A New Ergotaxonomy of the Order Urodela Duméril, 1805 (Amphibia, Batrachia)

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 ** Penclen, 56420 Plumelec, France

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 taxonominal 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 particularly 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 clara’’ or photonyms (Dubois, 2011a), the onomatophore of which is clearly taxonomically allocated. Haeckel’s (1866) term monophyletic has been used in different senses in the literature: to avoid confusions, we here use holophyletic (Ashlock, 1971) for ‘‘monophyletic sensu Hennig (1950)’’ (qualification of a group that is neither polyphyletic nor paraphyletic) and homophyletic (Dubois, 1986) for ‘‘monophyletic sensu Haeckel (1866)’’ (qualification of a group that is not polyphyletic).

Introduction

No taxonomy is, or most probably will be, the ‘‘final’’ one for any group of living organisms: this is due to the permanent increase of the organisms and characters we have 80 ALYTES 28 (3¢4) access to, to the improvement of our techniques of analysis and to the changes in our taxonomic paradigms. Therefore, the idea of ‘‘taxonomic and nomenclatural stability’’, widely supported by administrators and technocrats, is in fact praise for ignorance and should not be backed (Gaffney, 1977, 1979; Dominguez &Wheeler, 1997; Dubois, 1998, 2005b, 2010b;Benton, 2000; Schuh, 2003). Rather, any classification and nomenclature of a group of organisms may be viewed as a step along the progress of our knowledge, not as an achievement. Each of these steps results in a provisional ‘‘working taxonomy’’ or ergotaxonomy (Dubois, 2005b), which will be replaced, sooner or later, by a better one, or considered to be such. In the present paper, we explore again the questions posed by the taxonomy and nomenclature (or taxonominal hierarchy;Dubois, 2011c: 51) of the amphibian order including all the recent salamanders and their closely related fossil taxa. Six rather detailed and comprehensive ergotaxonomies of the salamanders were published in the recent years, by Dubois (2005c)1,Frost et al. (2006), Raffaëlli (2007), Zhang &Wake (2009), Pyron &Wiens (2011) and Blackburn &Wake (2011). The first, third and sixth ones were based on the then available published data dealing with the phylogenetic relationships among salamanders (the first and six including the fossil groups). The other three were based in part on new molecular data that led to modify these previous schemes, but only for recent taxa. Besides being based on different hypotheses regarding the phylogenetic relationships among the groups of salamanders, these six ergotaxonomies were relying on very different approaches regarding the nomenclature of higher taxa (i.e., above superfamily). This problem, which arises because the Code (Anonymous, 1999) does not provide Rules for this purpose, is becoming more and more important and frustrating as time passes and as the number of higher taxa increases with the multiplication of molecular phylogenetic works. As this question was discussed in detail in several recent publications (Dubois, 2005b, 2006a, 2011a), only a brief summary will be provided here. Other nomenclatural problems can be identified in these recent ergotaxonomies and will be discussed below. The purpose of the present paper is to present an updated complete (and of course provisional) classification of the salamanders, based on the most recent works dealing with their molecular phylogeny, morphology and life history, and using a consistent nomenclature based on explicit criteria and Rules. For this work, we used the same methodological approach, concepts, terms and tools as in our recent paper devoted to a single family of salamanders, the SALAMANDRIDAE (Dubois &Raffaëlli, 2009). We here adopted an ergotaxonomy of the salamanders (appendix 1) largely based on that of Zhang &Wake (2009). In agreement with several other recent works, they recognize three main groups within the URODELA: the SIRENIDAE, the CRYPTOBRANCHIDAE+HYNOBIIDAE and all other families. However, there is a discrepancy between these works and others regarding the relative phylogenetic positions of these three groups. Whereas several studies (e.g., Wiens et al., 2005; Roelants et al., 2007; Wiens, 2007; Pyron &Wiens, 2011) found the relationships

1. A warning is in order here. On the website AmphibiaWeb (AW) [http://amphibiaweb.org], this paper is mentioned and seems to be available through a link, but the link provided in fact points to another paper (Dubois, 2004a). Only the next two papers (2 and 3) of the special issue Amphibia Mundi 1 of the journal Alytes where this paper was published are available on this website. This ‘‘error’’ was not corrected although it was reported to the AW administrators. A reprint or a pdf of this paper can be provided by the author upon request (address above). It is also accessible on the Web at the address [http://www.cnah.org/pdf_ﬁles/1912.pdf]. Dubois &Raffaëlli 81

(CRYPTOBRANCHIDAE+HYNOBIIDAE)((SIRENIDAE)(all other families)), Zhang &Wake (2009) proposed the pattern (SIRENIDAE)((CRYPTOBRANCHIDAE+HYNOBIIDAE)(all other families)). We here follow these latter authors because their tree is based on complete mitochondrial genomes and above all on a good knowledge of these animals, of their anatomy and biology (not only of their molecular sequences), and because their work includes detailed discussions and makes sense in terms of phylogenetic hypotheses integrating molecular, morphological and life history data, and in terms of paleontology and biogeography. Several authors (Duellman &Trueb, 1986, Larson &Dimmick, 1993) already suggested that both morphological data (no hind limbs, no cloacal glands, no metamorphosis, no crown or pedicel on teeth, external fertilization) and molecular data were in favour of the hypothesis that this family is probably a sister taxon of all other salamanders. In the past, they were even referred by several authors to a distinct order of recent amphibians, under the nomina TRACHYSTOMATA (e.g., Goin &Goin, 1962) or MEANTES (e.g., Goin et al., 1978). Alternatively, sirenids and cryptobranchids¢hynobiids might have diverged nearly simultaneously from a common ancestor. Unfortunately, the nomenclatural aspects of several recent works dealing with the taxonomy of the salamanders is weak and sometimes deficient. Problems can be identified first above the rank family, especially regarding the nomen of the order, but also for nomina of unclear ranks used by various authors for the main groups of families. Problems exist also below the rank family, where several gymnonyms (‘‘nomina nuda’’) were recently introduced for new taxa, and for which we are bound to provide available nomina. The new ergotaxonomy presented here does not use unclear terms like ‘‘taxon’’ or ‘‘clade’’ but recognizes formal taxa designated by formal nomina. The allocation and validity of these nomina is not based on opinions or tastes but on an explicit rationale explained here for higher taxa, and strictly following the Code for the nomina at rank superfamily and below. We took this opportunity to introduce a few other changes in the taxonomy and nomenclature of salamanders, discussed in more details below. We included all fossil salamander taxa currently recognized by paleontologists (see Martín &Sanchíz, 2012), although we are fully aware that very little is known about many of them and that their higher taxonomic allocation is often questionable. We think this is useful in order to keep an eye on the whole group, even in its least explored aspects. This highlights the disparity of fossil material in the different groups, the fossils being more abundant in the most ancient families: for example, very few fossils of PLETHODONTIDAE are currently known, although it is by far the largest family of the order, but also the most recent one. In some cases we think that it is likely that some nomina, currently allocated with doubt to some ‘‘all-fossil’’ taxa, will later prove to designate the same taxa as some recent taxa, especially at genus level, and may in some cases even be the valid nomina for the latter. But our main aim here was to homogenize the treatment of ranks and nomina over the whole order, in line with our previous detailed proposals regarding the SALA- MANDRIDAE (Dubois &Raffaëlli, 2009). In the family HYNOBIIDAE, we implemented some taxonomic and nomenclatural changes implied by the comprehensive work of Poyarkov (2010) on the molecular phylogeny and morphology of these salamanders. We also recognized additional taxa in the AMBYSTOMATIDAE and PLETHODONTIDAE in order to have more balanced and consistent uses of nomenclatural hierarchies and a more 82 ALYTES 28 (3¢4) precise nomenclature for taxa at different levels of this hierarchy. More details on our rationale in these various respects are to be found in Dubois &Raffaëlli (2009) and below. In particular, some basic problems regarding taxonomy, nomenclature and the relationships between these two fields must be discussed again before going into the details of our proposals. A final aim of this work was to provide ‘‘well-thought’’ nomina for a few new taxa. We have had for a long time a great interest in nomina ‘‘per se’’. Nomina are labels for taxa, i.e., tools for the communication between humans, and we think some time and care should be devoted to their characteristics when coining them (Ng, 1994; Dubois & Raffaëlli, 2009; Dubois, 2010c). In some cases, a minimum attention given to some characteristics of the nomina would avoid some trivial nomenclatural problems, such as those posed by some authors who lack a classical background and have difficulties when it is necessary to modify the ending of a specific or subspecific epithet transferred from one genus to another whenever this is necessary because their nomina have different grammatical genders: this problem disappears if, when coining a new generic or subgeneric nomen for some species that are removed from an existing genus, the simple care is just taken to use the same grammatical gender for the new nomen as for the old one. This very simple precaution (Dubois &Raffaëlli, 2009: 21) was unfortunately often ignored by authors of new nomina, including in amphibians and including by one of us (e.g., Blommersia, Bourretia, Hightonia, Nanotriton, Oeditriton, Sahona, Taylorana, Wakea). But this is only one of the problems raised by the little care given by recent authors to the form and characteristics of nomina. The purpose of nomina is not to please their authors (or dedicatees) but to be used in texts, tables, lists and other written documents to refer unambiguously and universally to taxa. They are ‘‘labels’’, nothing more. For this reason, they should be as simple and short as possible, composed of simple syllables and easy to pronounce in most languages. This is much more important in our opinion than their being ‘‘meaningful’’, or of erudite formation based on complex etymologies. In fact, they can be simple ‘‘arbitrary combinations of letters’’, as allowed by the Code, provided that they are euphonious, easy to write, pronounce and remember, and, even better, that they can be easily associated with the nomina of similar or closely related taxa (e.g., through use of a common stem or ending). In the recent decades, we have been appalled by the repeated appearance in zoological nomenclature of nomina which are clearly too long, barbarian, pedant or aggressive to the reader (Dubois, 2010c). Well, how do you pronounce Tylototriton broadoridgus, Saevesoederberghia or Vandijkophrynus?Who wants to use nomina like Bolitoglossa huehuetenanguensis, PSEUDOPHLEGETHONTIIDAE or HYDATINOSALAMANDROIDEI? What is the purpose of imposing to colleagues the use of ‘‘labels’’ like Crotaphatrema tchabalmbaboensis, Hyalinobatrachium guairarepanensis or Batrachochy- trium dendrobatidis (which most biologists now mention as ‘‘Bd’’ because the ‘‘official’’ nomen is too long and unpalatable)? As some taxonomists nowadays clearly do not pay attention to such questions, on several occasions we decided to publish simple and short new nomina for taxa which we thought would sooner or later be recognized, in order to avoid the creation of these terrible nomina that biologists will have to carry for decades once they have been published. We did not regret having done so in some amphibian families (Dubois, 1992; Dubois &Raffaëlli, 2009), and Dubois &Raffaëlli 83 we rather regret not having done so in other families before their nomenclature was spoiled by nomina that won’t be impossible to ignore now. This is one of the (negative) consequences of ‘‘taxonomic freedom’’ for which we have always fought! This motivation was behind our action of ‘‘nomination’’ of taxa in the recent decades, but the motivation of being the ‘‘authors’’ of these taxa was not at stake. On the contrary, we think that the presence of the names of authors within the nominal-complexes of nomina is a real nuisance to zoological nomenclature. It is certainly largely responsible for the erection of unwarranted taxa by authors who wanted to ‘‘associate their names’’ to taxa in order to become ‘‘immortal’’. These nomina will increase the ‘‘synonymy load’’ of the taxa which they apply to. They will later have to be synonymized, thus increasing the burden of useless works by taxonomists, as if the latter had nothing better and more urgent to do in the ‘‘century of extinctions’’ (see Dubois, 2010b). The recent proposal (Dubois, 2008b) to suppress the author’s name in the nominal-complex was meant at reducing the impact of this problem. We hope it will eventually be adopted by the international community of taxonomists, and we implement it below for the nomina of species, exerges and subspecies in appendix 1. We already showed our lack of interest for this ‘‘authorship quest’’ by our previous publication of suggestions of nomina of plethodontid taxa for which we had on purpose not met the criteria of availability (Dubois, 2008e), leaving their formal nomenclatural founda- tion and authorship to subsequent authors working on these groups, but unfortunately they did not seize this opportunity. In the present work, we preferred to publish new nomina complying with the Code rather than still contributing to an increase of the nomenclatural chaos in this family in erecting taxa but not naming them properly.

The basic distinction between taxonomic categories and nomenclatural ranks

We are facing a very strange situation nowadays regarding the use of hierarchies and ranks in zoological taxonomy.On one hand, a number of authors agree that taxonomy should be ‘‘phylogenetic’’, i.e., that it should express in some ways our hypotheses regarding the cladistic relationships between organisms. Let us insist here on these being hypotheses of relationships, not observed facts: taxa are hypotheses, models or theories. Taxa are therefore defined, not ‘‘discovered’’, as some put it: this is well shown by the fact that different sets of data or the use of different methods of analysis of these data very often produce different, or even contradictory and incompatible, phylogenetic trees, even over a very short period of time ¢ as well illustrated by the recent works on the salamanders of Frost et al. (2006), Zhang &Wake (2009) and Pyron &Wiens (2011). Some of the supporters of ‘‘phylogenetic taxonomy’’ are in favour of the use of ‘‘phylogenetic nomenclatures’’ like the Phylocode, despite the major problems, both theoretical and practical, raised by this approach (see e.g. Dubois, 2005b). Thus doing, they propose to stop using nomenclatural ranks in their taxonominal hierarchies, arguing that these ranks are ‘‘meaningless’’ because they are not equivalent in different groups of taxa. This statement is misleading and based on an essentialist belief in absolute ranks that would convey some 84 ALYTES 28 (3¢4) information on the ‘‘nature’’ or ‘‘essence’’ of the taxa referred to them. In fact, in taxonomy ranks only have a relative function which is to inform on the internal structure of the taxonominal hierarchy (Dubois, 2007a, 2008e). Unfounded as it may be, this criticism of ranks has apparently convinced some other taxonomists who, although they do not follow the Phylocode, have stopped using ranks, at least for higher taxa, which they all refer to under the undifferentiated terms ‘‘taxon’’ or ‘‘clade’’. But they do so in an heterogeneous and inconsistent way, because they still use ranks (like superfamily, family, subfamily, genus, etc.) for the lowest supraspecific taxa and above all because they use different ranks or denominations for taxa that appear as sister-taxa in their trees, such as ‘‘taxon’’ and ‘‘family’’, ‘‘family’’ and ‘‘genus’’, ‘‘tribe’’ and ‘‘genus’’ or ‘‘genus’’ and ‘‘species-group’’. Such taxonominal schemes are pseudoranked (Dubois, 2007a, 2008e) and do not carry any reliable information on the phylogenetic hypotheses on which the taxonomy is supposed to be based. Despite the apparent agreement of many on the benefits of ‘‘phylogenetic taxonomy’’, their still frequent use in recent ergotaxonomies testifies to a strong persistence of a gradist conception of taxonomy, according to which the ranks of taxa would in some way provide information on their ‘‘degree of divergence’’ (whether morphological, genetic, cytogenetic, ethological, ecological, etc.) between these taxa, or on the ages of the splits between clades. However, despite the statements of the Phylocode supporters and others, it is quite possible, not to say easy, to use nomenclatural ranks to express hypothetic cladistic relationships in an ergotaxonomy: this only requires to always afford the same nomenclatural rank to the two (or more in case of unresolved polytomy) taxa that are considered sister- groups (Raikow, 1985; Sibley &Ahlquist, 1990; Dubois, 2007a, 2008e). This is in fact the only, but very useful indeed, role that ranks can play in taxonomy, as ranks ‘‘by themselves’’ carry indeed no information and are not equivalent by any criterion between different taxonomic groups, a fact that was stressed, often independently, in the recent decades by different authors who often ignored their respective works (e.g., Schaefer, 1976; Dubois, 1988, 2005b, 2007a, 2008e, 2011a,c;Smith, 1988; Sundberg &Pleijel, 1994; Minelli, 2000; Pleijel &Rouse, 2003; Kluge, 2005; Bertrand et al., 2006; Laurin, 2010; Avise &Liu, 2011). This problem has been emphasized by the long maintained confusion between two concepts: whereas taxonomic categories (such as mixiological species or mayron, phylogenetic species or simpson, klepton, superspecies, synklepton, mixogenus; for details see Dubois, 2008d, 2009b, 2011b, and below) do carry some biological or historical information about taxa, nomenclatural ranks convey no such meaning but just allow to express the structure of the taxonomy or of the tree on which it is based. This rejection of relative ranks is accompanied, in some recent works, by an apparent renewed belief in absolute ranks, such as ‘‘family’’ as a basic standard in taxonomy, as well illustrated in the recent special volume on ‘‘animal biodiversity’’ (Zhang, 2011a; ABD below). The editorial choice of this volume was to list only taxa at the most frequently used ranks (such as phylum, class, subclass, order, suborder, superfamily, family, subfamily), with the idea that the mere numbers of these taxa would provide ‘‘diversity estimates’’ (Zhang, 2011b: 7). This aim by itself is confusing and meaningless. At the species level, this would be true only if all authors were using the same ‘‘taxonomic species concept’’, which is far from being the case, except sometimes among closely related organisms. But this is even more irrelevant as far Dubois &Raffaëlli 85 as supraspecific classification is concerned. It then relies on the untold assumption that numbers of supraspecific taxa provide so to say a ‘‘measurement’’ of the taxonomic or phylogenetic diversity of a group, an idea which is based on the wrong belief that higher taxa sharing the same rank are equivalent by some criterion. Strangely, this basic problem was mentioned in only one contribution of ABD, even if it was not fully understood even there. Thus, Pape et al. (2011: 222) were fully right when they wrote: ‘‘While groups may have the same rank, they may not be equivalent and should not be used to make comparisons between similarly ranked taxa. For example, the family Limoniidae contains over 10,000 species and is over 235 million years old, whereas the family Braulidae contains only 7 species and is less than 30 million years old.’’ But they were wrong when they wrote: ‘‘We have reluctantly assigned the traditional Linnaean ranks to the groups in our list, with the caveat that this implies an equivalence that does not exist.’’ As a matter of fact, the use of nomenclatural ranks does not ‘‘imply’’ such equivalence at all. It does so only for those who adopt the gradist concept that ranks carry some information on the ‘‘amount of divergence’’ (measured by whatever criterion) or on the ‘‘time elapsed since separation’’ between taxa, i.e., who perpetuate the old confusion between nomenclatural ranks and taxonomic categories. In the present work as in our previous one (Dubois &Raffaëlli, 2009), we use nomenclatural ranks of taxa as a simple way to transcribe nomenclaturally a phylogenetic taxonomy, i.e., strictly as a tool to indicate hypothesized cladistic relationships between these taxa, but not in the least to indicate amount of divergence between them or time of separation between their ancestors. Therefore, within a given group, two taxa considered sister-taxa are always afforded the same rank, but the same rank in different groups does not have the same ‘‘meaning’’, and ranks cannot be used for any external comparison based e.g. on numbers of species or of supraspecific taxa.

Nomenclatural problems

Class-series nomenclature

In zoological nomenclature and as defined by Dubois (2000), the class-series (CS) accommodates the nomina of all taxa at ranks above superfamily. The Code does not provide Rules for these nomina, which results in a great confusion, if not a real chaos, in higher zoological nomenclature. For this reason, Dubois (2004a, 2005a¢b, 2006a), after a detailed discussion of various aspects of this question, provided a set of Rules for this management. As, since then, the International Commission on Zoological Nomenclature (ICZN) has not yet promulgated any improvement of the Code in this respect, we here follow largely these proposed Rules. We do not follow them exactly, because, in the last six years, our unpublished ‘‘experimental’’ application of these proposed Rules, in collaboration with several colleagues, to several zoological groups, has shown that they could be profitably simplified in several respects. These changes and their reasons for these changes will be developed in full elsewhere (Dubois, in preparation), but for the purpose of the present paper it is enough to explain the rationale we followed here to establish the valid nomina of the class-series taxa of salamanders. 86 ALYTES 28 (3¢4)

The nomenclatural system in force in zoological nomenclature relies on a three-step process (Dubois, 2005b, 2011a) that distinguishes ﬁrst the criteria of nomenclatural availability of nomina, then those of allocation of nomina to taxa as recognized in a given ergotaxonomy, and ﬁnally those of validity of nomina for given taxa in an ergotaxonomy. Let us examine these three steps successively.

Availability

The availability of class-series nomina relies first on the ‘‘usual’’ criteria of availability in zoological nomenclature highlighted by the Code: paper publication after 1757, presence of a definition, diagnosis, description or indication of characters of the taxon, etc. Two particular problems are posed in the class-series. The first one concerns the many early nomina that were first published in a non-latinized form. These nomina do not qualify as ‘‘vernacular names’’, because the term ‘‘vernacular’’ means ‘‘of one’s native country, native, indigenous, not of foreign origin or of learned formation’’ (Fowler &Fowler, 1929: 1370). On the contrary, these nomina were not names used in the common language of the country but new terms coined on purpose by zoologists to designate taxa in international scientific language, almost invariably derived from classical Greek or Latin roots and then transferred into the language of the author (French, German, English, etc.). These are not vernacular names but French, German, etc., scientific nomina. The Code (Article 11.7.2) recognizes that family-series nomina first published in a non-latinized form but subsequently latinized may under certain conditions be nomenclaturally available, their authorship and date being those of the original publication as non-latinized nomina. Even in the genus-series, some nomina that were not originally Latin may be ‘‘treated as Latin’’ and accepted as available from their first publication (see Kottelat, 2001; Dubois, 2009a). In such conditions, it would be completely unjustified to retroactively deny nomenclatural availability to scientific nomina of the class-series first published in a non-latinized form, all the more that class-series nomina have been kept until now outside the Rules of zoological nomenclature and therefore not submitted to any Rule. As long as authors’ names are mentioned in the nominal-complexes of zoological nomina, doing so would result in ‘‘stealing’’ many old CS nomina to their real authors ¢ who were not only the nomenclatural authors of their nomina but also the authors of the concepts of the taxa to which they apply ¢ to ‘‘offer’’ them to authors who sometimes were just translators or popularisators of their works and had not contributed in the least to the progress of ideas in the domain of taxonomy at stake. This is why Dubois (2006a) simply required in his Rule [R2] that, to be available, a CS nomen published in a language other than Latin ‘‘should have been latinized before 1st January 2007’’. Regarding this question, we think the Article 11.7.2 of the Code (1) is not fully clear as it relies on the vague concept of ‘‘generally accepted as valid’’ which is not defined in the Code, and (2) is inappropriate and should be modified to allow for availability of all scientific nomina published in Latin-form provided they have been latinized before 1900.

Application of this Rule to the nomina of higher taxa of AMPHIBIA has important nomenclatural consequences, as discussed already by Dubois (2004a, 2006a, 2009a): contrary to the statements of some recent authors, nomina like BATRACIENS Brongniart, 1800, ANOURES Duméril, 1805, URODÈLES Duméril, 1805 or AMPHYBIENS de Blainville, 1816, which have soon Dubois &Raffaëlli 87 been subsequently latinized, are nomenclaturally available with their original authors and dates, and are in fact the valid nomina for the taxa they designate. A second, important, problem regarding availability concerns the distinction between family-series and class-series nomina. As the Code does not provide any Rule or clue in this respect, different authors have applied different rationales (or more often no rationale at all), which has resulted in a variety of incompatible decisions regarding the allocation of some nomina to one of the two nominal-series, with of course important consequences regarding the valid nomina of some taxa. Dubois (2006a) proposed two Rules [R4] and [R5] that allow in all cases an objective distinction between the two situations, and Dubois &Bour (2010) provided further discussion and details. An important point to understand here is that the rank ‘‘family’’, as a rank intermediate between order and genus, and for nomina based on generic nomina, did not exist in Linnaeus (1758) and in the early days of zootaxonomy. It appeared only later, first in an isolate way in Batsch (1788, 1789, 1796), but became in current use in zoological taxonomy only after the first quarter of the 19th century. The current Code requires that, to be available in the family-series, a nomen should be a rhizonym (Dubois, 2006c), i.e., should have been coined by addition of a suffix denoting plural to the stem of an available, and then considered valid, genus-series nomen (an important distinction, because two nomina may ‘‘look similar’’ for being based on the same classical root, although not designating the same organisms or taxa). But this condition is not sufficient, as some unquestionable CS nomina (being explicitly referred to one of the ‘‘classical’’ Linnaean ranks phylum, class or order) are also rhizonyms, which may be a strong source of confusion. This complex problem is not discussed further here, but will be so in detail elsewhere (Dubois, in preparation). It results from this complex situation that, in the early texts of zootaxonomy, the fact that an author used the denomination ‘‘family’’ for a suprageneric taxon is not reliable evidence that this nomen should be referred to the family-series. There are two main situations. In the first one (S1), all ‘‘family’’ nomina in such a work are arhizonyms (Dubois, 2006a), i.e., are not based on the roots of generic nomina then considered valid: they must then be referred to the class-series, not to the family-series. Although this was not specified in any set of Rules before those of Dubois (2006a), such an ‘‘untold rule’’ was followed by most authors in the past. In contrast, in situation (S2), where all or even some or a single one of these nomina are rhizonyms, according to Rule [R5] of Dubois (2006a) they must all be referred to the family-series, but then those which are arhizonyms are nomenclaturally unavailable FS nomina (and not available CS nomina). These general ideas may now be applied to some early nomina of higher taxa in zoology. Thus, in his review of the animal kingdom, Duméril (1805) introduced 151 nomina of ‘‘families’’. They were all scientific nomina coined on the basis of Greek roots pointing to characteristics of the taxa, but most of them (131) appeared only under a French form. As they were all soon latinized subsequently, they are available in zoological nomenclature, dating from Duméril (1805). None of these 151 nomina is a rhizonym and therefore they should all be referred to the class-series, not to the family-series. These nomina are not unavailable FS nomina but fully available CS nomina, as acknowledged by a few recent careful authors working on different zoological groups (e.g., Dubois, 2004a, 2009a;Pape et 88 ALYTES 28 (3¢4) al., 2011: 222). This has an important bearing on the valid nomen of the order of salamanders, as we will see below.

Allocation Taxonomic allocation of CS nomina is also a very problematic question, possibly even more than the previous one. As it is a complex one, which was discussed in detail in several publications already (Dubois 2005a¢b,d, 2006a,c, 2009a, 2011a;Dubois &Ohler, 2009; Dubois &Bour, 2010), it will only be tackled briefly here. Recent examples, such as the inconsistent use of criteria for the allocation of CS nomina to taxa in ABD (Dubois, in preparation), show that it is impossible to produce an internally consistent higher nomenclature of any zoological group if no Rules or guidelines are followed for the allocation of nomina to taxa. The absence of any statement in this respect in the Code is a strong nuisance for the uniqueness, universality and robustness of zoological nomenclature. For this reason, in the present work we followed strict Rules for the allocation of CS nomina to taxa. These Rules are slightly modified compared to those of the original Ambiostensional Nomenclatural System (AONS) proposed by Dubois (2006a, 2011a). Although this new change of Rules has a few drawbacks from the rather abstract point of view of an ‘‘ideal’’ nomenclatural system, it so to say ‘‘imposed’’ itself to us because, in the last six years, the unpublished ‘‘experimental’’ application of the original Rules to several zoological groups showed us that the implementation of the part of the Rule [R11] dealing with the taxonomic allocation of nomina through extragenera as onomatostases was of complex and heavy use for individual zootaxonomists without the help of an appropriate computer software. Although such a software is now available (Gérard et al., 2006; Berkani &Dubois, in preparation), we think nomenclatural Rules should be simple enough to be usable by hand, without the help of computers. The improved system, the Metrostensional Nomenclatural System (MONS), will be presented in detail elsewhere (Dubois, in preparation), but for the purpose of the present paper it is enough to explain briefly this system of allocation of CS nomina which we applied here to the higher taxa of salamanders. This system simply relies on inclusive ostensional allocation of nomen to taxon through a collective onomatophore consisting in the set of genus-series nomina referred as valid by the original author to the nominal taxon for which the nomen was created. This set of conucleogenera is considered an indissoluble unit from which no member can be withdrawn ¢ with two exceptions: unavailable nomina (anoplonyms) and nomina unallocated to any current genus- series taxon in the ergotaxonomy of reference (aporionyms). The analysis therefore bears only on the conucleogenera which are both available (hoplonyms) and taxonomically allocated and identified (photonyms). Allocation of a CS nomen to an ergotaxon simply relies on the current taxonomic allocation of all these conucleogenera. The CS nomen applies then, within the ergotaxonomy at stake, to the metrotaxon (Dubois, 2006a), i.e., the least inclusive CS ergotaxon including all the conucleogenera of the protaxon. This system is fully objective, unambiguous and automatic. Its implementation is very quick for many old CS nomina that were introduced in the early times of zootaxonomy, for taxa that included only a few genera. It is slightly heavier to implement in the case of recent CS nomina proposed for taxa including dozens or hundreds of conucleogenera, but it remains manageable by hand by any careful Dubois &Raffaëlli 89 taxonomist, and even more so using an appropriate software (Berkani &Dubois, in preparation). During the long history of zoological nomenclature, many CS nomina have gone through drastic changes in their allocation to taxa. As explained in detail by Dubois (2006a), as long as such a change consisted in an addition of specimens or lower taxa to the original onomatophore of the CS nomen, no change of nomenclatural concept occurred and no problem arose, but as soon as it consisted in a subtraction from this original onomatophore, it resulted in fact in the introduction in zoological nomenclature of a new CS nomen, with its own distinct onomatophore. Normally, such a new nomen will simply have to disappear as a junior homonym of the first nomen published, but in some rare cases a different solution will have to be implemented, as we will now see.

Validity

The absence for two and a half centuries of any Rule regarding the nomenclature of higher taxa in zoology has had a major consequence: the incredibly high number of synonyms published for the same taxa, at least in some zoological groups, those that have been studied by a high number of zoologists. This growing chaos, which strongly hampers communication among zootaxonomists and between them and all other biologists and users of zoological classifications and nomina, needs to be remedied for, and there is no other serious solution for this than following universal and stringent Rules for the validity of nomina. Here also we follow the Rules proposed by Dubois (2006a), except from a few minor changes. The system based on the Principle of Priority of publication, complemented in case of simultaneous publication by the Principle of First-Reviser, has proved to be a very efficient, powerful and honest system for the clarification and stabilization of zoological nomenclature in the three nominal-series covered by the Code. It is more than time to implement this system in zoological class-series nomenclature. These two Principles should apply regarding homonymy, synonymy and parography (see Dubois, accepted). Concerning the Principle of First-Reviser (FRA), it is suggested that, in the class-series, be adopted the distinction introduced in Article 24.2.4 of the last edition of the Code between Internal First-Reviser Action (IFRA) and External First-Reviser Action (EFRA) (see Dubois, 2010a). These FRAs are limited in the Code to the choice of the valid spelling (lectoproto- graphs) among multiple original spellings (symprotographs). Full details on this matter were givenbyDubois (2010a) and it would be too long to repeat them here. Briefly, in the case of an EFRA, the notion of first-reviser is strictly defined: to be valid and definitive, the choice between symprotographs must be explicit, through mention of the competing spellings and clear choice between them. The conditions are not as strict in the case of an IFRA: here it is enough for the original author to have mentioned a single symprotograph in a subsequent publication for this to be deemed to be an explicit and valid FRA. We suggest to adopt this distinction in CS nomenclature, not only in the case of multiple original symprotographs (different spellings of a nomen), but also in that of allelonyms (different nomina for the same taxon). In both situations, we propose that both kinds of FRAs should be accepted as valid, priority being given in all cases to the first FRA published. However, the following condition should be added, because its absence in Article 24.2.4 of the Code poses problems: for the 90 ALYTES 28 (3¢4) purpose of IFRA, the term ‘‘original author’’ should be understood as meaning ‘‘all the persons that were members of the original signature’’ of the first publication. This means that the choice by IFRA of a valid spelling or nomen among symprotographs or allelonyms is valid only if its subsequent use, without mention of the alternative one(s), appeared in a publication the signature of which includes the names of all these original authors (not necessarily in the same order, and possibly with additional new authors). In the class-series, homonymy should be understood not as strict homography (see Dubois, 2012) but as identity of the root or stem on which the nomen was formed ¢ just like in the family-series. Thus, nomina like NUDA, NUDI and NUDES,orICHTYOIDA, ICHTHYOIDEA and ICHTHYODEI, if proposed for different taxa and not as emendations or apographs of a same nomen, should be considered homonyms, and the junior ones should be definitively invalidated, even if the senior one is not itself valid in the frame of a given ergotaxonomy. The same should apply to synonymy, whether it is objective (isonymy;Dubois 2000) for being based on the same onomatophore (i.e., the exactly same list of conucleogenera) or subjective (doxisonymy;Dubois 2000) for being based on a taxonomic interpretation of the allocation of their different onomatophores: only the first published nomen should be potentially valid, if it is not invalid itself for being a junior homonym. However, whereas isonymy results in the definitive invalidation of a nomen, doxisonymy is labile, as a nomen once invalidated for being considered a junior doxisonym in a given ergotaxonomy may become valid again under a new taxonomic arrangement. These two modes of invalidation of nomina, by junior homonymy or junior synonymy, are by no means new in zoological nomenclature, as they have been in force in the three nominal-series covered by the Code for more than a century. But here we are in a new situation, and special care and attention should be paid to the risks that could be carried by a too rigid implementation of the Principle of Priority to nomina that have never been submitted to it. Some CS nomina, such as AMPHIBIA, COLEOPTERA or CRUSTACEA, were proposed originally for taxa that were much more comprehensive that the taxa to which they apply nowadays. They have been in permanent use in thousands of publications for two centuries or more, for the same taxon or for very close taxa, and having to rename them because of newly implemented nomenclatural Rules could be very disturbing for taxonomy and even for the whole of biology. Particular Rules should therefore allow conservation of such nomina, but these Rules should be devised with care, in order to avoid falling in another trap, that of validating a poorly documented ‘‘usage’’ by a few zoologists in order to preserve their habits, tastes and peace of mind. This latter problem was discussed in detail elsewhere (Dubois, 2005a¢b, 2006a, 2010b), and the best solution appears to implement well-defined categories of usage of nomina. Three such categories are particularly useful and will be discussed here: those of sozonym, sozodiaphonym and distagmonym. The first and last ones were defined by Dubois (2005a) and the second one is new. Sozonyms are nomina that have had since 1900 a real massive usage in the general literature, i.e., not limited to the specialized taxonomic literature, while no other nomen had a large usage for the same taxon: this justifies their ‘‘protection against priority’’. Sozodiaphonyms are nomina that have also had a large usage, but alternatively to other nomina which also had a large usage. Distagmonyms are nomina which have not had a large usage since 1900. Dubois &Raffaëlli 91

The formula ‘‘large usage’’ must of course be clearly defined to be operational. Rule [R12] of Dubois (2006a) defines sozonyms as nomina that have been applied after 1899 by all or almost all authors to designate taxa traditionally recognized outside the narrow community of taxonomists, phylogeneticists and systematists, whereas no other nomen has been used significantly for the same taxon. For the purpose of this Rule, the term significantly is to be understood as qualifying a nomen that has been used for a taxon, either in its Latin form or under a non-latinized form, either (1) in the titles of at least 25 non-purely systematic books, written by at least 25 independent-authors and published in at least 10 different countries after 31 December 1899, or (2) in the titles of at least 100 non-purely systematic publications (books, book chapters or periodical articles) written by 100 independent-authors and published in at least 10 different countries after 31 December 1899. The details and justifica- tions of these conditions were discussed elsewhere (Dubois, 2005b, 2006a, 2010b). According to the Rule [R13] of Dubois (2006a), a sozonym, if it is not a junior homonym of another sozonym, must be conserved through sozonym validation. This is not the case for sozodiaphonyms and distagmonyms, which should follow the ‘‘normal’’ Rules of validity, with precedence fixed by Priority or FRA. Recent unpublished discussions of this question (Dubois, in preparation) have led to the conclusion that the detailed criteria of ‘‘agathonym validation’’ proposed in Rule [R16] of Dubois (2006a) should be much simplified. For the purpose of this Rule, it is enough to distinguish two categories of nomina: sozodiaphonyms and distagmonyms. Sozodiaphonyms are nomina that have had a large usage since 1900 and that meet the quantitative criteria required for sozonym validation, but that were not the only ones to meet these criteria, having been used alternatively for the taxon at stake by different authors. In the new simplified Rule [R16], sozodiaphonyms should always take precedence over distagmonyms. If sozodiaphonyms are available in a synonymy, precedence (through Priority or FRA) should be working only between them; if not, precedence should be working indiscriminately among distagmonyms. It is important to note that, among sozodiaphonyms, just like among distagmonyms, only the criteria of Priority and FRA can be used for establishing validity. It would be inappropriate, and so to say ridiculous, to compare the numbers of usages of each nomen in such cases, because nomenclature is not a football game or a movie award competition: the important question in such cases is simply to know whether a single nomen has been used in the general literature for the taxon at stake, or whether two or more nomina are in this case. As was noted by Dubois (2005b: 410), a purely quantitative approach of usage would be philosophically, ethically and scientifically untenable: ‘‘In all cultural domains including science, as soon as two different opinions exist, both have their philosophical justification and are entitled to consideration, irrespective of their numbers of supporters, of the institutional ‘importance’ of the latter, of their financial situation or other similar criteria: no quantitative criterion is useful to decide which opinion is ‘more important’ or ‘more significant’ and therefore should be followed (actually, the history of science is full of examples where the majority was later shown to have been completely wrong).’’ 92 ALYTES 28 (3¢4)

Nomenclatural ranks and the taxonomy of the URODELA

The hierarchy of nomenclatural ranks

As we have seen above, in zoological taxonomy, nomenclatural ranks (and nominal- series as well) do not have any ‘‘meaning’’ by themselves but are very useful, if used consistently, to provide unambiguous information on the structure of the phylogenetic tree on which a classification is based. For this, the only but strict Rule that should be followed is that taxa that are considered sister-taxa (resulting from a process of cladistic splitting) should be given the same nomenclatural rank. This point is central for a proper usage of ranks, so that Dubois (2011a: 21) proposed to incorporate this as a Rule in the Code and even suggested that works where new nomina were published without following this Rule should be declared nomenclaturally unavailable, just like for works not using binominal nomenclature. Nomenclatural ranks are arbitrary, but not all of them have the same importance: a few are so to say compulsory, whereas all the others are optional. This is because, beside their ‘‘explanatory’’ function (i.e., providing an evolutionary interpretation and explanation of the diversity of organisms, in particular through scientific hypotheses about their phylogenetic relationships), biological classifications have a second, ‘‘practical’’ function (i.e., providing a universal system of storage and retrieval of the taxonomic information) (Mayr, 1969, 1981, 1982, 1997). Ignoring the second of these functions to concentrate only on the first may seem appealing as a ‘‘purer’’ approach and may fascinate some professional taxonomists and theoreticians, but is not doing a service to taxonomy and its innumerable users in all domains of human activity (Cracraft, 1974; Ashlock, 1984; Benton, 2000; Dubois, 2005b). This is why Dubois (2006a: 217; 2007a: 50) and Kuntner &Agnarsson (2006) independently suggested that most animal taxa of the planet are or should be referable to seven primary key ranks (species, genus, familia, ordo, classis, phylum, regnum), which could thus be considered virtually ‘‘compulsory’’. This proposal follows a long tradition in zootaxonomy. This is illustrated in figure 1 of Dubois (2007a: 49): some taxa must be named not because this brings additional phylogenetic information but simply in order to have a representative of the group at stake in one of these primary key ranks. Such taxa may be ‘‘redundant’’ relative to some of their included taxa (often a single species or other terminal taxon) but they are useful for information storage and retrieval. A famous example is that of Orycteropus afer, which is the only recent species of the mammal genus Orycteropus, the latter the only recent genus of the family ORYCTEROPODIDAE and the latter the only family of the order TUBULIDENTATA. Because these taxa are redundant, they have the same diagnoses, as there exist no such things as ‘‘specific’’, ‘‘generic’’, ‘‘familial’’ or ‘‘ordinal’’ characters in taxonomy, the ranks being arbitrary (see Dubois, 1988). But this should not be used as a reason for abandoning the use of taxa at the ranks genus, family and order for this species (even if no fossils of this group were known) and for referring the latter directly to the infraclass PLACENTALIA. For the purpose of storage and retrieval of information, it is important to be able to see immediately, e.g. through a simple look at a taxonominal hierarchy like that of Wilson &Reeder (2011), that in this ergotaxonomy the order TUBULIDENTATA is one of the members of a currently unresolved polytomy including 20 other orders. Dubois &Raffaëlli 93

In contrast, additional ranks should be used only when this brings some additional phylogenetic information, which depends on the structure, more or less divided and balanced, of the tree. Ranks by themselves, including the primary key ranks, are arbitrary, and the decision to recognize a taxon as a family, an order or a class does not rest on scientific criteria but merely reflects the traditions in force in the taxonomic literature dealing with the group at stake. This is an important point and this should be maintained in order to leave some ‘‘basic landmarks’’ for all users of taxonomic data and nomina who are not phylogeneticists or taxonomists. There would be no scientific benefit in deciding all of a sudden that the AMPHIBIA are no more a class but a superorder, the ANURA no more an order but an infra-order, or the SALAMANDRIDAE no more a family but a tribe. Taxonomists should be left free to choose the taxa which they allocate to primary key ranks. But, for a clear use of nomina, nominal-series and ranks for communication about the structure of a tree or of a taxonomy, this freedom should stop here. The ranks of the taxa that occupy intermediate positions between the key ranks should follow some Rules or at least a clearly identified and ‘‘automatic’’ rationale. As an imaginary example, no taxonomist would consider appropriate to use a nomenclatural hierarchy using only the four following ranks in ascending order: subspecies (S), subgenus (G), superfamily (F) and class (C): probably all taxonomists would refer the taxa (S) to the rank species, (G) to the rank genus, (F) to the rank family and (C) to the rank class, and would also recognize an additional rank order (O) between (F) and (C). The same applies to many other situations, although usually in a less caricatural or obvious way.

Let us take one example among the AMPHIBIA ANURA.Sanchíz (1984), after an analysis of the relationships among the genera of the family then called DISCOGLOSSIDAE, considered that a taxon containing three genera including the genus Alytes should be separated taxonomically from a second taxon with the three other genera then recognized in this family including Discoglossus, and erected a ‘‘tribe ALYTINI’’, but no other suprageneric infrafamilial taxon in this family. This action was twice inappropriate, first because the erection of a tribe required the erection in parallel of a sister-tribe for the other genera, and second because tribes should not be erected directly below the rank family, but only below the rank subfamily (Dubois, 1987: 12). Therefore, whereas the allocation of primary key ranks should remain free and decided upon by taxonomists on the basis of tradition and of homogeneity and consistence with other closely related taxa, the ranks given to taxa above them (angiotaxa;Dubois, 2005b: 406) and below them (endotaxa;Dubois, 2005b: 406) should be automatically determined by the hierarchical structure of the taxonomy, reflecting that of the tree on which it is based. Some guidelines or, even better, Rules about this in the Code would no doubt be of a great help in this respect. Dubois (2006a: 208, 211¢225) proposed an ‘‘ideal’’ expanded nomenclatural hierarchy using 9 primary and 10 secondary key ranks (from forma to imperium), and 5 descending and 5 ascending subsidiary ranks (from infim-tosuprem-) for each of the 19 key ranks, the total providing up to 209 potential ranks. This is probably much more than what any taxonomist working at any given level of the zoological taxonomic hierarchy will ever need, especially because not all nodes of the hypothetical ‘‘Tree of Life’’ need to be named (Dubois, 2006a). But this system would be possible, at least compatible with the Code, only if the latter was 94 ALYTES 28 (3¢4) modified in order to allow the use of an undeterminate number of ranks in all nominal-series, which is far from being the case nowadays. The current Code only allows the use of two ranks in the genus-series and four in the species-series. In the family-series, the number of ranks is not limited but the ‘‘last’’ rank authorized upwards is superfamily. There is no theoretical or practical justification for these limitations, which clearly infringe upon ‘‘the freedom of taxonomic thought or action’’ and are therefore unacceptable according to the ‘‘Preamble’’ of the Code itself (Anonymous, 1999: 2). However, until these restrictions are cancelled (Dubois, 2011a), taxonomists who want to remain within the Rules of the Code are limited in the number of ranks they can use. Under the nomenclatural paradigm summarized above, whereas primary key ranks are compulsory, the use of additional ranks (secondary key ranks and subsidiary ranks) is warranted only if it provides some additional information on the structure of the taxonomic hierarchy and by the way of the phylogenetic tree, but additional ranks should not be used to carry a gradist message about the ‘‘divergence’’ or ‘‘age’’ of taxa. In this respect we disagree with quite frequent practices in taxonomy, as can be shown by an example in the salamanders. In their subgenus Pachymandra of the genus Bolitoglossa,Parra-Olea et al. (2004) recognized two monospecific ‘‘species groups’’ (alvaradoi and dofleini). The recognition of these two taxa, presumably aiming at expressing the fact that the two species at stake are ‘‘widely divergent’’, is taxonomically redundant with the recognition of the two species, as it does not point to a cladogenesis between two supraspecific taxa and therefore provides no additional taxonomic information. In each case, the species and the ‘‘species group’’ cannot have but the same diagnosis. Except in the case of primary ranks, where this can be useful for purposes of information storage and retrieval as explained above, two subordinate taxa should be recognized within a superordinate taxon only when one of them at least contains itself at least two species, otherwise it is meaningless.

Nomenclatural ranks in the order URODELA

Let us just consider here all the ranks that are necessary, under such constraints, to express in an expanded, unambiguous and rigorous way the hierarchical structure of the taxonomy of the URODELA that we adopted here (appendix 1), which is based on the last phylogenetic data and hypotheses currently available. From the rank subspecies to the rank order, these ranks are in the number of 17, including 3 ranks in the class-series, 7 in the family-series, 2 in the genus-series and 5 in the species-series.

Primary key ranks Let us start with the four primary key ranks involved in this hierarchy: order, family, genus and species.

The taxon including all recent salamanders, the valid nomen of which is URODELA but which has also been called CAUDATA, has been aﬀorded the rank order by a very vast majority of authors since the middle of the 19th century. As there exists no deﬁnition of what an order ‘‘is’’, there would be no point in challenging this tradition. This requires of course that the sister-taxon of this order be the order ANURA, and that both be included in a superorder BATRACHIA sister to a superorder GYMNOPHIONA (Dubois, 2004a). Dubois &Raffaëlli 95

At family level, a rather strong tradition and a good consensus have been in force for a long time among taxonomists (e.g., Gorham, 1974; Dubois, 1985, 2005c;Duellman & Trueb, 1985; Frost, 1985; Laurent, 1986; Frost et al., 2006; Zhang &Wake, 2009; Pyron &Wiens, 2011) to recognize the following 8 urodelan families as valid: AMBYSTOMATIDAE, AMPHIUMIDAE, CRYPTOBRANCHIDAE, HYNOBIIDAE, PLETHODONTIDAE, PROTEIDAE, SALAMANDRIDAE and SIRENIDAE. As the rank family does not mean anything by itself either, there would also be no reason to change this tradition if this is compatible with the phylogenetic tree adopted. In contrast, other families (DICAMPTODONTIDAE, RHYACOTRITONIDAE and several all-fossil families) have less regularly been recognized and their recognition will depend on the structure of the taxonomic hierarchy and their relationships with the other families. At genus level, the consensus between authors is also good, although less than for higher taxa. The rank genus also lacks a scientific ‘‘definition’’, so that it is acceptable to take some arbitrary decisions at this level: see e.g. the discussion of Dubois &Rafaëlli (2009: 13¢14) about the possible use of the crossability criterion at genus level. Finally, there is no consensual use regarding the ‘‘species concept’’ that should be used in salamanders. The situation at this level is made more difficult by the frequent confusion made between ‘‘species’’ as a nomenclatural rank (which is unique) and as taxonomic categories (which are multiple). From the latter point of view, many authors tend to follow a ‘‘biological’’, ‘‘mixiological’’ or mayron concept, others a ‘‘phylogenetic’’, ‘‘unified’’ or simpson concept, and others favour a plurality of concepts according to the evolutionary situation, for example in the case of complexes of gynogenetic polyploid forms that qualify as kleptons (see details in Dubois, 2008d, 2009b, 2011b). Besides, the completeness and reliability of the information published about gene flow, hybrid zones, ploidy, reproductive mode, fixed syna- pomorphies and molecular divergence, is very variable in different clusters of species. Concre- tely, as no author can have studied personally all the terminal taxa of a group, any author who prepares a checklist cannot but depend on this heterogeneous and lacunar information and has to make a compromise between different ergotaxonomies dealing with different subgroups, published by different authors using different ‘‘species concepts’’. Despite this however, often the concrete results in terms of species accepted as valid are quite similar and compatible even if based on different theoretical approaches of the ‘‘species problem’’. These four primary key ranks being given, the nominal-series and ranks of all the other taxa recognized in a given ergotaxonomy should not be offered the same ‘‘freedom’’ and should follow fixed general guidelines or Rules in all taxonomic groups.

Secondary key ranks and subsidiary ranks First of all, for anyone who follows the Code, the number of ranks is strictly limited in the genus- and species-series. In the genus-series, only two ranks are allowed: genus and subgenus. Ranks like supergenus or additional ranks below subgenus, sometimes used by a few recent authors (e.g., Hillis et al., 2001; Hillis &Wilcox, 2005; Vieites et al., 2007), are not Code-compliant and should not be used (Dubois, 2006d, 2007c). In the species-series, four ranks are allowed by the Code: ‘‘aggregate of species’’, species, ‘‘aggregate of subspecies’’ and subspecies. The three-word formulae starting by ‘‘aggregate’’ 96 ALYTES 28 (3¢4) are both unpalatable and unused in zootaxonomy, so we proposed (Dubois &Raffaëlli, 2009) to replace them respectively by the single terms supraspecies (Génermont &Lamotte, 1980) and exerge (Verity, 1925). It should be stressed that the term supraspecies designates a nomenclatural rank (a place in a hierarchy of taxa) and should not be confused with the term superspecies (Mayr, 1931) which designates a taxonomic category meaning a group of parapatric vicariant prospecies (taxa belonging to a taxonomic category of nomenclatural rank species) derived from a single ancestral species (Bernardi, 1980). Under this system arbitrarily limited to the use of six ranks below genus, only two ranks (subgenus and supraspecies) are allowed between genus and species, and two more ranks (exerge and subspecies) below species. However, as we have seen above, the allocation of taxa to the ranks genus and species are rather strongly constrained by ‘‘tradition’’ and ‘‘consensus’’, and cannot be changed easily. This may pose problems if the phylogenetic tree used as a basis for a taxonomy points to the need to recognize more than two ranks between genus and species. In such cases, and pending an improvement of the Code through introduction of taxonomic freedom in the use of ranks, it is impossible to name taxa at these necessary additional ranks in a Code-compliant manner. In such cases, we do not recommend the use of additional Latin ‘‘informal names’’ at these ranks, as was recently done by some (e.g., Deuve, 2004; Guayasamin et al., 2009), because such Latin names will ‘‘look like’’ nomina duly available and valid under the Code and are likely to be confused with them (and sometimes transferred later to other ranks where they could indeed appear as available and valid). We rather suggest the use of a completely distinct system using for example letters (A, B, C...) or codes (A1, A2, B1...) to designate taxa at such additional ranks (such as infragenus, hypogenus, hyperspecies, epispecies, infraspecies, hypospecies, or even hypergenus, supergenus or epigenus above genus). As we will see below, the current taxonomy of the URODELA (appendix 1) is not complex enough to impose the recourse to this poorly satisfying and hopefully transitory solution, except in three cases for epispecies, but this will be necessary in other taxonomic groups, like the ANURA, which are much richer in species and need much more ranks to express unambiguously their hypothesized phylogenetic relationships. In the family-series, the Code allows for an undeterminate number of ranks below the rank superfamily, but goes further in recommending the use of five standard ranks for which it provides standard endings: superfamily (¢OIDEA), family (¢IDAE), subfamily (¢INAE), tribe (¢INI) and subtribe (¢INA). One of these standard ranks is above family and three below. Below family, the use of three ranks only would not be sufficient to express in detail the phylogenetic relationships as currently understood within the URODELA, which require four ranks (appendix 1). As the Code does not allow the use of a rank (‘‘supergenus’’) above genus in the genus-series, a fourth rank must be added below family in the family-series, which is allowed by the Code. For this rank, among the two possibilities of infrafamily and infratribe (see Bour &Dubois, 1985: 83 and Dubois, 2006a: 211), we here chose to use the latter, with the ending ¢ITA proposed by Bour &Dubois (1985: 83). This rank was already used in the URODELA by Dubois &Raffaëlli (2009), and it has the advantage not to disturb the sequence subfamily-tribe-subtribe recommended by the Code. If more ranks had to be used in this nominal-series, the additional ranks proposed by Bour &Dubois (1985: 83) and adopted by Dubois (2006a: 211) could however also be useful, starting with the intermediate rank infrafamily and following with the four lowest ones, and possibly additional still lower ones in Dubois &Raffaëlli 97 case of absolute need in very speciose animal taxa. But we think that in general this multiplication of low family-series taxa should be avoided. This can be done through using rather a general upgrade of the ranks of all taxa, if this is possible without strongly challenging the use of very widely used nomina of families. Above family, the Code only recommends the use of one standard rank, superfamily. It also allows the use of additional subsidiary ranks between family and superfamily, but not above the latter rank: any rank above must then be referred to the class-series, and is therefore outside the ‘‘jurisdiction’’ of the Code. Whether a rank above superfamily belongs in the family- or class-series is, once again, fully arbitrary, and should preferably be regulated by tradition or consensus but, once the border between the two nominal-series has been so fixed, the subsequent ranks in the hierarchy should derive automatically from this decision like in the cases above.

The ergotaxonomy of the URODELA that we adopted here (appendix 1) requires the use of four ranks between family and order. The numbers of taxa needed at these different ranks are, in ascending order, 4, 2, 2 and 2, i.e., a total of 10. Although this can be discussed, we think that 10 CS taxa for this rather homogeneous order is too much, especially if compared with all the previous ergotaxonomies ever used for this taxon ¢ except the now obsolete one of Frost et al. (2006) ¢ and we suggest to limit this number to 6. This results in the need to incorporate the lowest of these ranks, with 4 taxa, in the family-series, and therefore to recognize a second rank below superfamily above the rank family. For this rank, Bour &Dubois (1985: 83) proposed the term epifamily and the standard ending ¢OIDAE, a proposal that has been adopted by various authors since then (e.g., Gaffney &Meylan, 1988; Dubois, 1992; Shaffer et al., 1997; Hooks, 1998; Vences &Glaw, 2001; Eggleton et al., 2007; Beccaloni &Eggleton, 2011) and which is followed here. Unfortunately, a regrettable confusion concerning the use of the rank epifamily was introduced by Dubois (2005c) and repeated in Dubois (2006a): in these works, the rank epifamily was transferred above the rank superfamily, with the ending ¢OIDIA. This proposal was not Code-compliant and should not be followed. For sake of homogeneity over the whole nomenclatural hierarchy, this further entails another change in the ascending hierarchy of prefixes denoting ascending subsidiary ranks of key ranks presented by Dubois (2006a: 208, 222-223): the prefix super¢ should become the second one above the key rank, and epi¢ should be intercalated between the key rank and super¢. But the prefix epi¢ should not be used in the absence of a rank denoted by the rank super¢, and should be so only whenever two or more subsidiary ranks above a key rank are used in an ergotaxonomy.

Miscellaneous nomenclatural problems

Priority The basic Principle of the Code for the establishment of the valid nomen of a taxon in case of conﬂict between two or more nomina is the Principle of Priority (Dubois, accepted). It applies both in cases of synonymy and of homonymy, and in the four nominal-series. Concerning synonymy, implementation of this Principle led us to replace a few invalid nomina by the valid ones. This concerns in particular the nomina of several supraspecies. 98 ALYTES 28 (3¢4)

Taxa at the latter rank, often called species-groups or species-complexes in the literature, correspond to what the Code calls ‘‘aggregates of species’’ (a term never or very seldom used in taxonomic publications). Although this is ignored by some taxonomists, the nomina of taxa of this rank are regulated by the Code and the Principle of Priority applies to them.

Nomenclatural ranks and the simultaneous publication of nomina competing for homonymy or synonymy

Within each nominal-series, all nomina are in permanent potential competition for precedence regarding homonymy and synonymy (Dubois, accepted). Precedence is usually determined by priority of publication, and this is irrespective of their original rank in the work where they were made available. But there is an exception to this situation, stated in Articles 24.1, 55.5, 56.3 and 57.7 of the Code: whenever homonyms or synonyms were introduced simultaneously, but at diﬀerent ranks, the nomen proposed at higher rank takes precedence over the other one(s). This situation is not very common but it applies to three cases in the taxonomy of the URODELA (see appendix 1), discussed in some detail below.

AMPHIUMIDAE and PROTEINA

The first one is the conflict between two family-series nomina AMPHIUMIDAE Gray, 1825 and PROTEINA Gray, 1825, for the valid nomen of the first superfamily of the infraorder which in our ergotaxonomy includes the epifamilies AMPHIUMOIDAE and PROTEOIDAE. The former was introduced by Gray (1825), for a taxon of explicit rank family, and the second for a taxon below the rank family, i.e., implicitly of rank subfamily. Therefore the valid nomen of the superfamily is AMPHIUMOIDEA.

PLETHODONTIDAE,DESMOGNATHINA and ENSATININA

A similar situation concerns the three family-series nomina PLETHODONTIDAE, DESMOGNA- THINA and ENSATININA introduced by Gray (1850), and which are all used as valid for diﬀerent taxa in the same family. Precedence of the former on the other two follows the hierarchy of their original ranks: explicit family for the ﬁrst and implicit subfamily for the second and third. So PLETHODONTIDAE is the valid nomen of the family and PLETHODONTINAE the valid nomen of the subfamily which includes the tribes PLETHODONTINI, DESMOGNATHINI and ENSA- TININI. Precedence among the last two is of no relevance here because the two nomina are not in competition as long as they are used as the nomina of two tribes of the same subfamily.

Subgenera of Eurycea The third case applies to two of the seven subgenera we recognize in the genus Eurycea. These two subgenera are meant at accommodating some of the species of this genus studied by Hillis et al. (2001). These authors distinguished two ‘‘groups’’ of species (i.e., subgenera) within this genus. Their work was concentrated on one of these two groups, in which they recognized 5 genus-series taxa, and for 4 of which they introduced new nomina: the ‘‘section’’ Paedomolge (which, being at the first rank below genus, may be considered to have been Dubois &Raffaëlli 99 introduced as a subgenus), containing two ‘‘clades’’, for which they proposed the new nomina Septentriomolge and Notiomolge (which would correspond to the rank ‘‘infragenus’’ if it was recognized by the Code), the latter including still two more ‘‘clades’’ (corresponding to the rank ‘‘hypogenus’’ if it was recognized by the Code), one for which they coined the nomen Blepsimolge and one for which they used the older nomen Typhlomolge Stejneger, 1896. This nomenclature falls in full under the criticisms exposed in detail by Dubois (2006b,d, 2007c) about another work from the same first author on the anuran genus Rana (Hillis &Wilcox, 2005). Under the Rules of the Code, the valid nomen for their subgenus Paedomolge would be Typhlomolge through simple application of the Principle of Priority, and the other three subordinate nomina would simply be invalid because the Code accepts only one rank (subgenus) below the rank genus. In Amphibian Species of the World (ASW below) [http:// research.amnh.org/herpetology/amphibia/], Frost noted that the use of these three nomina (and also of some of the subdivisions of Rana evoked above) was invalid but added incorrectly: ‘‘and presumably therefore unavailable’’. This statement results from a repeated confusion between the first and third steps of the nomenclatural process. Invalid does not mean unavailable and does no entail unavailability. These three nomina, which were accompanied in the original publication by diagnoses and by explicit designations of type-species, are fully available as subgeneric nomina, by virtue of Article 10.4 which states that a genus-series nomen proposed for a taxon at any rank below genus, ‘‘even if proposed for a secondary (or further) subdivision’’ and ‘‘even if the division is denoted by a term such as section or division’’, is deemed to be a subgeneric nomen. In other words, the four genus-series nomina proposed by Hillis et al. (2001) are nomenclaturally available, being deemed to have been published as subgeneric nomina, but their validity in zoological nomenclature will depend, among other factors, on the rank to which they are allocated: they will be invalid if used at a rank below subgenus, but they may be valid if used at the ranks subgenus or genus. Allocation of these four nomina to taxa does not depend in the least on the intension (‘‘phylogenetic definitions’’) or on the extension (contents) given for them in Hillis et al. (2001) but, as a general Rule in zoological nomenclature, it results automatically, within the frame of a given ergotaxonomic arrangement, from their onomatophores (type-species or nucleospecies). In our ergotaxonomy (appendix 1), we recognize three distinct subgenera for the species included by Hillis et al. (2001) in their ‘‘section’’ Paedomolge: [sG1] with the 7 species latitans, nana, neotenes, pterophila, sosorum, tridentifera and troglodytes; [sG2] with the 3 species chisholmensis, naufragia and tonkawae; and [sG3] with the 3 species rathbuni, robusta and waterlooensis. Now, the nucleospecies, by original designation, of the five nominal genera in competition here are: (NG1) rathbuni for Typhlomolge; (NG2) tonkawae for Paedomolge; (NG3) chisholmensis for Septentriomolge; (NG4) neotenes for Notiomolge; (NG5) nana for Blepsimolge. The taxonomic allocation of the five nomina is therefore straightforward: (NG1) to [sG3]; (NG2) and (NG3) to [sG2]; and (NG4) and (NG5) to [sG1]. In two of these cases we are confronted with a situation where two available nomina were published simultaneously for the same taxon. In both cases the situation of precedence between them is clarified automatically by Article 24.1, i.e., by their relative position in the original hierarchy: Paedomolge is valid and Septentriomolge is its invalid doxisonym; Notio- molge is valid and Blepsimolge its invalid doxisonym. 100 ALYTES 28 (3¢4)

Relative priority between generic and subgeneric nomina in case of current usage Although the Principle of Priority usually applies between all nomina in a given nominal-series, Article 23.9 of the last edition of the Code requires protecting a nomen that has been in ‘‘prevailing usage’’ for a taxon after 1899 against ‘‘resurrection’’ of a senior synonym that had been overlooked until then. As discussed by Crochet (2007), this Rule applies to the generic nomen Speleomantes Dubois, 1984, vs. its senior doxisonym Atylodes Gistel, 1868 that had been resurrected by Wake et al. (2005). If a single taxon (genus or subgenus) is recognized within this group, it should be called Speleomantes,butAtylodes may be used as a valid subgeneric nomen within Speleomantes if the latter taxon is given the rank genus. The same would apply if, following the suggestion of Dubois &Raffaëlli (2009) based on the taxonomic concept of mixogenus, one decided to revert to a situation that has long been in force where all European newts were placed in a single genus. Although we refrained from implementing this proposal in our ergotaxonomy of the SALAMANDRIDAE, and still refrain to do it here, this is not due to a disagreement with the concept of mixogenus, but merely to the fact that the Code currently forbids the use of more than two ranks between genus and species, which would then require to abandon some of the subgenera or supraspecies we recognized in the infratribe MOLGITA, or to treat them as unnamed epispecies (see below). But if this proposal was implemented, despite the fact that the nomen Ichthyosaura Sonnini & Latreille, 1801 is senior to the nomen Triturus Raﬁnesque, 1815, the genus should keep the latter nomen that has clearly been in ‘‘prevailing usage’’ since 1899, and Ichthyosaura could be used only as a subgenus of the latter.

Incorrect authorship Dubois (2008c) had already pointed out a mistake made on several occasions ; which was to credit some publications (and therefore the nomina made available therein) to misleading authorships, as they cited authors’ names that did not appear in the publications at stake, such as ‘‘Fischer von Waldheim’’ instead of ‘‘Fischer’’ or ‘‘Rafinesque’’ instead of ‘‘Rafinesque- Schmaltz’’. The purpose of citing the author’s name after a nomen is not to praise a person but to point to a bibliographic reference, so if the name of the author is cited differently from the way it is printed in the publication, and under which the latter is referenced in bibliographic databases, this citation will fully miss its aim (see Ng, 1994; Dubois, 2008b¢c). A case of incorrect authorship in ASW concerns the genus nomen Siren, which this site persists in crediting, in agreement with the ‘‘tradition’’, to Linnaeus (1767), although it had been shown by Dubois (1991b) to have been introduced one year earlier by Linnaeus’ student Österdam (1766) in his thesis. The latter was published as a small printed book and therefore qualifies as an available publication as defined by the Code. Of course, there exists nothing like ‘‘stability of authorship’’ in zoological nomenclature. This is fortunate, because this would mean that an author who ‘‘stole’’ a published nomen by claiming its authorship would remain credited with this authorship if this ‘‘robbery’’ had remained unnoticed for a while. Therefore, if an authorship has long been credited in error to an author and is shown to be wrong, it must simply be corrected, especially if this has no bearing on the nomen itself, like in this case. However, ASW tried to justify this decision by quoting an anomymous online reference [http://huntbot.andrew.cmu.edu/HIBD/Departments/Library/LinnaeanDiss.shtml] where Dubois &Raffaëlli 101 the following statements appear: ‘‘There were 186 academic dissertations defended by students of Carolus Linnaeus during the period 1743¢1776. (...) Unlike the common practice today in which doctoral students write their own theses and defend them, in 18th-century Sweden the respondents typically expounded and defended the ideas of the praeses. Linnaeus served as praeses for 185 of the 186 dissertations (...). Thus what are usually referred to as the Linnaean dissertations are attributed to Linnaeus, even though they are also associated with the respondents who defended them and whose names appear on their title pages.’’ These statements are interesting from the point of view of the history of science, but fully irrelevant from the point of view of zoological nomenclature. In nomenclature, stating the ‘‘authorship’’ of a nomen, nomenclatural act or publication does not point to a person but to a signature (Dubois, 2008c): once again, its purpose is not to praise a person but to point to a bibliographic reference. Probably a significant proportion of the taxonomic works published so far in zoology were in fact due to other persons than the unethical colleagues who signed them ¢ and these include not only directors of theses who wrote the theses of their students, but also those who signed or co-signed papers written by their students or based on the data obtained by the latter! But this is of no relevance from the point of view of nomenclature, which is based only on facts (e.g., authorships printed in publications) and should not embark into historical or police investigations on ‘‘who did what’’. If a control was to be introduced in this domain, it should be so by scientific institutions or by publishers, not by taxonomists. Therefore, both nomina Siren and Siren lacertina must be credited to Österdam (1766), not to Linnaeus (1767), who clearly mentioned Österdam’s thesis as the original reference of these nomina, so that, contrary to the statement of Frost et al. (2006: 173), there exists no nomen ‘‘Siren Linnaeus, 1767’’ distinct from Siren Österdam, 1766. Another problem of incorrect authorship (and date) occurs when the family-series and the class-series are treated as a single nominal-series, and when the Principle of Coordination is applied to nomina at different ranks some of which only in fact belong in the family-series. For example, Fei et al. (2006: 394¢398) recognized a ‘‘suborder CRYPTOBRANCHOIDEA Fitzinger, 1826’’ including a ‘‘family CRYPTOBRANCHOIDAE Fitzinger, 1826’’, as if these two nomina were coordinate in the family-series. In fact, Fitzinger (1826: 41) coined the rhizonym CRYPTO- BRANCHOIDEA for a taxon explicitly designated as a family including the genus Cryptobranchus, and this nomen therefore belongs in the family-series. In contrast, although it was originally given the rank ‘‘family’’ by Wagler (1828: 859), his nomen CRYPTOBRANCHI was coined independently for a taxon including the single genus Amphiuma but not the genus Crypto- branchus. It is therefore an arhizonym and a class-series nomen, which applies in our ergotaxonomy (appendix 1) to the infraordo PSEUDOSAURIA de Blainville, 1816, of which it is an invalid synonym. Finally, the nomen CRYPTOBRANCHOIDEA coined by Noble (1931: 465) for a suborder including the CRYPTOBRANCHIDAE and HYNOBIIDAE is a distinct class-series nomen which applies in our ergotaxonomy to the infraordo IMPERFECTIBRANCHIA Hogg, 1838, of which it is an invalid synonym. The nomina CRYPTOBRANCHOIDEA Fitzinger, 1826, CRYPTOBRAN- CHI Wagler, 1828 and CRYPTOBRANCHOIDEA Noble, 1931 are three distinct nomina belonging in two different nominal-series, which therefore are not coordinate, have different onomatophores, authorships and dates, and apply to different taxa. All this is incomprehensible in works where no clear distinction, based on objective criteria, is made between family-series and class-series nomina, like the database ASW. 102 ALYTES 28 (3¢4)

Problems of spelling

In preparing the list of taxa of appendix 1, we realized that the spellings traditionally given to some species-series nomina in publications and databases were incorrect for not respecting the Code’s requirement of grammatical agreement between the generic substantive and the ﬁnal epithet. The Code provides Rules for establishing the grammatical gender of genus-series substantives and these must be followed, which sometimes requires to emend the original spelling of the epithet. For example, the stem erpeton, borrowed from the Greek ε ρπετο´ ν, ‘‘reptile, snake’’, is neuter, and all amphibian generic nomina ending with this stem (e.g., Beiyanerpeton, Chunerpeton, Pangerpeton, Piceoerpeton, Regalerpeton, Sinerpeton)are also neuter, which required to modify the endings of some of the epithets associated with these nomina. Its stem is erpet¢, so family-series nomina based on such nomina must be based on this ending, not on erpetont¢ (e.g., NOTERPETIDAE, not NOTERPETONTIDAE,orSCAPHERPETI- DAE, not SCAPHERPETONTIDAE). On the other hand, the Code is quite clear in stating that in most cases the original spelling of a nomen must be kept unchanged, even in case of ‘‘incorrect transliteration or latinization, or use of an inappropriate connecting vowel’’ (Article 32.5.1). Thus for example, the endings of epithets based on names of persons should not be modiﬁed even if they seem incorrect because of the gender or number of the dedicatees, and changes introduced eventually in this respect by subsequent emendations are incorrect emendations and should not be used (Dubois, 2007b;Nemésio &Dubois, 2012).

Unusual taxonomic categories

As we have seen above, taxonomic categories and nomenclatural ranks are fully distinct concepts and tools. Nomenclatural ranks, if used consistently, can provide precious information on the structure of an ergotaxonomy, particularly within the frame of a phylogenetic taxonomic paradigm, but they can provide no biological or historical information on the organisms and the taxa at stake. In contrast, some information of this kind can be provided by the taxonomic categories, at least for the taxa of low ranks, mostly from genus to subspecies. A very well-known taxonomic category is that of superspecies, a Latin term proposed by Mayr (1931) to replace the German term Artenkreis of Rensch (1928), to designate a homophyletic group of vicariant species. Birula’s (1910) term prospecies is the first one available to designate the species that compose a superspecies. Bernardi (1980) reviewed the main taxonomic categories that had then been distinguished by evolutionary taxonomists. Since then, others have been defined, often in the case of particular evolutionary phenomena involving interspecific hybridization, such as the various categories of kleptons and klonons (Dubois &Günther, 1982; Dubois, 1991a, 2008d, 2009b, 2011b) or the category of mixogenus briefly tackled above (Dubois, 1981, 1988, 2004b;Dubois &Raffaëlli, 2009). These questions are too complex for being explained in detail here, so we refer to these works for a proper introduction to these peculiar biological systems. To distinguish the biological information provided by taxonomic categories from the nomenclatural information provided by ranks, some devices can be implemented, such as the interpolation of some symbols within the Latin nomina of the taxa. This practice is allowed Dubois &Raffaëlli 103 by the Code at least in two cases, for the naming of ‘‘aggregates of species’’ and ‘‘aggregates of subspecies’’ (see Article 6.2, its Recommendation and its Example). Generalization of its use was proposed at the nomenclatural rank species by Dubois &Günther (1982) for the taxonomic category of klepton, and the use of the symbol ‘‘kl.’’ for klepton has been adopted since then by various authors working on the European ranid green frogs of the genus Pelophylax. Kleptons are just taxa of a particular kind at the nomenclatural rank species, they do not represent a distinct nomenclatural rank. Such a system has been identified in the North American genus Ambystoma. It deserves a special taxonomic treatment, which will be discussed below. Dubois (1991a, 2008d, 2009b, 2011b) proposed other symbols for other species-rank categories. Such a mode of notation for unusual taxonomic categories can also be used at higher nomenclatural levels. Dubois &Günther (1982) proposed to use the symbol ‘‘synkl.’’, for synklepton, to designate a group of species including a klepton and the ‘‘biological species’’ or mayrons from which the klepton originated through hybridization, as well as other mayrons or kleptons with which they interfere genetically in some population systems. At even higher level, the category of mixogenus (see Dubois &Raffaëlli, 2009) which only applies in the rather rare cases of documented hybridization between species, could be referred to by the use of the symbol ‘‘mxg.’’ placed before the generic nomen. Such taxonomic categories may coincide only partially with nomenclatural ranks. For example, a given synklepton may be included within a genus or subgenus, or may cover the whole extension of a genus or subgenus. It may also straddle several, but not all, subgenera of a genus. A few taxa belonging to unusual taxonomic categories have been identified so far in the URODELA, but probably more remain to find. For example, the category superspecies was used to designate a group of species of the genus Triturus (the ‘‘Triturus cristatus superspecies’’). Following Article 6.2 of the Code, the prospecies that are the members of this superspecies could be designated by nomina of the form Triturus (supersp. cristatus) cristatus or even Triturus (supersp. cristatus) prosp. cristatus to expand Dubois &Günther’s (1982) proposal. In this system, the taxonomic category superspecies (for the use of which the Code is irrelevant) is just one possible category referred to the nomenclatural rank supraspecies (‘‘aggregate of species’’) recognized by the Code.

Proposed taxonomic and nomenclatural changes in the order URODELA

Implementation of the concepts, criteria and Rules discussed above leads us to propose some taxonomic and nomenclatural changes in the order URODELA. Some of these changes concern the class-series taxonomy and nomenclature of the order, whereas others are internal to some of its 10 recent families. We present these two kinds of changes successively.

Valid class-series nomina in the URODELA

As we have seen, in the class-series just like in the other nominal-series, the basic criterion that determines the valid nomen of a taxon is the Principle of Priority of Publication, 104 ALYTES 28 (3¢4) completed in case of simultaneous publication by the Principle of First-Reviser. In some special cases however, it is possible to ignore these Principles, in order to ‘‘protect’’ or ‘‘conserve’’ some nomina, the sozonyms, which have had a universal or almost universal use in zootaxonomy since 1900 for a given taxon, while no other nomen was used signiﬁcantly for the same taxon. If, for a given taxon, no nomen corresponds to this situation, all the available nomina that are synonyms are sozodiaphonyms and distagmonyms, and the ‘‘normal’’ Principle of Priority and First-Reviser will simply apply, under the revised conditions described above (ﬁrst among sozodiaphonyms, second among distagmonyms).

The situation in the URODELA is quite clear: no CS nomen available for taxa of this order qualiﬁes as a sozonym; two qualify as sozodiaphonyms; all others are distagmonyms. As these concepts are not yet in widespread use, we will illustrate them by discussing separately and in detail each of the ﬁve class-series nomina needed by our ergotaxonomy (appendix 1), starting by the least inclusive one and ending by the most inclusive.

Subordo MEANTES Linnaeus, 1767 The holophyly of this taxon was recently conﬁrmed both by Zhang &Wake (2009) and Pyron &Wiens (2011), although with diﬀerent relationships with the other salamanders. In our ergotaxonomy, it is a suborder that includes the single family SIRENIDAE with two recent and three fossil genera.

The nomen MEANTES was proposed by Linnaeus (1767) for a new order, the fourth one of his class AMPHIBIA, to accommodate the single genus Siren described the preceding year by his student Österdam (1766), which is therefore the nucleogenus of this nominal order. This was the ﬁrst class-series nomen published for a higher salamander taxon after 1757. All other available CS nomina all conucleogenera of which belong in the SIRENIDAE, to the exclusion of all other salamander families, also apply to this taxon. This CS taxon was intermittently recognized as valid during the history of the classiﬁ- cation of the salamanders, being sometimes merged with other families, and sometimes used for a fourth order of recent amphibians besides the salamanders, the frogs and the caecilians. Several nomina have been used for this taxon in the past, including MEANTES, TRACHYSTOMATA, SIRENOIDEA, SIRENOIDEI and others. None of these nomina meets the requirements of Rule [R12] of Dubois (2006a) for sozonymy or sozodiaphonymy, so the valid nomen of this CS taxon simply results from Priority of Publication among all the distagmonyms that apply to the taxon.

Infraordo IMPERFECTIBRANCHIA Hogg, 1838 The holophyly of this taxon was recently conﬁrmed both by Zhang &Wake (2009) and Pyron &Wiens (2011), although with diﬀerent relationships with the other salamanders. In our ergotaxonomy, it is an infraorder that includes the two families CRYPTOBRANCHIDAE and HYNOBIIDAE.

Several CS nomina are also available for this taxon. The ﬁrst published one is IMPERFEC- TIBRANCHIA Hogg, 1838, created for an order including the single nominal genus Menopoma Harlan, 1825 (a doxisonym of Cryptobranchus Leuckart, 1821), which is therefore its nucleogenus. Dubois &Raffaëlli 105

This CS taxon was rarely recognized as valid during the history of the classiﬁcation of the salamanders, sometimes under the rhizonyms CRYPTOBRANCHOIDEA or CRYPTOBRANCHOIDEI. These nomina do not meet the requirements of Rule [R12] of Dubois (2006a) for sozonymy or sozodiaphonymy, so the valid nomen of this CS taxon simply results from Priority of Publication among all the distagmonyms that apply to the taxon.

Infraordo PSEUDOSAURIA de Blainville, 1816

The holophyly of this taxon was recently confirmed both by Zhang &Wake (2009) and Pyron &Wiens (2011), although with different internal relationships. In our ergotaxonomy, it is an infraorder that includes the seven families AMBYSTOMATIDAE, AMPHIUMIDAE, DICAMPTO- DONTIDAE, PLETHODONTIDAE, PROTEIDAE, RHYACOTRITONIDAE and SALAMANDRIDAE. Many CS nomina are available for this taxon, none of which qualifies as a sozonym or sozodiaphonym, so the ‘‘normal’’ Rules for precedence should be applied to establish its valid nomen. The first ones ever published are two allelonyms proposed by de Blainville (1816) as possible alternatives to designate an order: PSEUDO SAURIENS and SALAMANDRES. The former is an arhizonym whereas the latter is a rhizonym based on the generic nomen Salamandra, which was the only genus included in the order and therefore the nucleogenus of both nomina. Although published as French scientific nomina, both are nomenclaturally available for having been latinized and used as valid by subsequent authors, first respectively as PSEUDO- SAURII by Gray (1825: 215) and as SALAMANDRINA by Müller (1831: 711), and then under other spellings. This taxon composed of these seven families was rarely recognized in the past. For example, Zhang &Wake (2009) used the nomen ‘‘Salamandroidea’’ for this taxon, but without precising its nominal-series, i.e., whether it designates a superfamily (which would then be SALAMANDROIDEA Goldfuss, 1820) or a suborder (which would then be SALAMANDROI- DEA de Blainville, 1816). If considered a CS nomen, the latter would be very far from meeting the criteria of Rule [R12] of Dubois (2006a) for sozonymy or sozodiaphonymy. Among the two allelonyms that compete for being the valid CS nomen of this taxon, the precedence was established by the IFRA of de Blainville (1835: 280), who only mentioned its aponym PSEUDO-SAURIA as valid for this taxon. This is fortunate, because using for the latter the rhizonym based on the generic nomen Salamandra would be highly confusing, as (1) it could be mistaken for a FS nomen, and (2) this nomen has been used in the past, under various spellings, for different CS taxa having widely different contents (from the sole family SALA- MANDRIDAE up to the whole order of salamanders) and would have no clear meaning for users, even for those who pay close attention to nomenclatural matters. The valid nomen of this taxon according to the guidelines above has been little used in the literature, and no spelling was used universally by taxonomists after its latinization. We think that the spelling PSEUDOSAURIA should be retained for this nomen, by parallelism with three of the other CS nomina of salamanders, which end with an ¢A. This spelling was first used by Cope (1889: 30). 106 ALYTES 28 (3¢4)

Subordo ICHTHYOIDEA Leuckart, 1821 The holophyly of this taxon was recently supported by Zhang &Wake (2009) but rejected by Pyron &Wiens (2011). We here follow the former authors for the reasons given aboveintheIntroduction. In our ergotaxonomy, this taxon is a suborder that includes the two families of IMPERFECTIBRANCHIA and the seven families of PSEUDOSAURIA listed above. Several nomina are available for this taxon. None of them meets the criteria for sozonym or sozodiaphonym validation mentioned, so the ‘‘normal’’ Rules of precedence must be used for establishing the valid nomen of this taxon. The first published nomen that applies to this taxon is ICHTHYOIDEA Leuckart, 1821, introduced for a taxon of unstated rank but clearly belonging in the class-series, and including the two genera Cryptobranchus Leuckart, 1821 (CRYPTOBRANCHIDAE) and Phanerobranchus Leuckart, 1821, a junior doxisonym of Necturus Rafinesque, 1819 (PROTEIDAE). It should become the valid nomen of this taxon. In the past, various authors intermittently recognized a higher taxon including all salamanders except the SIRENIDAE, but under various nomina, none of which had a regular or massive use. Priority is doubtless the best way to fix the valid nomen of this taxon.

Ordo URODELA Duméril, 1805

This taxon which includes all the families listed above as well as poorly known all-fossil families and genera of salamanders (see appendix 1) was recently conﬁrmed both by Zhang &Wake (2009) and Pyron &Wiens (2011) to be holophyletic and is here recognized as an order within the superorder BATRACHIA Brongniart, 1800 of the class AMPHIBIA de Blainville, 1816. Its valid nomen has long been the matter of a still unsettled dispute, so we will devote a detailed discussion to this question. Six diﬀerent reasons support considering the nomen URODELA Duméril, 1805 as the valid nomen of this order, and the nomen CAUDATA as invalid.

(1) As we have seen above, the nomen MEANTES was the ﬁrst one proposed for a taxon including only salamanders, but the least inclusive CS taxon that currently includes its nucleogenus covers only a part of the salamanders.

The nomen PNEUMOBRANCHIENS Sonnini & Latreille, 1801 was the next one coined for a class-series taxon including only salamanders. Its three conucleogenera are Proteus Laurenti, 1768 (PROTEIDAE), Ichthyosaura Sonnini & Latreille, 1801 (SALAMANDRIDAE) and Siren Öster- dam, 1766 (SIRENIDAE). In our ergotaxonomy, the least inclusive taxon that includes these three genera is the order itself. This nomen is nomenclaturally available for having been latinized by Hunter (1834: 145) as PNEUMOBRANCHIATA. However, it was not used as valid after the works of Hogg (1839a¢b) and it is therefore a distagmonym that cannot be kept for the taxon as sozodiaphonyms are available for it.

The second nomen that applies to this taxon, ‘‘BATRACIENS PISCIFORMES’’ Latreille, 1804, created for a ‘‘family’’ including the genera Proteus and Siren, is an anoplonym (unavailable nomen) for being a binomen, which furthermore has never been latinized before 1900. The next available nomina are the two allelonyms introduced for the ‘‘family’’ of salamanders by Duméril (1805). As explained by Dubois (2004a), this author was the first Dubois &Raffaëlli 107 one who distinguished the two taxa of frogs and salamanders (removed from the lizards), that together compose nowadays the higher taxon BATRACHIA Brongniart, 1800. For naming these two ‘‘families’’, Duméril (1805) used two pairs of allelonyms, respectively ANOURES and ECAUDATI for the frogs and URODÈLES and CAUDATI for the salamanders. Some recent authors (e.g., Frost et al., 2006, implicitly followed by Blackburn & Wake, 2011) considered the ‘‘familial’’ nomina in Duméril (1805) as unavailable FS nomina, and credited these nomina in the class-series to ‘‘Fischer von Waldheim, 1813’’ (in fact Fischer, 1813; see Dubois, 2008c). This decision was particularly ill-advised, because Fis- cher (1813) had merely copied the taxonomy of the BATRACII of Duméril (1805: 90¢96), without any change even in details except that he called them BATRACHII. This is clearly highlighted by his mention of the authorship ‘‘D.’’ (for Duméril) after several nomina in his table III (pages 58¢59). The statement of Frost et al. (2006: 356¢357) that the nomina CAUDATI, URODELI, ECAUDATI and ANURI were used by him for orders is simply not true: in his table III, Fischer (1813) did not state the rank of his taxa! Above in the same text, Fischer (1813: vii) announced that this table provided a ‘‘Tabula synoptica Ordinum’’, but no ranks were given for the taxa in the table itself. In this work, as well as in his previous book (Fischer, 1808), this author largely followed the nomenclatural hierarchy of Linnaeus (1758). As he afforded the rank classis to the taxon REPTILIA, the most reasonable interpretation of his hierarchy is to consider that he applied the rank ordo, like in Linnaeus (1758), to the two taxa immediately subordinate to it, i.e., PEDATA and OPHIDII. His taxon BATRACHII, just below PEDATA, and the taxa URODELI and ANURI, just below BATRACHII, are of unclear rank but certainly not of rank order: as this taxonomy was directly copied from Duméril (1805), the most logical treatment is to consider URODELI and ANURI as ‘‘families’’. The first author who used these nomina, under the aponyms ANURIA and URODELIA, for taxa explicitly above the rank family, was Rafinesque (1815: 78), who considered them as suborders (see Dubois, 2009a), but it would also be unwarranted to credit these nomina to him as he had also directly borrowed them to Duméril (1805), like many other nomina in his book. In fact, all these ‘‘interpretations’’ (which apply also to other nomina like AMPHIBIA, BATRACHIA or GYMNO- PHIONA; see Dubois, 2004a, 2009a) just tend to ‘‘steal’’ these nomina, in the sense that they now have in zoological nomenclature, from their real authors, for reasons that would be interesting to understand.

For the two ‘‘families’’ of his BATRACII,Duméril (1805) used two alternative pairs of allelonyms: ANOURES and ECAUDATI for the frogs (genera Bufo, Hyla, Pipa and Rana), and URODÈLES and CAUDATI for the salamanders (genera Protoeus, Salamandra, Siren and Triton). Both pairs include nomina which are junior homonyms of two nomina previously proposed by Scopoli (1777) for two orders of his REPTILIA: CAUDATA for the genera Draco, Lacerta, Siren and Testudo (i.e., salamanders, lizards and turtles) and ECAUDATA for the single genus Rana (frogs). The nomen ECAUDATA Scopoli, 1777 applies nowadays to the least inclusive CS ergotaxon including its nucleogenus, which is less inclusive than the CS ergotaxon including all the conucleogenera of ECAUDATI of Duméril (1805), so the latter is an invalid junior homonym of the former. The same is true for CAUDATI Duméril, 1805, which applies to the whole order of salamanders, whereas CAUDATA Scopoli, 1777, which has a diﬀerent, only partially overlapping, onomatophore, applies to a higher taxon of vertebrates referred to nowadays by some authors under the nomen NEOTETRAPODA (see e.g. Dubois, 2006a: 192), and not to a taxon composed exclusively of amphibians. Therefore, both nomina CAUDATI and 108 ALYTES 28 (3¢4)

ECAUDATI are invalid junior homonyms, which is not the case of ANOURES and URODÈLES,so there should have been no problem and discussion for the adoption of the two latter nomina as valid, as they were both latinized in the early 19th century (see Dubois 2009a) and massively used since then, with frequent mention of Duméril as their author. However, the subsequent fate of these nomina was very strange indeed, and in fact fully illogical. The nomen ECAUDATI Duméril, 1805, emended in ECAUDATA, was used for the order of frogs by various authors in the 19th century, particularly in the title of the famous catalogue of Boulenger (1882a), quoted in thousands of publications. However, it was replaced in the th 20 century by its allelonym ANURA Duméril, 1805 (apograph ﬁrst used by Hogg, 1839a: 270). The latter nomen, that has been used thousands of times until now in titles of publications of all kinds, whereas no other nomen was used largely for the same taxon, fully qualiﬁes as a sozonym and must be validated (see Dubois, 2004a, 2006a, 2009a).

For the order of salamanders, the nomen CAUDATI Duméril, 1805 cannot be used as it is a junior invalid nomenclatural homonym of CAUDATA Scopoli, 1777, a clearly available CS nomen. Consequently, the nomen URODELA Duméril, 1805 (apograph first used by Gray, 1825) should soon have been considered the valid nomen for this order. But the history of amphibian taxonomy made this simple situation unnecessarily confusing: until now, both nomina URODELA and CAUDATA have remained in use for the order in the literature, being used ‘‘in parallel’’ by different authors. (2) Although no official Rule for the nomenclature of class-series taxa exists in the Code, the Principle of Homonymy is a very old and common one, not only in biological nomenclatures but also in all domains of human language. Human communication would be very hampered, if not difficult, if different concepts or objects could be freely given the same name. In common language and especially in specialized technical terminologies, homonymy (which is often defined in a less strict way than in biological nomenclature; see Dubois, 2012) may be tolerated when it exists between usages of the same name or of similar names that occur in widely distant domains of activity (e.g., syzygy in astronomy, in the biology of gregarines, or in the study of meiosis) but it is not so when both homonyms belong in the same domain. In a scientific field like taxonomy and nomenclature, it is not acceptable that two different taxa bear the same nomen ¢ except in very specific and duly regulated cases when this concerns taxa in immediate hierarchical relationships in the same nominal-series, their nomina being submitted to the Principle of Coordination (see Dubois, 2012). (3) The fact that the earliest nomen is no more in use is of no relevance in this respect. First of all, this is liable to change at any time, if new ergotaxonomies are proposed, or simply as a result of a nomenclatural update like the one presented here for salamanders. And second and mostly, because if there were no Principle of Homonymy there would be no reason to impede authors from ‘‘reviving’’ or ‘‘recycling’’ some old nomina that are no more in use for some good or bad reason in zoological nomenclature. Thus, nomina like APODA, ECAUDATA, GRADIENTIA, INVERTEBRATA, NANTES, OSSEA, PISCES, QUADRUPEDIA, SALIENTIA or VERMES, which are currently considered invalid in zoological nomenclature, could be ‘‘resurrected’’ nowadays to designate taxa widely different, or even only slightly different, from those for which they had first been introduced. Refusing to use a Principle of Homonymy in class-series zoological nomenclature would be as inappropriate as it would be in the other nominal-series and it would open the way to increased chaos in a field that is already much too chaotic. Therefore, Dubois &Raffaëlli 109

once validly introduced in zoological nomenclature as a hoplonym, a class-series nomen should preoccupy forever its original spelling (its protograph) as well as all derived spellings based on the same stem (its apographs). (4) However, according to the MONS Rules mentioned above, a class-series nomen that is a junior homonym of another one can in some cases be validated. For this, two conditions must be met: (a) the senior homonym must be a distagmonym; (b) the junior homonym must be a sozonym. In the present case, only the first condition applies. The nomen CAUDATA is far from having been universally or quasi-universally used as valid for the order of salamanders in the literature since 1900. It was used alternatively to the nomen URODELA, and both nomina can easily be documented as appearing in more than 100 titles of non-specialized publications from many countries since that date. This is easy to check by browsing a few fascicles of the Zoological Record (ZR) or reference lists of papers or books dealing with the taxonomy of salamanders. Table 1 provides counts of usages of both nomina in the titles of the publications listed in the ZR every ten years from 1901 to 2011 (i.e., after the starting date fixed by Article 23.9 of the Code for the definition of the terms ‘‘nomen protectum’’ and ‘‘nomen oblitum’’). For most of the period covered (1901¢1971, 1991), the term URODELA had a wider use than CAUDATA, and the reverse occurred only three times (1981, 2001, 2011), but as stated above, the absolute numbers of usages are not relevant here. What is important is that no consensus ever appeared over the international community of zoologists for the use of either nomen. Therefore, none of these two nomina is a sozonym, both are sozodiaphonyms and precedence among them cannot be settled through sozonym validation.

(5) Both nomina URODELA and CAUDATA were made available in the same publication (Duméril, 1805) and were therefore published simultaneously. Irrespective of the fact that one is an invalid junior homonym, precedence among them was fixed by the choice made by the first-reviser2. As we have seen above, there are two kinds of first-reviser actions in nomenclature, IFRA and EFRA, and we suggested that both could be accepted as valid in the class-series to determine precedence between simultaneous alternative new nomina or spellings for new nomina. So, who was the first-reviser in this case? A thorough survey of the literature (Dubois, unpublished) shows that this action was quite late after the work of Duméril (1805). This is due to several distinct causes. One is the fact that some authors (e.g.,

2. The same would also apply between URODELI and CAUDATI in Fischer (1813) if he was the author of these nomina, although this author showed apparently a preference for the former by printing it in small capitals instead of lower case italics for the latter. The decision of Frost et al. (2006) to aﬀord priority among them to the latter was fully arbitrary: our table 1 shows that the criterion of ‘‘usage’’ which they referred to is not supported by the facts. 110 ALYTES 28 (3¢4)

Oppel, 1811a¢b;Giebel, 1846) unambiguously referred this taxon to the family-series and used either CAUDATA or URODELA as family nomina (being therefore now considered anoplonyms in this nominal-series), which has no bearing on their relative precedence in the class-series. The second fact is that few authors of that time followed Duméril (1805) in placing all salamanders in a single higher taxon, because the old taxonomy ‘‘resisted’’ this change for decades. Fischer (1813: 58) mentioned both nomina (as CAUDATI and URODELI), without choosing between them, as allelonyms for a taxon of unknown rank (but presumably of the class-series), with the exactly same content as in Duméril (1805). But then, for a long time most subsequent authors recognized two or even more orders for the salamanders. When they used the nomina CAUDATA or URODELA (or their paronyms), it was for an order including Salamandra and Triturus (or its isonyms) but excluding Proteus and Siren. Thus for example the order nomen URODELA in Gray (1825: 215), based on the genera Salamandra and Triton alone, did not include all the conucleogenera of URODELA Duméril, 1805: it is therefore a distinct nomen and an invalid junior homonym of the latter. Some subsequent authors included additional genera in the URODELA (e.g., Cryptobranchus in Ficinus &Carus, 1826, or Salamandrina in Bell, 1836), but as they kept Proteus and Siren outside this taxon they in fact used Gray’s (1825) and not Duméril’s (1805) URODELA. It was only in the years 1830s that some authors again grouped all salamanders in a single taxon sister to the frogs, and this practice gained progressively in popularity until the end of the 19th century. But even then, no first-reviser action was taken for a long time: (a) some authors (e.g., Duméril &Bibron, 1841) still mentioned both nomina as allelonyms without choosing between them; (b) others (e.g., Wiegmann &Rüthe, 1832: 202; Gravenhorst, 1843: 393; Boulenger, 1882b: vii, 1) used the nomen CAUDATA for this taxon, but, as they did not mention the nomen URODELA, they did not act as first-revisers; (c) others (e.g., Giebel, 1847: 184; Cope, 1889: 13) did the opposite choice, using URODELA as valid but then without mentioning CAUDATA. The first-reviser action was taken only at the end of the 19th century, but very clearly, by Zittel (1888: 412), who placed all recent and fossil salamanders in the same order, for which he used the nomen URODELA as valid and mentioned the nomina CAUDATA and ‘‘BATRACHIA GRADIENTIA’’ as invalid synonyms. First-reviser actions in zoological nomenclature are irrever- sible acts that cannot be challenged later by other authors (otherwise the concept of FRA would be completely useless). So, even if CAUDATA Duméril, 1805 was not an invalid junior homonym, it would be an invalid junior synonym of URODÈLES Duméril, 1805 following this first-reviser action. (6) Finally, mention should be made here of the Rule [R14] of Nomenclatural Consis- tency proposed by Dubois (2006a). This Rule was devised in order to deal with nomenclatural validity in the case of the existence of several distinct pairs or sets of sister-nomina or didymonyms, i.e., nomina coined together to designate the taxa resulting from the process of splitting an existing class-series taxon, in particular in the case where the nomina of the first published pair or set were later replaced by a new pair or set of nomina (e.g., as ‘‘nomina nova’’ or neonyms of the former ones). This proposed Rule stated that in such cases all members of a pair or set should be validated or invalidated together, and that no mixture between distinct sets should be allowed. However, implementation of this Rule at the level of the whole zoology would pose genuine problems in a few cases, and it seems better to abandon the idea of this being a Rule, but it should at least remain a strong Recommendation. Under such a Recommendation, either ANURA and URODELA,orECAUDATA and CAUDATA, should be Dubois &Raffaëlli 111 validated together, but no mixture between the two pairs of nomina (phobonyms;Dubois, 2006a) would be appropriate. As ECAUDATA has long ago been rejected as invalid, it is now a distagmonym which should not be validated against the sozonym ANURA Duméril, 1805, and its sister-nomen CAUDATA should not be validated either. Although this statement should only be given the value of a Recommendation, not of a Rule, it is striking to note that it points to the same conclusion as the five other points above. This conclusion is quite straightforward: the valid nomen of the order of tailed amphibians or salamanders is URODELA Duméril, 1805. As shown above, persistence in rejecting this conclusion would not be supported by any consistent rationale or set of Rules, nor by ‘‘usage’’, and could only be explained by a refusal to follow any set of universal a priori Rules in higher nomenclature, possibly because of a preference for ‘‘informal nomenclatural systems’’, in which every researcher is left free to follow only his/her personal opinions, tastes or habits, or groups of zoologists to act as ‘‘lobbies’’ to impose ‘‘their’’ nomenclature, a practice which has nothing to do with science. Although such a clear rejection of constraints carried by Rules was expressed in full words only rarely (e.g., Hillis, 2006), it may be a good explanation for the obstinate attitude of some zoologists to continue using the nomen CAUDATA Duméril, 1805 for this taxon, although it is a junior homonym, was rejected by a first-reviser action, and above all has never had a ‘‘universal usage’’ in zoology, and therefore does not in the least qualify as a ‘‘nomen protectum’’ or sozonym.

Taxonominal changes in the URODELA below the class-series

We present these changes separately for each family, by alphabetical order of families.

Familia Ambystomatidae Gray, 1850 Although more than 30 recent species are currently known in this family, they have long all been referred to a single genus Ambystoma. This is fully justified by the holophyly of the group (Zhang &Wake, 2009; Pyron &Wiens, 2011) and in light of the mixogenus concept, as many hybrids are known between various species of this group. However, several well identified subgroups can be identified morphologically and phylogenetically within this genus (Tihen, 1958; Jones et al., 1993; Shaffer &McKnight, 1996; Highton, 2000). We here recognize them taxonomically, as four distinct subgenera, one of which includes three supraspecies. Tables A2 and A3 of appendix 2 provide the nomina and short diagnoses for these taxa. It should be stressed that several nomina, senior to Ambystoma Tschudi, 1838, which would have been available for some of these subgenera, were suppressed by the ICZN (Riley &China, 1963). On the basis of the original description (Sager, 1858), we consider the nomen Des- miostoma maculatum as a synonym of Ambystoma mavortium, as suggested by Dunn (1940), and therefore Desmiostoma as a synonym of Heterotriton. In the subgenus Heterotriton, according to Shaffer &McKnight (1996), both species Ambystoma mavortium and A. tigrinum include several divergent genetic groups which should be recognized as species, as it was already done for several Mexican taxa of this subgenus. 112 ALYTES 28 (3¢4)

As we have seen, in North America a complex system of populations involving five distinct species and various ‘‘forms’’ of hybrid origin has been described in detail in the last decades. In this system, various kinds of hybrids with several genome compositions and up to five ploidy levels have been identified. They have unusual meioses (metameioses) and are liable to reproduce either by gynogenesis or by zygogenesis (fertilization) according to the local conditions (Bogart et al., 2007; Bi et al., 2009). To account taxonomically for the peculiar evolutionary situation of this complex, the concept of tychoklepton was proposed (Dubois, 2008d, 2009b, 2011b). This is a taxonomic category of nomenclatural rank species. The situation in these salamanders is different from that in European green frogs of the genus Pelophylax,wheredifferent taxa of rank species and of category zygoklepton, which have different and largely parapatric (non-overlapping) distributions, are taxonomically recognized (Dubois, 1998). In contrast, in the genus Ambystoma, all the ‘‘forms’’ regularly interact reproductively and genetically, so that there would be no point in recognizing taxonomically adifferent taxon of species rank for each genome composition and ploidy level; but on the other hand they must be distinguished taxonomically from their ‘‘parental’’ species. The latter are evolutionary independent and do not need these ‘‘forms’’ for their perpetuation, but the reverse is not true. Following Dubois (2008d), we here recognize a single tychoklepton for all the ‘‘forms’’ of this North American complex of Ambystoma. Two nomina have been proposed in the past for members of this taxon: Ambystoma platineum (Cope, 1868) and Ambystoma tremblayi Comeau, 1943. Following the Principle of Priority, we retain the nomen Ambystoma (Xiphonura) kl. platineum for this taxon. The five species that have been involved in hybridization events at the origin of this taxon belong in three distinct subgenera of Ambystoma as recognized here: Linguaelapsus (barbouri and texanum), Heterotriton (tigrinum) and Xiphonura (jeffersonianum and laterale). This is not contradictory with the concept of mixogenus, as the criterion of crossability applies within genera, not within subgenera. As all the members of this tychoklepton possess at least one genome inherited from the species Ambystoma laterale, which suggests that possibly this genome carries a factor that might be responsible for the special meioses of these organisms, we here refer the klepton Ambystoma kl. platineum to the subgenus Xiphonura. This is a rather arbitrary solution however. Another possibility could be to erect for this taxon a fifth subgenus in the genus Ambystoma.

Familia Cryptobranchidae Fitzinger, 1826 We hereby reinstate Cryptobranchus bishopi Grobman, 1943 as a valid species according to Crowhurst et al. (2011). In fact this species consists in two distinct genetically diﬀerenti- ated lineages which should be recognized as distinct taxa, one of which has still not been named.

Familia Dicamptodontidae Tihen, 1958 The generic nomen Bargmannia Herre, 1955 is invalid for being a junior homonym of Bargmannia Totton, 1954 (SIPHONOPHORA), and we therefore provide the neonym (‘‘nomen novum’’) Sanchizia nov. for this fossil genus. This taxon is dedicated to Borja Sanchíz (Madrid), who provided a major contribution to the knowledge of fossil recent amphibians. Dubois &Raffaëlli 113

Familia Hynobiidae Cope, 1859 (1856) The double date of this nomen (ignored by most authors) reﬂects the fact that the ﬁrst nomen available for this taxon was ELLIPSOGLOSSIDAE Hallowell, 1856. The generic nomen Ellipsoglossa Duméril, Bibron & Duméril, 1854 being a junior doxisonym of Hynobius Tschudi, 1838, the family and its coordinate taxa must bear the nomen HYNOBIIDAE Cope, 1859 (1856) by virtue of Article 40.2 of the Code (Dubois, 1984: 114). Our ergotaxonomy of this family is derived from the very comprehensive phylogenetic analysis of this taxon by Poyarkov (2010)3 based both on mitochondrial and nuclear DNA, as well as on morphology.

These data support recognition of two recent subfamilies in this family, HYNOBIINAE Cope, 1859 (1856) and ONYCHODACTYLINAE nov. The diagnoses of these subfamilies are provided in table A1 of appendix 2.

In the ﬁrst one, HYNOBIINAE, we distinguish three tribes, HYNOBIINI Cope, 1859 (1856), PACHYHYNOBIINI nov. and RANODONTINI nov. Their diagnoses are given in table A2 of appendix 2.

The ﬁrst tribe HYNOBIINI includes three subtribes, HYNOBIINA Cope, 1859 (1856), PROTO- HYNOBIINA Fei & Ye, 2000 and SALAMANDRELLINA nov. Their diagnoses are provided in table A2 of appendix 2.

In the subtribe HYNOBIINA, we hereby subdivide the genus Hynobius in four subgenera, including a new one Poyarius nov. (etymology: taxon dedicated to Nikolaï Poyarkov; grammatical gender masculine, like Hynobius) and two others that had diﬀerent statutes in the past (genus Pachypalaminus, genus or subgenus Satobius). Their diagnoses are given in table A3 of appendix 2. In the hyponymous subgenus Hynobius, we recognize six supraspecies, diagnosed in table A5 of appendix 2. One of them, long known under the nomen nigrescens but for which the nomen lichenatus has priority, accommodates 5 species, to which, according to Poyarkov (2010) a sixth one should be added (‘‘Hynobius sp. 8’’), which corresponds to the former Honshu population traditionally referred to Hynobius nebulosus but for which the nomen Hynobius peropus Boulenger, 1882 might be available. If the latter hypothesis is right, the supraspecies should be renamed peropus. In the supraspecies nebulosus, several new species from South Korea remain to be described and named.

In the subtribe PROTOHYNOBIINA, we recognize four genera, including a new one Tsinpa nov. (etymology: English transcription of the name of the locality where the species was originally reported from; grammatical gender feminine, like Liua). Their diagnoses are given in table A3 of appendix 2. In the genus Batrachuperus, we recognize two subgenera, Batrachuperus Boulenger, 1878 and Tibetuperus nov., which correspond only in part with the ‘‘groups’’ pinchonii and tibetanus

3. Since 2010, we repeatedly invited Nikolaï Poyarkov to join us for publishing the taxonomic and nomenclatural novelties in the HYNOBIIDAE presented here, but despite his original agreement with the proposal, he apparently found no time to follow this project out. It is to be hoped that the full text of his thesis (in Russian, a language mastered by one of us, JR) will soon be published and made available to the international community. 114 ALYTES 28 (3¢4) of Fei et al. (2006). The nomen Tibetuperus is derived from the country name Tibet and the ending ¢uperus of the generic nomen Batrachuperus. It has the same grammatical gender (masculine) as the latter. The diagnoses of these subgenera appear in table A1 of appendix 2. Within the genus Pseudohynobius Fei & Ye, 1983, we recognize two subgenera for which nomina are already available. Their diagnoses are given in table A1 of appendix 2.

In the tribe RANODONTINI, we recognize three genera, Ranodon Kessler, 1866, Afghanodon nov. and Iranodon nov., which are diagnosed in table A2 of appendix 2. The latter two nomina (grammatical gender masculine, like Ranodon) are based respectively on the country names Afghanistan and Iran, combined with the ending ¢odon of the generic nomen Ranodon. The nomen Iranodon is meant at providing an available nomen for the taxon called ‘‘Paradactylo- don’’ by a few recent authors (Zhang et al., 2006; Raffaëlli, 2007; Poyarkov, 2010). This nomen is unavailable under the Code because it does not respect the requirement of Article 13.1.2 to provide ‘‘a description or definition that states in words characters that are purported to differentiate the taxon’’. Despite its title, the original paper of Risch (1984) did not provide a single diagnostic character between this ‘‘new genus’’ and the genus Salamandrella Dybowski, 1870. The ‘‘diagnosis’’ of the former mentioned the existence of ‘‘differences’’ in ‘‘certain details’’ regarding the vomero-palatine teeth of the two genera, but these ‘‘differences’’ and ‘‘details’’ were not given, so that ‘‘Paradactylodon’’ is a gymnonym (‘‘nomen nudum’’).

In the genus Onychodactylus of the subfamily ONYCHODACTYLINAE, only two species were long recognized, but four additional ones were recently described by Poyarkov et al. (2012). The phylogenetic data of these authors support the recognition of two subgenera in this genus (see table A1 of appendix 2). In the hyponymous subgenus Onychodactylus, their data support the recognition of two supraspecies, corresponding to their two ‘‘species complexes’’. The species Onychodactylus zhangyapingi, which is unlocated to a species complex in their work, is here referred to the supraspecies japonicus based on their phylogenetic data. The taxonomic status of the fossil genus † Liaoxitriton Dong & Wang, 1998, that is diagnosed by the absence of a fontanelle (present in the two recent subfamilies of HYNOBII- DAE), is open to question and we provisionally treat this genus as incertae sedis at familial level.

Familia Plethodontidae Gray, 1850

With 437 species, i.e. 64.9 % of the 673 recent salamander species recognized here (see table 2), this family is by far the largest one of the order. It has shown a very quick evolutionary diversification and high rates of speciation, and accounting for this evolutionary history under a phylogenetic taxonomic paradigm requires a complex classification, using a high number of ranks and taxa. Important progresses in the understanding of phylogenetic relationships within this group were published in the recent years. Unfortunately, the taxonomic and nomenclatural treatment of these data has so far been partly defective, as discussed in detail by Dubois (2008e) who offered solutions to some of these problems, but taking the care not to provide available nomina for the new taxa that required taxonomic recognition, in order to leave this privilege to the colleagues working on the group. Unfortunately, the nomina ANEIDINI, HYDROMANTINI and BATRACHOSEPINI introduced by Vieites et al. (2011) and used again by Jockush et al. (2012) for the latter, do not conform to Articles 13.1.1 and 16.2 (and Dubois &Raffaëlli 115 to Recommendation 16B) of the Code and remain gymnonyms (‘‘nomina nuda’’) in their paper for missing a ‘‘definition that states in words characters that are purported to differentiate the taxon’’. The misleading idea that a phylogenetic tree (which is a hypothesis of relationship) can replace characters for the availability of nomina has been rejected on several occasions already (see Ohler &Dubois, 2012, and references therein), and does not need to be discussed again here. In order to have a complete and consistent ergotaxonomy of the URODELA reflecting the recent results, we are compelled to make these three nomina finally available here. In fact, in order to express in a more detailed way the available phylogenetic hypotheses concerning the family, we adopted here a more detailed ergotaxonomy using more ranks than Vieites et al. (2011), and consequently these three nomina are here validated at two different ranks, and with two different endings, as we will see below.

In the subfamily HEMIDACTYLIINAE, we adopt a taxonomic hierarchy more complex than in Vieites et al. (2011), with an additional rank subtribe, to account more closely for their phylogenetic tree. The tribe HEMIDACTYLIINI therefore consists in three subtribes BATRACHOSE- PINA nov., BOLITOGLOSSINA and HEMIDACTYLIINA (diagnosed in table A2 of appendix 2), whereas the tribe SPELERPINI accomodates two subtribes PSEUDOTRITONINA nov. and SPELERPINA (diagnosed in table A1 of appendix 2).

In the HEMIDACTYLIINI BOLITOGLOSSINA, the taxonomy of the speciose genus Bolitoglossa is still unstabilized. In the subgenus Bolitoglossa (Eladinea), for the time being, the two species aureogularis and robinsoni seem to represent a distinct lineage which appears as sister-taxon to both supraspecies cerroensis and subpalmata (Boza-Oviedo et al., 2012) and, pending the forthcoming results of studies under way, we treat them as incertae sedis at supraspecies level. In the subgenus Bolitoglossa (Magnadigita), according to Campbell et al. (2010), the supraspecies dunni and morio appear as sister-taxa. Taxonomically, this situation could be accounted for by merging the two taxa in a supraspecies morio and in recognizing two epispecies in this supraspecies, but, as we have seen above, the Code forbids naming taxa at this rank in a Code-compliant manner. A provisional solution to this shortcoming of the Code is to recognize these two epispecies but without naming them formally, e.g., designating them under the letters ‘‘A1’’ and ‘‘A2’’, as shown in appendix 1. In the subgenus Bolitoglossa (Pachymandra), as we have seen above, the two monospecific ‘‘species groups’’ (alvaradoi and dofleini) recognized by Parra-Olea et al. (2004) are redundant and useless, relying presumably on a gradist approach meant at expressing ‘‘degree of divergence’’ through ranks. Therefore in this case, contrary to the cases where at least one of two or more supraspecies includes more than one species, we do not recognize these two ‘‘species groups’’ as supraspecies. In the genus Chiropterotriton, many cryptic species should be soon described by the specialists of this group, clarifying a complex situation with two groups (subgenera) badly defined (Darda, 1994). In contrast to Pachymandra, we think that two distinct subgenera should be recognized in the genus Nototriton to account for the provisional phylogenetic data of García-París & Wake (2000), Sunyer et al. (2010), Townsend et al. (2010, 2011) and Boza-Oviedo et al. (2012). For the unnamed subgenus (diagnosed in table A1 of appendix 2) which includes the species Nototriton barbouri (Schmidt, 1936) and 6 related species (see appendix 1), we provide 116 ALYTES 28 (3¢4) the nomen Bryotriton nov. This term (grammatical gender masculine, like Nototriton)is derived from the classical Greek terms βρυ´ ον, ‘‘moss’’, and Τρι´των, ‘‘Triton’’, name of a Greek marine god, used after Laurenti (1768) as a root for many genus-series nomina of URODELA. It refers to a common habitat of the species of this group, in mosses, barks and low vegetation on the ground. In each of the subgenera of Nototriton, according to the published data (García-París &Wake, 2000; Sunyer et al., 2010; Townsend et al., 2010, 2011; Boza-Oviedo et al., 2012), we recognize two supraspecies (appendix 1). The genus Oedipina includes three subgenera (McCranie et al., 2008). In the hyponymous subgenus Oedipina, four lineages have been identified (García-París &Wake, 2000; Vieites et al., 2007; Sunyer et al., 2010; Boza-Oviedo et al., 2012; Brodie et al., 2012). They could be taxonomically recognized as four supraspecies collaris (valid nomen instead of poelzi), cyclocauda, taylori (valid nomen following the first-reviser action of Brame, 1960) and uniformis. However, three species of the subgenus (alfaroi, altura and paucidentata)havean unresolved position in this group, so we refrain to do so at the moment. The same problem exists within the subgenus Oedopinola which includes multiple cryptic species under the nomina complex and parvipes, even if four lineages have been identified (García-París & Wake, 2000; McCranie & Townsend, 2011; Boza-Oviedo et al., 2012). In the genus Pseudoeurycea Taylor, 1944, to account for the phylogenetic hypotheses of Parra-Olea (2002) and Wiens (2007), we also propose to recognize two subgenera, both of which include supraspecies. For the unnamed subgenus (diagnosed in table A1 of appendix 2) including the supraspecies bellii and cephalica (see appendix 1), we provide the nomen Isthmura nov. This term (grammatical gender feminine, like Pseudoeurycea) is based on the Greek terms ισθμο´ , isthmos, ‘‘isthmus’’, which evokes the constricted basis of the tail of these salamanders, and ου ρα´ , oura, ‘‘tail’’. The works of Parra-Olea (2002) and Wiens (2007) also suggest that the genera Ixalotriton and Parvimolge might be reinstalled inside Pseudoeurycea but we refrain to do so pending the works of the colleagues specialists of this group. Ixalotriton especially might be inserted inside the supraspecies unguidentis in a revised Pseudoeurycea or, alternatively, infratribes could be recognized within the subtribe BOLITOGLOSSINA. In the supraspecies gadovii of the hyponymous subgenus Pseudoeurycea, four species from Guatemala form a distinct lineage (Parra-Olea, 2002). This is another situation where the limitations of the Code impede the recognition and naming of two epispecies in this supraspecies, which can be provisionally solved through the informal recognition of epispecies designated as B1 and B2, as shown in appendix 1.

In the genus Gyrinophilus of the SPELERPINI SPELERPINA, we recognize three exerges (groups of subspecies) within the species Gyrinophilus porphyriticus: danielsi (with danielsi and dunni), duryi (with duryi) and porphyriticus (with porphyriticus and subterraneus, which should perhaps be considered only as a hypogeous population of porphyriticus rather than a taxon). All these taxa are not yet clearly taxonomically deﬁned (Bishop, 1943; Adams & Beachy, 2001). The genus Pseudotriton consists in three main entities, the species diastictus and two other species (montanus and ruber) that include two exerges. This follows partially Bishop Dubois &Raffaëlli 117

(1943) and Collins &Taggart (2009). In the species montanus, we refrain to elevate ﬂavissimus to the species rank because of the existence of a hybrid zone with montanus, even if these two taxa are highly divergent morphologically (Bishop, 1943; personal observations of JR). In the species ruber, the subspecies nitidus, ruber and schencki are very close and Martof(1975)suggestedtosynonymizethesetaxa,fromwhichvioscaiappearsmoredivergent. We provisionally maintained all these taxa pending the publication of molecular data.

In the SPELERPINI SPELERPINA, we propose to recognize provisionally seven subgenera (diagnosed in table A6 of appendix 2) in Eurycea, pending a clariﬁcation of their phylogenetic relationships between these groups, some of which are highly derived because of their cave habitat (Camp et al., 2009). The nomenclatural status of two of these genera (Notiomolge and Paedomolge) was discussed above.

In the subfamily PLETHODONTINAE, we recognize ﬁve tribes (diagnosed in table A4 of appendix 2): ANEIDINI nov., DESMOGNATHINI, ENSATINI, KARSENIINI nov. and PLETHODONTINI.

In the ANEIDINI, we recognize two subgenera (diagnosed in table A1 of appendix 2) in the genus Aneides. For the new genus accommodating the single species Aneides aeneus,we coined the nomen Castaneides nov. (grammatical gender masculine, like Aneides), derived from two scientific nomina: Castanea, the generic nomen of the American chestnut-tree Castanea dentata, and Aneides, the generic nomen of this salamander species. This is meant at recalling that this species depends on the presence of trees and particularly of this chestnut, whose decline in the Appalachians was apparently followed by that of the salamander and its restriction to rocky areas (Wilson, 2003). This subgenus in fact includes three cryptic species, two of which are awaiting description (Highton, personal communication to JR). In the supraspecies flavipunctatus of the hyponymous subgenus Aneides, we follow Lowe (1950), Rissler &Apodeca (2007) and Reilly (2009) to resurrect sequoiensis at the species level for the most northern population, a taxon highly divergent morphologically with a higher number of costal folds (16¢17) than the other species of the supraspecies, a different coloration (grey to olive with a few spots), paedomorphic traits and the habitat on a rocky substrate. Aneides flavipunctatus quercetorum Lowe, 1950, the ‘‘central lineage’’ of Reilly (2009), living east of Central Valley and with large white spots, might be resurrected too in the future on a morphological basis, but is not considered as divergent as the other populations of the former complex from the topotypic flavipunctatus.

In the DESMOGNATHINI, based on the works of Tilley &Mahoney (1996), Larson et al. (2003), Kozak et al. (2005), Tilley et al. (2008) and Beamer &Lamb (2008), we distinguish four subgenera (diagnosed in table A3 of appendix 2) in the genus Desmognathus, including two new ones. The nomina of the latter (of masculine grammatical gender) are combinations of the ending ¢gnathus (from the Greek γνα´ θο, ‘‘jaw’’), which is already common to several generic nomina of the tribe, with roots that indicate the preferred habitats and modes of life of the species at stake, i.e., respectively, terrestrial for Geognathus nov. (from γη˜ , ge, ‘‘earth’’) and aquatic for Hydrognathus nov. (from υ δωρ, hudor, ‘‘water’’). The subgenus Desmognathus (Leurognathus) is currently considered to include three species (e.g., Collins &Taggart, 2009). In fact, two of these species (marmoratus and quadramaculatus) consist in multiple divergent lineages (Martof, 1956; Voss et al., 1995; Jackson, 2005; Kozak et al., 2005; Jones et al., 2006). Nomina (aureatus and melianus)are 118 ALYTES 28 (3¢4) already available for two of these lineages and we resurrect them as valid species. D. aureatus is the southern lineage of the former marmoratus (Voss et al., 1995), whereas melianus from northern Carolina and adjacent Tennessee is very divergent morphologically from topotypic marmoratus and phylogenetically closer to D. quadramaculatus living in sympatry. It is much more terrestrial than other members of the former marmoratus complex (personal observations of JR). The genus Desmognathus is composed of multiple cryptic forms and many more species should be described in the near future. The overall diversity of this genus is in fact approaching that of Plethodon in the eastern United States, with at least 45 species of which only slightly more of 20 are formally described and named.

In the PLETHODONTINI, we reinstalled Plethodon larselli in the supraspecies vandykei,as suggested by Mahoney (2001), Pyron &Wiens (2011), Highton (personal communication), and according to biogeographical evidence. This taxon was considered close to P. neomexi- canus by the presence of only one phalanx at the ﬁfth toe, but this is probably a phenomenon of convergence. P. larselli might be the sister-taxon of all other western Plethodon (subgenus Hightonia). The supraspecies Plethodon glutinosus consists in several groups which have been clearly named (Highton et al., 1989; Highton &Peabody, 2000) and which correspond to the rank epispecies. This situation is similar to those of the supraspecies Bolitoglossa morio and Pseudoeurycea gadovii: because the Code forbids the use of a formal nomenclatural rank between supraspecies and species, we are compelled to use informal (C1 to C4) designations for these epispecies. Finally, we recognize two subtribes (diagnosed in table A1 of appendix 2) in the tribe KARSENIINI: HYDROMANTINA nov. and KARSENIINA nov. As both nomina are new, we give priority to the shortest one for reasons given by Dubois &Raffaëlli (2009: 17¢22).

Familia Proteidae Gray, 1825

We here follow Blackburn &Wake (2011: 47) in recognizing two subfamilies in this family, but then, because of the Principle of Priority, the subfamily including the genus Necturus must be called PHANEROBRANCHINAE Fitzinger, 1826, not NECTURINAE Fitzinger, 1843. In the genus Necturus, we recognize two subgenera, Necturus and Parvurus nov., diagnosed in table A1 of appendix 2. The nomen Parvurus (of masculine grammatical gender, like Necturus) derives from the Latin adjective parvus, ‘‘small’’, combined with the ending ¢urus of the nomen Necturus (derived from the Greek noun ου ρα´ , oura, ‘‘tail’’). We here resurrect Necturus lodingi, from the Dog River drainage in Mobile county, Alabama, as a valid species. This taxon, long known as Necturus cf. beyeri or cf. alabamensis, was already recognized as a distinct species under the nomen lodingi by Bishop (1943), and as a subspecies of N. punctatus by Hecht (1958), when Maxson et al. (1988), Guttman et al. (1990) and Bart et al. (1997) suggested than an undescribed taxon diﬀerent from N. beyeri and alabamensis occurs in the Gulf Coast area west of Mobile Bay (Alabama). It is highly divergent morphologically from the populations of beyeri living east of Mobile Bay. This small species with a dark colour, black spots and an unpigmented belly, is not restricted to Alabama: its distribution covers also western Florida and southern Georgia, in the Choc- Dubois &Raffaëlli 119 tawhatchee and Apalachicola drainages (Maxson et al., 1988; Petranka, 1998). It is sometimes called the Florida Waterdog.

Familia Salamandridae Goldfuss, 1820

We recognize four distinct species so far known under the nomen Cynops pyrrhogaster. We hereby resurrect Cynops immaculiventris (Krefft, 1898) for the populations of Kagoshima and Miyazaki (extreme south of Kyushu, Japan), which are very divergent both on a molecular basis and morphologically (small size, usually no black spots on the belly) from the other populations of the so-called ‘‘Hiroshima race’’ (Sawada, 1963; Hayashi &Matsui, 1988, 1990; Highton 2000). We also resurrect Cynops pyrrhogaster sasayamae (Mertens, 1969) on a morphological basis and because of its peculiar reproductive behaviour (Sawada, 1963; personal observations of JR). Other forms traditionally referred to C. pyrrhogaster, mainly the populations of Atsumi and Kanto, should be recognized as distinct species, and the population of Tohoku, itself polymorphic (Hayashi &Matsui, 1990), as a distinct subspecies of C. pyrrhogaster, but no nomen is so far available for these forms. In the genus Hypselotriton, we hereby recognize the former subspecies yunnanensis of the species cyanurus as a distinct species due to its larger size, granular skin, very different coloration and behaviour (personal observation of JR). Cynops yunnanensis is much larger than cyanurus (12.5 cm vs. 10 cm), spotted in black (vs. no black spots or very few), it has a more granular skin, lives at lower temperatures (optimum 18°C vs. 25°C) and the male displays a different mode of fanning during sexual courtship. The size of C. yunnanensis is intermediate between those of cyanurus and of the (now considered extinct) wolterstorffi. The sexual dimorphism is similar between wolterstorffi and yunnanensis, males being much smaller than females (personal observations of JR on specimens in the Paris Museum collection), whereas the sexual dimorphism in cyanurus is weaker. Cynops cyanurus chuxiongensis Fei & Ye, 1983, currently considered as a synonym of H. yunnanensis, might be in the future reinstalled as a valid taxon. The western Anatolian populations of Lissotriton vulgaris which were referred by some authors to the subspecies schmidtleri (sometimes misspelt schmidtlerorum; see Dubois & Raffaëlli, 2009) are not morphologically distinct from those of the hyponymous subspecies vulgaris, except for that of Efes near Izmir (Olgun et al., 1999). The latter could deserve to be recognized as a distinct subspecies, but for this a new nomen should be provided because the onymotope (‘‘type-locality’’; Dubois, 2005b: 404) of schmidtleri is Karacabey, not Efes.

Familia Sirenidae Gray, 1825

We here resurrect Siren intermedia texana as a valid subspecies. Siren texana was described by Goin (1957) from the Rio Grande Basin in Texas and this taxon was recognized by Martof (1973) and Petranka (1998) but not by Flores-Villela &Brandon (1992). Its morphology is very different from that of the other close forms. It has 36¢38 costal folds (vs. 31¢33 for intermedia and 35¢37 for nettingi), a very large size (up to 69 cm vs. 38 cm and 50 cm respectively) and a different coloration (usually no yellow stripe on the side of the head, belly light grey). It has a narrow isolated distribution at the Mexican-US border of Texas and Tamaulipas. Another unnamed form, different from texana, lives in Mexico along the Gulf of 120 ALYTES 28 (3¢4)

Mexico, from Tamaulipas to northern Veracruz. It is still not clear if texana is not closer to S. lacertina than to S. intermedia and the zones of intergradiation between the 3 subspecies of intermedia might not exist (Petranka, 1998). Pending molecular works of all taxa of Siren, we refrain to elevate to a species level S. i. intermedia and S. i. nettingi, but a general review of the group is clearly needed to clarify the status of all the forms of the genus (Moler in Petranka, 1998). There might be several unnamed taxa under the names intermedia and lacertina, and texana itself might be a good species.

Conclusion

Some quantitative data

The ergotaxonomy of the URODELA presented here (appendix 1) recognizes 1240 taxa (1054 recent and 186 all-fossil taxa) referred to 17 ranks (including an informal one). Table 2 gives details on their distribution in the four nominal-series and in the main higher taxa of the order.

The ten families that we recognize in the URODELA are highly diﬀerent in size and composition, which provides information on several aspects of the taxonomy of the order.

The largest family, the PLETHODONTIDAE, contains 437 species, i.e. 64.9 % of the 673 recent species of the order. However, this is doubtless still quite far from the total number of taxa of this family, as witnessed by several quantitative indicators and specialists of this group in North and Latin America. First, with 27 genera, i.e. 38.6 % of the 70 recent genera known in the order, and 27, i.e. 24.3 % of its 111 subspecies, the taxonomy of this family appears less ‘‘divided’’ than those of other families, especially the SALAMANDRIDAE for which many additional taxa were recently recognized (Dubois &Raffaëlli, 2009). No fossil taxa of PLETHO- DONTIDAE have been described so far, which is also a strong diﬀerence with several other families. Finally, the high number of new species described each year in the recent decades also suggests that the taxonomy of this family is still far from being stabilized, even if much has been done in the recent years (see e.g. Vieites et al., 2011).

The second largest family, the SALAMANDRIDAE, shows quite opposite characteristics. It harbours 107 recent species, i.e., 15.9 % of the 673 recent species of the order, but 31 (44.3 % of those of the order) of its genera and 62 (55.9 % of those of the order) of its subspecies: its taxonomy is therefore much more ‘‘divided’’ than that of the PLETHODONTIDAE. It also includes 29 all-fossil species, i.e., 24.8 % of the 117 all-fossil species of URODELA currently recognized. The other 8 families are much less speciose and allow little generalization, but their data support the following rough interpretation. It seems that the urodeles of Europe and Asia are better known than those of America, where many more species would remain to discover. Fossil urodeles have long been known and studied in the Old World, especially in the most ancient families (see Zhang &Wake, 2009), but their knowledge is in a much less advanced stage in the New World. Possibly an important factor for the discrepancies between the two groups of taxa could derive from a different approach of the ‘‘species problem’’. In the recent years, many works on Dubois &Raffaëlli 121 122 ALYTES 28 (3¢4) the New World urodeles were based on a phylogenetic approach using mostly nucleic acid sequencing. In the Old World, whereas this approach is also developing, it is often combined with detailed data on hybridization between natural entities in the field, on the structure and dynamics of hybrid zones and on gene flow between entities. Although there are exceptions, it would seem that on the whole the dominant ‘‘species concept’’ used in the New World tends to be the ‘‘phylogenetic species concept’’, ‘‘PSC’’ or simpson (Dubois, 2011b), whereas in the Old World the ‘‘mixiological species concept’’, ‘‘BSC’’ or mayron (Dubois, 2011b) would still be largely used. This might explain in part the apparent relative scarcity of subspecies in the New World and their much higher abundance in the Old World. In other words, the relatively high differences between the taxonomic patterns in the two regions would be in part or even largely an artifact, due to the use of different dominant taxonomic paradigms rather than to characteristics of the organisms themselves. This suggestion might be worth exploring on the basis of more complete and varied data. Anyway, for the future, the best solution to the ‘‘species-problem’’ probably lies in a harmonious combination of phylogenetic, genetic, ecological and other biological data, that would allow to account for the high variety of evolutionary situations that occur in nature, rather than in an artificial and rigid ‘‘unified species concept’’ (Dubois, 2011b).

Zoological taxonomy and nomenclature nowadays

In the recent years, zootaxonomy has witnessed a boost of quick and drastic improve- ments in several technical fields: computer sciences (size of equipment, speed and efficiency), electronic communication (including scanning of literature), molecular techniques (automatic sequencing, alignment, barcoding), cladistic analysis (softwares), geometric morpho- metry, etc. All these techniques are impressive and receive strong financial support from academic and private fund sources. But they also carry the risk of a fascination by the tool,at the expense of scientific questions (Dubois, 2008a). Taxonomy, long considered an ‘‘old-fashioned’’ discipline, was revolutionized by Hennig’s works which provided conceptual tools (mostly the distinction plesiomorphy/ apomorphy) to make it refutable and repeatable, and thus quit its previous functioning under an ‘‘argument of authority’’. In contrast, nomenclature is still expecting its ‘‘Hennigian revolution’’. The hopes that some had put in some projects of ‘‘phylogenetic nomenclature’’ have been deceived, for various reasons, the most important of which being (1) the fact that the new systems devised are theory-bound, do not respect the freedom of taxonomic thought and will not be able to adapt to future potential changes in our taxonomic paradigms, and (2) that they would require to rename millions of taxa, at a time when taxonomists should rather concentrate on the discovery and study of the vanishing biodiversity of our planet (Dubois, 2005b, 2010b). However, the situation of ‘‘traditional nomenclature’’, despite its many achievements in the past (it has allowed to name millions of taxa), is far from being satisfactory.Nomenclature is (and cannot be but) a very complex field but it is still in medieval times in some respects. It contains a mixture of useless rigidity (e.g., concerning the number of acceptable ranks) and of (sometimes apparent) laxity, in particular because it still heavily calls on an ‘‘argument of Dubois &Raffaëlli 123 authority’’ whenever problems appear: it still makes use of vague concepts like ‘‘consensus’’ and ‘‘usage’’, which act directly in the direction opposite to its needs of standardisation and repeatability. Some taxonomists and phylogeneticists do not hesitate to declare that they have ‘‘no interest’’ in zoological nomenclature, because they are more interested in evolution and phylogeny than in codes and rules This is their full right! All biologists can be grateful to them if they can contribute to an increase of our knowledge about organisms and evolution. But, as we have shown above, use of well-defined concepts, criteria and Rules allows zoological taxonomy and nomenclature to express in a clear and unambiguous way our data and ideas about species taxa and our data and hypotheses about phylogenetic relationships between species. This is crucial for proper communication between taxonomists, evolutionists and other biologists. In contrast, use of an unclear methodology, pre-eminence given to opinions and tastes over criteria and Rules, and lobbyism, act against this effort. We hope in the future taxonomists will choose objectivity and clarity of methods and concepts rather than subjec- tivity, traditions, habits and ‘‘group behaviours’’.

Acknowledgements

We are very grateful to Marc Alcher (Hanches), Roger Bour (Montgeron), Myrianne Brival (Paris), Pierre-André Crochet (Montpellier), Andrea Kourgli (Wien), Victoire Koyamba (Paris) and Annemarie Ohler (Paris) for their varied support and help, especially regarding bibliographic research, during the work on this paper. The rich and multiform contribution of Thierry Frétey (Saint-Maugan) to our work is particularly appreciated. Jean-Michel Collet (Saint-Pol-de-Léon), member of the French Urodela Group, helped us eﬀiciently for the preparation of diagnoses and for bibliographic research regarding fossil taxa. Stéphane Grosjean and two reviewers provided useful comments on the original manuscript.

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Appendix 2

Diagnostic comparisons among several groups of URODELA Duméril, 1805

The criterion of adaptability of the species to terrarium is based on the personal experiences of one of us (JR) and of several other breeders (personal communications). We recognize two categories (Dubois &Raffaëlli, 2009: 42): high adaptability in terrarium (HAT), for species that can be kept for several years in captivity under various conditions of temperature, humidity and food offer, in terraria where they can develop complete breeding behaviour and give birth to offspring, sometimes repeatedly; and low adaptability in terrarium (LAT), for species that do not easily reproduce in captivity and do not adapt well to variability for conditions of temperature, humidity, food offer and general husbandry; in the last case, animals must be kept under strict conditions of captivity which have to be determined on a permanent basis. This criterion expresses in a synthetic way several ethological, physiological and more generally biological characteristics and limitations of the organisms (requirements and constraints regarding temperature, humidity, space, shelter, etc.), that have not been analysed in detail yet. Other abbreviations: SVL, snout-vent length; TL, total length. Dubois &Raffaëlli 155 156 ALYTES 28 (3¢4) Dubois &Raffaëlli 157 158 ALYTES 28 (3¢4) Dubois &Raffaëlli 159 160 ALYTES 28 (3¢4) Dubois &Raffaëlli 161