A New Ergotaxonomy of the Order Urodela Duméril, 1805 (Amphibia, Batrachia)
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October 2012 Volume 28, No 3¢4 Alytes, 2012, 28 (3¢4): 77¢161. 77 A new ergotaxonomy of the order Urodela Duméril, 1805 (Amphibia, Batrachia) Alain Dubois* & Jean Raffaëlli** * Vertébrés: Reptiles & Amphibiens, UMR 7205 OSEB, Département Systématique & Evolution, Muséum national d’Histoire naturelle, CP 30, 25 rue Cuvier, 75005 Paris, France <[email protected]> ** Penclen, 56420 Plumelec, France <jean.raff[email protected]> Various recent works, particularly based on nucleic acid sequencing, have improved our understanding of the phylogenetic relationships among recent salamanders. Besides, new species and other taxa (subspecies, genera, subgenera) are regularly described in this group. Some of these data, but not all, have been the basis for taxonominal (taxonomic and nomenclatural) changes. Many taxonominal problems nevertheless remain unsettled in this group. We provide here a comprehensive review of the taxonominal hierarchy of this order, for which we strongly insist that the valid nomen is URODELA Duméril, 1805. In this taxon, we recognize a total of 1240 taxa, including two suborders, two infraoders, 10 recent and 5 fossil families, 70 recent and 64 fossil genera, 673 recent and 117 fossil species, and 111 recent subspecies. This works leads us to introduce 22 new nomina, at the following ranks: subfamilia (1), tribus (4), subtribus (5), genus (4) and subgenus (8). We also discuss several general questions of taxonomy and nomenclature, in particular regarding the structure of taxo- nominal hierarchies, the use of taxonomic categories and nomenclatural ranks in taxonomy, the nomenclatural Rules for ranks above superfamily, and the nomenclatural status of various nomina. We think the taxonominal hierarchy should reflect the best as possible the hypothesized phylogenetic relationships between species and other taxa, and not a gradist approach of taxonomy which is still prevalent in pseudoranked ergotaxonomies in which ranks are meant at expressing ‘‘degree of divergence’’ between taxa or the hypothesized age of cladogeneses. The use of ranks to express phylogenetic hypotheses on relationships is fully compatible with the Rules of the Code, provided a consistent and strict use of the hierarchy of nomenclatural ranks is followed. 78 ALYTES 28 (3¢4) Contents Abstract....................................................................... 77 Terminological note ............................................................. 79 Introduction .................................................................... 79 The basic distinction between taxonomic categories and nomenclatural ranks ....... 83 Nomenclatural problems ........................................................ 85 Class-series nomenclature ................................................... 85 Availability ............................................................ 86 Allocation ............................................................. 88 Validity................................................................ 89 Nomenclatural ranks and the taxonomy of the URODELA ...................... 92 The hierarchy of nomenclatural ranks ................................... 92 Nomenclatural ranks in the order URODELA .............................. 94 Primary key ranks ................................................. 94 Secondary key ranks and subsidiary ranks .......................... 95 Miscellaneous nomenclatural problems ...................................... 97 Priority................................................................ 97 Nomenclatural ranks and the simultaneous publication of nomina competing for homonymy or synonymy ............................... 98 AMPHIUMIDAE and PROTEINA ......................................... 98 PLETHODONTIDAE, DESMOGNATHINA and ENSATININA .................... 98 Subgenera of Eurycea.............................................. 98 Relative priority between generic and subgeneric nomina in case of current usage................................................................ 100 Incorrect authorship ................................................... 100 Problems of spelling.................................................... 102 Unusual taxonomic categories .......................................... 102 Proposed taxonomic and nomenclatural changes in the order URODELA ............. 103 Valid class-series nomina in the URODELA ................................ 103 Subordo MEANTES Linnaeus, 1767 .................................. 104 Infraordo IMPERFECTIBRANCHIA Hogg, 1838........................... 104 Infraordo PSEUDOSAURIA de Blainville, 1816 .......................... 105 Subordo ICHTHYOIDEA Leuckart, 1821 ............................... 106 Ordo URODELA Duméril, 1805 ...................................... 106 Taxonominal changes in the URODELA below the class-series ............... 111 Familia AMBYSTOMATIDAE Gray, 1850 ................................ 111 Familia CRYPTOBRANCHIDAE Fitzinger, 1826........................... 112 Familia DICAMPTODONTIDAE Tihen, 1958 ............................. 112 Familia HYNOBIIDAE Cope, 1859 (1856) .............................. 113 Familia PLETHODONTIDAE Gray, 1850................................. 114 Familia PROTEIDAE Gray, 1825 ...................................... 118 Familia SALAMANDRIDAE Goldfuss, 1820.............................. 119 Familia SIRENIDAE Gray, 1825 ....................................... 119 Dubois &Raffaëlli 79 Conclusion ..................................................................... 120 Some quantitative data ................................................. 120 Zoological taxonomy and nomenclature nowadays ....................... 122 Acknowledgements ............................................................. 123 Literature cited ................................................................. 123 Appendix 1 ..................................................................... 133 Appendix 2 ..................................................................... 154 Terminological note In the present work, we strictly respect the Rules of the International Code of Zoological Nomenclature (Anonymous, 1999; ‘‘the Code’’ below), but we sometimes use different terminologies to designate the concepts of the Code, for reasons explained in detail by Dubois (2000, 2005b, 2011a). We use the term nomen (plural nomina) for ‘‘scientific name’’ and the term nominal-series for the three ‘‘groups of names’’ recognized by the Code: the family-series (FS), genus-series (GS) and species-series (SS), and for a fourth one, established by Dubois (2000), the class-series (CS), accommodating all ranks above superfamily. These nomina are here printed in italics in the SS and GS, in ITALIC CAPITALS in the FS and in BOLD CAPITALS in the CS. Class-series nomina first published in a non-latinized form are presented in UNDERLINED BOLD CAPITALS. The terms hoplonym and anoplonym (Dubois, 2000) designate respectively a nomen that is or is not nomenclaturally available according to the Code. Anoplonyms are of two main kinds (Dubois, 2011a:18¢19 and accepted): gymnonyms (‘‘nomina nuda’’ in the Code) are nomina that are unavailable for failing to conform to Articles 12 and/or 13 of the Code, whereas atelonyms (unnamed category in the Code) are nomina that are unavailable for failing to conform to Articles 1, 3, 10, 11, 14, 15, 16, 19, 20, 24, 32, 33, 34, 50 and/or 79 of the Code.Agiven nomen, once introduced in the scientific literature, can appear under different avatars (paronyms), and particu- larly under different spellings or parographs (Dubois, 2010a): its original spelling or protograph and its subsequent apographs. In rather rare cases, ‘‘multiple original spellings’’ or symprotographs appear in the first publication, and a first-reviser action will then have to determine the correct original spelling or lectoprotograph (Dubois, 2010a).Adifferent and also quite rare situation is that of allelonyms (Dubois, 2006a), i.e., alternative nomina proposed as new altogether for a single taxon in the same publication. The term neonym (Dubois, 2000) is here used to designate the concept called ‘‘new replacement name’’, ‘‘nomen substitutum’’ or ‘‘nomen novum’’ in various successive editions of the Code, and the term archaeonym (Dubois, 2005a) to designate the nomen replaced by a neonym (unnamed category in the Code). The term hyponymous (Dubois, 2006b) is used to point to a subordinate coordinate nomen (‘‘nominotypical’’ in the Code) which is also the nomen of its superordinate taxon. The use of the term ‘‘type’’ in nomenclature may be misleading (Dubois, 2005b), and this term is appropriately replaced by the term onomatophore (Simpson, 1940). There are different kinds of onomatophores. Those of FS and GS nomina, termed in the Code respectively ‘‘type-genus’’ for the former and ‘‘type-species’’ for the latter, are nominal taxa respectively of rank genus and species. They are designated below respectively by the terms nucleogenus and nucleospecies (Dubois, 2005a¢b), which are not based on the inappropriate root ‘‘type’’. The term conucleogenera designates the nominal genera which are the collective onomatophore of a CS nomen (Dubois, 2005a¢b, 2006a). The term monophory (Dubois, 2005b) is here used instead of ‘‘monotypy’’ as used in the Code to designate a mode of designation of onomatophore. Having an onomatophore may not be enough for the unambiguous allocation of a nomen: ‘‘nomina dubia’’ unallocated to any currently recognized taxon in an ergotaxonomy, or aporionyms (Dubois, 2011a), contrast with ‘‘nomina