Shonfeld M (2005) Automated phylogenetic taxonomy: an example in the homobasidiomycetes (mushroom-forming fungi). Systematic Biology 54, 660–668.
CrossRef | open url image1

Holman EW (2002) The relation between folk and scientific classifications of plants and animals. Journal of Classification 19, 131–159.
CrossRef | open url image1

Holman EW (2005) Domain-specific and general properties of folk classifications. Journal of Ethnobiology 25, 71–91. open url image1

Hull DL (1965) The effect of essentialism in taxonomy—two thousand years of stasis. The British Journal for the Philosophy of Science 15, 314–326. open url image1

Hull DL (1976) Are species really individuals? Systematic Zoology 25, 174–191. open url image1

Hull DL, Snyder DP (1969) Contemporary logic and evolutionary taxonomy: a reply to Gregg. Systematic Zoology 18, 347–354. open url image1

Humphries CJ, Chappill JA (1988) Systematics as science: a response to Cronquist. Botanical Review 54, 129–144. open url image1

Jain R, Rivera MC, Lake JA (1999) Horizontal gene transfer among genomes: the complexity hypothesis. Proceedings of the National Academy of Sciences USA 96, 3801–3806.
CrossRef | open url image1

Janovec JP, Clark LG, Mori SA (2003) Is the neotropical flora ready for the Phylocode? Botanical Review 69, 22–43. open url image1

Johnson LAS (1968) Rainbow’s end: the quest for an optimal taxonomy. Proceedings of the Linnean Society of New South Wales 93, 8–45. open url image1

Jorgensen PM (2002) Two nomenclatural systems? Taxon 51, 737. open url image1

Jorgensen PM (2004) Rankless names in the Code? Taxon 53, 162. open url image1

Judd, WS , Campell, CA , Kellogg, EA , Donoghue, MJ ,  and  Stevens, PF (2003). ‘Plant systematics, a phylogenetic approach.’ 2nd edn . (Sinauer: Sunderland, Mass)

Keller RA, Boyd RN, Wheeler QD (2003) The illogical basis of phylogenetic nomenclature. Botanical Review 69, 93–110. open url image1

Knowlson, J (1975). ‘Universal language schemes in England and France, 1600–1800.’ (University of Toronto Press: Toronto)

Knox EB (1998) The use of hierarchies as organisational models in systematics. Biological Journal of the Linnaean Society 63, 1–49.
CrossRef | open url image1

Kojima J (2003) Apomorphy-based definition also pinpoints a node, and phylocode names prevent effective communication. Botanical Review 69, 44–58. open url image1

Kress WJ, Prince LM, Williams KJ (2002) The phylogeny and a new classification of gingers: evidence from molecular data. American Journal of Botany 89, 1682–1696. open url image1

Kron KA (1997) Exploring alternative systems of classification. Aliso 15, 105–112. open url image1

Kron KA, Judd WS, Stevens PF, Crayn DM, Anderberg AA, Gadek PA, Quinn CJ, Luteyn JL (2002) Phylogenetic classification of Ericaceae: molecular and morphological evidence. Botanical Review 68, 335–423. open url image1

Lamarck, JBPA ,  and  de, M de (1778). ‘Flore Françoise.’ 3 vols. (Imprimerie Royale: Paris)

Laurin M, Anderson JS (2004) Meaning of the name Tetrapoda in the scientific literature: an exchange. Systematic Biology 53, 68–80.
CrossRef | open url image1

Lee MSY (1999) Stability of higher taxa in phylogenetic nomenclature—some comments on Moore. Zoologica Scripta 28, 361–366.
CrossRef | open url image1

Lee MSY (2001) On recent arguments for phylogenetic nomenclature. Taxon 50, 175–180. open url image1

Lee MSY (2002) Species and phylogenetic nomenclature. Taxon 51, 507–510. open url image1

Lidén M, Oxelman B (1996) Do we need “phylogenetic taxonomy”? Zoologica Scripta 25, 183–185.
CrossRef | open url image1

Lidén M, Oxelman B, Backlund A, Andersson L, Bremer B , et al. (1997) Charlie is our darling. Taxon 46, 735–738. open url image1

Mahner, M ,  and  Bunge, M (1997). ‘Foundations of biophilosophy.’ (Springer: Berlin)

Malt BC (1995) Category coherence in cross-cultural perspective. Cognitive Psychology 29, 85–148.
CrossRef | open url image1

Mason-Gamer RJ (2004) Reticulate evolution, introgression, and intertribal gene capture in an allohexaploid grass. Systematic Biology 53, 25–37.
CrossRef | open url image1

Mayr, E (1942). ‘Systematics and the origin of species: from the viewpoint of a zoologist’. (Columbia University Press: New York)

Mayr, E (1957). Species concepts and definitions. In ‘The species problem’. pp. 1–22. (American Association for the Advancement of Science: New York)

Mayr, E (1969). ‘Principles of systematic zoology.’ (McGraw-Hill: New York)

Mayr, E (1982). ‘The growth of biological thought.’ (Harvard University Press: Cambridge, Mass)

Mayr E (1995) Systems of ordering data. Biology and Philosophy 10, 419–434.
CrossRef | open url image1

Mayr, E ,  and  Ashlock, PD (1991). ‘Principles of systematic zoology.’ 2nd edn . (McGraw Hill: New York)

Mayr E, Bock WJ (2002) Classifications and other ordering systems. Journal of Zoological Systematics and Evolutionary Research 40, 169–194.
CrossRef | open url image1

McNeill J (1979) Structural value: a concept used in the construction of taxonomic classifications. Taxon 28, 481–504. open url image1

Mishler, BD (1999). Getting rid of species. In ‘Species: new interdisciplinary essays’. pp. 307–315. (MIT Press: Cambridge, MA)

Mishler, BD ,  and  Fisher, KM (2004). Terminating species: a rank-free approach to terminal taxa. In ‘ First international phylogenetic nomenclature meeting’. [Abstracts]. pp. 6.

Monsch KA (2003) The use of apomorphies in taxonomic defining. Taxon 52, 105–107. open url image1

Moore G (1998) A comparison of traditional and phylogenetic nomenclature. Taxon 47, 561–579. open url image1

Moore G (2003) Should taxon names be explicitly defined? Botanical Review 69, 2–21. open url image1

Moore G (2005) A review of past and current debates in nomenclature: 250 years of progress or going around in a circle? Symbolae Botanicae Upsalienses 33(3), 109–117. open url image1

Moore G, Barkley TM, DePriest P, Funk V, Kiger RW, Kress WJ, Nicolson DH, Stevenson DW, Wheeler QD (2004) (065–067) Proposals to amend Article 3.1, Article 22.3, and Article 26.3. Taxon 53, 214. open url image1

Murphy, GL (2002). ‘The big book of concepts.’ (MIT Press: Cambridge, MA)

Nelson G, Murphy DJ, Ladiges PY (2003) Brummitt on paraphyly: a response. Taxon 52, 295–298. open url image1

Nixon KC, Wheeler QD (1990) An amplification of the phylogenetic species concept. Cladistics 6, 211–223. open url image1

Nixon KC, Carpenter JM (2000) On the other “phylogenetic systematics”. Cladistics 16, 298–318. open url image1

Nixon KC, Carpenter JM, Stevenson DW (2003) The phylocode is fatally flawed, and the Linnaean system can easily be fixed. Botanical Review 69, 111–120. open url image1

Nordal I, Stedje B (2005) Paraphyletic taxa should be accepted! Taxon 54, 5–8. open url image1

Olmstead, RG ,  and  Cantino, PD (2004). Phylogenetic nomenclature of Lamiales. In ‘First international phylogenetic nomenclature meeting’. [Abstracts]. pp. 14.

Padian K (1999) Charles Darwin’s view of classification in theory and practice. Systematic Biology 48, 352–364.
CrossRef | open url image1

Panchen, AL (1992). ‘Classification, evolution and the nature of biology.’ (Cambridge University Press: Cambridge)

Pennisi E (1996) Evolutionary and systematic biologists converge. Science 273, 181. open url image1

Philipson WR (1987) The treatment of isolated genera. Botanical Journal of the Linnean Society 95, 19–25. open url image1

Pleijel F, Rouse GW (2000) Least-inclusive taxonomic unit: a new concept for biology. Proceedings of the Royal Society of London. Series B. Biological Sciences 267, 627–630.
CrossRef | open url image1

Pleijel F, Rouse GW (2002) Ceci n’est pas une pipe: names, clades and phylogenetic nomenclature. Journal of Zoological Systematics and Evolutionary Research 4, 162–175. open url image1

Pryor, LD ,  and  Johnson, LAS (1971). ‘A classification of the eucalypts.’ (Australian National University: Canberra)

de Queiroz K (1988) Systematics and the Darwinian revolution. Philosophy of Science 55, 238–259.
CrossRef | open url image1

de Queiroz K (1992) Phylogenetic definitions and taxonomic philosophy. Biology and Philosophy 7, 295–313.
CrossRef | open url image1

de Queiroz K (1997) The Linnaean hierarchy and the evolutionization of taxonomy, with emphasis on the problem of nomenclature. Aliso 15, 125–144. open url image1

de Queiroz K (2000) The definitions of taxon names: a reply to Stuessy. Taxon 49, 533–537. open url image1

de Queiroz K (2005) Linnaean, rank-based, and phylogenetic nomenclature: restoring primacy to the link between names and taxa. Symbolae Botanicae Upsalienses 33, 127–140. open url image1

de Queiroz K, Gauthier J (1990) Phylogeny as a central principle in taxonomy: phylogenetic definitions of taxon names. Systematic Zoology 39, 307–322. open url image1

de Queiroz K, Gauthier J (1992) Phylogenetic taxonomy. Annual Review of Ecology and Systematics 23, 449–480.
CrossRef | open url image1

de Queiroz K, Gauthier J (1994) Towards a phylogenetic system of biological nomenclature. Trends in Ecology & Evolution 9, 27–31.
CrossRef | open url image1

de Queiroz, K ,  and  Gauthier, J (2004). Toward an integrated system of phylogenetically defined names. In ‘First international phylogenetic nomenclature meeting’. [Abstracts]. pp. 7.

Rapini A (2004) Classes or individuals? The paradox of systematics revisited. Studies in the History and Philosophy of Biology and Biomedical Sciences 35, 675–695.
CrossRef | open url image1

Raven PH, Axelrod DI (1974) Angiosperm biogeography and past continental movements. Annals of the Missouri Botanical Garden 61, 593–673. open url image1

Rieseberg LH, Brouillet L (1994) Are many plant species paraphyletic? Taxon 43, 21–32. open url image1

Sauquet H, Doyle JA, Scharaschkin T, Borsch T, Hilu KW, Chatrou LW, Le Thomas A (2003) Phylogenetic analysis of Magnoliales and Myristicaceae based on multiple data sets: implications for character evolution. Botanical Journal of the Linnean Society 142, 125–186.
CrossRef | open url image1

Schander C (1998a) Types, emendations and names—a reply to Lidén et al. Taxon , 401–406. open url image1

Schander C (1998b) Mandatory categories and impossible hierarchies—a reply to Sosef. Taxon 47, 407–410. open url image1

Schander C, Thollesson M (1995) Phylogenetic taxonomy—some comments. Zoologica Scripta 24, 263–268.
CrossRef | open url image1

Schuh R (2003) The Linnaean system and its 250-year persistence. Botanical Review 69, 59–78. open url image1

Sereno PC (1999) Definitions in phylogenetic taxonomy: critique and rationale. Systematic Biology 48, 329–351.
CrossRef | open url image1

Sereno, PC (2004). Notation, definitional rationale, and recall in phylogenetic taxonomy. In ‘ First international phylogenetic nomenclature meeting’. [Abstracts]. pp. 4.

Sereno PC (2005) The logical basis of phylogenetic taxonomy. Systematic Biology 54, 595–619.
CrossRef | open url image1

Simpson, GG (1961). ‘Principles of animal taxonomy.’ (Columbia University Press: New York)

Sitte, P , Weiler, EW , Kadereit, JW , Bresinsky, A ,  and  Körner, C (2002). ‘Lehrbuch der Botanik für Hochschulen.’ (Spektrum: Heidelberg)

Smedmark JEE, Eriksson T, Evans RC, Campbell CS (2003) Ancient allopolyploid speciation in Geinae (Rosaceae): evidence from nuclear granule-bound starch synthase (GBSSI) gene sequences. Systematic Biology 52, 374–385. open url image1

Soltis DE, Soltis PS (1990) Isozyme evidence for ancient polyploidy in primitive angiosperms. Systematic Botany 15, 328–337. open url image1

Sosef MSM (1997) Hierarchical models, reticulate evolution and the inevitability of paraphyletic supraspecific taxa. Taxon 46, 75–85. open url image1

Stace HM, Douglas AW, Sampson JF (1998) Did ‘paleo-polyploidy’ really occur in Proteaceae? Australian Systematic Botany 11, 613–629.
CrossRef | open url image1

van Steenis CGGJ (1978) On the doubtful virtue of splitting families. Bothalia 12, 425–427. open url image1

Stevens, PF (1991). George Bentham and the Kew Rule. In ‘Improving the stability of names: needs and options’. pp. 157–168. (Koeltz: Königstein)

Stevens, PF (1994). ‘The development of biological systematics.’ (Columbia University Press: New York)

Stevens PF (1997) How to interpret botanical classifications: suggestions from history. Bioscience 47, 243–250. open url image1

Stevens, PF (1998). What kind of classification should the practicing taxonomist use to be saved? In ‘Plant diversity in Malesia III’. pp. 295–319. (Royal Botanic Gardens: Kew)

Stevens, PF (1999). Conventions, standards and consensus in systematic practice: How far can (or should we go? In ‘Metadiversity’. pp. 43–50. (National Federation of Abstracting and Information Services

Stevens PF (2001a) Mark Ereshefsky, The poverty of the Linnaean hierarchy:...’ Journal of the History of Biology 34, 600–602. open url image1

Stevens PF (2001 b) ‘Angiosperm phylogeny website.’ Available at http://www.mobot.org/MOBOT/research/APweb/ (verified 28 November 2005)

Stevens PF (2002) Why do we name organisms? Some reminders from the past. Taxon 51, 11–26. open url image1

Stuessy, TF (1990). ‘Plant taxonomy: the systematic evaluation of comparative data.’ (Columbia University Press: New York)

Stuessy TF (2001) Taxon names are still not defined. Taxon 50, 185–186. open url image1

Takhtajan, A (1997). ‘The diversity and classification of flowering plants.’ (Columbia University Press: New York)

Thorne R (2002) How many species of seed plants are there? Taxon 52, 511–512. open url image1

Wallace (1856) Attempts at a natural arrangement of birds. Annals and Magazine of Natural History ser. 2 16, 184–196. open url image1

van Welzen PC (1997) Paraphyletic groups or what should a classification entail. Taxon 46, 99–103. open url image1

van Welzen PC (1998) Phylogenetic versus Linnaean taxonomy, the continuing story. Taxon 47, 413–423. open url image1

Whewell, W (1847). ‘The philosophy of the inductive sciences, founded upon their history.’ 2nd edn . (John W. Parker: London)

Wiley E (1979) An annotated Linnaean hierarchy, with comments on natural taxa and competing systems. Systematic Zoology 28, 308–337. open url image1

Wiley, E (1981). ‘Phylogenetics: the theory and practice of phylogenetic systematics.’ (Wiley: New York)

Williams JH, Friedman WE (2004) The four-celled female gametophyte of Illicium (Illiciaceae; Austrobaileyales): implications for understanding the origin and early evolution of monocots, eumagnoliids, and eudicots. American Journal of Botany 91, 332–351. open url image1

Winsor MP (2001) Cain on Linnaeus: the scientist-historian as unanalysed entity. Studies in the History and Philosophy of Biology and Biomedical Sciences 32, 239–254.
CrossRef | open url image1

Winsor MP (2003) Non-essentialist methods in pre-Darwinian taxonomy. Biology and Philosophy 18, 387–400.
CrossRef | open url image1









1 Hennig (1966) had used the term monophyly, and its redefinition by phylogenetic systematists did not sit well with some evolutionary systematists, who preferred to call Hennigian monophyly, holophyly (e.g. Ashlock 1971; Mayr and Ashlock 1991). Their definition of monophyly was broader: a monophyletic taxon was one whose members descended from another taxon of the same or lower rank (e.g. Simpson 1961; Mayr 1969; Mayr and Ashlock 1991). This would include paraphyletic taxa as recognised here.

2 Strictly speaking, Linnaean nomenclature refers to Linnaeus’s generic names and the polynomials he used for species, e.g. Linnaeus (1751).

3 3Of course, some vigorously reject the value or applicability of ideas of individuality in systematics (e.g. Nixon and Carpenter 2000 and refs) or at least classification (Rapini 2004), or that species are clades (Nixon and Wheeler 1990). Others have redefined ‘class’, removing some connotations that are perceived to be negative (e.g. Mahner and Bunge 1997; Boyd 1999; Keller et al. 2003).

4 This is connected to a confusing debate as to whether (and how) taxon names are defined (e.g. de Queiroz 1997; Stuessy 2001; Monsch 2003; Schuh 2003; Laurin and Anderson 2004). Of course, clades can be recognised only if there are apomorphic characters, whether morphological or molecular, supporting their existence. In PC apomorphy-based definitions such features are used to diagnose the name, and they can also be used in mL diagnoses or characterisations. It is unhelpful to equate apomorphies and essences.

5 The comparable numbers in de Candolle (1844) are 6000 generic names, 1000 epithets, and 80 000 species, but millions of species could be unambiguously named using those 7000 words.

6 Note that if more than one apomorphy is used in an apomorphy-based definition, the one denoting the least inclusive clade may be chosen (Pleijel and Rouse 2002). Parallelisms will not cause problems since the apomorphy is linked to its occurrence in a particular taxon, e.g. in Magnoliaceae, sheathing stipules as found in Magnolia virginiana (Pleijel and Rouse 2002; see also PCn for complex apomorphies). Nevertheless, assigning particular apomorphies to an unambiguous position on the tree can be difficult (e.g. Stevens 2001b), even given a stable hypothesis of phylogeny. Thus one possible apomorphy of angiosperms, triploid endosperm (see e.g. Nixon and Carpenter 2000), is currently of uncertain position (Williams and Friedman 2004; for similar problems with apomorphies of Magnoliales, cf. Sauquet et al. 2003).

7 There have been extensive arguments whether monotypic taxa with empty ranks are permissible in a Linnaean hierarchy (‘Gregg’s paradox’: Gregg 1950, 1968; Buck and Hull 1969; Hull and Snyder 1969). Within families, the arrangement of genera remains alphabetical.

8 Alternatively, one could hire a nomenclator, somebody who could whisper in your ear the full clade address whenever a uninomial was mentioned. Cicero would have understood.

9 A reorganisation of part of a larger herbarium (the Harvard University Herbaria at Cambridge, MA) along phylogenetic principles was carried out as some 1 400 000 specimens were moved in 1997.


Export Citation Cited By (10)