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The polytypic Straight-billed Woodcreeper Dendroplex picus (J. F. Gmelin, 1788) is one of the most complex species groups of Dendrocolaptidae (Aves: Passeriformes), from both the nomenclatural and morphological standpoints. Firstly, its alpha taxonomy is debatable. Virtually all recent works (e.g. Aleixo 2002; Marantz et al. 2003; del Hoyo & Collar 2016) have recognized just two species in the group—Dendroplex picus and Zimmer’s Woodcreeper Dendroplex kienerii (Des Murs, 1856)—although some of the other described taxa possess singular morphological characters and well-defined ranges somewhat isolated from their geographically closest relatives (e.g. Plain-throated Woodcreeper Dendroplex picirostris Lafresnaye, 1847). Secondly, the correct genus to which to allocate taxa presently included in this group (vide Aleixo 2002) has been controversial. There is a considerable confusion as to which nominal species should be regarded as the type of Dendroplex Swainson, 1827b. Three species are involved in the dispute (Cory & Hellmayr 1925; Peters 1951; Aleixo et al. 2002; Marantz et al. 2003; Aleixo et al. 2007): Oriolus picus J. F. Gmelin, 1788; Dendrocolaptes guttatus M. H. C. Lichtenstein, 1818; and Dendrocolaptes ocellatus Spix, 1824. Here, we re-examine the nomenclatural issue and show that application of the nomen Dendroplex to the clade comprising the species-group D. picus (Aleixo et al. 2007) is based on a misunderstanding of the application of Article 70.3 of the Code (Anon. 1999) and that Dendrocolaptes ocellatus Spix, 1824, is its real type species. Consequently, the genus Dendroplex Swainson, 1827b, must be considered a junior synonym of Xiphorhynchus Swainson, 1827a. Because no generic nomen is currently available for them, we propose a new genus nomen to encompass the species originally described as Oriolus picus J. F. Gmelin, 1788, Dendroplex picirostris Lafresnaye, 1847, and Dendrornis kienerii Des Murs, 1856.
Accepted by A. Minelli: 6 Nov. 2018; published: 20 Dec. 2018
ISSN 1175-5326 (print edition)
(online edition)
Copyright © 2018 Magnolia Press
Zootaxa 4532 (4): 561
Synonymization of the genus nomen Dendroplex Swainson, 1827 and description
of a new genus of woodcreeper (Aves: Passeriformes: Dendrocolaptidae)
with remarks on Articles 67.5 and 70.3 of the Code
Setor de Ornitologia, Departamento de Vertebrados, Museu Nacional/UFRJ, Quinta da Boa Vista s/n, Rio de Janeiro, RJ, 20940-040,
Brazil. E-mail: (MAR)
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
E-mail: (AD), (RS)
Research Associate, Field Museum of Natural History, 1400 South Lakeshore Drive, Chicago, IL 60605, USA
E-mail: (GMK)
Universidade de São Paulo and Research Associate of the Museu de História Natural de Taubaté, Rua Juvenal Dias de Carvalho 111,
12070-640 Jardim do Sol, Taubaté, SP, Brazil. E-mail: (EH)
Departamento de Biologia, FFCLRP, Universidade de São Paulo, Av. dos Bandeirantes 3900, 14040–901 Ribeirão Preto, SP, Brazil.
E-mail: (RS)
The polytypic Straight-billed Woodcreeper Dendroplex picus (J. F. Gmelin, 1788) is one of the most complex species-
groups of Dendrocolaptidae (Aves: Passeriformes), from both the nomenclatural and morphological standpoints. Firstly,
its alpha taxonomy is debatable. Virtually all recent works (e.g. Aleixo 2002; Marantz et al. 2003; del Hoyo & Collar
2016) have recognized just two species in the group—Dendroplex picus and Zimmer’s Woodcreeper Dendroplex kienerii
(Des Murs, 1856)—although some of the other described taxa possess singular morphological characters and well-defined
ranges somewhat isolated from their geographically closest relatives (e.g. Plain-throated Woodcreeper Dendroplex pici-
rostris Lafresnaye, 1847). Secondly, the correct genus to which to allocate taxa presently included in this group (vide
Aleixo 2002) has been controversial. There is a considerable confusion as to which nominal species should be regarded
as the type of Dendroplex Swainson, 1827b. Three species are involved in the dispute (Cory & Hellmayr 1925; Peters
1951; Aleixo et al. 2002; Marantz et al. 2003; Aleixo et al. 2007): Oriolus picus J. F. Gmelin, 1788; Dendrocolaptes gut-
tatus M. H. C. Lichtenstein, 1818; and Dendrocolaptes ocellatus Spix, 1824. Here, we re-examine the nomenclatural issue
and show that application of the nomen Dendroplex to the clade comprising the species-group D. picus (Aleixo et al. 2007)
is based on a misunderstanding of the application of Article 70.3 of the Code (Anon. 1999) and that Dendrocolaptes ocel-
latus Spix, 1824, is its real type species. Consequently, the genus Dendroplex Swainson, 1827b, must be considered a ju-
nior synonym of Xiphorhynchus Swainson, 1827a. Because no generic nomen is currently available for them, we propose
a new genus nomen to encompass the species originally described as Oriolus picus J. F. Gmelin, 1788, Dendroplex pici-
rostris Lafresnaye, 1847, and Dendrornis kienerii Des Murs, 1856.
Key words: Aves, Dendrocolaptidae, Nomenclature, New genus, Dendroplex
In the original description of Dendroplex, Swainson (1827b: 354) provided only a brief diagnosis of his new genus:
Rostrum rectissimum. Alæ mediocres, rotundatæ; remigibus 3tiá, 4tá, et 5tá longissimis”. Concerning its type
species, he wrote: “I know not whether the type of this genus has been described”. Following the International
Code of Zoological Nomenclature (Anon. 1999, the Code below) the new nomen Dendroplex is therefore available
(Art. 12), but the new genus taxon was established without originally included species (Art. 67.2) and without any
type species fixation (Art. 69).
Zootaxa 4532 (4) © 2018 Magnolia Press
Cuvier (1829
) was the second author to at least indirectly refer to this genus and could be interpreted as having
fixed its type species by subsequent designation, as he referred just one species to the genus, that first mentioned by
Buffon (1783: pl. 605) under the French name “Le Talapiot”. However, Cuvier (1829: 583) wrote: “je crois que le
Talapiot est son [Swainson’s] genre Dendroplex”. The phrase “je crois” [‘I believe’] used here corresponds to the
formulae “doubtfully or conditionally included” of Art. 67.2.5 and “ambiguous or conditional manner” in Art.
67.5.3 of the Code, and precludes considering this species as having been originally included in the genus, much
less to have been fixed as its type species
. It is worth mentioning that even if Cuvier had not mentioned his doubts,
he could not be considered to have fixed a type species by subsequent monotypy, as he wrote (1829: 431) about this
species: “(3) Le talapiot, Buff. (Oriolus picus, Gm. et Lath.; Gracula picoides, Sh., enl. 605, ou Dendrocolaptes
guttatus, Spix. 91, i.)”. Thus, he had mentioned three distinct nominal species, which currently correspond to two
taxonomic species, as synonyms of the talapiot. Art. 69.3 expressly states that subsequent monotypy applies solely
“If only one nominal species was first subsequently included in a nominal genus or subgenus established without
nominal species”, so Cuvier (1829) did not fix a type species for Dendroplex, and even failed to provide a list of
subsequently included species for this genus because the three nominal species mentioned were only doubtfully
referred to the genus.
Subsequently, Griffith et al. (1829: 351)
referred again to this bird, citing the mention of Cuvier (1829: 431),
but as Le Galapiot instead of Le talapiot: “Le Galapiot. Buff. Oriolus Picus. Gm. et Lath. Gracula Picoides. Sh.
Enl. 605, or Dendrocolaptes Guttatus. Spix. 91. 1.” However, as they did not mention the generic nomen
Dendroplex, the authors failed to clarify the nomenclatural status of the genus with respect to its type species or its
subsequently included species. One year later, Swainson (1830: 689)
, referring to Griffith et al. (1829: 351),
wrote: “351. Le Galapiot. We omitted to cite this as the type of our sub-genus, Dendroplex. (Zool. Journ. 10, p.
354.)”. Despite the formula “the type of our subgenus”, this is still not a valid type species designation, as Le
Galapiot is not an available nomen (Art. 12.3).
Swainson (1837: 313–314) provided essentially the same diagnosis as his original description, but it was
accompanied by an illustration of the straight culmen with lateral compression. At the end of his characterization,
Swainson added: “D. guttatus. Spix, i. 91. f. 1.”. According to Art. 69.3, this is indeed a type species fixation by
subsequent monotypy, for a nominal genus established without included species.
However, according to all subsequent authors (Cory & Hellmayr 1925; Peters 1951; Aleixo et al. 2002;
Marantz et al. 2003; Aleixo et al. 2007), the plate “Spix, i. 91. f. 1.” referred to by Spix (1824) and Swainson
(1837) as Dendrocolaptes guttatus is in fact Dendrocolaptes ocellatus Spix, 1824, and not Dendrocolaptes guttatus
M. H. C. Lichtenstein, 1818. Spix (1824: 89) himself made a very clear distinction between Dendrocolaptes
guttatus M. H. C. Lichtenstein, 1818, and his “Dendrocolaptes ocellatus (guttatus)” described in association with
1. Published on 11 April 1829 according to Dickinson et al. (2011).
2. Regarding our interpretation of “je crois” as a doubtfully or conditionally included statement, the verb‘croire’ in French
refers to a belief, not a certitude. The leading French dictionary Le Robert defines this verb as follows: “croire. 1. Penser
que (quelque-chose) est vérifiable, donner une adhésion de principe ... 4. croire que, considérer vraisemblable ou
probable...”. Therefore use of this verb strongly implies a degree of doubt or uncertainty. Similarly, to say “I believe the
Earth is flat” is not the same as “the Earth is flat” or “I know that the Earth is flat”. The former permits the existence of
some doubt as to the ‘veracity’ of the statement. This perspective is corroborated by the rest of pages 582–584 in Cuvier
(1829), who listed a number of other bird genera and, in some cases, their ‘type species’ (although he rarely employed this
term). For all of them, except Dendroplex, he wrote in the form: “[l’espèce X] est [le genre Y]” or a similar formulation. If
no available nominal species was previously referred to genus Y, this amounts to a type species designation for Y by
subsequent monotypy. Examples of this format occur in relation to the following genera: Gyps, Aegyptius (for which he
used “type”), Hierax, Nauclerus, Ictinia, Tyrannula, Mylagra, Ripidura, Setophaga, Stenura, Pachycephala, Seisura,
Aglaia, Sterm agra, Monarcha, Tropidorhynchus, Megalurus, Aegotheles, Brachonyx, Macronyx, Dolichonyx, Dasyornis,
Sittasomus, Oxyglossus, Xiphorhynchus, Todiramphes, Colaptes, Peristera and Ectopistes. In only a few cases (Harpagus,
Anthochaera, Eudynamys, Chamaepelia), he mentioned more than one nominal species for a genus, which means that in
these cases he cannot be construed as having designated a type species by subsequent monotypy. For still other genera, he
mentioned diagnoses or vernacular names, but did not list the included (or type) species. All of the statements by Cuvier
(1829) listed above are clearly affirmative, i.e. without any element of doubt. The sole use of “je crois” pertains to the
genus Dendroplex. It is made even more striking because the relevant sentence commences with an affirmative statement
(“est le genre”) but terminates in this clause that evidences a lack of certainty.
3. Published on 21 May 1829 according to Dickinson et al. (2011).
4. Published on 23 January 1830 according to Dickinson et al. (2011).
Zootaxa 4532 (4) © 2018 Magnolia Press
the plate referred to by Swainson (1837). In providing a diagnosis for “Dendrocolaptes ocellatus (guttatus)”, Spix
also compared his new species with Dendrocolaptes guttatus M. H. C. Lichtenstein, 1818, stating that it “differt a
D. guttato corpore minore…” (Spix 1824: 88) as well as with O. picus (“a D. Pico pectore non late albo…”).
The reason that Spix’s plate (i, 91, f. 1) carries the legend Dendrocolaptes guttatus despite being used as the
basis for his description of Dendrocolaptes ocellatus (guttatus) is probably merely editorial (i.e., due to the fact that
the plate had already been engraved before the text was ready), explaining why the nomen “guttatus” appears in
brackets after the species nomen in the text. This also demonstrates that Spix (1824) was aware of the
misidentification of his own plate. We have examined the specimen on which the plate is based, housed at the
Zoologische Staatssammlung München (ZSM B96) and it is not only a specimen of Dendrocolaptes ocellatus, but
its putative holotype.
Nomenclatural rationale
In such cases, the Code (Anon. 1999, Art. 69.2.4, deliberately cited misidentifications) states explicitly that: “If an
author subsequently designates as type species a species originally included [Art. 67.2.1] as an expressly stated
misidentification or misapplication of a previously established nominal species, the species so designated is the
nominal species denoted by the name of the taxonomic species actually involved (and not the nominal species
cited).” So Dendrocolaptes ocellatus Spix, 1824, is the type species of the genus Dendroplex Swainson, 1827b,
designated by Swainson (1837).
Aleixo et al. (2007) applied Art. 70.3 to Dendroplex Swainson, 1837, but this article cannot be used for type
species fixed by deliberately cited misidentifications (see Anon. 1999: 74). Reconciling Art. 69.2.4 (deliberately
cited misidentifications) and Art. 70.4.2 (identification of type species by deliberate misapplication for the
subsequent fixation as the type species) with the correct usage of the woodcreeper genus Dendroplex becomes
quite evident when all of the literature is considered. The fact that Spix (1824) described both Dendrocolaptes
guttatus M. H. C. Lichtenstein, 1818, and Dendrocolaptes ocellatus Spix, 1824, and that the reference made by
Swainson (1837) is exclusively to the plate and not to the descriptions makes this sufficiently clear.
It is worth mentioning that Art. 11.10 and 67.13 of the Code, also refer to type species cited as deliberately
used misapplications. Art. 11.10 states that “If an author employs a specific or subspecific name for the type
species of a new nominal genus-group taxon, but deliberately in the sense of a previous misidentification of it, then
the author’s employment of the name is deemed to denote a new nominal species and the specific name is available
with its own author and date as though it were newly proposed in combination with the new genus-group name.”.
Following this rationale, Dendrocolaptes guttatus Swainson, 1837, should be considered indeed a new available
name, albeit, permanently invalid, given the fact it is a junior primary homonym (see Art. 53.3, 57.2) of
Dendrocolaptes guttatus M. H. C. Lichtenstein, 1818. In addition, Dendrocolaptes guttatus Swainson, 1837, is also
invalid because it is a junior objective synonym (Art. 61.3.4) of Dendrocolaptes ocellatus Spix, 1824, given the
fact its holotype (Spix 1824: 91, figure 1, ZSM B96) is also the holotype of Dendrocolaptes ocellatus. Regarding
Art. 67.13, because the designation of the type species is subsequent (Art.67.13.2), the action is transferred to the
already mentioned Art. 69.2.4.
But even if Art. 70.3 was applicable, following a correct interpretation of the Code, only Dendrocolaptes
guttatus M. H. C. Lichtenstein, 1818 (the nominal species) and Dendrocolaptes ocellatus Spix, 1824 (taxonomic
species) could be fixed as the type species of Dendroplex, and not Oriolus picus J. F. Gmelin, 1788, the putative
intention of Swainson when describing his new genus according to Aleixo et al. (2007). Its use highlights a highly
specific misinterpretation that can occur with respect to Art. 70.3.2 of the Code. This article states that “If an
author discovers that a type species was misidentified… the author may select, and thereby fix as type species, the
species that will, in his or her judgment, best serve stability and universality, either: the nominal species previously
cited as type species [Arts. 68, 69], or the taxonomic species actually involved in the misidentification…”.
As stated above, Aleixo et al. (2007) defended that the species illustrated by the bill and re-description of
Swainson (1837) was, in fact, Oriolus picus J. F. Gmelin, 1788, and that this is the taxonomic species actually
involved in the misidentification (Art. 70.3.2). In this case, it appears that the article was not perfectly understood
by the authors. Art. 70.3 refers to misidentified type species of genera. Unlike issues regarding types of nominal
species, the type species of genera are not based directly on specimens but on nominal species, i.e. the types of
Zootaxa 4532 (4) © 2018 Magnolia Press
genera are nomenclatural, not physical, entities. Therefore, when the Code establishes the options to fix a given
genus it invariably points to the nominal species (Dendrocolaptes guttatus) or the taxonomic species directly
involved in the case of a misidentification. The taxonomic species in the present case has no direct relation to the
description presented by the author but to the precise identity of the cited nominal species. Following this
reasoning, the nominal species previously cited by Swainson (1837) is Dendrocolaptes guttatus (Art. 70.3.1) and
the taxonomic species actually involved in the misidentification” (Art. 70.3.2) of the nominal species is
Dendrocolaptes ocellatus Spix, 1824. The taxonomic species referred to under Art. 70.3.2 is necessarily attached to
the nominal species (Art. 70.3.1) and to no other aspect of the description (figures, author’s intention, given
characters etc.). The description and the illustration of the bill presented by Swainson (1837) are not included in the
meaning of Art. 70.3, and lack any nomenclatural value. The nomenclatural act by Swainson (1837) was strictly to
fix the type species of the genus, while the original description of the genus is Swainson (1827b). Therefore, in
both cases (Dendrocolaptes guttatus M.H.C. Lichtenstein, 1818 [Art. 70.3.1] or Dendrocolaptes ocellatus Spix,
1824 [Art. 70.3.2]), in the case of application of Art. 70.3, the nomen Dendroplex would be necessarily fixed to a
species currently included in the genus Xiphorhynchus Swainson, 1827a. Interpretation of the historical reasons
that led Swainson (1837) to mention the plate of D. guttatus (Spix, 1824, currently D. ocellatus) and not to make
direct reference to his O. picus, are subjective and less important when the Code is rigorously followed, especially
when we consider that it was expressly constructed to deal with such cases.
Therefore, considering that the type species of the genus Dendroplex Swainson, 1827b (published between
September and 31 December 1827) is Dendrocolaptes ocellatus Spix, 1824 (under Art. 69.2.4, 70.4.2 or even if
Art. 70.3.2 is applied) and that the type species of the genus Xiphorhynchus Swainson, 1827a (published on 1 June
1827) is Xiphorhynchus flavigaster Swainson, 1827a (an originally included nominal species subsequently fixed as
the type of the genus Xiphorhynchus by Oberholser, 1905), the genus Dendroplex Swainson, 1827b must be
considered a junior synonym of Xiphorhynchus Swainson, 1827a. Because no generic nomen is currently available
for the species comprising the group of Dendroplex picus (see Aleixo 2002), we propose the following:
Paludicolaptes genus nov.
Type species: Dendrornis kienerii Des Murs, 1855.
Diagnosis: The genus Paludicolaptes differs immediately from those genera in its putative sister clade
(Campylorhamphus and Lepidocolaptes, see Aleixo 2002) by its straight and strong bill, and from all other
Dendrocolaptidae by the combination of a straight pale bill and whitish pectoral streaks broadly bordered black or
dark brown that generally do not reach the abdomen. It conforms to the usual pattern of Dendrocolaptidae syrinx
and skull, differing from other families by presenting a combination of the following characters: Cartt. accessoriae
named Processus vocalis with lateromedial projections (horns of the Processi vocales); strong reduction of the
tracheal rings (elements) of the Mem. tracheosyringealis region; two pairs of intrinsic muscles (M. vocalis ventralis
and M. vocalis dorsalis); and holorhine nares (see Feduccia 1973). According to Raposo et al. (2006),
Paludicolaptes differs from other woodcreeper genera (except Xiphorhynchus, Campylorhamphus and
Lepidocolaptes) by details of the syringeal elements. Paludicolaptes presents well-marked A elements in the
tracheosyringeal membrane, also different from Hylexetastes, Drymornis and Xiphocolaptes where the elements
are almost invisible; in Dendrocincla, the B4 element has a larger diameter in relation to B1, B2 and B3, which
have identical sizes; Hylexetastes has the A2 element greatly developed and A3 almost absent, which modifications
are not observed in Paludicolaptes. From Nasica, Paludicolaptes differs immediately by the number of T elements
(vide Raposo et al. 2006)—12 in Nasica and six in Paludicolaptes. From Deconychura, Certhiasomus and
Sittasomus, Paludicolaptes differs in the number of visible elements in the Membrana trachealis, five in those
genera and six in Paludicolaptes. Paludicolaptes also differs from Sittasomus and Glyphorynchus by the
proportions of the A elements sited caudally on the Membrana trachealis. Paludicolaptes differs from
Dendrocolaptes by the extremely large “horns” of the Processi vocales in the latter. The species are further
characterized by their habitat choice, shunning the interior of humid forests, in favour of deciduous types, including
mangrove and desert formations, where they breed in cacti. Coastal populations inhabit islands in river deltas and
mangrove, and, where associated with forested environments, they clearly prefer borders and gallery forests.
Zootaxa 4532 (4) © 2018 Magnolia Press
Phylogeny: Monophyly of the genus Paludicolaptes is corroborated by the molecular phylogeny recovered by
Aleixo (2002) which placed it as sister group to that comprising the genera Campylorhamphus and Lepidocolaptes
(see Aleixo 2002).
Etymology: L. paludicola, “marsh-dweller” < palus, paludis, “swamp”; cola, “dweller” < colere “to inhabit”;
Gr. Κολαπτης, kolaptēs, “chiseller” < κολαπτω, kolaptō, “to chisel, to peck, to strike”. This nomen points to the
ability of the species P. picus and, especially, of its type species P. kienerii, to occupy water-associated habitats, like
igapós. Paludicolaptes picus is also commonly present in mangroves and gallery forests.
Grammatical gender: Masculine.
Species included: Based on our taxonomic analysis, Paludicolaptes contains at least three species, those
already widely considered valid, namely Paludicolaptes picus (J. F. Gmelin, 1788) and Paludicolaptes kienerii
(Des Murs, 1855), as well as Paludicolaptes picirostris (Lafresnaye, 1847). Although the last species has been
considered a subspecies in virtually all of the literature and species lists since Peters (1951), no formal revision has
justified this treatment. A more comprehensive review points to its validity as a species (Raposo et al. in prep.).
Concluding remarks
It is important to bear in mind an important fact. Under the Code, the taxonomic allocation of nomina, the second
floor of the ‘nomenclatural house’ (Dubois 2005), is not made by intension, i.e. through definitions based on
characters, as in other nomenclatural systems such as the Phylocode (see Dubois 2005), but by ostension, directly
via the type specimen(s). Article 70.3 deviates from this general and clear philosophy of the Code in affording pre-
eminence to taxonomic interpretation of the status of nomina over a strict nomenclatural interpretation. We
consider that this Rule, like others in the Code (see Dubois 2011), tends to weaken the internal consistency of the
Code and in so doing sends a wrong message to taxonomists regarding its epistemological foundation, with
deleterious consequences for all zoological nomenclature (see Dubois 2010). For the time being, however, this
Rule forms part of the Code and should be followed, but very strictly, by following not only its ‘spirit’ but also the
‘letter’. In other words, the term ‘misinterpretation’ should not be used in a loose sense, meaning ‘different
opinions’, but only where misinterpretation is a demonstrable fact, which can be the case only when the original
type specimen of a nominal taxon is extant. This does not apply in the present case, where we should strictly abide
by the Code: the type species of a genus is the nominal species cited as such in the work where it was fixed.
We are indebted to CNPq for support given to MAR for his project ‘Catálogo dos tipos de espécies de aves
brasileiras’ (310384/2017-6) and to FAPESP for financial support of to post-doctoral fellowship by RS (2013/
26609-1 and 2016/18963-8) and CPA (2014/10914-2). We thanks M. Unsöld and J.-F. Voisin for their help in the
analysis of the type specimens at Munich and Paris Museums of Natural History. We are also grateful to
Alessandro Minelli and two anonymous reviewers for their constructive comments, which helped us to improve the
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The scientific names (nomina) of higher-ranked taxa (above the superfamily) of animals are not regulated by the International Code of Zoological Nomenclature, but by "consensus" among workers. However, when there exists no real consensus, a frequent situation, some criteria must be used to establish which nomen should be considered valid for any given taxon. With the multiplication of taxa that follows the development of cladistic analyses, the implementation of such rules will become more and more necessary and important. To be acceptable by all zoologists worldwide, today and tomorrow, these rules should be independent from the philosophy of taxonomy adopted, but should allow unambiguous, automatic and universal allocation of a single nomen to each higher taxon, within the frame of any taxonomy, including "phylogenetic" ones. This first paper is devoted to the detailed discussion of general theoretical and terminological problems related with this question. It is here argued that it is misleading and dangerous to try and make nomenclature artificially "simple". The problems posed by the naming of millions of kinds of organisms, related through evolution and that have been studied for two and a half century under different approaches, are indeed complex: this complexity should be acknowledged, and the discipline in charge of this study should be recognized as a specific technical field, with its own methods, concepts and terms. Among various proposals made in this paper, it is suggested to definitely abandon the misleading term "type" in taxonomy and nomenclature, objective categories for the "usage" of nomina are defined for the first time, and a distinction is made between taxonomic "categories" and nomenclatural "ranks". © Publications Scientifiques du Muséum national d'Histoire naturelle.
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At the beginning of the century of extinctions, science has only inventoried a very small proportion of the living species of the globe. In order to face the taxonomic urgency that results from this taxonomic gap combined with the biodiversity crisis, zootaxonomy needs efficient, rigorous and automatic nomenclatural Rules, that allow to spend a minimal time on nomenclatural problems—rather than investing time, energy and money in renaming millions of already named taxa in order to follow alternative nomenclatural systems, e.g., “phylogenetic” ones, that furthermore do not show theoretical superiority to the current Linnaean-Stricklandian one. The current Code, result of a 250-year improvement process, is based on very sound and healthy Rules, being theory-free regarding taxonomy, relying on objective allocation of nomina to taxa by a system of ostension using onomatophores, and on an objective basic Principle, priority, for recognizing the valid nomen of a taxon in case of synonymy or homonymy. Nevertheless, this nomenclatural system is certainly not perfect. It should be modified at least in nine directions: (1) it should adopt a technical terminology avoiding possible misinterpretations from outsiders of the field and even from specialists, and allowing a precise formalisation of its mode of functioning; (2) its plan should be drastically modified; (3) its Principles should be redefined, and some added; (4) material evidence for the allocation of nomina to taxa through specimens deposited in permanent collections should be given more weight; (5) it should incorporate all nomina of higher taxa, providing clear and strict universal Rules for their naming, whereas conserving the traditional nomina largely used in non-specialized systematic literature; (6) it should allow for the recognition of many more ranks at lower nomenclatural levels, i.e., just above genus, between genus and species, and below species; (7) it should provide much more stringent Rules for the protection against priority of “wellknown” nomina or sozonyms; (8) various “details” should be addressed, various Rules and Recommendations changed before a new edition of the Code is published; (9) the procedure of implementations of changes in the Code should be modified in order to involve zootaxonomists of the whole world in the decisions. In several instances, the Rules of the Code should become much more compulsory for all zoologists, editors and publishers, to avoid the publication of endless and sometimes most detrimental discussions among taxonomists which give a poor image of nomenclature and taxonomy among the biological sciences, such as bitter discussions about the “best” nomen to be used under a so-called “usage” philosophy, or about nomina to be applied to higher taxa. Code-compliance in zootaxonomic publications should be highlighted, and editors and publishers should require from authors who follow alternative nomenclatural Rules (or no rule at all) to make it clear by using particular modes of writing their nomina. It is argued here that if the Code of the 21st century does not evolve to incorporate these changes, it will prove unable to play its role in front of several important recent theoretical and practical developments of taxonomy and run the risk of being abandoned by a part of the international community of zootaxonomists. The latter could then adopt alternative “phylogenetic” nomenclatural Rules, despite the severe practical problems and theoretical flaws posed by such projects. This would be most detrimental for all comparative biological disciplines including systematics, and even for the unity of biology. In the course of this discussion, a few recommendations are given concerning the standards and guidelines suggested by recent authors for a good, modern, integrative taxonomy.
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Since 1951, various authors (Peters 1951, Clements 2000, Marantz et al. 2003, Dickinson 2003) have placed the taxa based on Oriolus picus J. F. Gmelin, 1788 (Straight-billed Woodcreeper), and Dendrornis kienerii Des Murs, 1856 (Zimmer's Woodcreeper), in the genus Xiphorhynchus Swainson, 1827 (Dendrocolaptidae). Earlier authorities (Sclater 1890, Hellmayr 1925, Zimmer 1934, Todd 1948) used Dendroplex Swainson, 1827, for these taxa. Swainson's original characterisation of Dendroplex (1827: 354) provided only a brief diagnosis of the new taxon ('Rostrum rectissimum. Alae mediocres, rotundatae; remigibus 3 tiá , 4 tá , et 5 tá longissimis'), and no reference to a type species, as he explicitly acknowledged being unsure whether the specimen upon which he based the new genus, had already been described as a species or not. He further stressed the diagnostic bill shape of the new genus and its close affinity with other Dendrocolaptid genera, stating: 'The living bird has all the manners of a Picus. Except in its perfectly straight bill, it differs not from Dendrocolaptes'. Ten years later, Swainson (1837: 313–314) provided essentially the same diagnosis of the original description: 'Bill moderate, very straight, perfectly conic in profile; the sides much compressed.' This time the diagnosis was accompanied by a depiction of the straight culmen and lateral compression (Fig. 1). But, at the end of the characterisation, Swainson added: 'The scansorial type D. guttatus Spix i, 91, f. 1', which refers to fig. 1 on pl. 91 in Spix (1824). The term scansorial here relates to Swainson's quinarian concept of re-occurring types within the animal kingdom; nevertheless, the fact remains that he included a single nominal species, thereby satisfying the requirements of Art. 69.3 for designation by subsequent monotypy (ICZN 1999). In his review of the Dendrocolaptidae, Hellmayr (1925: 288) pointed out that Swainson's diagnoses of 1827 and 1837, and bill outline (Fig. 1), correspond to the characters of Oriolus picus, though the only species mentioned (D. guttatus Spix i, 91, f. 1) 'belongs to the genus Xiphorhynchus Swainson'. Hellmayr continued that Swainson obviously followed Lesson (1830: 313) in misidentifying Spix's plate with Oriolus picus which, he believed, had to be regarded as the genotype of Dendroplex, being the only species then known with these generic characters. Hellmayr (1925) referred readers also to Gray (1840: 17), and Lafresnaye, Rev. Mag. Zool. (2)2, 1850, p. 959, where they merely grouped O. picus J. F. Gmelin, 1788, in Dendroplex without providing valid genus type designations. Alexandre Aleixo et al. 242 Bull. B.O.C. 2007 127(3) boc1273-070716.qxp 7/16/2007 10:05 AM Page 242 Following Hellmayr (1925), Peters (1951: 36) recognised that 'D. guttatus Spix i, 91, f. 1' depicts a bird now known as Xiphorhynchus ocellatus (Spix 1824), and stressed that under Opinion 65 (Schenk & McMasters 1948: 54) the case of misidentification had to be formally presented to the ICZN for ruling, and that until a decision was reached, Xiphorhynchus ocellatus ocellatus = Dendrocolaptes ocellatus Spix 1824 must continue as the type of Dendroplex. Aleixo (2002) demonstrated, using molecular data, that the genus Xiphorhynchus (sensu Peters 1951) is paraphyletic, and that the sister taxa X. picus = Oriolus picus J. F. Gmelin, 1788, and X. kienerii = Dendrornis kienerii Des Murs, 1856, are the only species which do not belong to a clade with the remaining Xiphorhynchus species. He suggested that they might be grouped in a different genus, in which case the name Dendroplex Swainson, 1827, would be available if problems with its type species designation could be resolved. The fourth edition of the International code of zoological nomenclature (ICZN 1999) states: 67.2.2. If a nominal genus or subgenus was established before 1931… without included nominal species…, the nominal species that were first subsequently and expressly included in it are deemed to be the only originally included nominal species.
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Syrinx anatomy of the Dendrocolaptidae (Aves, Passeriformes). The syrinx is the vocal organ of the birds and it corresponds to a modification of the apparatus respiratorius, being at most of the cases, localized at the caudal end of the trachea and the cranial portion of the bronchi. The main purpose of this paper is to describe the syrinx anatomy of the Dendrocolaptidae, with the analysis of 11 from the 13 genera accepted to this family. Besides the two series of cartilaginous elements ("A" and "B") previously recognized, we adopted a third series of elements designated as "T" elements, which correspond to those cranial to the Membrana tracheosyringealis. The structural variation of the Dendrocolaptidae syrinx is considered small when compared with other well known suboscine taxa, such as the Pipridae. We found out intra-group diagnostic characters, as well as some phylogenetically informative characters. Contrary to the current knowledge, the woodcreeper's syrinx is dorsoventrally assimetric, a characteristic that should be taken into account in future studies of other suboscine groups, especially in the case of using syringeal anatomy for systematic studies.
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Se reconstruyó la filogenia de todas las especies conocidas y de muchas de las subespecies de Xiphorhynchus (Dendrocolaptidae) para evaluar los límites de las especies en este género taxonómicamente complejo y para investigar el rol del ecotono entre “várzea” (bosque de inundación) y “terra-firme” (bosque de tierras altas) del Amazonas en su diversificación. Las filogenias fueron inferidas a partir de 2,430 pares de bases de los genes de ADN mitocondrial ND2, ND3 y citocromo b. Todas las estimaciones filogenéticas avalaron la monofilia de todas las especies vivientes de Xiphorhynchus, con excepción del par de especies hermanas X. picus y X. kienerii. Se encontró fuerte respaldo para incluir a Lepidocolaptes fuscus en Xiphorhynchus, confirmando estudios moleculares y anatómicos previos. Los niveles de divergencia en las secuencias entre algunas subespecies de X. guttatus, X. ocellatus y X. spixii alcanzaron o excedieron aquellos encontrados entre especies biológicas cercanamente emparentadas de Xiphorhynchus. Los altos niveles de diferenciación en las secuencias y la parafilia de algunas especies de Xiphorhynchus indicaron que los siguientes taxones deberían ser reconocidos como especies: X. guttatoides, X. chunchotambo y X. elegans. Todas las especies de Xiphorhynchus restringidas a las áreas de bosque de terra-firme de las tierras bajas del Amazonas formaron un grupo monofilético fuertemente respaldado, mientras que las especies restringidas a bosques de várzea aparecieron en la base del clado que contenía a aquellas encontradas en una amplia variedad de hábitats (X. obsoletus) o pertenecieron a un linaje separado que probablemente pueda ser considerado como un género separado (X. kienerii). Estos resultados falsifican la relación de hermandad esperada entre las especies de várzea y terra-firme que se esperaría si el ecotono de várzea y terra-firme hubiera jugado un rol importante en la diferenciación entre poblaciones y en la especiación de Xiphorhynchus. En cambio, las estimaciones filogenéticas sugirieron que la especialización de hábitat de várzea y terra-firme evolucionó temprano en la historia evolutiva de Xiphorhynchus y que las diferenciaciones subsecuentes ocurrieron principalmente en el hábitat de terra-firme.
The phylogeny of all known Xiphorhynchus (Dendrocolaptidae) species and many of its subspecies was reconstructed to evaluate species limits in this taxonomically challenging genus and investigate the possible role played by the Amazonian “várzea” (floodplain forest)–“terra-firme” (upland forest) ecotone in its diversification. Phylogenies were inferred based on 2,430 bp of the mitochondrial DNA genes ND2, ND3, and cytochrome b. All phylogeny estimates supported the monophyly of all extant Xiphorhynchus species to the exclusion of the sibling species pair Straight-billed (X. picus) and Zimmer's (X. kienerii) woodcreeper. Confirming findings of previous molecular and anatomical studies, strong support was found to include the Lesser Woodcreeper (Lepidocolaptes fuscus) in Xiphorhynchus. Levels of sequence divergence among some subspecies of Buff-throated (X. guttatus), Ocellated (X. ocellatus), and Spix's (X. spixii) woodcreepers reached or exceeded those found between closely related, undisputed biological species of Xiphorhynchus. High levels of sequence differentiation and the paraphyly of some Xiphorhynchus species indicated that the following taxa should be recognized as species: Lafresnaye's (X. guttatoides), Tschudi's (X. chunchotambo), and Elegant (X. elegans) woodcreepers. All Xiphorhynchus species restricted to terra-firme forest in lowland Amazonia formed a well supported monophyletic group, whereas species restricted to várzea forest were either basal to a clade containing species found in a wide variety of habitats (Striped Woodcreeper [X. obsoletus]) or belonged to a distinct lineage likely to be regarded as a separate genus (X. kienerii). These findings falsified an anticipated sister relationship between várzea and terra-firme species, as expected if the várzea–terra-firme ecotone had played a decisive role in population differentiation and speciation within Xiphorhynchus. Instead, phylogeny estimates suggested that the várzea–terra-firme habitat specialization evolved early on in the evolutionary history of Xiphorhynchus and that subsequent differentiation occurred mostly within the terra-firme habitat.
Taxonomy is currently facing a major crisis and is likely to have strong difficulties to reduce significantly the taxonomic gap before the biodiversity crisis has wiped out a large proportion of the living species of the earth. In this context, taxonomists should pay great attention to the nomenclatural Rules, and care for them to help them in this urgent task, rather than diverting their time and energy to secondary or useless questions or debates. A major purpose of the Code is to promote nomenclatural stability in zoology. This requires stability in the Rules, or at least that a great care be taken, when establishing new Rules, to avoid that they can have unexpected deleterious consequences for stability. In particular, in most cases, it is crucial to deny retroactivity to the new Rules. Several examples of problems created in zoological nomenclature by introduction of changes in Articles dealing with the spellings of nomina are examined in detail. These Articles were modified, with retroactive value, in the 1985 edition (Art. 32, 33, 35 and 39) and in the 1999 edition (Art. 24) of the Code. It is shown that these changes, which have no clear "philosophical" or practical justifications and which result in no clear benefits, have in fact had negative impacts on nomenclatural practice. Their implementation requires heavy useless additional work from taxonomists and has negative results in nomenclatural stability that had clearly not been anticipated by the ICZN when promulgating them. In a few sets of nomina tested below, the changes in the 1985 edition resulted in spelling changes for 10.0 to 22.2 % of the nomina, and those in the 1999 edition for 21.7 to 33.3 % of the nomina, roughly a quarter of them on the whole (24.5 %). Among others that are less emblematic, a striking case is that of the fish generic nomen Tetraodon, widely used especially since the genome of a species of this genus has been sequenced, and which should be changed to Tetrodon because of the unwarranted introduction of the new Art. 24.2.4 into the Code. It is suggested that these changes should be cancelled, or at least denied retroactivity from the years of their promulgations. In order to make this discussion easier, a "taxonomy" of the different kinds of spellings of nomina, and a dichotomic key to such situations, are provided. This stresses the fact that detailed discussions on very precise aspects of the functioning of nomenclatural Rules, as well as the computerization of nomenclatural data for online databases, require to use a specialized technical terminology to designate the nomenclatural concepts and tools, not vague "common language" terms like "name" or "type": "keep the Rules, but change the terms". The problems outlined here should be kept in mind by the ICZN before implementing drastic changes in the Rules of nomenclatural availability, as recently suggested.