DIFFERENCES BETWEEN ZOOLOGICAL & BOTANICAL CODES

1. DIFFERENCES BETWEEN ZOOLOGICAL & BOTANICAL CODES IN COMMON: stability, priority, hierarchy, types, Latin names DIFFERENCES: • Suprageneric name endings • Italicization • Ranks covered • Infraspecific connecting terms • Principle of Coordination & autonyms • Different terminologies • Different kinds of type specimens • Recombining author • Square brackets • Tautonyms • Illegitimacy • Recent vs date for priority • Hyphens allowed in genus & species names

2. SUPRAGENERIC NAME ENDINGS These are very different for equivalent ranks in the two codes. Examples of “homonyms”: ZOOLOGICAL CODEBOTANICAL CODE -idae Family Subclass -inae Subfamily Subtribe ITALICIZATION ZOOLOGICAL CODE: Genus & species ranks only BOTANICAL CODE: All ranks; however, this is not yet mandatory, only encouraged.

3. RANKS COVERED ZOOLOGICAL CODEBOTANICAL CODE (Kingdom) Kingdom (Phylum) Division or Phylum (Class) Class (Order) Order Family Family Tribe Tribe Genus Genus - Section - Series Species Species - Variety - Form [plus sub-categories of all [plus subcategories of all] & super-categories above Genus] [“( )“ indicates: not regulated by the code except for certain basic principles]

4. INFRAGENERIC & INFRASPECIFIC CONNECTING TERMS Because there are multiple infrageneric & infraspecific ranks in Botanical nomenclature, these are specified in the name. E.g.: Saxifraga aizoon subf. surculosa Engl. & Irmsch. Because there are intermediate ranks between subform and species, the taxon can be referred to in full in a combination of ‘name plus classification’ as: Saxifraga aizoon var. aizoon subvar. brevifolia f. multicaulis subf. surculosa Engl. & Irmsch. In Zoological nomenclature, they are unnecessary. Subgenera are placed in round brackets. Infrasubspecific names are not available, unless before 1961 they were termed variety or form, in which case they are deemed subspecific, or unless before 1985 they were adopted as subspecific names.

5. PRINCIPLE OF COORDINATION In Zoological nomenclature: each subordinate rank within a given rank group (family, genus, species) takes the same author & date (the prior one). E.g.: Subfamily Microchoerinae was originally erected as family Microchoeridae Lydekker,1887. The subfamily keeps the same author and date despite its rank being changed by another author. Likewise, if a superfamily Microchoerioidea were to be erected it would also be attributed to Lydekker, 1887 and without brackets. In Botanical nomenclature: priority is established within each rank, with individual authors & dates for each. The Principle of Coordination is not a part of the Botanical Code.

6. AUTONYMS If a new subsidiary rank of a particular taxon is named where no nominate equivalent previously existed, that nominate equivalent is deemed to have been thereupon named automatically with the same author and date as the non-nominate one. • Hypothetical scenario: • Family Planteaceae Smith (1754) has never been divided into subfamilies. • Bloggs (2006) names a subfamily Botanioideae. • In naming this new subfamily, he is considered to have automatically named a nominate subfamily too, which is the autonym Planteoideae Bloggs (2006).

7. DIFFERENT TERMINOLOGIES ZOOLOGICAL CODEBOTANICAL CODE Junior homonym Later homonym Objective synonym Nomenclatural synonym Subjective synonym Taxonomic synonym Available Validly published Valid name Correct name Specific name Specific epithet Binomen, name of a species Specific name

8. DIFFERENT KINDS OF TYPES • HOLOTYPE, SYNTYPE & NEOTYPE are essentially the same as in the Zoological code, but the first 2 must be from a single gathering. • ISOTYPE is a duplicate of the HOLOTYPE from one and the same gathering. (A specimen cited in the original work but from e.g. a different horizon and/or locality is a PARATYPE, not an ISOTYPE). • ISOSYNTYPE is likewise a duplicate of SYNTYPES. • PARATYPE is an originally cited additional specimen that is neither ISOTYPE nor SYNTYPE nor ISOSYNTYPE. It can exist alongside these unlike in Zoology. • LECTOTYPE can be selected from ISOTYPES, SYNTYPES or, if these are lost, from ISOSYNTYPES or PARATYPES. • EPITYPE is selected when original types are ambiguous (= NEOTYPE designation for Nomen Dubium in Zoology Code).

9. RECOMBINING AUTHOR ZOOLOGICAL CODEBOTANICAL CODE E.g.: E.g.: Motacilla troglodytes L., 1758 Petrophiloides richardsonii Bowerbank (1840) Viellot (1806) makes recombination: Chandler (1964) makes recombination: Troglodytes troglodytes (L., 1758) Platycarya richardsonii (Bowerbank) Chandler (1964) Original author in ( ) with date, no Original author in ( ) without date, recombining author. recombining author added. [N.B. in botany, brackets round date not mandatory. Abbreviated author not followed by comma.]

10. SQUARE BRACKETS ZOOLOGY CODE: Used to enclose author if cited when external evidence indicates original anonymity. BOTANY CODE: Used to denote pre-starting point authority citation; the starting point for palaeobotany is Sternberg 1820, substantially after the 1753 starting point for modern botany.

11. TAUTONYMY Troglodytes troglodytes, i.e. genus and species with same name not allowed in Botanical Code. If e.g. it resulted from recombination, the species would have to be changed to the next oldest legitimate name.

12. ILLEGITIMACY “A name of a family, genus or species, unless conserved, or sanctioned, is illegitimate if it is a later homonym, that is, if it is spelled exactly like a name based on a different type that was previously and validly published for a taxon of the same rank” (ICBN Article 53.1). Thus a name can be rejected on the grounds of junior homonymy (and for other reasons), unlike in the Zoological Code.

13. PRIORITY RECENT VS. FOSSIL FOSSIL Petrophiloides Bowerbank, 1840 Hexaprotodon Falconer & Cautley, 1836 Priority to date RECENT Platycarya Siebold & Zucc., 1843 Priority to Recent & date Choeropsis Leidy, 1853

14. This priority rule only applies, however, if a plant fossil name is being used as a biological genus or species (now regarded as exceptional). • Fossil diatoms (Bacillariophyceae) are treated as biological taxa. • But plant fossil names (excluding diatoms, but including other fossil algae, even Cyanobacteria) are normally treated as MORPHOTAXA.

15. MORPHOTAXA “Fossil taxa may be treated as morphotaxa. A morphotaxon is defined as a fossil taxon which, for nomenclatural purposes, comprises only the parts, life-history stages, or preservational states represented by the corresponding nomenclatural type.” Spinizonocolpites: pollen of the palm genus Nypa. Araucarioxylon: wood of the family Araucariaceae

16. PRIORITY IN MORPHOTAXA Plant fossil taxa do not compete with modern plant taxa when they are treated as morphotaxa. E.g., if Petrophiloides is treated as a catkin morphogenus of family Juglandaceae (walnuts), it is not synonymised with recent Platycarya. Often, there are several different names for different parts/organs of a fossil plant, e.g. the giant lycopod ‘Lepidodendron’: Lepidodendron – stem Knorria – older bark Lepidophylloides – leaves Lepidostrobus – free-sporing mono- or bisporangiate cones Lepidocarpon – megasporangiate cones Stigmaria – rhizophores (roots) Lycospora – microspores. All these different names can be used concurrently even though they may belong to the same organism.

17. ALTERNATIVE NOMENCLATURES

18. INTERNATIONAL CODE FOR PHYTOLITH NOMENCLATURE Madella et al. 2005

19. THE PHYLOCODE • New system of biological nomenclature to provide rules to govern the naming of clades across all of biology. • PREMISE: There should be congruence between phylogenetic hypotheses & nomenclature. • The system is in draft form but expects to go live in June 2006. Current plans cover taxa above the species level, but are intended to extend to species later. It is intended to run concurrently with pre-existing codes, or to replace them once it is extended to species, if the scientific community decides that it should. FEATURES: 1) It ignores rank. Formal endings denoting different ranks have no hierarchical significance. 2) It is aimed at reflecting phylogenetic hypotheses through names, whose usage should be explicit, unambiguous & stable.

20. PHYLOCODE PRINCIPLES 1) REFERENCE. The primary purpose of taxon names is to provide a means of referring to taxa, as opposed to indicating their characters, relationships, or membership. 2) CLARITY. Taxon names should be unambiguous in their designation of particular taxa. Nomenclatural clarity is achieved through explicit definitions. 3) UNIQUENESS. To promote clarity, each taxon should have only one accepted name, and each accepted name should refer to only one taxon. 4) STABILITY. The names of taxa should not change over time. As a corollary, it must be possible to name newly discovered taxa without changing the names of previously discovered taxa. 5) PHYLOGENETIC CONTEXT. The phylocode is concerned with the naming of taxa and the application of taxon names within a phylogenetic context. 6) The PhyloCode permits freedom of taxonomic opinion with regard to hypotheses about relationships; it only concerns how names are to be applied within the context of a given phylogenetic hypothesis.

21. 3 WAYS OF NAMING A CLADE: Must be defined by at least 2 specifiers (like Linnaean types) Forey (2001) PHYLOGENY: NODE-BASED 2 included names STEM-BASED 1 included & 1 excluded name APOMORPHY-BASED 1 included name & 1 apomorphy

22. NEW HYPOTHESIS: NAMES DON’T CHANGE Forey (2001) NO HOMONYMY allowed between node-based, stem-based & apomorphy-based methods NO SYNONYMY SYNONYMY with revision

23. PERCEIVED PROBLEMS w. LINNAEAN SYSTEM 1) Ranks are artificial groupings, not reflecting exact position in hierarchy. 2) Paraphyletic groups like Reptilia are named. 3) Redundancy of ranks through extinction or asymmetry Forey (2001)

24. PERCEIVED PROBLEMS w. LINNAEAN SYSTEM 4) Ranks cause instability when relationships change. Forey (pers. com.)

25. PERCEIVED PROBLEM SOLVED Forey (pers. com.)

26. EXAMPLE OF LINNAEAN INSTABILITY Forey (pers. com.)

27. PROBLEM WITH APOMORPHY-BASED DEFINITIONS IN THE PHYLOCODE Homoplasy by reversal vs convergence. In (a) the apomorphy “fingers & toes” defines a clade. In (b) it reflects polyphyly. Forey (2001)

28. LINNAEAN SYSTEM: SAVING GRACES 1) Even though ranks are artificial, they are useful boxes and much used in modern biodiversity studies. 2) Paraphyletic groups don’t have to be used under the Linnaean system. 3) There is potentially as much instability following revisions with the 3-system naming procedure under the PhyloCode. 4) In the Linnaean system you don’t have to name every rank. New classifications commonly list e.g. a genus undifferentiated within an order. Thus Archaeopteryx need not have its own monotypic infraclass, supercohort, cohort, subcohort and order.

29. METHODS OF NAMING IN THE PHYLOCODE The PhyloCode proposes a registration system whereby clade names are submitted electronically. The following information is needed: 1) DEFINITION TYPE: node- stem- or apomorphy-based (mandatory) 2) PHYLOGENETIC DEFINITION: (mandatory) 3) LIST OF SPECIFIERS: (at least 2 mandatory) 4) QUALIFYING CLAUSE 5) REFERENCE PHYLOGENY: bibliographic reference, URL, or accession no. in public repository

30. Linnaean names already being published according to PhyloCode guidelines (Wyss & Flynn 1993). New node-based homonyms of superfamilies Ursoidea Fischer de Waldheim, 1817 & Phocoidea Gray, 1821. Superfamily Arctoidea Flower, 1869 with node-based definition at different hierarchical position. Stem-based definition for Carnivoramorpha

31. IMPLICATIONS OF PARALLEL SYSTEMS OF NOMENCLATURE • Not too serious unless or until the PhyloCode extends its scope to species, which are intended to be uninominal. • One suggestion is to add suffix ‘P’ to names that are homonyms between Linnaean & PhyloCode systems. • However, if authors of databases (including MOA) wish to retain only Linnaean nomenclature, any PhyloCode classification would have to be interpreted in terms of Linnaean nomenclature.

32. PROBLEMS FOR CHARACTER DIAGNOSES PhyloCode: 2 out of 3 ways of defining a taxon excludes characters Cladistics: stresses synapomorphies. Diagnosis could differentiate types of characters. E.g.:

33. DIAGNOSES: STAND ALONE OR DIFFERENTIAL?

34. DIAGNOSES: STAND ALONE & DIFFERENTIAL

35. Pseudoneothoracophorus Psilastephanocolporites Pseudopromerycochoerus Ekgmowechashala Kukusepasutanka Intratriporopollenites

36. REFERENCES Bengtson, P. 1988. Open nomenclature. Palaeontology, 31: 223-227. Candolle, Alph. De. 1867. Lois de la Nomenclature Botanique adoptées par le Congrès International de Botanique tenu à Paris en août, 1867, suivies à un deuxième édition de l’introduction historique et du commentaire qui accompagnaient la rédaction préparatoire présentée au Congrès. H. Georg, Genève & Bale; J.-B. Baillière et fils, Paris, 64 pp. Forey, P.L. 2001. The PhyloCode: description and commentary. Bulletin of Zoological Nomenclature, 58: 81-96. Greuter, W. et al. 2000. International Code of Botanical Nomenclature (St Louis Code). Regnum Vegetabile 138, Koeltz Scientific Books, Königstein. Hughes, N.F. 1989. Fossils as Information; New recording and Stratal Correlation Techniques. Cambridge, Cambridge University Press, 136 pp. International Commission on Zoological Nomenclature. 1999. International Code of Zoological Nomenclature, 4th ed., ITZN c/o NHM, London, 306 pp. Linnaeus, C. 1753. Species Plantarum. Vol. 1, Laurentius Salvius, Stockholm, 560 pp. Linnaeus, C. 1758.Systema Naturae. Vol. 1, Regnum Animale, 10th ed., revised. Laurentius Salvius, Stockholm, 824 pp. Madella, M., Alexandre, A. & Ball, T. 2005. International Code for Phytolith Nomenclature 1.0. Annals of Botany, 96: 253-260. Matthews, S.C. 1973. Notes on open nomenclature and on synonymy lists. Palaeontology, 16: 713-719. Opinion 1894. 1998. Regnum Animale …, Ed. 2 (M.J. Brisson, 1762): rejected for nomenclatural purposes, with the conservation of the mammalian generic names Philander (Marsupialia), Pteropus (Chiroptera), Glis, Cuniculus and Hydrochoerus (Rodentia), Meles, Lutra and Hyaena (Carnivora), Tapirus (Perissodactyla), Tragulus and Giraffa (Artiodactyla). Bulletin of Zoological Nomenclature, 55: 64-71. Strickland, H.E., Darwin, C., Owen, R. & Westwood, J.O. 1843. Series of propositions for rendering the nomenclature of zoology uniform and permanent, being the report of a committee for the consideration of the subject, appointed by the British Association for the Advancement of Science. Annals & Magazine of natural History, (1)11: 259-275. Wyss, A.R. & Flynn, J.J. 1993. A phylogenetic analysis and definition of the Carnivora. In: Szalay, F.S., Novacek, M.J. & McKenna, M.C. (eds), Mammal Phylogeny, 2: Placentals. New York, Springer: 32-52.