The Nomenclatural Status Hypsaplesia-DUBIOS 2017

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The nomenclatural status of Hysaplesia, Hylaplesia, Dendrobates and related

nomina (Amphibia, Anura), with general comments on zoological
nomenclature and its governance, as well a...
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DOI: 10.11646/bionomina.11.1.1
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Alain Dubois
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Bionomina, 11: 1–48 (2017) ISSN 1179-7649 (print edition)
http://www.mapress.com/j/bn Article BIONOMINA
Copyright © 2017 • Magnolia Press ISSN 1179-7657 (online edition)
https://doi.org/10.11646/bionomina.11.1.1
http://zoobank.org/urn:lsid:zoobank.org:pub:021A0840-2236-4067-AE88-39A11F52D13A
The nomenclatural status of Hysaplesia, Hylaplesia, Dendrobates and related

nomina (Amphibia, Anura), with general comments on zoological nomenclature
and its governance, as well as on taxonomic databases and websites
Alain DUBOIS
Institut de Systématique, Évolution, Biodiversité, ISYEB – UMR 7205 – CNRS, MNHN, UPMC, EPHE, Muséum national
d’Histoire naturelle, Sorbonne Universités, 57 rue Cuvier, CP 30, F-75005, Paris, France. <adubois@mnhn.fr>.
Table of contents
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Key words . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Terminology, abbreviations and symbols . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Categories of new nomina . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Chronology of the case . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Discussion and conclusion. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
A comprehensive solution to all the nomenclatural problems at stake . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
Zoological nomenclature and its governance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
Zoological nomenclature . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
The Commission . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
The Code. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
Taxonomic databases and websites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
[D1] Bibliographic databases and websites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
[D2] Phylogenetic databases and websites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
[D3] Taxonomic (s. str.) databases and websites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
[D4] Nomenclatural databases and websites . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
Appendix 1. Extracts of the paper of Dubois (1987b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
Abstract
Dozens of publications, mostly in the last 45 years, have been devoted to nomenclatural problems concerning the status
of the zoological nomina Hysaplesia, Hylaplesia, Dendrobates and related nomina. The Commission finally voted on
this case in 2009, but this vote shows a misunderstanding of several of the problems at stake, as it contains a double and
contradictory decision: the change in the type species of Hysaplesia and its suppression, although one only of these two
acts would have been necessary and sufficient to solve this case, whereas taking them both together has other unforeseen
and negative nomenclatural consequences. A final comprehensive solution to these nomenclatural problems is presented
here, which does not require any more action from the Commission. However, the fact that the Commission, as well as
the whole international community, have proved to be unable to understand fully the nomenclatural problems at stake
and to solve them truthfully, while ignoring deliberately some contributions to the discussion, calls attention. This case
suggests that there is a strong risk that nomenclature might become a domain where intellectual fairness and competence
1
2 • Bionomina 11 © 2017 Magnolia Press DUBOIS
are secondary and where problems are ‘solved’ through the medieval ‘Principle of Authority’, through relying on the
‘opinions’ of a few persons, committees or websites rather than on rational discussions based on a knowledge of
taxonomic publications and an understanding of the Code. This course would result in dragging zoological nomenclature
so to say outside the field of science and contribute to weakening still more a domain, taxonomy, which is already facing
major problems, at the time of the crisis of biodiversity. Additional problems concerning taxonomic databases and
websites are pointed out, and suggestions are offered in this respect, including the distinction between the concepts of
‘nomenclatural status’ and ‘taxonomic status’ of nomina.
Key words: Taxonomy; nomenclature; Code; Commission; nomen novum; incorrect spelling; Plenary Powers; Principle
of Authority; databases; websites; nomenclatural status of nomen; taxonomic status of nomen.
Introduction
The nomenclatural problems posed by the amphibian nomina Hysaplesia, Hylaplesia, Hyloplesia and
Dendrobates, and their related family-series nomina, have already been discussed at length in various
publications, and, after a long delay, have finally been the matter of a decision by the International
Commission of Zoological Nomenclature (‘the Commission’ below). However this decision, based on a
wrong interpretation of some facts, still leaves this question partly unsolved. It is, therefore, necessary to
come back to it in detail again, in the light of all the relevant publications and of the International Code of
Zoological Nomenclature (‘the Code’ below; Anonymous 1999, 2012), as several recent authors still have not
understood, or deliberately ignore, some aspects of the controversy. The best way to examine this case is
through a chronological presentation of the facts, followed by a discussion and a conclusion.
Terminology, abbreviations and symbols
In what follows, for sake of both brevity and more clarity in the presentation of the facts and problems, the
following terms, abbreviations and symbols are used:
Terms
Alloneonym ● Neonym having a partially or totally different etymology from its archaeonym, i.e., not directly derived
from it through unjustified emendation (Dubois 2000).
Ameletograph ● Spelling of a nomen used inadvertently in a publication by an author, editor or publisher (Dubois 2011).
Amendment ● Taxonomic concept designating a modification in the status of a taxon, concerning its intension
(definition) or its extension (content, circumscription) (Dubois 2012, as ‘amendation’).
Apograph ● Any subsequent spelling of a nomen. (Dubois 2010a).
Aponym ● Any subsequent paronym of a protonym, modified in spelling, rank and/or combination (Dubois 2000).
Arbiter ● Name(s) of the person(s) responsible of the publication of a ‘First Reviser’ action settling an ambiguous
nomenclatural situation regarding a nomen, a spelling or a nomenclatural act (Dubois 2013).
Archaeonym ● Original nomen that has been replaced by a neonym (Dubois 2006a).
Assignment ● Nominal-series assignment of a nomen: the nominal-series to which this nomen is referred according to a
given system of assigment of nomina to nominal-series (e.g., through original statement of the author of the nomen
or through objective criteria) (Dubois 2015b).
Auctor ● Name(s) of the person(s) (‘author’) responsible of the publication of a new nomen or nomenclatural act
(Dubois 2013).
Authorship ● In the context of zoological nomenclature, name(s) of the auctor(s), or author(s), of a published work,
nomen or nomenclatural act, which is usually followed by the year of its publication.
HYSAPLESIA, HYLAPLESIA, DENDROBATES AND RELATED NOMINA Bionomina 11 © 2017 Magnolia Press • 3
Autoneonym ● Neonym having the same etymology as its archaeonym, i.e., directly derived from it through unjustified
emendation (Dubois 2000).
Class-series (CS) ● In the nomenclatural hierarchy, the nominal-series ranked above the family-series, which is not fully
regulated by the Code. It includes nomina of taxa at the ranks of phylum, class, order, and any additional ranks that
may be required (Dubois 2000).
Emendation ● Nomenclatural concept designating a modification in the spelling of a nomen, which may be [1] either
voluntary (meletograph) or involuntary (ameletograph), and [2] either justified (nomograph), thus devoid of
independent nomenclatural availability, or unjustified (autoneonym), thus being an available nomen independent
from its archaeonym and having its own authorship and date.
Ergotaxonomy ● Any classification considered valid in a certain work by a given author (Dubois 2005).
Family-series (FS) ● In the nomenclatural hierarchy, the highest-ranking nominal-series fully regulated by the Code. It
includes nomina of taxa at the ranks of family, subfamily, tribe, superfamily, and any additional ranks that may be
required (Dubois 2000).
Genus-series (GS) ● In the nomenclatural hierarchy, the nominal-series ranked between the species-series and the
family-series. It includes taxa at the ranks of genus and subgenus (Dubois 2000).
Meletograph ● Spelling of a nomen used intentionally in a publication by an author (Dubois 2011).
Monophory ● Qualification of a nomen created with and supported by an onomatophore composed of a single specimen
(in the species-series) or taxomen (in the three other nominal-series) (Dubois 2005).
Neonym ● Nomen proposed expressly to replace an available nomen (its archaeonym), and having the same
onomatophore (nomen novum or new replacement name in the Code) (Dubois 2000). Two categories: alloneonym
and autoneonym.
Nomen (pl. nomina) ● Scientific name as defined and regulated by the Code (Dubois 2000).
Nominal-series ● Any of the sets of coordinated nomina interacting for priority regarding synonymy and homonymy
(species-series, genus-series, family-series or class-series) (Dubois 2000).
Nomograph ● Spelling that is imposed as ‘correct’ by the Code to a given nomen, superseding the protograph if
necessary, in two situations: either mandatory spelling correction (‘justified emendation’) because the protograph is
an ‘incorrect original spelling’; or mandatory ending correction (‘mandatory change’) because the ending of the
protograph must be corrected as a result of combination change in the SS or of rank change in the FS (Dubois
2013).
Onomatophore ● Objective standard of reference determining the taxonomic allocation of a nomen: the nomen can be
potentially applied to any taxon that includes the onomatophore (Simpson 1940).
Parograph ● Any spelling of a nomen, whether original (protograph) or subsequent (apograph) (Dubois 2010a).
Paronym ● Any of the avatars of a nomen, either original (protonym) or subsequent (aponym), and concerning its
spelling, rank and/or combination (Dubois 2000).
Poieonym ● Brand new nomen directly introduced for an unnamed brand new taxon, not as a neonym for an already
available nomen for an already defined taxon. New term introduced by Dubois (2017a) in replacement of the term
‘euhoplonym’ (Dubois 2012) having the same meaning but an inappropriate etymology to point to this concept.
Protoallelonym ● One of two (or several) nomina having the same onomatophore proposed for the same taxon (same
content) in the same publication (Dubois 2011).
Protograph ● A category of protonym: original parograph (spelling) of a nomen (Dubois 2010a).
Protonym ● Original spelling, rank and/or combination of a nomen (Dubois 2000).
Scriptor ● Name(s) of the person(s) responsible of the fitst publication of an aponym (see Dubois 2013).
Species-series (SS) ● In the nomenclatural hierarchy, the lowest-ranking nominal-series which is fully regulated by the
Code, ranked below the genus-series. It includes nomina of taxa at the ranks of species, subspecies, species
aggregate and subspecies aggregate (Dubois 2000).
Symprotograph ● One of two or more alternative original spellings of a nomen (Dubois 2010a).
Symprotonym ● One of two or more alternative original protonyms of a nomen (Dubois & Ohler 2009).
Synchronous ● In the context of zoological nomenclature, the fact that two publications were distributed at the same date
(Dubois 2013).
Taxomen ● The permanent association between a nomen and an onomatophore, allowing objective, non-ambiguous and
stable allocation of nomina to taxa (Dubois 2000).
Abbreviations and symbols
Article [followed by a number] ● Article of the Code.

ASW ● Website Amphibian Species of the World <http://research.amnh.org/vz/herpetology/amphibia/>, version 6.0 of 8
August 2016.
BZN ● Bulletin of zoological Nomenclature, published by the Commission.
The Code ● The International Code of Zoological Nomenclature (Anonymous 1999, 2012).
The Commission ● The International Commission on Zoological Nomenclature.
≡ ● Objective synonym of.
≈ ● Subjective synonym of.
◄ ● Neonym (nomen novum) for.
Nomen between double straight quotes " " ● Unavailable nomen.
Nomen between single straight quotes ' ' ● Nomen invalidated by the Commission for the purposes of synonymy but not
of homonymy: e.g. 'Hysaplesia' H. Boie in Schlegel, 1826a.
Part of text between double curved quotes “ ” ● Exact quotation (‘sic’) from a cited publication.
Part of text between simple curved quotes ‘ ’ ● Word or expression highlighted but not being an exact quotation from a
cited publication.
Categories of new nomina
In order to fully understand what follows, it is important to realise that, under the Rules of the Code, there are
different categories of ‘new nomina’ in the taxonomic literature. It would be very cumbersome and redundant
to repeat here the detailed analysis of this question provided elsewhere (Dubois 2010a), which should be
consulted for more details.
A large part of the difficulties which many taxonomists face to master this question is due to its illogical
presentation in the Code, and particularly to the fact that ‘new replacement names’ (or nomina nova) and
‘unjustified emendations’ are presented in different parts of this document (in Articles 12–13 and 33
respectively), as if these concepts had nothing to do with each other, although they are closely related and
even quite difficult to distinguish in some cases (see e.g. the case of the nomen Triturus Rafinesque, 1815
analysed by Dubois 1985: 67–68). In both cases, an author decides, either for good or for bad reasons, to
replace an existing nomen by another one, which differs from the former only by a few letters (unjustified
emendation) or is completely different (nomen novum). In both cases, the Code considers the spelling or
nomen proposed to replace the original one as a distinct nomen, with its own author and date, but with the
same onomatophore ‘(name-bearing type’) and date. Although in the early days of taxonomy the new spelling
or nomen could be adopted as valid by some authors, nowadays in both cases the junior nomen is invalid for
being a junior objective synonym of the original one—except when the senior nomen itself proves to be
invalid, for being a junior homonym of another zoological nomen or for having been invalidated by the
Commission. In order to show that both categories of nomina are much more closely related than it appears in
the Code, Dubois (2000) proposed the general term neonym for both categories, with a distinction between
autoneonyms (unjustified emendations) and alloneonyms (new replacement nomina). Furthermore, as the
Code provides no term for the ‘replaced’ nomen in such cases (except the unclear formula ‘prior nominal
taxon’ in Article 67.8), the term archaeonym was proposed for this purpose (Dubois 2006a). Additionally, for
more clarity in the explanation of the rather complex Rules of the Code in these matters, the introduction of
some special terms not used in the Code (Dubois 2011) proved to be very efficient. These include the terms
protograph (original spelling) and apograph (subsequent spelling), as well as meletograph (intentional
spelling) and ameletograph (unintentional spelling).
The result of this analysis was summarised in a dichotomic key to different nomina and spellings
proposed by Dubois (2010a: 29–30). The interest of such a dichotomic key is that it so to say ‘prepares the
ground’ for a potential conception and implementation of a software allowing a partial automatisation of the
Rules of the Code, which would allow an important improvement of the work of taxonomists. The original
key is here modified and simplified as follows:
1. The writer uses a new nomen for a taxon that he/she considers new: the nomen (poieonym) and the
nominal taxon are new, and must all be credited to the author of the publication. Nowadays, the new nomen is
usually presented with an indication (‘gen. nov.’, ‘n. sp.’, etc.) which makes it clear that a new nomen and
nominal taxon are created, and, according to Art. 16.1 of the 1999 edition of the Code, since 2000 this
statement is compulsory for nomenclatural availability. In older texts however, indications like ‘mihi’ or
‘nobis’ were sometimes used, but in many other cases the fact that the nomen and nominal taxon were created
as new in the paper must be inferred from other direct or indirect sources of evidence. Regarding the spelling
of the new nomen, two possibilities exist:
1.1. The new nomen appears in the original publication under a single spelling, its protograph. Two cases
must be distinguished:
1.1.1. The protograph is correctly formed and must therefore be considered the ‘correct original spelling’
of the nomen according to Art. 32.2 of the Code.
1.1.2. There exists, in the original publication itself, evidence of an ‘inadvertent error, such as a lapsus
calami or a copyist’s or printer’s error’ (Art. 32.5.1) in the protograph, which is therefore an ‘incorrect original
spelling’. In such a case, it is necessary to ‘correct’ the spelling of the nomen. The new spelling, called
‘justified emendation’ in the Code, is just an apograph, which has no independent nomenclatural status: it is
the same nomen, has the same onomatophore, auctor (‘author’) and date as the protograph. In this case the
‘correct spelling’ of the nomen is not the protograph, but one of its apographs, specifically its nomograph.
1.2. The new nomen appears in the original publication under two or more parographs, its ‘multiple
original spellings’. The ‘correct original spelling’ (nomograph) must then be fixed by a ‘First Reviser’ action
by an arbiter under either Art. 24.2.3 or 24.2.4. The rejected spelling(s) become(s) incorrect original
spelling(s) of the same nomen, devoid of independent nomenclatural status (Art. 19.3).
2. The writer deals with a nomen which is credited to a previous auctor, even if the intension (definition) or
extension (content) of the taxon was modified (amended) in part, e.g. by retiring specimens or taxa from it or
by adding others. This reference to an already existing nomen is usually explicit, but in some cases must be
inferred from the context: for example, in many ancient publications, nomina like Rana or Hyla were used
without any mention of their auctors, but it was however clear that they were not new, homonymous nomina.
This is particularly frequent regarding family-series nomina, as the tradition in zootaxonomy is not to mention
the auctors of these nomina. Regarding the spelling of this nomen, two possibilities exist:
2.1. The writer uses exactly the same spelling (protograph) as the auctor who had first recognised the taxon
and named it. In such situations, neither the nomen nor the nominal taxon are new, even if the taxon or its rank
have been amended (e.g., a subgenus raised to generic rank).
2.2. The writer uses a spelling slightly or totally different from that of the original nomen. Two
possibilities again appear here:
2.2.1. The new spelling differs slightly (e.g., by one letter or a few letters) from the protograph, and this
difference in spelling is not intentional from the part of the writer: it may be due to a misspelling on the part of
the auctor (e.g., a mistake in copying the original text) or of the printer (misprint). Such a modified spelling or
apograph is an ‘incorrect subsequent spelling’ or ameletograph which has no independent nomenclatural
status (Art. 19.1).
2.2.2. The writer intentionally uses a different spelling or emendation (meletograph), or even a brand new
different nomen, because he/she thinks, for some reason, that the protograph is incorrect or invalid and must
be modified or replaced. Two main categories exist here:
2.2.2.1. The protograph was ‘incorrect’ and had to be changed into its nomograph to become Code-
compliant. Two cases must then be distinguished:
2.2.2.1.1. The protograph resulted from an ‘inadvertent error’ (see 1.1.2 above): the new spelling is a
‘justified emendation’ of the same nomen, which has no independent nomenclatural status (Art. 19.2).
2.2.2.1.2. The protograph was ‘correct’ at the original rank and/or combination in which it was originally
published, but has to be modified in its ending because of a change in rank (in the family-series) and/or
combination (in the species-series): the new spelling is a ‘mandatory change’ of the same nomen, which has
no independent nomenclatural status (Art. 19.4).
2.2.2.2. The protograph was ‘correct’ under the Rules of the Code, and a change in its spelling was not
warranted. In this case it is clear that the nominal taxon is not new but that the nomen is. This new nomen or
neonym is called by the Code ‘new replacement name’ or ‘nomen novum’. It has an independent
nomenclatural status, with its own auctor and date, but keeps the same onomatophore as the replaced nomen
or archaeonym. Nowadays, this new nomen is usually first published with the indication ‘nom. nov.’. In older
times however, whereas it was sometimes presented as a new nomen in an explicit sentence, in many other
cases the fact that it was such a nomen must be inferred from other direct or indirect sources of evidence,
analysed in detail by Dubois (1987b: 35–38). This latter category may again be subdivided into two
categories, although, a discussed elsewhere (Dubois 2010a: 11 and references therein), no objective or reliable
criteria currently exist to distinguish between them in all cases:
2.2.2.2.1. The new nomen was coined as an intentional emendation of the protonym of the original nomen,
from which it was ‘clearly’ derived. It is then an ‘unjustified emendation’ or autoneonym for the original
nomen.
2.2.2.2.2. The new nomen is a ‘new replacement name’ in the narrow sense of the term, i.e., excluding
‘unjustified emendations’. It was not ‘clearly’, or not at all, derived from the original nomen. It is then an
alloneonym for the original nomen, i.e., a completely new nomen. In practice however, as discussed with
some examples above and in Dubois (1985, 1987b), it is not always easy to establish whether a neonym was
derived from the replaced nomen, or not, i.e., whether it is an autoneonym or an alloneonym.
Chronology of the case
Note. For sake of brevity, in the text below the following nominal species will be designated by numbers in
the order of their appearance in the text. In the following list, each nominal species is followed by the sign >,
and then by the nomen of the genus to which this species is currently referred and to the current nomen of its
family.
[S01] "Hylaplesia borbonica" Boie in Schlegel, 1826a (nomen nudum).

[S02] "Hylaplesia achatina" Boie in Schlegel, 1826a (nomen nudum).
[S03] Hyla trivittata Spix, 1824 > Ameerega Bauer, 1986 (DENDROBATIDAE).
[S04] Hyla nigerrima Spix, 1824 > Ameerega Bauer, 1986 (DENDROBATIDAE).
[S05] Rana tinctoria Cuvier, 1797 > Dendrobates Wagler, 1830 (DENDROBATIDAE).
[S06] Calamita punctatus Schneider, 1799 > Boana Gray, 1825 (HYLIDAE).
[S07] Hyla luteola Wied-Neuwied, 1824 > Phyllodytes Wagler, 1830 (HYLIDAE).
[S08] Hylaplesia borbonica Tschudi, 1838 > Leptophryne Fitzinger, 1843 (BUFONIDAE).
[S09] Hylaplesia picta Tschudi, 1838 > Ameerega Bauer, 1986 (DENDROBATIDAE).
[S10] Microhyla achatina Tschudi, 1838 > Microhyla Tschudi, 1838 (MICROHYLIDAE).
[S11] Phyllobates bicolor Bibron in de la Sagra, 1840 > Phyllobates Bibron in de la Sagra, 1840
(DENDROBATIDAE).
[S12] Dendrobates obscurus Duméril & Bibron, 1841 > Ameerega Bauer, 1986 (DENDROBATIDAE).
[S13] Colostethus latinasus Cope, 1866 > Colostethus Cope, 1866 (DENDROBATIDAE).
[S14] Rana boans Linnaeus, 1758 > Boana Gray, 1825 (HYLIDAE).
[C01] Gray (1825)
Gray (1825) introduced the new subgeneric nomen Boana. Its status is discussed in detail below under
[C34b].
[C02] Schlegel (1826a)
The nomen Hysaplesia first appeared, credited to ‘Boïé’, in a paper in Dutch by Schlegel (1826a) published in
the October 1826 issue of the Bulletin des Sciences naturelles et de Géologie. This paper reported about an
unpublished manuscript by Heinrich Boie entitled Erpétologie de Java that was never published subsequently
(see Adler 1989). The entry for this genus, on page 239, was as follows:
“N. Gen.: HYSAPLESIA Boïé. Esp.: H. borbonica K. & V. H. n. esp. – H. achatina K. & V. H. n. esp. – H.
trivittata Spix. – H. nigerrima Spix. – H. punctata Daudin. – H. tinctoria Daud. – H. luteola P. Max.”
No diagnosis or description was provided for this new genus, but before 1931 this does not by itself
impede nomenclatural availability of a generic nomen provided at least one available specific nomen is
referred to the genus (Article 12.2.5). Among the seven nomina listed, two, [S01] "Hylaplesia borbonica" and
[S02] "Hylaplesia achatina", both credited to ‘K[uhl] & V[an] H[asselt]’, designating new species discovered
in Java by Boie and that Kuhl & Van Hasselt had planned to describe, but which were still undescribed in
publications, were nomina nuda in 1826 and therefore cannot be considered as potential type species, but the
other five constitute the originally included nominal species, among which a type species may be selected.
The nominal species at stake are currently referred to two distinct families. Three of them are now members
of the DENDROBATIDAE: [S03] Hyla trivittata Spix, 1824, now a member of the genus Ameerega Bauer, 1986
(see Dubois et al. 2016); [S04] Hyla nigerrima Spix, 1824, now considered a junior invalid synonym of [S03]
since the ‘First Reviser’ action of Peters (1872: 226); and [S05] Rana tinctoria Cuvier, 1797, now a member
of the genus Dendrobates Wagler, 1830. The other two are now members of the HYLIDAE: [S06] Calamita
punctatus Schneider, 1799, referred here (see [C34b] below) to the genus Boana Gray, 1825; and [S07] Hyla
luteola Wied-Neuwied, 1824, now a member of the genus Phyllodytes Wagler, 1830 (see Duellman 1977:
157).
No explanation was provided by Schlegel (1826a) for the origin of the word Hysaplesia, or can be
guessed nowadays, and this nomen must therefore be considered as an ‘arbitrary combination of letters’
(Article 11.3). As it was combined in the original publication with seven epithets in the feminine form, this
generic nomen is of feminine grammatical gender by virtue of Article 30.2.3.
[C03] Schlegel (1826b)
A very similar paper in German by Schlegel in Isis von Oken was apparently published in 1826 and not 1827
(as had long been believed) according to Savage et al. (2007: 256) who credited it with the default date 31
December 1826. In this text, Schlegel (1826b: 294) mentioned the same nominal species in the same order,
but here the nomen of the genus was spelt Hylaplesia. No explanation was given for this difference, but it
indeed makes sense: the word so spelt would correspond to an accretion of the generic nomen Hyla Laurenti,
1768 with the Greek word πλησιός (plesios), ‘near, close by’, thus meaning ‘close to Hyla’. This would be
logical as this genus was erected for five species previously referred to the genus Hyla. It is thus very likely
that the spelling used in the original manuscript of Boie was Hylaplesia and that the spelling Hysaplesia was
a misprint, that was ‘corrected’ in Schlegel (1826b). In fact it was probably not really a ‘correction’, as both
manuscripts may well have been submitted to both journals at approximately the same date, the misprint
Hysaplesia, possibly due to the printer of the Bulletin des Sciences naturelles et de Géologie and not to
Schlegel, creeping in one of them only, which happened to be published first by pure chance. However, in
zoological nomenclature, a one-letter difference between the spellings of two generic ‘names’ is enough to
consider them as two distinct nomina (Article 56.2). Because of the chronology of publication of the two
papers, from a nomenclatural viewpoint it would be quite logical to consider Schlegel’s spelling Hylaplesia as
an emendation, i.e. a particular case (autoneonym) of new replacement nomen or neonym (see Dubois
2010a), of Hysaplesia, or even the latter as an ‘incorrect original spelling’. However, formally Hysaplesia
does not meet the requirements of the current Article 32.5.1 for being considered as an incorrect original
spelling and Hylaplesia does not meet those of Article 33.2.1 for being interpreted as an intentionally
modified spelling or emendation. The latter Article is quite restrictive as it recognises only three situations for
such an interpretation: either an explicit statement of intention, or the mention of both the original and
changed spelling, or two or more nomina in the same work being treated in a similar way. But, especially
when ancient works are concerned, there can be other kinds of evidence for the fact that a new spelling was
proposed intentionally to replace the original nomen which, for some reason, was considered incorrect by an
author. I discussed this question elsewhere in details on two occasions (Dubois 1987b: 35–38; 2010a: 10–13,
29–31; see Appendix 1 below) and I suggested possible improvements of the current wording of Article
33.2.1 to include other possibilities. In my opinion, such possibilities should be recognised in the Code, at
least before a date to be fixed (e.g. 1930). However, under a strict adherence to the current Article 33.2.1,
Hylaplesia in Schlegel (1826b) must be treated as an incorrect subsequent spelling. As we will see, the
difference between the two interpretations, [I1] unjustified emendation (autoneonym) and [I2] incorrect
subsequent spelling (ameletograph), is not trivial and has drastic nomenclatural consequences.
[C04] F. Boie (1828)
Heinrich Boie’s brother Friedrich Boie (1828: 363), in a footnote, mentioned the genus Hylaplesia, which he
placed in the family HYLADAE, for which he provided a diagnosis and to which he referred only five nominal
species among the seven listed in Schlegel (1826a–b), i.e. [S01] to [S05] above.
The status of this nomen is open to discussion. Despite the fact that in this long paper he provided in
footnotes a number of references to previous works, including several in Isis von Oken (F. Boie 1822, 1826,
1827; Fitzinger 1827), he mentioned nowhere the two papers by Schlegel (1826a–b), and he presented the
genus Hylaplesia as a new one, described by his brother in the unpublished manuscript of the Erpétologie de
Java. This statement supports the assumption expressed above that Hylaplesia was most probably the original
spelling of the new genus in his brother’s book. It is therefore not logical to consider that he used an ‘incorrect
subsequent spelling’ of a nomen published by Schlegel in a French journal for which there exists no evidence
that he ever saw it. He must rather be considered to have established independently a new genus, on the basis
of the same unpublished manuscript of H. Boie used by Schlegel, but with a slightly different list of included
species. The status of this new nomen then depends on the status afforded to Hylaplesia in Schlegel (1826b).
If, under [I1], Hylaplesia H. Boie in Schlegel, 1826b is considered an intentionally changed spelling (then an
autoneonym) of Hysaplesia H. Boie in Schlegel, 1826a, then the nomen Hylaplesia H. Boie in F. Boie, 1828
is a junior homonym of the former, therefore invalid for this mere reason. But if, under the strict application of
the Code [I2], Hylaplesia in Schlegel (1826b) is considered simply as an incorrect subsequent spelling of
Hysaplesia H. Boie in Schlegel, 1826a, it has no separate nomenclatural existence and therefore does not
preoccupy the spelling Hylaplesia, which is then not a junior homonym in F. Boie (1828). In both cases
however, Hylaplesia H. Boie in F. Boie, 1828 is a distinct, independent nomen, having its own auctor and
date, but not necessarily the same type species as Hysaplesia.
[C05] Wagler (1830)
Wagler (1830: 202) proposed the nomen Dendrobates, clearly as a neonym (nomen novum) for “Hylaplesia
H. Boie Isis 1826b”. Therefore, just like Hyloplesia Agassiz, 1846 and Dendromedusa Gistel, 1848 (see
below under [C11] and [C12]), it must now be considered a neonym of Hylaplesia H. Boie in Schlegel,
1826b under [I1] and of Hysaplesia H. Boie in Schlegel, 1826a under [I2]. He gave a Greek etymology for
this generic nomen: “Δενδρος arbor, et βαινω incedo”, i.e. ‘tree’ and ‘I walk’. In classical Greek, the ending
-βάτης (-bates), derived from the verb βαίνω, is of masculine grammatical gender—e.g. in the word
‘ιπποβάτης (hippobates), ‘horseman’—, so this also applies to the generic nomen Dendrobates and all other
generic nomina having the same ending.
Wagler cited three nominal species, originally included in Hysaplesia, as belonging to this genus
Dendrobates: [S03] Hyla trivittata Spix, 1824, [S04] Hyla nigerrima Spix, 1824 and [S05] Rana tinctoria
Cuvier, 1797. He expressly mentioned that [S01] “Hylaplesia borbonica” H. Boie in Schlegel, 1826a and
[S02] “Hylaplesia achatina” H. Boie in Schlegel, 1826a were unknown to him. He removed the species [S07]
Hyla luteola Wied-Neuwied, 1824 from this genus and erected for it the genus Phyllodytes, of which it is
therefore the type species by original monophory (monotypy). As for the species [S06] Calamita punctatus
Schneider, 1799, he did not cite it in his book, so that one may presume that he did not have an opinion on its
validity and its generic allocation.
[C06] Bonaparte (1831)
Within the genus Hyla Laurenti, 1768, Bonaparte (1831: 76) erected a subgenus Eubaphus for the single
species “Rana tinctoria, Shaw”, i.e. Rana tinctoria Cuvier, 1797. This nomen, which was ignored by all
authors until Dubois (1982b: 272), is an invalid junior isonym (objective synonym) of Hysaplesia, Hylaplesia
(in Schlegel 1826b) and Dendrobates following Duméril & Bibron’s (1841) designation discussed below
under [C09].
[C07] Tschudi (1838)
Tschudi (1838: 27, 70) did not adopt the nomen Dendrobates and used the nomen Hylaplesia as published by
Schlegel (1826b) for a genus in which he included three nominal species: Hylaplesia borbonica Tschudi,
1838, a nomen to which he provided nomenclatural availability as [S08] through giving a diagnosis and
referring to Boie’s specimens in the Leiden Museum, now a member of the genus Leptophryne Fitzinger,
1843 (BUFONIDAE) [not MICROHYLIDAE as stated by Savage et al. 2007: 256]; “Hylaplesia tinctoria Boje”, i.e.
[S05] Rana tinctoria Cuvier, 1797, of which he considered that [S03] and [S04] were synonyms; and [S09]
“Hylaplesia picta Bibr. Mus. Par.”, i.e. a new nomen based on a specimen so named by Bibron in the Paris
Museum and to which he provided nomenclatural availability while becoming its nomenclatural auctor
[contrary to the statement in ASW which credits this nomen to “Bibron in Tschudi, 1838”, although the brief
diagnosis on page 28 which makes it available was clearly written by Tschudi himself], a species currently
referred to the genus Ameerega. He did not designate a type species for Hylaplesia. He also provided
nomenclatural availability to the nomen “Hylaplesia achatina Boje” [S10] through providing on page 71
descriptive notes on the species he called Microhyla achatina, which was the only species of his new genus
Microhyla (now referred to the MICROHYLIDAE), of which it is therefore the type species by original
monophory (monotypy).
[C08] Bibron in de la Sagra (1840)
Bibron (in de la Sagra, 1840: pl. 29) erected a new genus Phyllobates for the single species Phyllobates
bicolor [S11], stated to be from Cuba, which is therefore the type species of this genus by original monophory
(monotypy).
[C09] Duméril & Bibron (1841)
Duméril & Bibron (1841: 649) adopted the generic nomen Dendrobates Wagler, 1830 for a genus including
three species: Dendrobates tinctorius, i.e. the species [S05] above, of which they considered with doubt [S03]
Hyla trivittata Spix, 1824 as a synonym; Dendrobates pictus, i.e. [S09] Hylaplesia picta Tschudi, 1838; and
the new species [S12] Dendrobates obscurus, currently considered a synonym of [S03]. Most importantly, on
page 651, they clearly designated “la Hyla tinctoria de Daudin”, i.e. the nominal species Rana tinctoria
Cuvier, 1797, as “l’espèce type de notre genre Dendrobate”, i.e., the type species of Dendrobates. As
Dendrobates is an alloneonym for “Hylaplesia” and therefore for Hysaplesia of which the latter is either an
autoneonym (unjustified emendation) [I1] or an ameletograph (incorrect subsequent spelling) [I2], and as the
species Rana tinctoria was part of the originally included nominal species of Hysaplesia, this designation is
valid and applies to these two (or three) nominal taxa, according to the following Article of the Code:
“67.8. Type species of nominal genus-group taxa denoted by new replacement names (nomina nova). If
an author publishes a new genus-group name expressly as a new replacement name (nomen novum) for a
previously established name, or replaces a previously established genus-group name by an unjustified emendation
(…), both the prior nominal taxon and its replacement have the same type species, and type fixation for either
applies also to the other, despite any statement to the contrary (…).
67.8.1. The type species must be a nominal species eligible (…) for fixation as the type species of the prior
nominal genus-group taxon.”
[C10] Fitzinger (1843)
[C10a] Fitzinger (1843: 31) explicitly designated “Hylaplesia achatina Boie” as type species of “Hylaplesia
Boie”. As this nominal species was a nomen nudum, it is not available for type fixation of this genus and this
designation would be invalid even if it was not so for being subsequent to that of Duméril & Bibron (1841)
mentioned above.
[C10b] Not realising that Dendrobates had been proposed to replace Hylaplesia, in the same work Fitzinger
(1843: 32) designated “Dendrobates nigerrimus Wagler”, i.e. Hyla nigerrima Spix, 1824 [S04] as type
species of Dendrobates. As this nominal species was one of the originally included species of Hysaplesia, this
designation would be valid, and the nomen Dendrobates would have to replace the nomen Ameerega, if it was
not invalid for being subsequent to that of Duméril & Bibron (1841).
[C10c] Finally, Fitzinger (1843: 32) erected a family PHYLLOBATAE for four genera including Dendrobates
Wagler, 1830 and Phyllobates Bibron in de la Sagra, 1840. The latter is therefore its type genus by ‘implicit
etymological designation’ (see Dubois 1984: 24).
[C11] Agassiz (1845, 1846)
[C11a] Louis Agassiz published many lists of supraspecific zoological nomina, in which he often emended
their spellings in order to respect better their classical etymology in Greek and Latin. In 1845, on page 23 he
wrote: “Hylaplesia Boie Isis, 1827. Hyla; πλησιος, vicinus”. On page 24, he did not mention the spelling
Hysaplesia. In 1846, on page 189 of his ‘Index Universalis’, he wrote: “Hylaplesia Boie Rept. 1827. (Scr.
Hyloplesia)”. And on page 190: “Hyloplesia – (V. Hylaplesia Boie) Rept”. On page 192, he again did not
mention the spelling Hysaplesia, which he most likely considered an incorrect original spelling published by
inadvertence and having no nomenclatural existence, a logical interpretation (see above). Thus, in 1846, he
clearly provided an emendation Hyloplesia of H. Boie in Schlegel’s (1826b) Hylaplesia, which he considered
the original spelling of the nomen but which he interpreted as being ill-formed. Under interpretation [I1]
above, he indeed provided an emendation of the nomen Hylaplesia H. Boie in Schlegel, 1826b, but under
interpretation [I2] he in fact published an emendation of Hysaplesia H. Boie in Schlegel, 1826a, as in this case
the nomen Hylaplesia has no nomenclatural existence. In both cases, the nomen Hyloplesia is an ‘unjustified
emendation’ under the current Code, i.e. an autoneonym, having its own auctor and date (Agassiz, 1846).
Agassiz (1845, 1846) did not mention the type species of this genus.
[C11b] In the same works, Agassiz (1845: 14; 1846: 118) mentioned the generic nomen Dendrobates Wagler,
1830 with its etymology but without comments, i.e. without stating that it had been proposed to replace
Hylaplesia and without mentioning its type species.
[C12] Gistel (1848)
Gistel (1848: xi) published the neonym Dendromedusa for Hylaplesia of H. Boie in Schlegel (1826b). This
nomen was ignored by all subsequent authors (including Nieden 1923 and Duellman 1977) until Dubois
(1987b: 46).
[C13] Bonaparte (1850)
Bonaparte (1850) erected a family EUBAPHIDAE and a subfamily EUBAPHINA based on the type genus
Eubaphus Bonaparte, 1831 (an invalid junior isonym of Dendrobates Wagler, 1830), by implicit etymological
designation. The existence of this nomen was ignored by all authors until Dubois (1982b: 273).
[C14] Günther (1858, 1859)
In contrast to Fitzinger, Günther had understood that Dendrobates was a nomen proposed to replace
Hylaplesia and logically used the latter, credited to “Boie, Isis, 1827, p. 194”, as valid (Günther 1859: 341).
He erected a ‘section’ HYLAPLESINA (Günther 1858: 345, 1859: xiv, 122) and a family HYLAPLESIDAE (Günther
1858: 341, 346, 1859: xiv, 124) based on this generic nomen. The ‘name’ HYLAPLESINA is mentioned in the
website ASW as a synonym of ANURA, with the following comment: “Coined as a Section, at the
suprafamilial or infraordinal level (unclear in context), to contain Hylaedactylidae, Brachymeridae,
Hylaplesidae”. This statement is misleading, as the nomina of all the ‘sections’ in Günther (1858) are based
on the stems of generic nomina referred as valid to the section and are nomenclaturally coordinated with the
nomina of one of their subordinate families, so that the rank ‘section’ in this work belongs without doubt in
the nomenclatural family-series and is clearly equivalent to our current rank superfamily. Both ‘names’
HYLAPLESIDAE and HYLAPLESINA are therefore parographs of a single nomen, having the same auctor, date and
type genus, and which should be collectively referred to under its highest rank, i.e. as HYLAPLESINA (Article
24.1). This nomen originally thus applied to the family now called DENDROBATIDAE.
[C15] Cope (1865)
Cope (1865: 104) adopted the generic nomen Dendrobates and did not even mention the nomina Hysaplesia
and Hylaplesia. He also ignored Günther’s (1858) nomen HYLAPLESINA and erected (p. 100, 104) a family
DENDROBATIDAE, thus still complicating an already complex nomenclatural situation.
[C16] Cope (1866)
Cope (1866: 130) erected the genus Colostethus for a single new species, [S13] Colostethus latinasus, which
he explicitly designated as type species of this genus. He referred this genus to a “Group I of the family
Ranidae” for which he proposed a diagnosis but no nomen.
[C17] Cope (1867)
[C17a] Cope (1867: 191, 197) erected the family COLOSTETHIDAE for the single genus Colostethus which is
therefore its type genus by implicit etymological designation.
[C17b] In the same work, Cope (1867: 197) again recognised the family DENDROBATIDAE and the genus
Dendrobates as valid. Unlike in 1865 [C15], he duly mentioned the nomen Hylaplesia as a synonym of
Dendrobates, but credited it without justification to “Gthr. Catal. (not of Boie)” and failed again to mention
the nomen HYLAPLESINA.
[C18] Mivart (1869)
Mivart (1869: 293) introduced the spelling Calostethus for the nomen Colostethus, which he also mentioned
(1869: 284, 285) under its original spelling. As he adopted Calostethus as the valid nomen of this genus, this
situation complies with one of the criteria of Article 33.2.1 for recognition of a modified spelling as being
intentional and deserving the status of ‘emendation’: “when both the original and the changed spelling are
cited and the latter is adopted in place of the former”. Therefore Calostethus Mivart, 1869 is an available
junior objective synonym of Colostethus Cope, 1866, and the nomen CALOSTETHINA which was also
introduced by Mivart (1869: 293) for a subfamily of the family POLYPEDATIDAE is also available as a junior
isonym (objective synonym) of COLOSTETHIDAE Cope, 1867.
[C19] Boulenger (1882)
[C19a] Although he recognised that Hylaplesia had priority over Dendrobates, and HYLAPLESIDAE priority
over DENDROBATIDAE, Boulenger (1882: 140, 142), who had little respect for nomenclatural priority, in both
cases adopted the junior nomen as valid, without explaining his reasoning for doing so. Since then, most
authors followed this choice. Silverstone (1971: 262) cited 15 uses of Dendrobates and Phyllobates in the
literature (without distinguishing them) and Dubois (1982b: 275) cited 25 uses of Dendrobates, 22 uses of
DENDROBATIDAE and 6 of DENDROBATINAE.
[C19b] Boulenger (1882: 140) still referred the genus Colostethus to the family RANIDAE, and (1882: 194)
included the genus Phyllobates in the family CYSTIGNATHIDAE. Both genera were later referred to the
DENDROBATINAE and/or the DENDROBATIDAE (see e.g. Griffiths 1959; Savage 1968; Lynch 1973; Lynch &
Ruíz-Carranza 1982; Silverstone 1975, 1976; Myers et al. 1978; Duellman & Trueb 1985) where they still
stand.
[C20] Lutz (1925)
Lutz (1925: 139) described two new species (nigriventris and flavopicta) referred to the genus Hylaplesia but
did not discuss the nomenclatural status of this generic nomen or its type species. The two species described
by Lutz (1925) are currently referred respectively to the genera Ischnocnema Reinhardt & Lütken, 1862,
placed by Frost et al. (2006) in the family BRACHYCEPHALIDAE, and Ameerega Bauer, 1986 of the family
DENDROBATIDAE (see Grant et al. 2006: 164—not ‘Frost et al. 2006 by implication’ as stated in ASW1).
[C21] Stejneger (1937)
[C21a] Stejneger (1937) was the first author to mention the spelling Hysaplesia, that he considered as an
incorrect original spelling of Hylaplesia.
[C21b] Ignoring the fact that Dendrobates was a neonym for Hylaplesia and the fact that Duméril & Bibron’s
(1841) designation of [S05] Rana tinctoria as type species for Dendrobates also applied to its archaeonym
Hylaplesia, Stejneger (1937) designated “Hyla punctata Daudin”, i.e. [S06] Calamita punctatus Schneider,
1799, as type species of Hylaplesia (as used by Schlegel 1826b). He explained as follows the reason for this
strange choice: “By this action Hylaplesia becomes a synonym of Hyla and will cease to be a source of future
confusion in herpetological nomenclature.” However this action, being posterior to that of Duméril & Bibron
(1841), was invalid. A few subsequent authors (e.g. Duellman 1977: 23) accepted this designation as valid,
but this nomen was never used as valid for a genus of the family HYLIDAE.
[C21c] Although in 1937 the species [S06] Calamita punctatus Schneider, 1799 was considered as a member
of the genus Hyla Laurenti, 1768 (see e.g. Nieden 1923: 307) and remained so for a long time (see e.g.
Duellman 1977: 89), this situation ended with the work of Faivovich et al. (2005), who placed this species,
along with many others, in the genus Hypsiboas Wagler, 1830. By doing so, they seemingly rejected
Stejneger’s (1937) designation as invalid, because otherwise they should have used the nomen Hysaplesia, or
Hylaplesia, dated 1826, for this genus.
[C22] Silverstone (1971)
In an application to the Commission opening a new Case Z.N.(S.)1930, Silverstone (1971) ignored the fact
that Dendrobates was a neonym for Hylaplesia, as well as Duméril & Bibron’s (1841) designation of [S05]
Rana tinctoria as type species for Dendrobates, and he accepted Fitzinger’s (1843) invalid designation of
[S04] Hyla nigerrima as type species for this genus. In order to maintain the prevailing usage, he
consequently requested the Commission to invalidate Fitzinger’s designation and to replace it by that of
“Calamita tinctoria Schneider, 1799”, an unnecessary requirement for being redundant with the action of
Duméril & Bibron (1841).
[C23] Myers & Daly (1971), Cuellar et al. (1972), Peters et al. (1972)
While discussing some aspects of Silverstone’s (1971) proposal, the three contributions quoted above,
published in the BZN under Case Z.N.(S.)1930, followed this author in ignoring the fact that Dendrobates was
a neonym as well as Duméril & Bibron’s (1841) designation of [S05] Rana tinctoria as type species for this
nominal genus, and in crediting Schneider (1799) with the authorship of the nomen of this species as
1. The concept of nomenclatural act ‘by implication’ does not exist in the Code. It appeared first in the book of Frost (1985) and
was used repeatedly in publications involving this author (e.g., Faivovich et al. 2005, Frost et al. 2006) as well as on the site ASW,
despite warnings against the use of this unclear formula which is meaningless in taxonomy and nomenclature (Dubois 1987a: 141;
Ohler et al. 2015: 596).
“Calamita tinctoria”. No further comments on Silverstone’s (1971) application was published in the BZN
before 1982, and the Commission never voted on this application.
[C24] Duellman (1977)
Duellman (1977: 23) adopted as valid the designation of Calamita punctatus as type species of “Hylaplesia
Boie (non Schlegel, 1826 = Anura)” (!) and therefore included the latter nomen in the synonymy of Hyla.
[C25] Lescure (1982)
In a new contribution concerning Case Z.N.(S.)1930, Lescure (1982) pointed to Duméril & Bibron’s (1841)
very clear designation of the species [S05] as type species of Dendrobates before Fitzinger’s (1843)
designation, making the latter invalid, and to the precedence of the nomen Rana tinctoria Cuvier, 1797 over
all other usages of this nomen, including “Calamita tinctorius Schneider”, “Rana tinctoria, Shaw” and “Hyla
tinctoria Daudin”, which are just new combinations and therefore aponyms of Cuvier’s nomen, devoid of
independent nomenclatural status [contrary to the statement which still currently appears in the website ASW
regarding the former].
[C26] Dubois (1982b)
In a second contribution concerning Case Z.N.(S.)1930 published in the same issue of the BZN (Dubois
1982b), I summarised most of the facts detailed above and, in order to maintain nomenclatural stability, I
proposed to stabilise the nomen Dendrobates as the valid one for the genus earlier named Hysaplesia and
Hylaplesia, and DENDROBATIDAE as the valid one for the family earlier named PHYLLOBATAE, EUBAPHIDAE and
HYLAPLESIDAE. For this purpose, concerning the generic nomina, I proposed the Commission to use its
Plenary Powers to invalidate the nomina Hysaplesia and Hylaplesia. Concerning the family nomina, the
situation could be solved in part without having to require the use of the Plenary Powers. As for the priority of
DENDROBATIDAE (1865) over EUBAPHIDAE (1850), I proposed to establish it through the use of Article 40.a of
the 1985 Code then in force (now Article 40.2 of the 1999 Code) which stated that in case of a replacement of
a family nomen by a junior synonym before 1961, and of this replacement having “won general acceptance”,
this replacement was to be maintained but then the junior nomen should take the date of the senior one and be
afforded precedence over it. This proposal was a bit borderline, as in fact EUBAPHIDAE had never been
‘replaced’ by DENDROBATIDAE, but simply ignored by all authors, but this ‘trick’ allowed precedence to be
given to DENDROBATIDAE, then ‘re-dated’ 1850 instead of 1865, not only over EUBAPHIDAE (1850) but also
over HYLAPLESIDAE (1858), even before the proposed invalidation of Hylaplesia, without having to ask the
Commission to use its Plenary Powers for this purpose. This proposal left only one problem which could be
solved only by the Commission, that of the priority of PHYLLOBATAE (1843) over DENDROBATIDAE (1850), for
which the use of the Plenary Powers by the Commission was requested. If adopted then by the Commission,
this complete proposal would have solved all the nomenclatural problems discussed above and below, one of
which remains still unsolved today (see below under [C38]).
[C27] Dubois (1983a, 1984)
In the first comprehensive checklists of anuran amphibian family-series nomenclature ever published (Dubois
1983a, 1984), I used the nomenclature proposed in Dubois (1982b) and I briefly summarised the problems
discussed above.
[C28] Holthuis (1983)
Holthuis (1983), then a member of the Commission, was the only author to comment on Dubois’ (1982b) text
in the BZN, still under Case Z.N.(S.)1930. He denied that Dendrobates was a replacement nomen for
Hylaplesia, for two reasons: the fact that Wagler (1830) had not expressly stated that it was a replacement
name and the fact that he had not mentioned as members of Dendrobates all the nominal species cited by
Schlegel (1826a) as members of Hysaplesia, using fictitious examples to support his arguments—which are
both refuted below under [C37d]. He also discussed the question of the authorship of the nomen Hysaplesia,
of which he considered Hylaplesia to be an incorrect subsequent spelling.
I immediately (Dubois 1983b) replied to the comments of Holthuis (1983) in the BZN under Case
Z.N.(S.)1930. I showed that the replacement of Hylaplesia by Dendrobates was only one among several
similar cases of replacement nomina in Wagler (1830), and I commented on the authorship of Hysaplesia and
on the status (emendation or incorrect subsequent spelling) of Hylaplesia relative to the latter (see [C03]
above). Neither Holthuis nor any other zoologist published any answer to my 1983 paper in the BZN, but
Holthuis’ (1983) opinion alone seems to have been efficient to ‘block’ any action of the Commission
regarding this Case, as my 1982b contribution was never mentioned in any further document published before
2007 by the Commission and Case Z.N.(S.)1930 was never voted upon, despite my repeated requests to the
Commission’s secretariat.
[C30] Frost (1985)
Frost (1985) made an extensive use of my checklists (Dubois 1983a, 1984) for all the anuran amphibian
family-series nomenclature and followed my proposal (Dubois 1982b) regarding the valid nomen of the
family DENDROBATIDAE through an action of the Commission giving precedence to this nomen over
PHYLLOBATAE, but failed to mention that, for this proposal to be complete and efficient, the nomen
DENDROBATIDAE should be afforded the date 1850 instead of 1865.
[C31] Savage (1986)
In a very severe but ill-inspired criticism of Dubois (1984), Savage (1986), then another member of the
Commission, adopted the interpretation of Holthuis (1983) but did not cite it, which avoided him to have to
cite also Dubois’ (1983b) rebuttal and to have to reply to the arguments it contained. In this paper, Savage
(1986) presented his own analysis of the distinction between new replacement name (nomen novum),
emendation and incorrect subsequent spelling.
I replied in detail to Savage’s (1986) criticism, providing again a refutation of his arguments repeated from
Holthuis’ (1983) concerning the status of Dendrobates vs. Hylaplesia. My paper (Dubois 1987b) included a
detailed rebuttal of the analysis of Savage (1986) regarding the distinction between nomen novum,
emendation and incorrect subsequent spelling. I submitted this reply to the journal Copeia, which had
published Savage’s (1986) paper but denied me the right to reply to a criticism published in its columns, so
that I had to publish it in another journal. After publication I sent a copy of this work to the secretariat of the
Commission in London where, several years later, I had the opportunity to check that it had been properly
archived. Nevertheless, subsequently this paper was never cited, let alone refuted, by any of the authors who
again published on the Dendrobates and DENDROBATIDAE controversies, and was ignored by the Commission
when it finally addressed this case. In order to call again attention on this ‘invisible’ paper, I reproduce in
Appendix 1 below a copy of some parts of it which deal with the distinction between neonym and incorrect
subsequent spelling, and with Wagler’s neonyms. These questions were again discussed later from a more
general viewpoint in Dubois (2010a).
[C33] Duellman (2001)
Duellman (2001: 173, 776) accepted the designation of Calamita punctatus as type species of Hylaplesia
(credited to H. Boie in Schlegel, 1826b) and therefore logically concluded that the latter was a junior
synonym of Boana Gray, 1825 (see [C34b] below).
[C34] Faivovich et al. (2005)
[C34a] Faivovich et al. (2005: 86) disagreed with this statement of Duellman (2001). They accepted Dubois’s
(1982b) statement that Dendrobates was a replacement nomen of Hylaplesia (credited to H. Boie in Schlegel,
1826b) and therefore considered both nomina as objective synonyms, but did not discuss further the
nomenclatural implications of this fact.
[C34b] Until the end of the 20th century, the genus Hyla, just like the genera Rana, Bufo and
Eleutherodactylus (see Dubois 1977), was very heterogeneous and counted hundreds of species: for example,
in the checklist of Duellman (1977), the genus Hyla Laurenti, 1768 was credited with 30 synonyms and 270
species.
Faivovich et al. (2005) proposed a new phylogeny and taxonomy of the family HYLIDAE, which resulted
in dismantling the genus Hyla to recognise instead 15 distinct genera, with 353 species altogether. For one of
these genera, distributed in Central and South America, they ‘resurrected’ the nomen Hypsiboas Wagler,
1830, and they considered the nomen Auletris Wagler, 1830, published in the same work, as its invalid
synonym. These authors did not state on which ground they established this precedence, and this question will
be addressed elsewhere, but it is not central to the present discussion and is ignored here. Anyway, it turns out
that neither the nomen Hypsiboas nor the nomen Auletris can be the valid one for this genus, which requires
discussion.
The Code provides Rules for the nomenclatural availability (and hence potential validity) of nomina first
published as invalid synonyms of nomina then treated as valid. Article 11.6 states that “A name which when
first published in an available work was treated as a junior synonym of a name then used as valid is not
thereby made available”, but then Article 11.6.1 adds: “However, if such a name published as a junior
synonym had been treated before 1961 as an available name and either adopted as the name of a taxon or
treated as a senior homonym, it is made available thereby but dates from its first publication as a synonym
(...)”. In amphibians, a famous example of this situation is the nomen Tomopterna Duméril & Bibron, 1841,
which was first published as a synonym of Pyxicephalus Tschudi, 1838 but then validated by its use as a valid
generic nomen by Günther (1859) and subsequent authors (Dubois 1981: 251).
Faivovich et al. (2005) suggested that the availability of the generic nomen Boana Gray, 1825 had also to
be established through Article 11.6, but this statement is wrong.
These authors recognised that the generic nomen Boana Gray, 1825, which had traditionally been cited in
the synonymic list of Hyla Laurenti, 1768, was a synonym of Hypsiboas, but they rejected it as invalid, for the
following reason: “Gray (1825) suggested the name Boana as a synonym of Hyla, and Stejneger (1907)
subsequently designated Hyla boans as its type species. Unfortunately, as far as we can determine from
literature research, the name Boana has never been used in combination with an active species name,
therefore failing to fulfill the criterion established by article 11.6.1 (ICZN, 1999) for the availability of names
originally proposed as synonyms.” However, the main statement in this sentence is wrong: Boana was not
proposed as a synonym but as a subgenus of Hyla, and therefore its availability does not depend on Article
11.6.1.
The entry of the genus Hyla in Gray (1825: 213–214) reads as follows:
“HYLA, Laur. Calamita, Schneid.

Body slender; skin mostly smooth; toes all dilated at the end, the fourth one of the hind feet, of a moderate
length.
*H. tinctoria; Laur, Rana tinctoria, Shaw. **C. intermixtus, Merrem. ***Calamita, hind feet semipalmate.
H. arboreus, Schneid. Rana arboreus; Lin. ****Boana, Gray. Granulated feet palmate, β maxima. Rana Boans,
Lin.”
In order to interpret correctly these lines, it is necessary to compare them with the whole of Gray’s (1825)
paper. It is then clear that the same conventions of presentation were used very consistently throughout this
work for similar information. Thus, numbers (I, II, etc.; 1, 2, etc.), symbols (*, **, etc.; †, ††, etc.) or
combinations of both (§ 1, § 11, etc.; § 1, § 2, etc.) introduced different taxa subordinate to the same taxon,
whereas ‘synonyms’ (or in fact in some cases aponyms) were just presented one after another, without any
intermediate sign. Therefore, in the first line above, “HYLA” and “Calamita” are synonyms, just like “H.
tinctoria” and “Rana tinctoria” or “H. arboreus” and “Rana arboreus” below. In contrast, in the third
paragraph, “H. tinctoria”, “C. intermixtus”, “Calamita” and “Boana” designate four distinct taxa subordinate
to the genus Hyla. For each of the former two taxa, Gray mentioned only the nomen of one included species,
but for the latter two, he proposed nomina of supraspecific taxa: one (Calamita) borrowed from Schneider
(1799) and one (Boana) introduced as new in this work. The rank of these taxa is clearly subgenus, a term
which was not in common use then but which is mentioned on page 197 (under Agama) of Gray’s paper. The
situation of these two subgenera is similar to several other ones in the same work, e.g., the subgenera
Caudiuerbera in Uroplates, Coronella and Homalopsis in Coluber, Ibiba in Hurria, Rapara in Emys or
Oxyrhychus in Bufo. It is thus clear that the new nomen Boana was introduced by Gray (1825) not as a
synonym of Hyla but as the nomen of a subgenus of the latter genus. This nomen, which has never been
‘forgotten’ by taxonomists but appeared in all detailed synonymic lists of the genus Hyla since Stejneger
(1907: 76), is therefore fully available and cannot be rejected for the reason invoked by Faivovich et al.
(2005) and repeated in ASW and by Duellman et al. (2016).
Concerning the type species of Boana, Gray (1825: 214) associated this nomen with two specific nomina,
“Rana Boans, Lin.”, and “β maxima”. The latter doubtless refers to the nomen Rana maxima Laurenti, 1768,
currently considered a subjective synonym of Rana boans Linnaeus, 1758 (see above). Stejneger (1907: 76)
designated the latter as type species of Boana, which is therefore a strict objective synonym of Auletris and a
subjective synonym of Hypsiboas and should replace the latter as the valid nomen of this South American
hylid genus. This had already been suggested by Duellman (2001), Savage (2002) and originally by Wiens et
al. (2005), although none of these authors used indeed the new combinations resulting from the adoption of
this genus as distinct from Hyla, and although the latter stated on page 807 that they had changed their mind
after the publication of Faivovich et al.’s (2005) work. The statement of ASW according to which some of
these combinations were created ‘by implication’ in the work of Wiens et al. (2005) is unsubstantiated2,
because these authors used everywhere the combinations based on the generic nomen Hyla, as if they
considered Boana as a subgenus of the latter.
2. See note 1 above.

[C35] Frost et al. (2006)
Frost et al. (2006: 212), following the same reasoning concerning the status of Hylaplesia (and Hysaplesia),
rightly considered HYLAPLESIDAE as a synonym of DENDROBATIDAE. They accepted the latter nomen as valid
for the family but failed to mention that, in order for this to be valid, this nomen must be dated 1850 and not
1865 (see [C26]).
[C36] Grant et al. (2006)
[C36a] In the same year and in the same journal, Grant et al. (2006) published a revision of the family
DENDROBATIDAE. Although both papers [C35] and [C36] shared three authors (Frost, Haddad and Wheeler)
and were both published within a few months, in the second one they introduced some modifications in their
nomenclatural interpretations without explaining the reasons for this discrepancy. On page 173, they accepted
the designation by Stejneger (1937) of Calamita punctatus as type species of Hysaplesia (and Hylaplesia) as
valid—but nevertheless maintained the two latter nomina in the synonymy of Dendrobates, which is
inconsistent. But on page 210, they stated that the type species of Hysaplesia was Rana tinctoria, seeming to
imply that they accepted it as a neonym for Dendrobates. There is no clue to an explanation of this
contradiction in this paper, but the fact that on page 156 they placed the nomen HYLAPLESIDAE in the
synonymy of DENDROBATOIDEA suggests that the latter solution had perhaps their final preference.
[C36b] Grant et al. (2006: 168) erected a new subfamily HYLOXALINAE of the DENDROBATIDAE for the single
genus Hyloxalus Jiménez de la Espada, 1870. The nomenclatural availability of this nomen HYLOXALINAE is
questionable, for two different reasons.
First, Grant et al. (2006: 168) did not explicitly designate Hyloxalus as ‘type genus’ of this subfamily,
although they mentioned the ‘type genus’ of all the other family-series nomina cited in their work. Yet,
Article 16.2 of the Code requires that, for a new family-series nomen published after 1999 to be available, its
‘type genus’ must be cited. Grant et al. (2006: 168) stated that this subfamily included only one genus,
Hyloxalus, but added: “Although this name is currently redundant with Hyloxalus, we anticipate that the
available names Cryptophyllobates and Phyllodromus will be resurrected in the near future, making
Hyloxalinae an informative name (for species groups, see Comments for Hyloxalus, below). Moreover,
recognition of Hyloxalinae is necessitated by the recognition of Dendrobatinae for the five genera of brightly
colored and highly toxic species.” It is therefore quite clear from these statements that Hyloxalus, being at the
time of publication of this paper the only genus recognised as valid in the subfamily, was its ‘type genus’. In
fact, Article 16.2 of the Code is strangely written because, although its title includes the statement ‘type genus
to be cited’, the Article itself nowhere states explicitly that this genus nomen must be cited as the type genus
of the new nominal taxon. It writes: “In addition to satisfying the provisions of Articles 13–15, a new family-
group name published after 1999 must be accompanied by citation of the name of the type genus (i.e. the
name from which the family-group name was formed).” Grant et al. (2006) can therefore be argued to comply
with the ‘mild’ conditions of this Article, as they cited the nomen Hyloxalus although they did not state in full
words that it was the type genus of the new subfamilial nomen. This generic nomen can therefore be
considered to have been designated as type genus of this nominal subfamily by ‘implicit etymological
designation’, as most family-series nomina before 2000. This would not be the case if Article 16.2 was clearer
and stated, in parallel with the formulation of Article 13.3, that the new family-series nomen must “be
accompanied by the explicit fixation of the type genus in the original publication”.
The second problem comes from the absence in this work of “a description or definition that states in
words characters that are purported to differentiate the taxon”, as required by Article 13.1.1 for any new
nomen published after 1930. In this respect, Grant et al. (2006: 168) wrote: “CHARACTERIZATION, DIAGNOSIS,
AND SUPPORT: Branch length = 52. Bremer support = 35. All unambiguously optimized synapomorphies for
this clade are from DNA sequences (see appendix 8).” A ‘branch length’ or a ‘Bremer support’ do not qualify
as “a description or definition” stating diagnostic characters in words. They just provide information about the
reasons considered by the authors as sufficient for erecting a taxon for the organisms included in this
taxon—the diagnostic characters of which are not given. As the nomen HYLOXALINAE was considered by these
autors as ‘redundant’ with the nomen Hyloxalus (which in fact it is not, for reasons given e.g. in Dubois 2007:
48–50), one could expect that a diagnosis of this taxon would be given in the entry concerning the genus
Hyloxalus. This entry writes (p. 169): “CHARACTERIZATION, DIAGNOSIS, AND SUPPORT: As for Hyloxalinae,
above”, but this is followed by a list of 17 ‘other characteristics’ of the genus, which can be considered to
apply also to the subfamily as long as no other genus is formally recognised as valid in the latter. Finally,
Appendix 8 provides a list of “unambiguous DNA sequence transformations to diagnose the named clades
addressed in the monophyletic taxonomy presented above”, and such sequence transformations diagnostic of
Hyloxalus and HYLOXALINAE are given on pages 259–260. These sequence transformation data cannot be
considered to be “a description or definition that states in words characters that are purported to differentiate
the taxon”, but they provide some characterisation of this taxon. The combination of these molecular data
with the 17 ‘other characteristics’ of the genus Hyloxalus given above in the work can be considered to
provide nomenclatural availability to the nomen HYLOXALINAE, but admittedly this is a very borderline
situation, which could have been avoided e.g. by giving a ‘polythetic’ (Sneath 1962; Van Regenmortel 2016:
6) morphological diagnosis of the subfamily relatively to the other subfamilies of DENDROBATIDAE (see
examples in Dubois & Frétey 2016).
[C37] Savage et al. (2007)
[C37a] After a long period of silence, the BZN suddenly published a long paper (Savage et al. 2007)
‘resurrecting’ again this case and making new proposals. Quite interestingly, although Case Z.N.(1930),
opened by Silverstone (1971) and which had been the matter of seven comments published in the BZN from
1971 to 1983, had never been voted upon and therefore had not been closed, this new paper was registered as
opening a new Case numbered 3345. This had the effect of avoiding to have to compare in detail the new
proposal with the anterior contributions to Case Z.N.(S.)1930 when the ‘new’ Case was submitted to the
discussion and vote of the Commission, whose composition had largely changed compared to the 1971–1983
period. In particular, neither these authors nor the Commission’s members made any reference to the detailed
rebuttal of Savage’s (1986) ideas published by Dubois (1987b) although they could not have ignored them3.
[C37b] Regarding the respective status of Hysaplesia and Hylaplesia, Savage et al. (2007: 256) first stated
that Hylaplesia could be “considered an unjustified emendation” but “can also be interpreted as an incorrect
subsequent spelling” without choosing between both options, but below chose the second option without
further explanation except for a non-explicit reference to Article 33.5.
[C37c] Savage et al. (2007: 257) introduced the new spelling Hylapesia, which is clearly an incorrect
subsequent spelling of Hylaplesia, and therefore of Hysaplesia.
[C37d] Savage et al. (2007) repeated the arguments of Holthuis (2003) for considering Dendrobates as the
nomen of a new genus instead of a nomen novum for Hylaplesia. So doing, they did not mention and refute
the detailed arguments published against this interpretation (Dubois 1982b, 1983b, 1987b).
3. As stated above, this paper was archived in the Commission’s secretariat. As for the authors of the 2007 paper, the institutions
in which the first three of them worked were subscribers to the journal where it had appeared and had received the issue at stake
(personal communication of Thierry Frétey, General Secretary of ISSCA, the association publisher of the journal Alytes, 17 August
2016) and another paper in a preceding issue of this journal (Dubois 1987a) was cited in their paper. Furthermore, Darrel Frost, the
author of the website ASW, cited this 1987b paper in this website under the nomen Bombitator in the entry on Bombina, thus showing
that he had indeed seen it.
Their argumentation (following that of Holthuis 2003) presents an original misinterpretation of the
concept of nomen novum. About my statement that the nomen Dendrobates was a nomen novum for
Hylaplesia, they wrote (p. 257): “the fact that it [the genus Dendrobates in Wagler 1830] contained only three
of the species originally included in Hylaplesia does not support that notion. That Wagler did not
automatically include Hylaplesia borbonica, Hylaplesia achatina or Hyla luteola in Dendrobates indicates
that the latter is not a new replacement name but a new taxon based on a provisionally different concept.”
Beside the mysterious reference to the unclear notion of “provisionally different concept”, this interpretation
suggests that, in zoological nomenclature, a nomen is permanently attached to a taxonomic concept—i.e., that
it is defined either ‘intensionally’, as is the case in intensional nomenclatural systems like the Phylocode or
‘extensionally’ as it would be the case if it was attached to a fixed content of the taxon (for detailed
explanations, see e.g. Dubois 2005, 2006b)—which is completely wrong.
In zoological nomenclature, a nomen is not ‘defined’ taxonomically, but permanently attached by
‘ostension’ to its ‘name-bearing type’ or onomatophore (see e.g. Dubois 2005). In the case of generic nomina,
the onomatophores are nominal taxa, their ‘type species’. The taxonomic amendment of a taxon, through
inclusion or exclusion of other species in or from the taxonomic genus designated by this nomen, is a matter
of taxonomic interpretation and decision of any practicing taxonomist but does not belong in the field of
nomenclature and is not regulated by the Code. A particular case is that of a nominal genus established with a
set of ‘originally included nominal species’ among which none was designated as ‘type species’. As long as
this situation persists, there is no ‘type species’ but only a set of nominal species that are all liable to be
designated as such, a nomenclatural act which is valid only if this designation is explicit (Articles 67.2, 69.2).
Until this designation has been made, the mere fact of citing only one of the originally included species does
not result in a type species fixation (Article 69.4) and similarly that of citing only some of them as members of
the genus does not result in the elimination of the missing ones from the ‘originally included species’ or in the
erection of a new nominal genus (see also in this respect the entry 2 in the dichotomic key to different nomina
and spellings presented on pages 5–6 above under ‘Categories of new nomina’).
The establishment of a new generic replacement nomen is not a taxonomic but a purely nomenclatural
act. This act consists in the express proposal of “a new replacement name (nomen novum) for an available
name, whether or not required by any provision of the Code” (Articles 12.2.3 and 13.1.3). This definition only
refers to scientific names (nomina) but not in the least to taxa or their contents.
This can be easily illustrated in amphibians by two well-known examples. The nomen Triton Laurenti,
1768 was proposed for a genus that included 11 nominal species (gesneri, wursbaini, utinensis, alpestris,
carnifex, zeylanicus, palustris, cristatus, parisinus, americanus and salamandroides). Merrem (1820: 185)
proposed the new generic nomen Molge expressly as a nomen novum for Triton Laurenti, 1768 because the
latter was preoccupied by Triton Linnaeus, 1758. He mentioned 11 valid species in this genus (striata, rubra,
cinerea, punctata, palmata, wursbainii, alpestris, palustris, geitie, gigantea and tridactylus) and their
synonyms. Although the content of his genus Molge overlapped only very partially with that of Triton
Laurenti, 1768, no author has ever so far suggested to consider Molge as the nomen of a brand new genus
instead of a nomen novum for Triton. In the same works, Laurenti (1768: 35) established a genus Proteus with
three originally included species (raninus, tritonius and anguinus), only the third of which was retained by
Merrem (1820: 188) in this genus for which he proposed expressly the new replacement nomen Hypochthon.
This has not impeded all subsequent authors to recognise the latter as a nomen novum for the former. I think
no more examples are necessary to illustrate this fact well-known to all competent practicing taxonomists. It
is therefore quite surprising that the BZN, official organ of the Commission, accepted to publish Savage et
al.’s (2007) bizarre statement, and the fact that this did not raise any comment among the members of the
Commission or of the international community of zoologists (see below) is quite troubling. So far, the
international community has always accepted that neonyms (nomina nova) can be proposed to designate more
restrictive or more inclusive taxa than those designated by their archaeonyms (the nomina that they replaced).
If this had to change, this would mean that the Code would metamorphose from an ostensional to an
extensional nomenclatural system, a very drastic revolution indeed!
What is relevant to establish whether a modified spelling of an existing nomen is a nomen novum or not is
not the content of the taxa but the presence or absence of an evidence that the change of spelling (in the case
of unjustified emendation or autoneonym) or of nomen (in the case of nomen novum or alloneonym) was
intentional. As discussed already above under the items [I1] and [I2] and in Dubois (1982b, 2010a; see
Appendix 1), the current wording of the Code can be argued to be much too restrictive and a modification of
this Article may be considered desirable. But let us stick here to the current wording of this Article. Contrary
to the case of Hylaplesia vs. Hysaplesia, concerning Wagler’s nomen Dendrobates there may be no doubt that
it complies with the condition of ‘express proposal’ of Article 12.2.3 for being recognised as a nomen novum.
This was explained in detail in Dubois (1987b) in the text reproduced in part here in Appendix 1: Wagler
(1830: 17, footnote 2), following Linnaeus, stated clearly that generic nomina which do not have Greek or
Latin etymologies must be rejected. This is clearly the reason why he did not adopt Laurenti’s (1768) generic
nomen Hyla for the European treefrog Rana arborea Linnaeus, 1758 and proposed instead the generic nomen
Hyas Wagler, 1830 for this species. The same statement appeared also in Wagler’s manuscript later published
by Michahelles (1833) where he had rejected both nomina Hyla and Calamita for this genus and proposed for
it the nomen Discodactylus. Having clearly rejected Hyla for an explicit reason, Wagler could not accept
Hylaplesia as a valid nomen and was bound to replace it, which he did with the nomen Dendrobates. The fact
that he did not mention some of the original nominal species of this species is fully irrelevant in this respect.
This reasoning applies to several new nomina proposed by Wagler (1827, 1830), four of which are
nomina nova or alloneonyms (Asterodactylus for Pipa, Dendrobates for Hylaplesia, Enydrobius for Hylodes
and Systoma for Engystoma) and three unjustified emendations or autoneonyms (Calamites for Calamita,
Megalophrys for Megophrys and Bombitator for Bombinator). Strangely however, although all these couples
of nomina are in the same situation, the site ASW treats them differently (see below under [C39g]).
[C37e] Savage et al.’s (2007) interpretation of Dendrobates as the nomen of a new taxon and not as a neonym
for Hylaplesia had the consequence that these two nomina should be considered to have different type species
(respectively [S05] tinctoria and [S06] punctatus) and are therefore not synonyms. Their following statement
(p. 257) is then quite strange: “The usage of Hylapesia [sic] (Lutz, 1925, p. 139), the equivalent of
Hysaplesia, negates the possibility of applying Article 23.9.2 of the Code (the nomen oblitum option) for
suppression of Hysaplesia”. This sentence testifies to a double misunderstanding of Article 23.9. First, this
Article only applies in the case of synonymous or homonymous nomina, to maintain the usage of the junior
one, but it cannot be used in the case of nomina which are not synonyms or homonyms: so if Dendrobates is
not a synonym of Hysaplesia, as argued by these authors, it is irrelevant to refer to this Article here. Second,
the use of this Article does not in the least result in the ‘suppression’ of the senior nomen but in the reversal of
precedence between both nomina, which both remain available. In case an author is not satisfied with this
reversal of precedence and wishes to ‘suppress’ (invalidate) the senior one, an application for this purpose
should be submitted to the Commission (Recommendation 23A). These misinterpretations have escaped the
attention of the Commission’s secretariat who published Savage et al.’s (2007) paper in the BZN, and
apparently also to the members of the Commission, none of which commented on them (see below).
[C37f] Savage et al. (2007: 258) discussed the authorship and date of the nomina Phyllobates and Phyllobates
bicolor, which they credited to Bibron in ‘Sagra’ (1840) instead of Duméril & Bibron (1841) as most previous
authors.
[C37g] In conclusion, Savage et al. (2007: 258–259) supported Dubois’s (1982b) request to the Commission
to afford priority of DENDROBATIDAE Cope, 1865 (1850) over PHYLLOBATAE Fitzinger, 1843 whenever both
nomina are considered synonyms. They also supported the request to suppress the nomen Hysaplesia but did
not mention Hylaplesia and did not seize this opportunity to ask the Commission to use its Plenary Powers to
clarify the respective status of Hysaplesia vs. Hylaplesia and of the latter vs. Dendrobates. They just asked
the Commission to place Hysaplesia on the Official List of Rejected and Invalid Names in Zoology with the
type species Calamita punctatus Schneider, 1799 by subsequent designation of Stejneger (1937) as if this was
a proven fact, ignoring the controversy reminded above on this matter and the existence of an alternative
interpretation supported by strong and detailed arguments (Dubois 1982b, 1983b, 1987b). Furthermore, this
proposed action raised other problems, discussed below under [C38].
[C38] Opinion 2223 (2009)
[C38a] Unlike Case Z.N.(S.)1930, which was opened in 1971, discussed from 1971 to 1983 in seven papers,
and never closed, ‘Case 3345’ was opened in the December 2007 issue of the BZN and was hastily closed less
than one year later on 1 December 2008 by a vote of the Commission. During this period, no comment from
non-members of the Commission was published in the BZN. The proposals of Savage et al. (2007) were
adopted without modification, as Opinion 2223, by 17 positive votes, two negative ones, one abstention and
two absences of vote (Anonymous 2009).
This decision of the Commission was accompanied by the following comment:
“Bouchet, voting FOR, cautioned that the name of the work containing the description of Phyllobates
should be cited as de la Sagra, and not Sagra, as it was cited in the original application. He also mentioned, as
he has done elsewhere, that he dislikes conditional reversals of precedence, as they are a potential source of
nomenclatural instability. However, he voted in favour of the proposal as there was no alternative offered to
conserve the name DENDROBATIDAE. He also suggested that to give credit where it is due, it would have been
diplomatic for the authors to invite Dubois to join their application. Failing that, upon receipt of Case 3345,
the Secretariat might have considered reviving Dubois’s initial application of 1982, which was legitimate and
well-founded, though never went to vote due to a perceived shortcoming of one part by a single
Commissioner. Voting FOR, Brothers commented that the name Hylaplesia (incorrect subsequent spelling of
Hysaplesia) should probably also be placed on the Index (with the appropriate wording and attributions). Also
voting FOR, Štys pointed out that DENDROBATIDAE is one of the most intensely studied families of the
anurans. Consequently, he was unpleasantly surprised that no one in the batrachological community cared to
comment about its correct name.”
As these comments were brought to the knowledge of the Commission at the same time as the result of its
vote on this Case, they could not have any effect on the vote itself, which is a basic problem in the mode of
functioning of the Commission (see Dubois & Aescht 2016: 39, 42). The fact that the final decision of the
Commission does not mention at all the spelling or nomen Hylaplesia is indeed one of the defects of this
decision, but it is not the only one.
[C38b] As a matter of fact, in this Opinion the Commission used its Plenary Powers for two nomenclatural
acts only: the ‘suppression’ for the purposes of the priority among synonyms (but not among homonyms) of
the nomen Hysaplesia, and the conditional precedence given to the nomen DENDROBATIDAE Cope, 1865
(1850) over the nomen PHYLLOBATAE Fitzinger, 1843. The Plenary Powers were not used to modify the
nomenclatural status of the nomina Hysaplesia and Dendrobates or to clarify the status of the spelling or
nomen Hylaplesia, matters for which Savage et al.’s (2007) interpretations were taken for granted, despite
published evidence that they were wrong or could be considered to be so.
[C38c] In fact, given that the Commission did not take a stand on the fact that Dendrobates is, or not, a nomen
novum for Hysaplesia, the Code should have simply but fully applied for the interpretation of this case. In the
absence of an explicit action of the Commission in this respect, under a strict respect of the Code the detailed
evidence published in Dubois (1987b), which has never been shown to be wrong by anyone, still holds, and
Dendrobates is indeed a nomen novum for Hysaplesia. Consequently, both nomina are bound to have the
same type species. The fact of merely citing Stejneger’s (1937) designation of Calamita punctatus as type
species of this nominal genus, which is invalid for being subsequent to that of Duméril & Bibron (1841), as if
it was valid, is not enough to modify this situation as long as the Commission does not use its Plenary Powers
to nullify the latter designation. Of course, as Opinion 2223 mentions Calamita punctatus as type species of
Hysaplesia, which in fact was only due to an uncritical acceptation of Savage et al.’s (2007) analysis, it could
be considered that this Opinion ‘implicitly’ or ‘surrepticiously’ used its Plenary Powers to circumvent the
Code and render this designation effective, but admittedly this is quite a strange procedure. Furthermore, as
will be shown below, this has other, unexpected, nomenclatural consequences.
[C38d] As a matter of fact, Opinion 2223 can be stated to be at the same time ‘too much’ and ‘too little’. In
order to understand this, one must come back to the source of the problem which this Opinion was supposed
to solve. Why did Dubois (1982b) propose to invalidate the nomen Hysaplesia (as well as Hylaplesia which
he considered available)? It was because, Dendrobates being a nomen novum for Hylaplesia and therefore for
Hysaplesia, the latter, not being invalid for being a junior homonym of another generic nomen, should have
been adopted as the valid nomen of the genus then universally known as Dendrobates. Suppression of
Hysaplesia was then enough for solving this nomenclatural problem. But if the type species of Hysaplesia is
modified, explicitly by use of the Plenary Powers or ‘surreptitiously’ by adoption of the interpretation of
Holthuis (1983) and Savage et al. (2007), then there is no need to invalidate this nomen! The nominal species
Calamita punctatus Schneider, 1799 [S06] is currently referred by all authors to a genus of the family
HYLIDAE, called Hypsiboas by Faivovich et al. (2005) and Duellman et al. (2016), but for which the valid
nomen is shown above under [C34b] to be Boana Gray, 1825. The type species of Boana, by subsequent
designation of Stejneger (1907: 76), being Rana boans Linnaeus, 1758 [S14], if Calamita punctatus was
designated, under the Plenary Powers, as the type species of Hysaplesia, the latter nomen would be a junior
subjective synonym of Boana. But it could then be potentially available for a taxon corresponding to the
‘Hypsiboas punctatus group’ of Faivovich et al. (2005: 88), or slightly less or more inclusive than the latter
but anyway including the species Calamita punctatus but excluding Rana boans, in case the genus Boana
would later have to be split into several subgenera or genera. In such a case, the invalidation of Hysaplesia
(and consequently of Hylaplesia) by the Commission would prove to have been not merely useless (being
redundant with its change of type species), but even harmless from the viewpoint of ‘nomenclatural
parsimony’ if it was then necessary to coin a new nomen for the taxon including the species Calamita
punctatus.
In conclusion of this point, in using its Plenary Powers for Opinion 2223, the Commission should have
made a clear choice between two possible solutions to the problem at stake: either invalidating Hysaplesia
(and Hylaplesia) or changing its type species. But taking both actions together testifies to a misunderstanding
of the nomenclatural situation at stake and could even be counter-productive. Furthermore, whereas the
‘invalidation’ option would indeed have allowed to solve all the problems posed by this case, the second
option leaves some of these problems unsolved, as will now be shown.
[C38e] As a matter of fact, Opinion 2223 failed to address the question of the status of three other nomina,
namely Hylaplesia H. Boie in F. Boie, 1828, Hyloplesia Agassiz, 1846 and Dendromedusa Gistel, 1858.
The status of the second and third of these nomina is fully clear. The first one was explicitly proposed by
Agassiz, a famous and very careful nomenclaturalist of this period, as an emendation of Hylaplesia, the latter
being credited to H. Boie in Schlegel (‘1827’, i.e. 1826b here). Being an autoneonym for Hylaplesia and
therefore for Hysaplesia, the status of Hyloplesia depends on that of the latter. If Hysaplesia is considered an
objective senior synonym of Dendrobates, then Hyloplesia is a junior, and therefore invalid, synonym of the
latter. But if its type species is accepted as being Calamita punctatus, Hyloplesia is a subjective junior
synonym of Boana. Its potential validity would then depend on the type species designation of the nomen just
discussed below.
The exactly same analysis applies to the nomen Dendromedusa Gistel, 1848, which was also introduced
explicitly as a neonym for Hylaplesia.
As for the nomen Hylaplesia H. Boie in F. Boie, 1828, its status as a nomen distinct from both
Hysaplesia H. Boie in Schlegel, 1826a and Hylaplesia H. Boie in Schlegel, 1826b is presented for the first
time in the present paper and it could be discussed. But if the analysis given above under [C04] is accepted,
Hylaplesia H. Boie in F. Boie, 1828 is in a situation similar to that of Hyloplesia and Dendromedusa as just
discussed, except that in this case no type species has yet been designated for it. If Calamita punctatus was
chosen for this purpose, Hylaplesia H. Boie in F. Boie, 1828 would become a senior objective synonym of
Hyloplesia and Dendromedusa. Since Hysaplesia has been suppressed by Opinion 2223, and since this
Opinion, by not mentioning Hylaplesia as it appears in Schlegel (1826b), must be interpreted as meaning that
the latter is an ‘incorrect subsequent spelling’ devoid of nomenclatural status, Hylaplesia H. Boie in F. Boie,
1828 would then be liable to be the valid nomen of a subgenus or genus including the nominal species
Calamita punctatus but excluding the nominal species Rana boans.
All these potential consequences of the change in the type species of Hysaplesia were not at all
considered by Savage et al. (2007) and in Opinion 2223.
[C38f] Regarding the status of the spelling or nomen Hylaplesia in Schlegel (1826b), as pointed out by
Brothers (in Anonymous 2009), since Opinion 2223 did not mention it at all, some doubt and unclarity persist.
It is true that, despite the fact that it was most probably the original spelling in Boie’s unpublished manuscript,
under a strict (not to say rigid) interpretation of the current Article 33.2.1, Hylaplesia is an incorrect
subsequent spelling of Hysaplesia. However, the fact that Hysaplesia was never used as valid after its original
publication and even ignored by all authors until Stejneger (1937), while Hylaplesia was extensively
mentioned in the literature and used as the basis of the family-series nomen HYLAPLESINA (and its subordinate
parograph HYLAPLESIDAE) should have retained attention from the part of the Commission, and the possibility
to afford it availability while invalidating it at the same time should have seriously been considered.
In fact, it could be questioned whether in this case Article 33.2.3.1 applies. This Article states that “when
an unjustified emendation is in prevailing usage and is attributed to the original author and date it is deemed to
be a justified emendation”. Under this Article, Hylaplesia would have to be treated as a ‘justified emendation’
and Hysaplesia as an ‘incorrect original spelling’. However, because of the unclear writing of the Code in this
case like in others, it is ambiguous whether this Article indeed applies here. The ‘Glossary’ of the Code
defines “prevailing usage” of a nomen as “that usage of the name which is adopted by at least a substantial
majority of the most recent authors concerned with the relevant taxon, irrespective of how long ago their work
was published”. Such a definition is not operational. To be so it would have to rely on precise definitions of
‘substantial majority’, of ‘most recent authors’, of ‘authors concerned with the relevant taxon’ and even of
‘usage’: should the latter be understood as applying only to publications where the nomen is ‘used as valid’ or
merely ‘mentioned’? In the present case, the situation is further complicated by the fact that the nomen at
stake was used as valid by only a few authors and in a few works (Schlegel 1826a–b; F. Boie 1828; Tschudi
1838; Fitzinger 1843; Günther 1858, 1859) and in all cases except the first one as Hylaplesia. This spelling
was the only one used until 1937 when Stejneger ‘resurrected’ the spelling Hysaplesia, but from that date one
or the other spelling, or both, were mentioned. If the ‘most recent authors’ are understood as covering only the
last 20 (or 10? or 30?) years, Hylaplesia cannot be considered to have been ‘adopted by at least a substantial
majority’ of them, as it would have been before 1937.
As the Dendrobates problem could be solved only through use by the Commission of its Plenary Powers,
the best solution in 2008 would have been to use this opportunity to rule that Hysaplesia is to be interpreted as
an incorrect original spelling (thus devoid of nomenclatural independent status) of Hylaplesia and at the same
time to ‘suppress’ the latter for the purposes of priority for synonymy but not of homonymy, while
recognising that Dendrobates was a nomen novum for Hylaplesia, having therefore the same type species.
Consequently, it would have been clear that the nomen HYLAPLESINA (and its parograph HYLAPLESIDAE),
which Opinion 2223 ignored, would automatically have become an invalid senior synonym of
DENDROBATIDAE and could have been placed on the Official Index of rejected nomina without any need for
further action under the Plenary Powers.
This solution would have had another strong advantage: it would have prevented ipso facto any
subsequent introduction of a new nomen Hylaplesia to create a new nomenclatural confusion. As a matter of
fact, as long as Hylaplesia is not deemed to be available although invalidated, this spelling ‘does not exist’ in
zoological nomenclature and any zoologist is free to introduce it again as the nomen of a new genus, which
cannot but result in a new nomenclatural ambiguity and problem. This is indeed the case in the interpretation
presented above under [C04] of Hylaplesia H. Boie in F. Boie, 1828 as being a brand new nomen (poieonym),
distinct from the spelling Hylaplesia appearing in Schlegel (1826b). Today, as the Commission did not
‘suppress’ the 1826 Hylaplesia, the 1828 Hylaplesia is not preoccupied and not rejected as a junior homonym,
with the nomenclatural consequences discussed above in [C38e].
[C38g] As shown in the comments above, the hastily voted Opinion 2223 did not solve all the nomenclatural
problems posed by this case, despite the fact that the latter had been discussed in detail for almost 40 years. As
the Commission has proved to be unreliable to solve this case, a complete solution to it is presented below
which avoids having recourse to its vote.
[C39] Amphibian Species of the World (2016)
[C39a] The well-known website ASW does not have the status of a scientific publication: its content is not
fixed and permanent but regularly modified, it does not have an ISSN or ISBN and is not registered in
Zoobank (see Anonymous 2012), so it cannot be a source for nomenclatural acts. This website is the work of
a single person, who benefits from information provided permanently by colleagues worldwide, but the
content of this website is not submitted to any kind of peer review and is just the expression of his personal
opinions. However, and despite its unquestionable shortcomings (see e.g. Dubois & Raffaëlli 2009: 23–25
and Dubois 2017a), ASW is a useful source of information (which then needs verification before acceptance
as valid) and it is considered by many zoologists as a reliable source about the taxonomy and nomenclature of
recent amphibians, so it deserves some words here. I here refer to the version 6.0 of this website, as made
available online on 18 August 2016, which, as shown above, contains various errors concerning the
Dendrobates case. Although the author of this website was one of the authors of the Savage et al. (2007)
application, he did not implement in this website all the consequences of Opinion 2223 (Anonymous 2009),
as shown below.
[C39b] Both Savage et al. (2007) and the Opinion 2223 adopted Dubois’ (1982b) treament of EUBAPHIDAE
Bonaparte, 1850 as an invalid senior synonym of DENDROBATIDAE Cope, 1865 by virtue of Article 40.2, but
then the nomen of this family should be written ‘DENDROBATIDAE Cope, 1865 (1850)’, which is not the case in
ASW.
[C39c] The nomen CALOSTETHINA Mivart, 1869, based on the unjustified emendation Calostethus Mivart,
1869, is an available but invalid junior synonym of COLOSTETHIDAE Cope, 1867 (see [C17] above), not an
incorrect subsequent spelling of the latter as stated in ASW.
[C39d] The website ASW follows Opinion 2223 in accepting Stejneger’s (1838) type species designation of
Calamita punctatus Schneider, 1799 for Hysaplesia and therefore (although not explicitly) rejects the
interpretation of Dendrobates as a nomen novum for the latter. Writing that Duméril & Bibron (1841)
designated the type species of Dendrobates but not of Hylaplesia is meaningful only in the latter case,
because if Dendrobates is understood as a nomen novum a designation of type species for this genus also
holds for Hysaplesia according to Article 67.8.
But the fact of considering Calamita punctatus as the type species of Hysaplesia has another
consequence, which was not drawn in ASW: the latter nomen cannot remain in the synonymy of Dendrobates!
As shown above, it should then be referred to the synonymy of the genus Boana Gray, 1825 of the family
HYLIDAE—or of Hypsiboas Wagler, 1830, the nomen used in error in ASW to designate this genus. The same
applies to Hylaplesia in Schlegel (1826b) if the latter is treated as an incorrect subsequent spelling of
Hysaplesia as it is the case in ASW.
[C39e] For the same reason, under the nomenclatural interpretation of the status of Hylaplesia adopted in
ASW, the nomen HYLAPLESIDAE must be listed among the synonyms of HYLIDAE, not of DENDROBATIDAE as it
is the case in ASW.
[C39f] Without explanations, ASW did not adopt the two corrections of Savage et al. (2007) concerning the
dates and authorships of two works: Schlegel (1826b) instead of ‘1827’, and Bibron in de la Sagra (1840)
instead of Duméril & Bibron (1841).
[C39g] The website ASW adopted the interpretation of Holthuis (1983) and Savage et al. (2007) according to
which Dendrobates is not a neonym for Hysaplesia but a brand new nomen created for a new taxon. If this is
the case, simple intellectual consistency requires that the same interpretation be also adopted for the other six
nomina listed above under [C37d] which had been proposed by Wagler (1827, 1830) to replace already
existing nomina in the exactly same manner and under the same format in a footnote. But here the surprising
fact is that the site ASW treats these seven cases of ‘couples’ of archaeonyms/neonyms in a heterogeneous
manner, with four different interpretations according to the nomen at stake, although simple consistency
would require to recognise only two different situations, autoneonym and alloneonym. In the following two
cases, the junior nomina by Wagler are (correctly) treated in ASW as cases of nomina nova (alloneonyms):
Asterodactylus for Pipa and Enydrobius for Hylodes. In the following two cases they are (incorrectly) treated
in ASW as new nomina proposed for brand new taxa although the junior one is an alloneonym: Dendrobates
vs. Hylaplesia and Systoma vs. Engystoma. In the case of Bombitator for Bombinator, Waglers’s nomen is
(correctly) considered as an unjustified emendation (autoneonym). Finally, in the following two cases, they
are (incorrectly) treated as incorrect subsequent spellings (ameletographs), although the junior one is an
autoneonym: Calamites for Calamita and Megalophrys for Megophrys. This heterogeneous treatment of a
small set of exactly similar cases throws a harsh light on the nomenclatural reliability of this renowned
website.
[C39h] Regarding the nomina discussed above, ASW displays various other errors concerning the authorships
of nomina—Hylaplesia borbonica Tschudi, 1838 (see [C07])—, their nomenclatural availability—“Calamita
tinctorius Schneider, 1799” (see [C23])—, their nominal-series assignment (see Dubois 2015)—HYLAPLESINA
(see [C13])—, not to mention others which I did not note. Unfortunately, this is a general phenomenon
throughout this website.
Discussion and conclusion
A comprehensive solution to all the nomenclatural problems at stake
As we have seen, despite a long discussion, mostly in the last 45 years, on this case, it seems hopeless to ever
reach a consensual interpretation of the nomenclatural problems at stake, based on a thorough analysis of the
original publications, on a full understanding of the Code and on a fair and rational discussion of all the
arguments presented.
In its Opinion 2223, the Commission took a double and contradictory action which showed a basic
misunderstanding of the nomenclatural problems encountered and leaves some of them unanswered: [A1] to
‘surreptitiously’ accept to deny the indubitable fact that Dendrobates was a nomen novum for Hylaplesia, and
therefore for its archaeonym Hysaplesia, but without using its Plenary Powers to invalidate this original
nomenclatural act, and to accept Stejneger’s (1937) type species designation for Hylaplesia as valid, thus
removing Hylaplesia from the synonymy of Dendrobates; but also [A2] to explicitly use its Plenary Powers to
‘suppress’ the nomen Hysaplesia (and, ‘surreptitiouly’ also, its widely used autoneonym Hylaplesia, not
mentioned in the Opinion 2223), seemingly to ‘protect’ the nomen Dendrobates—although following the
decision [A1] the latter was not threatened any more by Hysaplesia and Hylaplesia—thus unnecessarily
‘suppressing’ a nomen that might be wanting later in case of changes in the generic taxonomy of the family
HYLIDAE.
Rather than continuing to argue rationally on this case, the time has now come to accept the irrational
decision of the Commission and to complete it by a few additional nomenclatural acts to limit its unclarities
and negative effect.
I therefore propose to the international community: [P1] to accept that in its Opinion 2223 the
Commission, in placing the nomen 'Hysaplesia' H. Boie in Schlegel, 1826a on the Official Index with the
statement that Calamita punctatus Schneider, 1799 is its type species by subsequent designation of Stejneger
(1937), has in fact ‘surreptitiously’ used its Plenary Powers to invalidate the original nomenclatural act of
Wagler (1830) establishing the nomen Dendrobates as a nomen novum of Hysaplesia, and has therefore so to
say ‘disconnected’ these two nomina which had historically been linked from the beginning; [P2] to accept
that, by not mentioning in this Opinion the spelling Hylaplesia which appeared in Schlegel (1826b), the
Commission has treated the latter as an incorrect subsequent spelling (ameletograph) of 'Hysaplesia' H. Boie
in Schlegel, 1826a, devoid of nomenclatural availability; [P3] to recognise that the nomen Hylaplesia that
appeared in F. Boie (1828) without any mention of the works of Schlegel (1826a–b) is a distinct nomen,
nomenclaturally available.
To complete proposal [P3], I hereby formally designate the nominal species Calamita punctatus
Schneider, 1799, one of the five originally included nominal species of Hylaplesia H. Boie in F. Boie (1828), as
type species of the latter nominal genus. This nomen is currently a subjective synonym of Boana Gray, 1825
but would be available, in case the latter genus is later dismantled into several subgenera or genera, for one of
them including the species Calamita punctatus Schneider, 1799 but excluding Rana boans Linnaeus, 1758.
The following synonymic lists are based on these proposals. They concern the valid nomina of four taxa
currently recognised by all authors: the genus Boana Gray, 1825 and its subfamily COPHOMANTINAE
Hoffmann, 1878, and the genus Dendrobates Wagler, 1830 and its family DENDROBATIDAE Cope, 1865
(1850). The synonymy presented here for the latter applies in the case where no subordinate family-series
taxon is recognised as valid in this family.
Genus Boana Gray, 1825
Boana Gray, 1825: 214. ● Type species, by subsequent designation of Stejneger (1907: 76): Rana boans Linnaeus, 1758.
● Comment: nomen stated by Faivovich et al. (2005: 85–86) to be unavailable for having been originally published
as a synonym and never used as valid subsequently, which is wrong as it had clearly been published as a new
nomen to designate a subgenus within the genus Hyla Laurenti, 1768.
'Hysaplesia' H. Boie in Schlegel, 1826a: 239. ● Type species, by subsequent designation (ex errore) of Stejneger (1937),
validated by the Commission in Opinion 2223 (Anonymous 2009): Calamita punctatus Schneider, 1799. ● Nomen
partially ‘suppressed’ (for the purposes of priority among synonyms but not among homonyms) by the Commission
in Opinion 2223 (Anonymous 2009).
'Hylaplesia': H. Boie in Schlegel 1826b: 294. ● Junior ameletograph (incorrect subsequent spelling).
'Hylapesia': Savage et al. 2007: 257. ● Junior ameletograph (incorrect subsequent spelling).
Hylaplesia H. Boie in F. Boie, 1828: 363. ● Type species, by present designation: Calamita punctatus Schneider, 1799.
Hypsiboas Wagler, 1830: 200. ● Type species, by subsequent designation (ex errore) of Duellman (1977: 24): Hyla
palmata Bonnaterre, 1789 ◄ Rana maxima Laurenti, 1768 [≈ Rana boans Linnaeus, 1758]. ● Precedence over
Auletris Wagler, 1830 fixed by ‘First Reviser’ action of Faivovich et al. (2005: 85).
Auletris Wagler, 1830: 201. ● Type species, by subsequent designation of Stejneger (1907: 76): Rana boans Linnaeus,
1758. ● Subservience to Hypsiboas Wagler, 1830 fixed by ‘First Reviser’ action of Faivovich et al. (2005: 85).
Lobipes Fitzinger, 1843: 30. ● Type species, by original designation: Hyla palmata Bonnaterre, 1789 ◄ Rana maxima
Laurenti, 1768 [≈ Rana boans Linnaeus, 1758].
Hypsipsophus Fitzinger, 1843: 30. ● Type species, by original designation: Hyla xerophila Duméril & Bibron, 1841 [≈
Hyla crepitans Wied-Neuwied, 1824].
Phyllobius Fitzinger, 1843: 30 [nec Schoenherr in Germar, 1824]. ● Type species, by original designation: Hyla
albomarginata Spix, 1824.
Hyloplesia Agassiz, 1846: 189, 190. ● Autoneonym (unjustified emendation) for Hysaplesia H. Boie in Schlegel, 1826a.
Dendromedusa Gistel, 1848: xi. ● Alloneonym (nomen novum) for Hysaplesia H. Boie in Schlegel, 1826a.
Centrotelma Burmeister, 1856: 97. ● Type species, by subsequent designation (ex errore) of Duellman (1977: 24): Hyla
infulata Wied-Neuwied, 1824 [≈ Hyla albomarginata Spix, 1824].
Hylomedusa Burmeister, 1856: 102. ● Type species, by subsequent designation (ex errore) of Duellman (1977: 24):
Hyla crepitans Wied-Neuwied, 1824.
Hyla Burmeister, 1856: 104 [nec Laurenti, 1768: 32]. ● Type species, by present designation: Rana boans Linnaeus,
1758.
Cinclidium Cope, 1867: 200 [nec Blyth, 1842]. ● Type species, by original specific monophory (monotypy): Cinclidium
granulatum Cope, 1867 [≈ Rana boans Linnaeus, 1758].
Cophomantis Peters, 1870: 650. ● Type species, by original specific monophory (monotypy): Cophomantis punctillata
Peters, 1870 [≈ Hyla geographica Spix, 1824].
Cincloscopus Cope, 1871: 554. ● Alloneonym (nomen novum) for Cinclidium Cope, 1867.
Included species ● Boana aguilari; Boana albomarginata; Boana albonigra; Boana albopunctata; Boana alemani;
Boana alfaroi; Boana almendarizae; Boana atlantica; Boana balzani; Boana bandeirantes; Boana beckeri; Boana
benitezi; Boana bischoffi; Boana boans; Boana botumirim; Boana buriti; Boana caingua; Boana caipora; Boana
calcarata; Boana callipleura; Boana cinerascens; Boana cipoensis; Boana cordobae; Boana crepitans; Boana
curupi; Boana cymbalum; Boana dentei; Boana diabolica; Boana ericae; Boana exastis; Boana faber; Boana
fasciata; Boana freicanecae; Boana fuentei; Boana geographica; Boana gladiator; Boana goiana; Boana
guentheri; Boana helprini; Boana hobbsi; Boana hutchinsi; Boana jaguariaivensis; Boana jimenezi; Boana
joaquini; Boana lanciformis; Boana latistriata; Boana lemai; Boana leptolineata; Boana leucocheila; Boana liliae;
Boana lundii; Boana maculateralis; Boana marginata; Boana marianitae; Boana melanopleura; Boana
microderma; Boana multifasciata; Boana nympha; Boana ornatissima; Boana palaestes; Boana paranaiba; Boana
pardalis; Boana pellucens; Boana phaeopleura; Boana picturata; Boana poaju; Boana polytaenia; Boana pombali;
Boana pugnax; Boana pulchella; Boana pulidoi; Boana punctata; Boana quoyi4; Boana raniceps; Boana
rhythmica; Boana riojana; Boana roraima; Boana rosenbergi; Boana rubracyla; Boana rufitela; Boana secedens;
Boana semiguttata; Boana semilineata; Boana sibleszi; Boana steinbachi; Boana stellae; Boana stenocephala;
Boana tepuiana; Boana tetete; Boana tuberculosa; Boana varelae; Boana wavrini; Boana xerophylla.
Subfamily COPHOMANTINAE Hoffmann, 1878
HYLAPLESINA Günther, 1858: 345 [section]. ● Type genus, by implicit etymological designation: 'Hysaplesia' H. Boie in
Schlegel, 1826a: 239, cited under its junior ameletograph (incorrect subsequent spelling): 'Hylaplesia' (as appearing
in H. Boie in Schlegel, 1826b: 294) [≈ Boana Gray, 1825: 214]. ● Comment: invalid nomen under Article 39 for
being based on a type genus suppressed by the Commission acting under its Plenary Powers in Opinion 2223
(Anonymous 2009).
HYLAPLESIDAE [family]: Günther 1858: 341, 346.
HYLAPLESIINA [unspecified family-series rank]: Günther 1868: 148.
HYSAPLESIIDAE [family]: Cope 1875: 8.
HYLAPLESINA [family]: Hoffmann 1878: 614.
HYLAPLESIDAE [subfamily]: Hoffmann 1878: 614.
HYLAPLESIIDA [family]: Knauer 1878: 112.
HYSAPLESIDAE [family]: Grant et al. 2006: 156.
COPHOMANTINA Hoffmann, 1878: 614 [family]. ● Type genus, by implicit etymological designation: Cophomantis
Peters, 1870: 650 [≈ Boana Gray, 1825: 214].
COPHOMANTINI [tribe]: Faivovich et al. 2005: 3, 75.
COPHOMANTINAE [subfamily]: Duellman et al. 2016: 29.
4. For the status of this specific nomen, see Ohler & Dubois (2017).
Genus Dendrobates Wagler, 1830
Dendrobates Wagler, 1830: 202. ● Type species, by subsequent designation of Duméril & Bibron 1841: 30: Rana
tinctoria Cuvier, 1797: 295.
Eubaphus Bonaparte, 1831: 76. ● Type species, by original specific monophory (monotypy): Rana tinctoria Cuvier,
1797: 295.
Family DENDROBATIDAE |Bonaparte, 1850|-Cope, 1865
PHYLLOBATAE Fitzinger, 1843: 32 [family]. ● Type genus, by implicit etymological designation: Phyllobates Bibron in
de la Sagra, 1840: pl. 29. ● Comment: invalid nomen for the family (but not for a subfamily or lower family-series
taxon) for having been juniorised relatively to DENDROBATIDAE Cope, 1865 by the Commission acting under its
Plenary Powers in Opinion 2223 (Anonymous 2009).
PHYLLOBATIDAE [family]: Parker 1933: 12.
PHYLLOBATINAE [subfamily]: Ardila-Robayo 1979: 385.
EUBAPHIDAE Bonaparte, 1850 [family]. ● Type genus, by implicit etymological designation: Eubaphus Bonaparte, 1831:
76 [≡ Dendrobates Wagler, 1830: 202]. ● Comment: invalid nomen under Article 40.2 for having been replaced by
its junior synonym DENDROBATIDAE Cope, 1865 before 1961, and this replacement having “won general
acceptance” (Dubois 1982b).
EUBAPHINA [subfamily]: Bonaparte 1850.
DENDROBATIDAE Cope, 1865: 100, 104 [family]. ● Type genus, by implicit etymological designation: Dendrobates
Wagler, 1830: 202. ● Comment: nomen placed on the Official List by the Commission in Opinion 2223
(Anonymous 2009).
DENDROBATINAE [subfamily]: Gadow 1901: xi, 272.
DENDRONATINAE [subfamily]: Bauer 1988: 6.
DENDROBATOIDAE [epifamily]: Dubois 1992: 309.
COLOSTETHIDAE Cope, 1867: 191, 197 [family]. ● Type genus, by implicit etymological designation: Colostethus Cope,
1866: 130.
COLOSTETHINAE [subfamily]: Bauer 1987: 5.
CALOSTETHINA Mivart, 1869: 293 [subfamily]. ● Type genus, by implicit etymological designation: Calostethus Mivart,
1869: 293 [≡ Colostethus Cope, 1866: 130].
CALOSTETHIDAE [family]: Cope 1875: 7.
HYLOXALINAE Grant et al., 2006: 4, 168 [subfamily]. ● Type genus, by implicit etymological designation: Hyloxalus
Jiménez de la Espada, 1870: 59.
Zoological nomenclature and its governance
Zoological nomenclature
Some taxonomists consider that nomenclatural problems are ‘secondary’ in taxonomy and that trying to solve
them is just ‘waste of time’ (personal conversations and e-mail communications with colleagues).
Nevertheless, just like orthography and grammar in any published text, if you do not care for nomenclatural
accuracy, after some time the latter will care for you, or at least for your works. As there is a single Code for
all zoologists (except those who explicitly declare not to follow it, e.g. for adopting alternative nomenclatural
rules like the Phylocode—but with some unavoidable consequences regarding their works: see e.g. Muona
2006), if a work contains nomenclatural mistakes, sooner or later these will be corrected by some member(s)
of the zootaxonomic community. The sooner this correction is published, the better, as this will avoid a long
use of an incorrect nomenclature before a necessary change which will be considered by some ‘adepts’ of the
‘sacro-sanct’ nomenclatural stability to cause taxonomic problems although only a few persons in the world,
specialists of the taxonomic group at stake, are aware of them (see Dubois 2010b). Usually, when a
nomenclatural error is discovered, its correction occurs within a short period and leaves no long-term memory
(see e.g. the case of the replacement of Ixalus by Philautus discussed by Dubois & Ohler 2001). But
sometimes a ‘cramping’ occurs from the part of a few authors, who apparently feel ‘personally offensed’ by a
nomenclatural correction, and may then ‘struggle’ hard to maintain their original interpretation against all
evidence. This may become a real problem if these authors are, or consider themselves to be, ‘top authors’ in
the taxonomy of a given zoological group: in such cases they may use various methods which should have no
place in science, including lobbying, censorship or falsification of the facts, to defend ‘their’ view and they
may sometimes gain the silent support of the community, mostly through the use of what clearly qualifies as
the medieval ‘Principle of Authority’: ‘this is true because I, a very important person, declare that it is true,
and I do not need to provide evidence or even arguments to support my interpretation’.
The statement that striving for nomenclatural accuracy, e.g. through a careful examination of ancient
publications, is a ‘waste of time’ and that zootaxonomists have ‘better things to do’, cannot be accepted.
There is a strong difference here between publications in taxonomy and publications in other fields of
biology. Of course, nobody is expecting from the author of a work on the ecology, behaviour, physiology,
conservation or even phylogeny of an animal species or group of species to have looked at its/their original
description(s) or to have solved nomenclatural problems concerning it/them—as long as new nomina or
nomenclatural acts are not proposed in the work. Such authors are simply using nomina either because they
are well-known or because they are considered the valid nomina for the taxa in a recent monograph, fauna,
key, or even website (but see below). But taxonomists, whose activity consists in studying, differentiating,
diagnosing and naming taxa, should not ignore the past literature on the group they are studying. This is
because so many taxonomists do indeed ignore it that so many junior synonyms are regularly created, thus
contributing to the artificial nomenclatural inflation which is one of the nuisances of taxonomic research, as it
was discussed in several recent publications (see e.g. Dubois 2008 and references therein). This is indeed
‘poor taxonomy’.
Of course, in most zoological groups, some very old or rare publications may be difficult to obtain
(although this is less and less the case with the growth of the Biodiversity Heritage Library website), and in
such cases the only solution may be to rely on some subsequent revisionary works by authors who have seen
the original works (or claim having seen them), but this should remain an exception, duly acknowledged
through quoting these works in the bibliography followed by the mention ‘Not seen’. At least this shows that
the author has cared for seeing this work, but unfortunately has not had the possibility to see it, which is not
the case if the reference does not appear in the reference list. Many old references are often copied from work
to work without anyone having cared for looking at the publication. When one finally does so, one has often
strange surprises, such as finding that the original nomen had a spelling different from that currently used, or
is a nomen nudum, or was coined for a taxon having an onomatophore, a diagnosis or a figure pointing to
another taxon than that currently understood by this nomen, etc. It is the responsibility of taxonomists who
describe new taxa or revise groups to look for these original publications and to study them, as nobody else
will do it for them. And the later an old publication is re-examined in the light of our current knowledge and
concepts, the later such early mistakes are discovered and eventually corrected.
The case examined in detail above stresses the fact that nomenclatural accuracy requires to pay close
attention to the details of the Rules of the Code and to the particulars of the publications analysed, especially
when old texts of the 18th and 19th centuries are at stake, and that particular attention should be paid to the
following points, already outlined elsewhere (e.g., Dubois 1987a–b, 2008, 2010b, 2011, 2015a):
[P1] The Rules of the Code are Rules that must be respected, not merely indications, guidelines or
advice that taxonomists are free to follow or not whether they suit them or not.
[P2] Nomina are just labels meant at designating unambiguously and universally the taxa, not
descriptions of the taxa, theories, living beings (that could get extinct or ‘resurrect’) or devices for the
glorification of their authors. Accepting synonymisation of a nomen whenever it is shown to be an invalid
synonym or homonym is a basic prerequisite for a good taxonomic work. Refusing it for simple reasons of
vexation or ‘bruised ego’ is unacceptable as it drags zoological nomenclature outside the domain of science.
[P3] Nomenclatural problems are likely to be permanently solved only when all taxonomists involved in
their discussion are fair, make all efforts to be informed of all contributions to a discussion, do not ignore
some arguments that do not support their own interpretations but care to reply them honestly, and do not hide
information or references.
[P4] Three strong nuisances for nomenclature are the worship for ‘usage’ (even for very recent or
obscure nomina which are familiar only to a handful of specialists), the undue importance given to the
authorship of nomina and its companion ‘mihilism’ (Dubois 2008), and the persistent role still played in
nomenclature by the medieval ‘Principle of Authority’. Nomenclatural problems will be much more easily
solved when all involved parties agree to ‘depassionate’ and ‘de-individualise’ the field and to simply follow
the Rules without always trying to circumvent them.
[P5] Zoological nomenclature is a very technical and complex field. It is meaningless and hopeless to try
to ‘simplify’ it at the expense of accurateness and reliability. No other scientific field would accept to abandon
its basic concepts and methods because they are ‘too complex’ for uninformed people. In all domains, to be
part of the game, one has to do the effort to learn and practice. As nomina of taxa are ‘Latin-like’ terms, often
derived from Greek, a basic knowledge of Latin and Greek vocabulary and grammar are necessary to handle
correctly the forms of nomina. Those we do not have this knowledge should seek information in this respect
from knowledgeable colleagues, or keep outside the field.
[P6] When analysing ancient texts of the 18th and 19th centuries, one should always bear in mind that
there was no Code then and that publication standards were very different from what they are today.
Application of the Rules of the current Code to such texts should therefore be made with ‘care, understanding
and intelligence’ (Bour & Dubois 1984), not in a rigid, mechanistic and blind way, using our current concepts
and standards. For example, in such old texts the rather frequent fact that a new nomen was introduced as a
nomen novum for an existing taxon and not as a brand new nomen for a brand new taxon must sometimes be
established through the use of indirect criteria.
Be it as it may, even if some consider that working for ‘nomenclatural accuracy’ is just a ‘waste of time’,
it must be acknowledged that such cases can occupy many persons, over many pages in many papers and
during a long period of time—an energy which would certainly be used more meaningfully in doing genuine
taxonomic research based on the organisms, especially in our ‘century of extinctions’ (Dubois 2003, 2010c,
2015a–b). As shown above under [C01] to [C39], so far the Dendrobates case and closely related
nomenclatural questions have been discussed in 44 main papers (not mentioning many other ones in which
the nomina at stake were cited) published by 60 authors from the whole world over a period of 190 years
(from 1826 to 2016). If this was ‘waste of time’, then the taxonomic community may indeed be viewed as
particularly inefficient and irrelevant in its work, which certainly should raise concern.
The Commission
Evidence was largely presented above that being a member of the Commission is not a guarantee that a
taxonomist fully masters the Code and some of the subtelties of zoological nomenclature. This fact has
nothing surprising or shocking in itself, as ‘nobody is perfect’ and ‘everybody can make mistakes’. But it
becomes a problem when some commissioners believe that this status confers them some ‘power of
infallibility’ and allows them to deliberately ignore the arguments presented by others and to censor their
expression, and are uncritically followed by the Commission as a whole as if they were ‘experts’ of the
question at stake.
The Dendrobates case is interesting in showing that, in zoological nomenclature, manipulation and
rigging can be practiced in broad daylight not only by individual zoologists but also by a collective body like
the Commission without any reaction of the community of scientists working on the scientific subject
debated. The fact is that, after a long period of silence about this case, the Commission suddenly decided to
‘re-publish’ this case under a new number and in an application deliberately ignoring several important
arguments published about it but proposing a totally inacceptable misinterpretation of the concept of nomen
novum (see above under [C37d]), and to hurridly vote for a decision that was shown above to be flawed5. This
case (as well as many others) raises a question: is the role of this international Commission to act as a neutral
and objective ‘court’ to solve controversies on a basis of a complete, fair and competent examination of all
arguments produced, or to accept as granted the arguments produced by a few of its members or former
members without comparing them with the competing ones? By behaving according to the latter principle as
in this case, the Commission cannot but lose much of its credit in the eyes of the zootaxonomic community.
Why do we need a Commission? This body has a double role: one of updating the Code, and one of
settling the nomenclatural situation in problematic and controversial cases. In both of these functions, it is
supposed to work on the basis of information as complete as possible, analysed in a depassionate, objective
and competent manner, and of a full understanding and mastering of the Code. Otherwise it cannot but appear
as being ‘at the service’ of the private interests, opinions or tastes of a few zoologists for whom it is of utmost
importance to obtain confirmation that they ‘were right’ from the start, despite all evidence to the contrary.
‘Nomenclatural stability’ is not in itself a sufficient reason for having a Code and a Commission (see
Dubois 2010b)6. In no other scientific field, ‘stability’ of accepted ideas, theories and practices would be
considered a valid reason for rejecting new information showing that these ‘accepted truths’ were wrong.
‘Nomenclatural accuracy’ is much more important than ‘nomenclatural stability’, because it is the only real
basis for genuine stability. Stability obtained at the expense of intellectual rigour and fairness is ill-based and
will be easily dismissed or ignored by some—while both the Code and the Commission will appear useless or
harmful to some practising taxonomists. This is already apparent in the lack of respect for the Code shown in
the recent decades from the part of many taxonomists, referees, editors and journals (Dubois 2003, Dubois et
al. 2013). This attitude opens the door to nomenclatural chaos. If every group of zoologists (gathered on the
basis either of the taxonomic group studied, or of their country or native language) decides to have its own
Code, or ‘sub-Code’ for specific questions (e.g. the decision of the Societas Europaea Lepidopterologica to
ignore the grammatical gender agreement between generic substantives and specific epithets; see Löbl 2015),
the universality of nomenclature will be challenged, and with it the universality of taxonomy and
consequently of biology: if the same pest, parasite or invasive species is given different nomina in China,
Brazil, the USA and Europe, the negative effects of this fact may not be limited to communication between
taxonomists but may concern biology, and even society, as a whole.
The Commission was created as a result of a complex history (see Melville 1995). As its members are not
elected but co-opted by the members still in activity, this body cannot claim to be ‘representative’ per se of
the international community of zootaxonomists (see Dubois et al. 2016). The only ‘legitimacy’ it could
pretend to would be if an overwhelming majority of authors, referees, editors and publishers of zootaxonomic
works followed the Code and the Commission’s decisions. As it is quite clear that this is currently not the
case, this ‘legitimacy’ is highly questionable. This situation might change if the Commission decided to
modify its procedures (see Dubois 2011, Dubois et al. 2016). If not, it cannot be excluded that an alternative
mode of governance of zoological nomenclature be proposed by another group of zootaxonomists and
eventually adopted also by others.
5. At the time when this occurred, this case was not the only one in which my opinion and arguments had been ignored or censored
by the Commission. Fourteen other manuscripts that I had submitted to the BZN from 1982 to 1997 were never published in this
bulletin: six were published elsewhere and eight were never published, so that the nomenclatural problems they raised are still pending
(see Dubois 2005: 417–418, 423–424). So when the Savage et al. (2007) paper was published I refrained from intervening in its
discussion. I did the same regarding another case, concerning a major and universally known work of early herpetology, containing
several nomina that had been used as valid in thousands of publications and which should no doubt have been ‘protected’ if the
Commission had really cared for ‘nomenclatural stability’ as it pretends to do: the ‘suppression’ (Anonymous 2005) of the work of de
la Cepède (1788a–b) brought no benefit to zoological nomenclature, quite to the contrary, but complied with the request of one
member of the Commission, an open adversary of the “tyranny of the past” (Savage 1990a–b, 1991), while deliberately ignoring a
warning against “a rigid application of the Rules to old, well-known zoological works” (Bour & Dubois 1984) as well as “strong
objection to the structure and content of the application” by one commissioner—the nature of this objection remaining mysterious, not
being mentioned in the final Opinion 2104 (see Dubois & Raffaëlli 2009: 26–27).
The Code
Articles 33.2.1 and 33.2.3 of the Code list the conditions that must be complied with for a new nomen to be
interpreted as an intentionally modified spelling but not a ‘justified emendation’—i.e., an ‘unjustified
emendation’ or autoneonym. As we have seen above, Article 33.2.1 is quite restrictive as it allows to
recognise that a nomen is an emendation only in three situations (explicit statement of intention, mention of
both the original and the changed spelling, and two or more nomina in the same work being treated in a
similar way). But, especially when ancient works are concerned, there can be other kinds of evidence for the
intentionality of spelling change (Dubois 1987b: 35–38; 2010a: 10–13, 29–31). A suggested improvement of
the current wording of this Article recognises six possibilities [Cr1] to [Cr6] listed in Appendix 1 below.
Criteria [Cr1] to [Cr3] correspond to those already in force in this Article, but [Cr4] to [Cr6] are new.
Furthermore, to avoid confusions, four criteria [Cr7] to [Cr10] are explicitly formulated that allow to
recognise situations which do not qualify as autoneonyms but as ‘incorrect subsequent spellings’ or
ameletographs. In my opinion, these suggestions should be duly considered for possible inclusion in the Code,
at least for modified spellings published before a date that would have to be fixed (e.g. 1930).
Taxonomic databases and websites
A new problem for zoological nomenclature has developed in the recent decades: that of the appearance and
multiplication of taxonomic databases and websites, supposed to provide useful information to all interested
persons, be them biologists or not, about the taxonomy and nomenclature of some zoological groups or of all
animals. The problem with such websites is that none of them provides complete and reliable information in
any of the fields that it is supposed to cover: bibliography, phylogeny, classification and nomenclature. This
has nothing surprising of course: for most zoological groups, the bibliography is enormous and covers
hundreds or thousands of references scattered over more that 250 years, and there exists no universal
consensus on the phylogeny and consequently on the taxonomy and nomenclature of the group. Such sites
would be extremely useful if they limited their role to the presentation of the published works and of the
existing alternative interpretations supported by different taxonomists or groups of taxonomists. But most of
them ambition to do more, to choose one among the different points of view and to promote this choice as a
kind of ‘truth’. As such, they tend to become a modern version of the medieval ‘Principle of Authority’.
This is well exemplified by the website ASW discussed above. Many zoologists and laymen use this site
to find information about the taxonomy and nomenclature of amphibians, and a tendency has developed in the
recent years to quote it in scientific papers as if it was a permanent bibliographic reference. This is
problematic because websites, being labile in their content, cannot constitute permanent scientific
6. As first formulated by Gaffney (1979) and much repeated (e.g., Dominguez & Wheeler 1997; Benton 2000; Dubois 2005,
2011), in taxonomy like in all other sciences, “stability is ignorance”. Taxonominal stability is not a scientific aim, but may correspond
to a social demand (e.g., for the purpose of ‘official lists’ or at the request of scientists using organisms for their research but not
interested in their classification and nomina). But taxonomists have been ill-inspired to comply with this social demand, as this has
been at the expense of their own scientific discipline. Whatever estimate is used for the total amount of speciodiversity, it is clear that
taxonomists have not collected, studied, described and named half of the species still existing on earth, possibly less than one tenth,
and these species are now engaged in an accelerated process of extinction in front of which science has very little possibility to act
(Dubois 2003), but the scientific community has not put the inventory of the earth species on top of its priorities, concentrating its
studies of biodiversity on much less urgent matters such as phylogeny (Wheeler et al. 2004). The causes of nomenclatural instability
are not mysterious: it results from incomplete or inaccurate nomenclatural work by taxonomists. Three measures could be taken to
reduce it drastically: [M1] increasing considerably the number of professional taxonomists and the fundings for field and laboratory
taxonomic research; [M2] developing theoretical and practical training in taxonomy worldwide; and [M3] resisting to the pressure
exerted on taxonomists to impose them to publish ‘quick and dirty’ taxonomic work in order to comply with the criteria of
‘evaluation’ and so-called ‘excellence’ required by administrative authorities. Unfortunately, instead of fighting to pass this message
to their hierarchy, in the recent decades taxonomists have ‘pleaded guilty’ for nomenclatural instability and have adopted the non-
scientific, or more exactly anti-scientific concept of ‘usage’ as a basic guide for their activity, thus contributing to nomenclatural chaos
and arbitrariness and to a poor image of taxonomy as a scientific discipline (see Dubois 1998).
bibliographic references (Dubois & Raffaëlli 2009). As this site always appear on top in any ‘Google search’,
many people think it is more of less ‘official’ and has the strong status of a basic, unavoidable reference, but
this is not true. This site is certainly very interesting and helpful to everybody, mostly as a source of
bibliographic references, but the information it contains should never be taken for granted and uncritically
accepted as valid or authoritative without having been checked from the original references and in the light of
the Code. It is the work of a single individual and its content has not been submitted to the critical control of
independent editors or referees as it is the case for any genuine scientific publication. Furthermore, it contains
some statements (e.g., subsequent type species designations for nominal genera) that would constitute
‘nomenclatural acts’ if validly published on paper or online with a proper registration and archiving, but
which are unavailable for having appeared only on a website which does not comply with the conditions of
Article 8 of the 2012 Amendment (Anonymous 2012): these unavailable ‘nomenclatural acts’ should never be
cited in genuine scientific works (see e.g. Ohler & Dubois 2017). All of this would be of relatively minor
consequences if some editors and referees did not consider that this website provides a ‘taxonomic and
nomenclatural truth’ which must be followed in all publications, and did not try to impose authors of
manuscripts submitted for publication to follow them on account of so-called ‘stability’ and ‘consensus’.
The same is true of all the other taxonomic websites that I have examined so far (see e.g. Dubois
2017a–b), whether limited to some taxonomic groups or more ‘general’, whether the work of a single person
or the result of a collective work—although the level of accuracy of the information provided is very variable.
One of the main reasons for this is that many of these websites tend to copy each other: thus, many of the
nomenclatural errors of ASW are copied ne varietur in various other websites—which in the eyes of many
supports the idea that this information is solid and reliable. Another major reason is that the authors or
moderators of these websites do not have the appropriate background for being able to make their own
analysis of the nomenclatural situations and to identify and correct the errors in this domain (see e.g. Dubois
2017b).
In fact, except in very small zoological groups counting only few species and supraspecific taxa, building
a complete taxonomic and nomenclatural website for a group must rely on the existence upstream of a solid
database devoted to this group—which means that one or several competent zootaxonomists have already
dedicated years of research to this subject (and then often published paper monographs presenting these data).
Creating such a database ex nihilo for the purpose of ‘feeding’ such a website is a very hazardous endeavour
unless cared for by very competent taxonomists having already a deep knowledge of the zoological group at
stake, of the relevant bibliography and of the history of its taxonomy and nomenclature, and ready to devote
much time and energy to this work.
All these comments are not meant at saying that all these online databases are not useful, which they
certainly are for some purpose (at least for that of bibliographical research) but that they are much less useful
than they could be if conceived differently. In fact, most of these websites are too ambitious. They strive at
providing a lot of information in several domains but they are incomplete or/and unreliable in several or most
of them. It would be much more efficient and useful if the developers of each database concentrated on a
single kind of information and gave themselves the means of providing it in a reliable manner.
Taxonomic (s. l.) databases dealing with zoological groups can provide information in four main distinct
domains: [D1] bibliography; [D2] phylogeny; [D3] taxonomy (s. str.), i.e. classification; and [D4]
nomenclature. Let us consider separately the kind of information each of them can provide.
[D1] Bibliographic databases and websites
For each zoological group, the existence of a comprehensive and duly indexed bibliographic database would
in itself be very useful. In this context, ‘indexed’ means that each reference be tagged with several key words
taken from a general list and concerning particularly the three fields [D2] to [D4]. Another important
information, useful mainly for nomenclature, concerns the actual date of publication, i.e. of public
distribution, of each work, which may be different from, or more precise than, the date printed on the work
itself. Such dates may be crucial for the determination of relative priority between works, nomina or
nomenclatural acts published over a short period of time. Rather than leaving each taxonomist alone in charge
of seeking such data, which may be difficult to trace, it would be very useful to have a comprehensive
bibliographic database providing this information, with mention of its source, at least for each major group, if
not for the whole of the animal kingdom.
[D2] Phylogenetic databases and websites
Phylogeny is a research field in itself. Phylogenetic hypotheses may be proposed on the basis of morpho-
anatomical or molecular data, or on a combination of both, and of other kinds of data (e.g. cytogenetic,
bioacoustic, behavioural). Currently, for any given group, most websites only present one or a few
phylogenetic hypothese(s), often the ‘last published one’, or sometimes the ‘preferred’ one of the person(s) in
charge of the database. However, since many of the analyses from which these trees were built were based on
different sets of specimens, species and characters (including genes), and on different methodologies for tree
construction, they are often not readily comparable, and choosing one of them for presentation in a website is
largely arbitrary. Rather than presenting a ‘truth’ in this respect (a ‘truth’ which in most cases will only have a
very short life), it would be much more useful if phylogenetic databases presented comprehensive surveys of
the phylogenetic works dealing with a zoological group and of the morpho-anatomical, molecular and other
characters they used. The simplest way to present the outcomes of a phylogenetic analysis is through a ‘tree’,
but the latter takes full signification only if accompanied by descriptions of the characters, by detailed
information on the methods of analyses used, by matrices of raw data, by quantitative data on the significance
of the nodes and by comments.
[D3] Taxonomic (s. str.) databases and websites
Taxonomy is the science of the classification of taxa. As such, it is largely independent from nomenclature: it
is fully possible to build up a classification of taxa without naming them, just using numbers, codes or other
devices to designate them. Nowadays, most taxonomists base their classifications on a critical evaluation and
acceptance of phylogenetic trees, the succession of nodes being ‘transcribed’ under the form of a hierarchy of
taxa of different ranks. However, in most cases, there is a loss of information between the phylogenetic
hypothesis adopted and the taxonomic hierarchy derived from it, because, for sake of taxonomic and
nomenclatural parsimony, not all nodes are recognised as taxa: therefore, the relationship between a
phylogenetic hypothesis and a taxonomic hierarchy is rarely bijective.
Of course, in order to communicate about a classification, the taxa must be unambiguously designated,
and for this the common (and most efficient) device is through nomina (scientific names ruled by a code of
nomenclature), but it should be clear that both stages (classification and nomenclature) are distinct and
successive. Although this does not seem to have existed so far, it would be fully conceivable to have
databases presenting several alternative classifications of a single zoological group, based either on different
trees or even on different taxonomic interpretations of a single tree, and showing them simply as ‘skeletons’
of hierarchies without attaching nomina to the branches except for the terminal taxa (species and subspecies).
[D4] Nomenclatural databases and websites
The undeniable difficulty of the discipline of nomenclature is due in part to its complex relationship with the
discipline of taxonomy. Nomenclature is in part independent from taxonomy and in part dependent from it.
This will be easier to understand by distinguishing two distinct aspects in what can be designated as the
‘status’ of a nomen: its nomenclatural status and its taxonomic status.
[D4a] Purely nomenclatural databases and websites dealing with the nomenclatural status of nomina
The nomenclatural status of a nomen is completely independent from taxonomy, i.e. from the classification
(ergotaxonomy) adopted as valid for the zoological group at stake. It consists in several pieces of information:
[i] The nomenclatural assignment of the nomen to a nominal-series.

[ii] Whether the nomen is available or unavailable under the Code.
[iii] Whether the nomen is a poieonym (brand new nomen directly introduced for an unnamed brand new
taxon), a neonym (nomen proposed expressly to replace an available nomen) or a protoallelonym (one of several
nomina having the same onomatophore proposed for the same taxon in the same publication).
[iv] The protonym or symprotonyms of the nomen.
[iv] The original spelling of the nomen, and whether it is its correct spelling (nomograph) or not.
[v] The grammatical gender of the nomen in the case of genus-series nomina.
[vi] The onomatophore (‘name-bearing type’) of the nomen if designated in the original publication.
[vii] Whether the nomen is, or not, an invalid junior homonym of another available nomen of the same
nominal-series.
[viii] Whether the nomen is, or not, an objective junior synonym of another available nomen of the same
nominal-series.
[ix] Whether the nomenclatural status of the nomen, as defined above under [i] to [vi], was left partly
ambiguous or erroneous in the original work, and was subsequently clarified or corrected by a nomenclatural act.
The latter can be either a ‘First Reviser’ action, a nomenclatural correction or a decision of the Commission:
[α] ‘First Reviser’ actions by arbiters are of various kinds, e.g.: subsequent designation of an
onomatophore (designation of a lectotype or neotype for a nominal species; subsequent designation of a
type species among the originally included nominal species of a nominal genus); choice of nomenclatural
precedence among synchronous objective synonyms, homonyms or nomenclatural acts; choice of the
correct original spelling among the multiple original spellings (symprotographs) of a nomen; etc.
[β] Nomenclatural corrections concern spellings which qualify as ‘incorrect original spellings’ under
the Code and should be replaced by their ‘correct spellings’ (nomographs).
[γ] Decisions of the Commission under its Plenary Powers may concern any aspect of the
nomenclatural status of the nomen.
[D4b] Taxonominal databases and websites dealing with the taxonomic status of nomina
The taxonomic status of a nomen depends on taxonomy, i.e. on the ergotaxonomy (classification) adopted as
valid for the zoological group at stake. It is therefore at the interface between taxonomy (s. str.) and
nomenclature. It may consists in several pieces of information:
[i] Whether the nomen is valid or invalid under the Code for a given taxon in the ergotaxonomy adopted as
valid.
[ii] If invalid, its invalidity may stem from its being:
[α] a junior objective or subjective synonym, or
[β] a junior primary or secondary homonym, or
[γ] a nomen rejected because of original rank precedence or
[δ] a nomen rejected because of ‘First Reviser’ action among synchronous objective synonyms,
homonyms or nomenclatural acts or
[ε] a nomen rejected through conditional rejection by decision of the Commission under its Plenary
Powers.
[iii] A mandatory change of ending following a change of generic allocation for a species-series nomen (new
combination) or of rank for a family-series nomen.
[iv] The recourse to Article 23.9 on reversal of precedence.
[v] The recourse to Article 33.2.3.1 resulting in an ‘unjustified emendation’ having to be treated as a ‘justified
emendation’.
[vi] The recourse to Article 40.2 to validate a family-series being a junior synonym.
[vii] A few other rare situations in which an invalid nomen or an incorrect spelling is validated through special
Rules of the Code.
[D4c] General comments and conclusion
The different data listed under [D4a] and [D4b] are of a very different nomenclatural nature.
The nomenclatural status of a nomen [D4a] does not depend at all on taxonomic data, decisions or
opinions. It is established once and for all in the original publication where the nomen is made available, or
slightly modified later through a nomenclatural act which then is definitive. Because this status is fixed and
immutable (often from the start and sometimes after a nomenclatural act), it can be ‘set in stone’. Building
purely nomenclatural databases concerning this information for all the available nomina concerning a given
zoological group would therefore be a very useful endeavour and should be encouraged. This would in part
correspond to the project of Lists of Available Names (LANs) proposed in Article 79 of the Code, with three
major differences: [Di1] such lists should not only include the available nomina but also information on their
onomatophores with their modes of designation, including through ‘First Reviser’ actions, as these data
concern the nomenclatural status of the nomina; [Di2] on the other hand, in such purely nomenclatural
databases it would be irrelevant to include information that depend on the ergotaxonomy adopted as valid, as
this concerns the taxonomic status of the nomen, not its nomenclatural status, and is liable to change over
time; [Di3] a procedure should be devised allowing to revise and update such lists in order to correct the
mistakes and include the missing data that will unavoidably exist in such lists, as in any important human
endeavour.
Nomenclatural databases of the [D4a] kind would be of great utility for all taxonomists, as they would
provide immutable and solid information that would remain valid under any ergotaxonomic frame. However,
to play fully this role, their preparation should be entrusted to competent taxonomists, not to people having an
approximate understanding and mastering of the Code or difficulties in the interpretation of ancient
publications, particularly in languages other than English. Evidence was presented above that this is not the
case of all taxonomists, even among members of the Commission. So it might be more appropriate for this
purpose to establish an ad hoc working group composed of taxonomists having demonstrated in their works
their ability to carry out such works. This working group could establish standards for the analysis of
taxonomic publications, including old ones, and for the collection and presentation of information relevant for
establishing the nomenclatural status of nomina. It might be a good idea to gather under a single web
domiciliation various [D4a] databases dealing with different taxonomic groups under a single ‘label’ for
which this working group would be in charge.
As concerns the taxonomic status of a nomen [D4b], this information makes sense only within the frame
of an ergotaxonomic frame adopted as valid. It would not be appropriate to include it in purely nomenclatural
databases as those just discussed above, but it should be incorporated in taxonomic databases as discussed
above under [D3].
For taxonomists, the existence of complete, detailed and accurate databases of the category [D4a] would
be of the greatest usefulness. These data being stable and inalterable, they would be useful for any taxonomic
work. Then, and only after databases of this first kind are established, it will be useful to build databases of the
kinds [D3] and [D4b], but, as they rely on subjective and potentially labile taxonomic decisions, the pieces of
information stored in such databases should be regularly updated and corrected, they should never been taken
for granted and their validity should be critically evaluated before each utilisation.
What is common to all these situations is that, for a proper use of the information provided in [D3] and
[D4] databases, a basic knowledge in taxonomy and nomenclature is indispensable to avoid small or gross
taxonomic or nomenclatural mistakes in taxonomic or taxonomy-based works.
Acknowledgements
I am very grateful to Thierry Frétey (Saint Maugan) and Annemarie Ohler (Paris) for their help in
bibliographic research for this paper, and to them as well as to Erna Aescht (Linz), Aaron Bauer (Villanova)
and Ivan Löbl (Genève) for their useful comments on the original manuscript.
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Appendix 1. Extracts of the paper by Dubois (1987b)
The paper by Dubois (1987b) was ignored by all authors after its publication, even in papers dealing with
some of the points it discussed in detail. Two parts of it (pages 35–36 and 40–44), which deal with the criteria
allowing to distinguish between unjustified emendation and incorrect subsequent spelling and with Wagler’s
generic new replacement names (including emendations), are largely reproduced below as originally
published (except for the format which was modified to fit that of the present journal). Other parts of the work
included an introduction, a discussion of the rules proposed by Dubois (1984) for the nomenclature of higher
taxa (including corrections of translation errors in Savage 1986), a detailed discussion about the distinction
between different kinds of generic nomina and spellings (a discussion deepened in Dubois 2010a),
discussions about other problems raised by Savage (1986) in generic nomina and about valid family-series
nomina, a few words about taxonomy and a conclusion which included the sentence: “After this detailed
analysis, I regret to say that I must reject as wrong all the corrections proposed by Savage (1986) to my list of
suprageneric and generic names of anuran”. This article having been universally ignored, none of its
statements has been discussed, let alone refuted, so far.
[p. 35]
Unjustified emendation versus incorrect subsequent spelling
(…)
[In order to distinguish an emendation from an incorrect subsequent spelling], some evidence that the
intention to change the spelling must exist in the original texts, and in this respect it is useful to give an
indicative, поп limiting, list of such potential evidence. To be deemed an emendation, a name must meet at
least either of the following criteria:
[Cr1] the name is presented in words as such;
[Cr2] both the original and the modified spellings are given, and the second one is retained by the writer
as the valid one;
[p. 36]
[Cr3] several names are treated in a similar way, e.g. corrected according to the etymology, or according
to some, possibly arbitrary, rule which is evident from the context;
[Cr4] the modified spelling is clearly etymologically justified and correctly formed, while the original
spelling was not, or could be considered not to be (see below for additional comments);
[Cr5] the modified spelling is introduced by the very author of the original spelling, either in a
‘corrigenda’, as stated by the Code (which however does not precise if the latter must have been published in
the same time as the original name, e.g. as an addenda to it, or may have appeared later), or in later
publications, especially when the new spelling is used repeatedly in subsequent works and the original
spelling definitively abandoned by the author;
[Cr6] the modified spelling is introduced by a subsequent author and used repeatedly in the same or,
better, in subsequent works, whereas the original spelling is definitively abandoned by this author (especially
when before introducing the new spelling he had made use of the original spelling in previous publications).
(…)
Now that we have discussed the major types of evidence for the fact that a new spelling was intentionally
created by an author, it may be useful to see which are the principal clues to the reverse situation, i.e. to
‘incorrect subsequent spellings’:
[Cr7] no explanation is given for the modified spelling, and the latter has no clear etymological
justification (on the contrary, it may often be incorrectly formed while the original spelling was correctly
formed);
[p. 37]
[Cr8] the original spelling is not mentioned, or, if it is, this is in a different part of the text, as a valid
name, and no choice is made between both spellings;
[Cr9] the modified spelling appears only once in the text, and is never used again by the same author in
subsequent publications, while this author may revert to the use of the original spelling in these papers;
[Cr10] the author of this modified spelling is known for having made numerous such mistakes (because
of his carelessness regarding these matters, of his bad handwriting, which was difficultly read by the printer,
of the carelessness of the printer, etc.).
(…)
[p. 40]
Wagler’s generic replacement names
Now that we have seen the things rather generally, let us look more carefully at the problems raised by some
of the generic names used by Wagler (1830). Savage (1986), following Holthuis (1983), is mainly concerned
with the problem of the name Dendrobates, because in my first paper on this question (Dubois 1982b) I had
presented an interpretation at variance with that accepted in all the previous applications published in the
Bulletin of zoological Nomenclature on this case, one of which was even co-signed by Savage (Silverstone
1971; Myers & Daly 1971; Cuellar et al. 1972; Peters et al. 1972; Lescure 1982). For some reason (...),
Holthuis (1983) and Savage (1986) want to preserve this first interpretation, but unfortunately they are wrong.
Because of the relative importance of this case as an exemplary one (since it has now been discussed by
several authors and that no agreement seems to be currently reached among the supporters of opposed
opinions), this question is dealt with here in some detail.
In Wagler’s times, the phrases ‘new replacement name’ or ‘nomen novum’, with the precise meaning
they now have in the Code, did not exist. If the word ‘expressly’ is taken literally (i.e. if these phrases must
appear as such in the text), it is clear that no name published at the time of Wagler, or even later, will ever
qualify as a ‘nomen novum’. But if it is taken, as suggested above, in the sense that the author must clearly
show, somehow or other, that his intention is to propose a new name for an already existing taxon, and not for
a new taxon, it will be clear that many names of this period qualify as such.
As I had already pointed out (Dubois 1982b), the names proposed by Wagler (1830) to replace already
existing names are all presented in a similar way, with a footnote giving (1) the etymology of the new name,
(2) the replaced name, with its author and sometimes its date and reference, and (3) sometimes, additional
comments. What is important in my argumentation (Dubois 1982b) is that I stressed this similarity of
presentation and considered that all pairs of names which appear in this way in Wagler’s text (e.g., in
Amphibia, Asterodactylus for Pipa, Dendrobates for Hylaplesia, Enydrobius for Hylodes, Systoma for
Engystoma, Bombitator for Bombinator) are to be treated similarly as a couplet composed of a replacement
name and of a replaced name. Strangely however, neither Holthuis (1983) nor Savage (1986) discuss this
aspect of my argument. Both of them refuse to consider Dendrobates as a replacement name for Hylaplesia,
and Savage (1986) further refuses to consider Asterodactylus and Bombitator as replacement names for
respectively Pipa and Bombinator. None of them however discusses the cases of Systoma or of Enydrobius
(for which Myers 1962 and Lynch 1971 had already adopted the same interpretation as me).
Savage’s (1986) insistance on the fact that Asterodactylus is not a replacement name for Pipa is all the
more incomprehensible that I had already stressed (Dubois 1983j) that this name had first been proposed by
Wagler, not in his 1830 book, but in a 1827 paper, where he wrote: “(Asterodactylus m. Pipa Auctor)”. To
refuse to interpret such a presentation as a statement that the first of these names is “proposed expressly as a
new replacement name” for the second one makes really no sense, since this mode of presentation for new
replacement names was very common in these times. To take examples dealing with amphibians, a similar
presentation appears in the works of Rafinesque-[Schmaltz] (1814, 1815) and of Gistel (1848) (…). If Savage
refuses to consider Asterodactylus as a replacement name of Pipa, he should logically also refuse to consider
e.g. Triturus Rafinesque, 1815 /
[p. 41]
/ as a replacement name for Triton Laurenti, 1768, or Philautus Gistel, 1848 as a replacement name for
Orchestes Tschudi, 1838 (see below). Since in these cases no diagnoses were given for the supposedly new
taxa, and no included species mentioned, Triturus and Philautus should be considered as nomina nuda,
without status in nomenclature. (…)
One may wonder why Wagler felt necessary to replace some existing generic names by others, coined by
him. Let us note in passing that even if we had no clue to help us to understand why he did so, that would not
in the least allow us to reject the evidence, and in this respect the discussion given by Savage (1986) about the
name Cacopus is completely irrelevant: no matter if Günther (1864) was wrong in believing the name
Uperodon to be preoccupied, the fact is that he clearly and ‘expressly’ presented Cacopus as a replacement
name for the latter! However and fortunately in this case, Wagler himself gave us the explanation. To be sure,
to find this it is necessary to read Latin and German in Wagler’s text, but I am of the opinion that it is not
possible to deal correctly with nomenclatural problems when one is unable to read, or at least decipher, Greek,
Latin, German and French! (…)
Wagler (1830: 17) gives us a first indication in a footnote concerning the mammalian generic name
Tapirus, for which he proposes the new replacement name Rhinochoerus. He then quotes Linnaeus, as
follows : “Nomina generica, quae ex graeca vel latina lingua radicem non habent, rejicienda sunt. Linné
Philos. bot. stud. Spreng. p. 265.” (‘Generic names, which do not have Latin or Greek roots, must be
rejected’). It is thus clear that he rejected as invalid all generic (not specific) names for which he did not find
a root in Latin or Greek languages. Careful examination of the whole book of Wagler bears this interpretation
out: in all those cases in which an existing generic name was neither Latin, nor Greek, /
[p. 42]
nor based on Latin or Greek roots, Wagler proposed a new replacement name. In some cases, he even
expressed his perplexity as to the possible etymology of a generic name, which for this reason he replaced by
a nomen novum. Thus, the name Systoma is proposed to replace Engystoma, which is presented as follows in
a footnote on p. 205: “Gen. Engystoma (quid?) Fitzing.”.
After Wagler’s death in 1832, Michahelles (1833) reproduced in Isis von Oken an unpublished manuscript
of Wagler, which had been written before the 1830 book of this author. In this text we can find additional
explanations of the reasons why Wagler rejected some generic names as invalid. In a footnote (in Michahelles
1833: 888), Wagler states that he found no evidence of the use of the names Hyla and Calamita in classical
Latin texts, and that for this reason these names must be rejected. He states that he found the name Calamites in
Plinius, as the name of the common European treefrog. He adds that unfortunately Fitzinger had already given
the name Calamita, “(das in Calamites umgeändert werden muß)” (‘which must be changed into Calamites’) to
another genus of frogs, and that, in order not to upset Fitzinger’s work, he refrains from using the name
Calamites for the European treefrog. In consequence he proposes the new replacement name Discodactylus for
Hyla, the latter not being of classical Latin or Greek origin. This footnote clearly shows: (1) that, as a general
rule, Wagler did propose new replacement names for generic names considered by him invalid, because they
were not of strictly classical Greek or Latin origin (i.e. a name used as such in classical Latin or Greek, or a
name based on classical Latin or Greek roots); (2) that nevertheless Wagler made clearly the distinction
between new names for already known taxa, and for new taxa, and that he respected the works of previous
authors (his refusal to use the name Calamites for a different genus than that called Calamita by
Fitzinger—after Schneider, 1799—is a rare example, for this epoch, of respect of the works of others); (3) more
specifically, that Savage’s (1986) analysis of the status of the names Calamites and Dendrobates is in error.
This leads us now to discuss the case of the unjustified emendations of generic names proposed by
Wagler (1830), such as Bombitator, Megalophrys or Calamites. The name Calamites is expressly presented
by Wagler (in Michahelles 1833) as a replacement name for Calamita, and could either be considered as a
nomen novum or as an unjustified emendation of this latter name (see discussion above). Strangely, Savage
(1986) considers it as the name of a new genus, overlooking the fact that Wagler (1830: 200) expressly wrote:
“CALAMITES (…) Fitzing.”, thus clearly acknowledging Fitzinger as the author of the taxon, even though
he emended his name. In reality, in Savage’s logic, if Calamites is not accepted as a replacement name or an
unjustified emendation for Calamita, it should not be considered as the name of a new genus (since it is
credited to a previous author), but as an incorrect subsequent spelling, without status in nomenclature. It is
true that the spelling Calamita does not appear in Wagler’s (1830) book, but it appears in the manuscript of
Wagler, anterior to this 1830 book, which was later published by Michahelles (1833). I contend that this case
is a typical example of type [Cr3] of evidence for emendations presented above, which is also acknowledged
in Art. 33(b)(i) of the new Code.
Savage’s (1986) too rapid way of working has another, quite funny indeed, consequence. Savage (1986)
states that Calamites is not an unjustified emendation of Calamita, but the name of a new taxon. He adds that
Fizinger’s (1843) designation of Hyla cyanea as type species of Calamites, rediscovered by Dubois (1984:
19), is valid, but he does /
[p. 43]
/ not discuss the consequences of this fact: these would be that Calamites Wagler, 1830 would be a senior
synonym of the well-known and much used generic name Litoria Tschudi, 1838! Fortunately, Calamites
Wagler, 1830 is preoccupied by Calamites Guettard, 1770 (see Dubois 1984: 14), so that, even if Savage
(1986) was right, no nomenclatural disruption would result. However, these facts and their consequences
should all be considered and discussed when such question are tackled: this clearly shows that in
nomenclature a very slight divergence of opinion as to the status of a name may have considerable
consequences in the long run.
As for the name Bombitator, Savage (1986) is still less excusable to consider it as the name of a new
genus, because Wagler (1830) mentions the name “Bombinator Merr.” in a footnote on page 206 and in the
index on page 346. In Latin, the name “bombitator” (bee) only did exist, while the names “bombina” and
“bombinator” did not exist but were coined by XVIII and XIX century authors on the basis of the name
“bombus” (buzzing). That Bombitator cannot either be considered as a fortuitous incorrect subsequent
spelling is also shown by the fact that this name alone (and not Bombinator) appears on several occasions in
Wagler’s book (pages 132, 206, 294, 296, 301, 302, 303, 305, 306, 346). It is thus clear that the spelling
Bombitator was purposedly used by Wagler instead of the spelling Bombinator, but for the same taxon as
Merrem (1820). If this is not an unjustified emendation, I wonder which name of this epoch will qualify for
this category.
To come back finally to the name Dendrobates now, it was clearly presented by Wagler (1830) as a
replacement name for Hylaplesia because the latter name was not acceptable according to Wagler’s
conceptions, being based on the root “Hyla” which was rejected by Wagler (1830 and in Michahelles 1833) as
invalid since it did not exist in classical Latin. The fact that Wagler (1830) only recognized some of the
species originally included in Hysaplesia (and Hylaplesia) by Boie (in Schlegel 1826, 1827) is of no
relevance here, because, as remembered above (…), an author may perfectly, while accepting a taxon created
by a previous author and crediting it to him, modify in part the diagnosis or content of the taxon: by doing so
he does not create a new taxon, because otherwise any new modification of the diagnosis or content of a taxon
would result in the creation of a new taxon and of a new name, and no stability of taxonomy and nomenclature
would ever be possible.
As for the status of the name Hylaplesia itself, it is certainly open to discussion. It might be possible to
consider it very formally as an incorrect subsequent spelling. However, as I have shown (Dubois 1982b,
1983j), this name has an etymological justification (being based on Hyla), while Hysaplesia has none and is
clearly the result of a misprint for Hylaplesia. Since the spelling Hylaplesia has been used by various authors
since its creation, while the spelling Hysaplesia has remained ignored until the paper by Stejneger (1937)
where it was resurrected, this name, on which is based the family-group name Hylaplesidae, is better
considered as having an independent status in nomenclature (for more detailed discussions of other similar
cases, see Dubois 1982a). It is therefore justified to consider Hysaplesia Boie in Schlegel, 1826, which was in
reality clearly a misprint in the original publication, as the “correct original spelling” of the name in the sense
of the Code, and Hylaplesia Boie in Schlegel, 1827 as an unjustified emendation of the latter. Finally,
Dendrobates Wagler, 1830 is without possible doubt a new replacement name for Hylaplesia. Therefore the
problems raised by Dubois (1982b) concerning the validity of the names Dendrobates and Dendrobatidae
remain, and must be solved by an action of the Commission.
[p. 44]
In conclusion the generic names Asterodactylus, Dendrobates, Enydrobius, Systoma, Bombitator,
Calamites and Megalophrys are all new replacement names (or, if one prefers, unjustified emendations, for
the last three ones) for existing generic names which were believed by Wagler (1827, 1830) to be invalid
because they were not of strict classical Latin or Greek origin. They have therefore by definition the same
type species as the replaced names, and Savage’s (1986) analysis is in error.
Submitted: 21 November 2016. Accepted: 17 January 2017. Published: 4 March 2017.

Corresponding Editor: Erna Aescht.
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The nomenclatural status of Hysaplesia, Hylaplesia, Dendrobates and related

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The nomenclatural status of Hysaplesia, Hylaplesia, Dendrobates and related

Terminology, abbreviations and symbols

Abbreviations and symbols

Article [followed by a number] ● Article of the Code.

Categories of new nomina

Chronology of the case

[S01] "Hylaplesia borbonica" Boie in Schlegel, 1826a (nomen nudum).

[C01] Gray (1825)

[C02] Schlegel (1826a)

[C03] Schlegel (1826b)

[C04] F. Boie (1828)

[C05] Wagler (1830)

[C06] Bonaparte (1831)

[C07] Tschudi (1838)

[C08] Bibron in de la Sagra (1840)

[C09] Duméril & Bibron (1841)

[C10] Fitzinger (1843)

[C11] Agassiz (1845, 1846)

[C12] Gistel (1848)

[C13] Bonaparte (1850)

[C14] Günther (1858, 1859)

[C15] Cope (1865)

[C16] Cope (1866)

[C17] Cope (1867)

[C18] Mivart (1869)

[C19] Boulenger (1882)

[C20] Lutz (1925)

[C21] Stejneger (1937)

[C22] Silverstone (1971)

[C24] Duellman (1977)

[C25] Lescure (1982)

[C26] Dubois (1982b)

[C27] Dubois (1983a, 1984)

[C28] Holthuis (1983)

[C29] Dubois (1983b)

[C30] Frost (1985)

[C31] Savage (1986)

[C32] Dubois (1987b)

[C33] Duellman (2001)

[C34] Faivovich et al. (2005)

“HYLA, Laur. Calamita, Schneid.

2. See note 1 above.

[C35] Frost et al. (2006)

[C36] Grant et al. (2006)

[C37] Savage et al. (2007)

[C38] Opinion 2223 (2009)

[C39] Amphibian Species of the World (2016)

Discussion and conclusion

A comprehensive solution to all the nomenclatural problems at stake

Genus Boana Gray, 1825

Subfamily COPHOMANTINAE Hoffmann, 1878

Genus Dendrobates Wagler, 1830

Family DENDROBATIDAE |Bonaparte, 1850|-Cope, 1865

Zoological nomenclature and its governance

Taxonomic databases and websites

[D1] Bibliographic databases and websites

[D2] Phylogenetic databases and websites

[D3] Taxonomic (s. str.) databases and websites