Abstract
This paper explores two aspects of the question of the allocation of nomina to taxa in zoological nomenclature.
The widespread belief that this allocation is effected only through onomatophores (‘types’) is shown to be wrong: if onomatophores only were involved, each zoological nomen would apply to all the taxa including the specimen(s) designated as onymophoront of each species-series nomen, up to the animal kingdom. Onomatophores do not provide the limit(s) of the taxon/taxa to which they apply. These limits are provided by two other pieces of information: the nominal-series to which the nomen of a taxon is assigned and the onomatostasis of this nomen, i.e. the onomatophore of the sister-taxon of the taxon under consideration.
In species-, genus- and family-series nomenclature, the onomatostasis of a nomen is not fixed but depends on the ergotaxonomy (working classification) adopted, which is ideally based on at least one accepted phylogenetic hypothesis. The situation is more complex in the class-series nomenclature. So far, the stages of taxonomic allocation and nomenclatural validity in this nomenclature have not been regulated by formal Rules shared by all zootaxonomists, so that chaos and miscommunication are in order regarding the Criteria to be used in these domains, which is detrimental to zoological nomenclature and its perception by the international scientific community. The adoption of clear and strict Rules in this respect appears as an urgent need. These Rules should allow both to provide clear objective Criteria for the allocation of class-series nomina to taxa, but also to validate the few very well known and consensually adopted class-series nomina (pansozonyms). For this purpose, five possible nomenclatural systems of allocation of nomina are compared. The best solution appears to have two different systems, one for almost all nomina (distagmonyms), and one only for pansozonyms. The latter system relies on a special kind of onomatostases, which are fixed and do not depend on the classification adopted: this allows to attach permanently these nomina to some well-known higher taxa, whatever changes are brought to the cladistic hypotheses and ergotaxonomic frames.
References
Anonymous [International Commission on Zoological Nomenclature] (1999) International code of zoological nomenclature. ‘Fourth edition’. London (International Trust for zoological Nomenclature): i–xxix + 1–306.
Alonso-Zarazaga, M. A. (2005) Nomenclature of higher taxa: a new approach. Bulletin of zoological Nomenclature, 62 (4): 189–199.
Amiet, J.-L. (2012) Les rainettes du Cameroun (Amphibiens Anoures). Nyons (J.-L. Amiet) & Saint- Nazaire (La Nef des Livres): 1–591.
Amorim, D. S., Santos, C. M. D., Krell, F.-T., Dubois, A., Nihei, S. S., Oliveira, O. M. P., Pont, A., Song, H., Verdade, V. K., Fachin, D. A., Klassa, B., Lamas, C. J. E., Oliveira, S.S., de Carvalho, C. J. B., Mello-Patiu, C. A., Hajdu, E., Couri, M. S., Silva, V. C., Capellari, R. S., Falaschi, R. L., Feitosa, R. M., Prendini, L., Pombal, J. P. Jr., Fernández, F., Rocha, R. M., Lattke, J. E., Caramaschi, U., Duarte, M., Marques, A. C., Reis, R. E., Kurina, O., Takiya, D. M., Tavares, M., Silva Fernandes, D., Franco, F. L., Cuezzo, F., Paulson, D., Guénard, B., Schlick-Steiner, B. C., Arthofer, W., Steiner, F. M., Fisher, B. L., Johnson, R. A., Delsinne, T. D., Donoso, D. A., Mulieri, P. R., Patitucci, L. D., Carpenter, J. M., Herman, L. & Grimaldi, D. (2016) Timeless standards for species delimitation. Zootaxa, 4137 (1): 121–128. <https://doi.org/10.11646/zootaxa.4137.1.9>.
Baird, S. F. (1850) Revision of the North American tailed-Batrachia, with descriptions of new genera and species. Journal of the Academy of natural Sciences of Philadelphia, (2), 1: 281–294.
Blainville, H., de (1816) Prodrome d’une nouvelle distribution systématique du règne animal. Bulletin des Sciences de la Société philomatique de Paris, juillet 1816: ‘105–112’ [113–120] + 121–124.
Bonaparte, C. L. (1837) Synopsis vertebratorum systematis. 1–30.
Brongniart, A. (1800) Essai d’une classification naturelle des reptiles. Ière partie. Etablissement des ordres. Bulletin des Sciences, par la Société philomatique, 2 (35): 81–82.
Bruun, A. F. (1950) The Systema Naturae of the twentieth century. Science, 112: 342–343. <https://doi.org/10.1126/science.112.2908.342-a>.
Cope, E. D. (1877) Descriptions of extinct Vertebrata from the Permian and Triassic formation of Texas. Proceedings of the American philosophical Society, 17: 182–195. <https://doi.org/10.5962/bhl.title.104569>.
Cope, E. D. (1878) Descriptions of extinct Batrachia and Reptilia from the Permian formation of Texas. Proceedings of the American philosophical Society, 17: 505–530. <https://doi.org/10.5962/bhl.title.104569>.
Cope, E. D. (1895) A batrachian armadillo. The American Naturalist, 29 (347): 998. <https://doi.org/10.1086/276277>.
Dubois, A. (1983) Classification et nomenclature supragénérique des Amphibiens Anoures. Bulletin mensuel de la Société linnéenne de Lyon, 52: 270–276. <https://doi.org/10.3406/linly.1983.10607>.
Dubois, A. (1984) La nomenclature supragénérique des Amphibiens Anoures. Mémoires du Muséum national d’Histoire naturelle, (A), Zoologie, 131: 1–64.
Dubois, A. (1987) Miscellanea taxinomica batrachologica (I). Alytes, ‘1986’, 5 (1–2): 7–95.
Dubois, A. (1997) An evolutionary biologist’s view on the science of biology. Alytes, 15 (3): 133–136.
Dubois, A. (2000) Synonymies and related lists in zoology: general proposals, with examples in herpetology. Dumerilia, 4 (2): 33–98.
Dubois, A. (2004) The higher nomenclature of recent amphibians. Alytes, 22 (1–2): 1–14.
Dubois, A. (2005a) Les règles de la nomenclature familiale en zoologie In: A. Dubois, O. Poncy, V. Malécot & N. Léger (ed.), Comment nommer les taxons de rang supérieur en zoologie et en botanique?, Biosystema, 23: 17–40.
Dubois, A. (2005b) Propositions pour l’incorporation des nomina de taxons de rang supérieur dans le Code international de nomenclature zoologique. In: A. Dubois, O. Poncy, V. Malécot & N. Léger (ed.), Comment nommer les taxons de rang supérieur en zoologie et en botanique?, Biosystema, 23: 73–96.
Dubois, A. (2005c) 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 (2): 365–426. <https://doi.org/10.11646/zootaxa.1337.1.1>.
Dubois, A. (2005d) Proposals for the incorporation of nomina of higher-ranked taxa into the Code. Bulletin of zoological Nomenclature, 62 (4): 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 (1): 165–258. <https://doi.org/10.11646/zootaxa.1337.1.1>.
Dubois, A. (2006b) New proposals for naming lower-ranked taxa within the frame of the International Code of Zoological Nomenclature. Comptes rendus Biologies, 329 (10): 823–840. <https://doi.org/10.1016/j.crvi.2006.07.003>.
Dubois, A. (2006c) Incorporation of nomina of higher-ranked taxa into the International Code of Zoological Nomenclature: some basic questions. Zootaxa, 1337: 1–37. <https://doi.org/10.11646/zootaxa.1337.1.1>.
Dubois, A. (2007a) Phylogeny, taxonomy and nomenclature: the problem of taxonomic categories and of nomenclatural ranks. Zootaxa, 1519: 27–68. <https://doi.org/10.11646/zootaxa.1519.1.3>.
Dubois, A. (2007b) Nomina zoologica linnaeana. In: Z.-Q. Zhang & W. A. Shear (ed.), Linnaeus tercentenary: progress in invertebrate taxonomy, Zootaxa, 1668: 81–106. <https://doi.org/10.11646/zootaxa.1668.1.9>.
Dubois, A. (2008a) A partial but radical solution to the problem of nomenclatural taxonomic inflation and synonymy load. Biological Journal of the Linnean Society, 93: 857–863. <https://doi.org/10.1111/j.1095-8312.2007.00900.x>.
Dubois, A. (2008b) Le Code international de nomenclature zoologique: présentation, philosophie, règles majeures, problèmes actuels. In: D. Prat, A. Raynal-Roques & A. Roguenant (ed.), Peut-on classer le vivant? Linné et la systématique aujourd’hui, Paris (Belin): 355–402.
Dubois, A. (2008c) Phylogenetic hypotheses, taxa and nomina in zoology. In: A. Minelli, L. Bonato & G. Fusco (ed.), Updating the Linnaean heritage: names as tools for thinking about animals and plants, Zootaxa, 1950: 51–86. <https://doi.org/10.11646/zootaxa.1950.1.7>.
Dubois, A. (2011a) The International Code of Zoological Nomenclature must be drastically improved before it is too late. Bionomina, 2: 1–104. <https://doi.org/10.11646/bionomina.2.1.1>.
Dubois, A. (2011c) A zoologist’s viewpoint on the Draft BioCode. Bionomina, 3, 45–62. <https://doi.org/10.11646/bionomina.3.1.4>.
Dubois, A. (2013) Zygoidy, a new nomenclatural concept. Bionomina, 6: 1–25. <https://doi.org/10.11646/bionomina.6.1.1>.
Dubois, A. (2015a) Zoological nomina in the century of extinctions: new proposals. Bionomina, 8: 11–53. <https://doi.org/10.11646/bionomina.8.1.2>.
Dubois, A. (2015b) The Duplostensional Nomenclatural System for higher zoological nomenclature. Dumerilia, 5: 1–108.
Dubois, A. (2016) The Duplostensional Nomenclatural System for higher zoological nomenclature: additional comments. Dumerilia, 6: 5–16.
Dubois, A. (2017a) A few problems in the generic nomenclature of insects and amphibians, with recommendations for the publication of new generic nomina in zootaxonomy and comments on taxonomic and nomenclatural databases and websites. Zootaxa, 4237 (1): 1–16. <https://doi.org/10.11646/zootaxa.4237.1.1>.
Dubois, A. (2017b) The nomenclatural status of Hysaplesia, Hylaplesia, Dendrobates and related nomina (Amphibia, Anura), with general comments on zoological nomenclature and its governance, as well as on taxonomic databases and websites. Bionomina, 11: 1–48. <https://doi.org/10.11646/bionomina.11.1.1>.
Dubois, A. (2017c) The need for reference specimens in zoological taxonomy and nomenclature. Bionomina, 12: 4–38. <https://doi.org/10.11646/bionomina.12.1.2>.
Dubois, A. & Aescht, E. (2016a) The Linz Zoocode Committee. Dumerilia, 6: 35–37.
Dubois, A. & Aescht, E. (ed.) (2016b) Sessions 1‒6 of the Linz Zoocode Committee (February‒June 2016). Dumerilia, 6: 38–70.
Dubois, A. & Aescht, E. (ed.) (2017a) Sessions 7‒14 of the Linz Zoocode Committee (June 2016‒April 2017). Dumerilia, 7: 18–49.
Dubois, A. & Aescht, E. (ed.) (2017b) LZC Session 11. Proposal GEN-05. The Principles of the Zoocode. [3] The Principle of Nominal-Series. Dumerilia, 7: 29–31.
Dubois, A. & Aescht, E. (ed.) (2019a) Sessions 15‒38 of the Linz Zoocode Committee (May 2017‒October 2019). Dumerilia, 8: 1–169.
Dubois, A. & Aescht, E. (ed.) (2019b) LZC Session 17. The Principles of the Zoocode. 4. The Principle of Binomina. Dumerilia, 8: 35–36.
Dubois, A. & Aescht, E. (ed.) (2019c) LZC Session 19. The Principles of the Zoocode. 5. The Principle of Coordination. Dumerilia, 8: 42–47.
Dubois, A. & Aescht, E. (ed.) (2019d) LZC Session 29. The Principles of the Zoocode. 11. The Principle of Priority. Dumerilia, 8: 110–113.
Dubois, A. & Aescht, E. (ed.) (2019e) LZC Session 32. The Principles of the Zoocode. 14. The Principle of Nomography. Dumerilia, 8: 119–132.
Dubois, A. & Aescht, E. (ed.) (2019f) LZC Session 34. The Principles of the Zoocode. 16. The Principle of Archoidy. Dumerilia, 8: 143–146.
Dubois, A. & Aescht, E. (ed.) (2019g) LZC Session 37. Diagrams of the Nomenclatural Process. Dumerilia, 8: 159–168.
Dubois, A., Aescht, E. & Dickinson, E. C. (2016) Burning questions of zoological nomenclature. The Linz International Workshop on Zoological Nomenclature (9–10 July 2014). Dumerilia, 6: 24–34.
Dubois, A., Bauer, A. M., Ceríaco, L. M. P., Dusoulier, F., Frétey, T., Löbl, 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, 17: 1–111. <https://doi.org/10.11646/bionomina.17.1.1>.
Dubois, A. & Berkani, M. (2013) The misleading use of the terms parent, child, ancestor and descendant in databases dealing with biological evolution and taxonomy. Bionomina, 6: 52–56. <https://doi.org/10.11646/bionomina.6.1.4>.
Dubois, A. & Malécot, V. (2005) Glossaire: termes de taxinomie et de nomenclature utilisés dans ce volume. In: A. Dubois, O. Poncy, V. Malécot & N. Léger (ed.), Comment nommer les taxons de rang supérieur en zoologie et en botanique?, Biosystema, 23: 97–103.
Dubois, A. & Ohler, A. (2019) The nomina Anura, Urodela, Ecaudata and Caudata, credited to ‘Fischer von Waldheim, 1813’, do not exist, with comments on the nomenclature of higher zoological taxa and on the authorships and dates of other amphibian nomina. Bionomina, 14: 1‒68. <https://doi.org/10.11646/bionomina.14.1.1>.
Dubois, A. & Raffaëlli, J. (2012) A new ergotaxonomy of the order Urodela Duméril, 1805 (Amphibia, Batrachia). Alytes, 28 (3–4): 77–161.
Duméril, A. M. C. (1805) Zoologie analytique, ou méthode naturelle de classification des animaux, rendue plus facile à l’aide de tableaux synoptiques. Paris (Allais), ‘1806’: i–xxxii + 1–344. <https://doi.org/10.5962/bhl.title.44835>.
Duméril, [A. M. C.] (1839) Mémoire sur la classification et la structure des Ophiosomes ou Céciloïdes, famille de reptiles qui participent des Ophidiens et des Batraciens, relativement à la forme et à l’organisation. Compte rendu des Séances de l’Académie des Sciences, 9: 581–587.
Etheridge, R. (1866) On the discovery of several new labyrintodont reptiles in the coal-measures of Ireland. The Geological Magazine, 3: 4–5. <https://doi.org/10.1017/S0016756800165094>.
Ford, L. S. & Cannatella, D. C. (1993) The major clades of frogs. Herpetological Monographs, 7: 94–117. <https://doi.org/10.2307/1466954>.
Frétey, T. & Dubois, A. (2019) The authorship and date of five generic nomina of Squamata and Amphibia published by Cuvier (1816, 1829), with recourse to and comments on twenty Articles of the Code. Bionomina, 15: 1‒36. <https://doi.org/10.11646/bionomina.15.1.1>.
Gadow, H. (1898) A classification of Vertebrata, recent and extinct. London (Adam & Charles Black): i‒xvii + 1‒82. <https://doi.org/10.5962/bhl.title.24205>.
Gadow, H. (1901) Amphibia and Reptiles. London (Macmillan & Co.): i–xiii + 1–668, 1 pl. <https://doi.org/10.5962/bhl.title.160212>.
Gérard D., Vignes-Lebbe R., Dubois A. (2006) Ziusudra, de la nomenclature à l’informatique: l’exemple des Amphibiens. Alytes, 24 (1‒4): 117–132.
Haeckel, E. (1866) Generelle Morphologie der Organismen. Zweiter Band. Allgemeine Entwickelungsgeschichte der Organismen. Berlin (Georg Kramer): i–clx + 1–462, pl. 1–8. <https://doi.org/10.1515/9783110848281>.
Hillis, D. M. (2006) Constraints in naming parts of the tree of life. Molecular Phylogenetics & Evolution, 42: 331–338. <https://doi.org/10.1016/j.ympev.2006.08.001>.
Hogg, J. (1838) [On the classifications of the Amphibia]. Annals of natural History, (1), 1 (2): 152. <https://doi.org/10.1080/00222933809512254>.
Keller, R. A., Boyd, R. N. & Wheeler, Q. D. (2003) The illogical basis of phylogenetic nomenclature. The botanical Review, 69: 93–110. <https://doi.org/10.1663/0006-8101(2003)069[0093:TIBOPN]2.0.CO;2>.
Kuntner, M. & Agnarsson, I. (2006) Are the Linnean and phylogenetic nomenclatural systems combinable? Recommendations for biological nomenclature. Systematic Biology, 55: 774–784. <https://doi.org/10.1080/10635150600981596>.
Lataste, F. (1878) Division en familles naturelles des Batraciens Anoures d’Europe. Revue internationale des Sciences, 2 (42): 488–499.
Lawrence, T., Pelkey, N. & Soares, S. (2010) Comment on the proposed conservation of usage of Testudo gigantea Schweigger, 1812 (currently Geochelone (Aldabrachelys) gigantea; Reptilia, Testudines). (Case 3463; see BZN 66: 34–50, 80–87, 169–186, 274–290, 352–357; 67: 71–90, 170–178). Bulletin of zoological Nomenclature, 67 (3): 246–254.
Linnaeus, C. (1758) Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio decima, reformata. Tomus 1. Holmiae (Laurentii Salvii): [i–iv] + 1–824. <https://doi.org/10.5962/bhl.title.542>.
Linnaeus, C. (1767) Systema Naturae. Editio duodecima, reformata. Tomus 1, Pars 2. Holmiae (Laurentii Salvii): 533–1327 + [i–xxxvii].
Marjanović, D. & Laurin, M. (2015) Reevaluation of the largest published morphological data matrix for phylogenetic analysis of Paleozoic limbed vertebrates. PeerJ PrePrints, 3:e1596v1: 1–343. <https://doi.org/10.7287/peerj.preprints.1596v1>.
Merrem, B. (1820) Versuch eines Systems der Amphibien. Tentamen systematis Amphibiorum. Marburg (Iohann Christian Krieger): [i–xv + 1–191] × 2, 1 pl. <https://doi.org/10.5962/bhl.title.5037>.
Meyer, A. (1926) Logik der Morphologie im Rahmen einer Logik der gesamten Biologie. Berlin (Julius Springer): i–vii + 1–290. <https://doi.org/10.1007/978-3-642-50733-5>.
Pyron, R. A. & Wiens, J. J. (2011) A large-scale phylogeny of Amphibia including over 2800 species, and a revised classification of extant frogs, salamanders, and caecilians. Molecular Phylogenetics & Evolution, 61: 543–583. <https://doi.org/10.1016/j.ympev.2011.06.012>.
Queiroz, K. de & Gauthier, J. (1992) Phylogenetic taxonomy. Annual Review of Ecology & Systematics, 23: 449‒480. <https://doi.org/10.1146/annurev.es.23.110192.002313>.
Rafinesque-Schmaltz, C. S. (1814) Fine del prodromo d’erpetologia siciliana. Specchio delle Scienze o Giornale enciclopedico di Sicilia, 2 (10): 102–104.
Reig, O. A. (1958) Proposiciones para une nueva macrosistemática de los anuros. (Nota preliminar). Physis, 21: 109–118.
Ritgen, F. A. (1828) Versuch einer natürlichen Eintheilung der Vögel. Nova Acta physico-medica Academiae Caesareae Leopoldino-Carolinae Naturae Curiosorum, 14: 245–284.
Rowe, T. & Gauthier, J. (1992) Ancestry, paleontology, and definition of the name Mammalia. Systematic Biology, 41 (3): 372‒378. <https://doi.org/10.1093/sysbio/41.3.372>.
Ruta, M., Coates, M. I. & Quicke, D. L. J. (2003) Early tetrapod relationships revisited. Biological Review, 78: 251–345. <https://doi.org/10.1017/S1464793102006103>.
Sanchíz, B. (1984) Análisis filogenético de la tribu Alytini (Anura, Discoglossidae) mediante el estudio de su morfoestructura ósea. In: H. Hemmer & J. P. Alcover (ed.), Història biològica del ferreret, Mallorca (Moll): 61–108.
Simpson, G. G. (1940) Types in modern taxonomy. American Journal of Science, 238: 413–431. <https://doi.org/10.2475/ajs.238.6.413>.
Sonnini, C. S. & Latreille, P. A. (1801) Histoire naturelle des Reptiles. Tome 4. Seconde partie. Serpens. Paris, Deterville: [i–iii] + 1–410, 14 pl. <1801-son-234>.
Wright, E. P. & Huxley, T. H. (1866) On a collection of fossils from the Jarrow Colliery, Kilkenny, Ireland. The Geological Magazine, 3: 165–171.