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Article Herpetological higher taxa nomina. 8. Amphibia Blainville, 1816
Abstract
Under the Duplostensional Nomenclatural System, the valid nomen of the class including all recent amphibians and all the Palaeozoic groups of anamniote tetrapods subsequent to the ‘lissamphibian-amniote phylogenetic split’ is the sozodiaphonym Amphibia Blainville, 1816. This corresponds to the usage that has been in force for two centuries in thousands of publications, and it should not be challenged, as this would entail instability and confusion.
... This paper is the second one of a series of articles devoted to the nomina of the higher taxa (above the rank superfamily) of amphibians and 'reptiles' as established by the Duplostensional Nomenclatural System (DONS). The first paper of this series (Dubois & Frétey 2020) provides a presentation of this nomenclatural system, of its rationale and methodology, and a Glossary of its technical nomenclatural terms, that should be consulted for a full understanding of what follows. ...
... Our purpose here is to establish the valid nomen, with its authorship and date, of this taxon under DONS-2 Criteria (Dubois & Frétey 2020). This means that what we call here, as well as in all the forthcoming papers of this series, the 'authorship and date' of a class-series nomen, is that of the first work where this nomen was used with a given intension (circumscription), irrespective of the authorship credited (or not) to this nomen in subsequent works. ...
A taxon, traditionally referred to the rank order, encompassing all recent taxa of frogs and their close fossil relatives, is highly supported as holophyletic in all recent phylogenetic analyses of Amphibia.
Under the Duplostensional Nomenclatural System, among more than thirty nomina available for this taxon in the literature, two only qualify as sozodiaphonyms: Anura Duméril, 1805 and Salientia Merrem, 1820. The conflict for validity between these two nomina is easily solved by publication dates. The valid nomen for this taxon is therefore Anura Duméril, 1805, which corresponds to the overwhelming majority of uses in the literature, and which should replace all other nomina, authorships and dates sometimes credited to this taxon in publications, catalogues and databases.
... Exomonyms are homonyms between nomina that designate taxa referred to different higher taxa, such as Caudata in insects and in amphibians (see Dubois & Frétey 2020b), Apodes in fishes and Apoda in amphibians (see Dubois & Frétey 2020c) or Amphibia in amphibians and in mammals (see Dubois & Frétey 2020f). This kind of homonymy is similar to that which occurs in the three nominalseries covered by the Code. ...
This work introduces a series of papers devoted to the ascertainment of the valid nomina, under the
revised Duplostensional Nomenclatural System (DONS-2), of the highest taxa (from the rank order to
the rank class) of recent amphibians and ‘reptiles’. The main general features of the DONS approach
to class-series nomenclature (above the rank superfamily) are explained in detail, the DONS-2
procedure for ascertaining the availability, allocation, validity and correctness of class-series nomina
is described, and a Glossary of all the technical nomenclatural terms used in this work is provided.
A taxon, traditionally referred to the rank order, encompassing all recent taxa of frogs and their close fossil relatives, is highly supported as holophyletic in all recent phylogenetic analyses of Amphibia. Under the Duplostensional Nomenclatural System, among more than thirty nomina available for this taxon in the literature, two only qualify as sozodiaphonyms: Anura Duméril, 1805 and Salientia Merrem, 1820. The conflict for validity between these two nomina is easily solved by publication dates. The valid nomen for this taxon is therefore Anura Duméril, 1805, which corresponds to the overwhelming majority of uses in the literature, and which should replace all other nomina, authorships and dates sometimes credited to this taxon in publications, catalogues and databases.
In July 2014, the international meeting ‘Burning questions and problems of zoological nomenclature’ was held in Linz (Austria). It acknowledged the presence in the current International Code of Zoological Nomenclature of a number of severe problems, and accordingly decided the creation of a new international body, the Linz Zoocode Committee (LZC), in charge of writing the Linz Zoocode, a set of new proposals regarding the terminology, the Principles and Rules of zoological nomenclature. Here we present the first report of the activities of this Committee, covering the period 2014‒2019. It contains the presentation of our work, and the first documents adopted by the Committee: the Preamble and Principles of the Zoocode, the description of the structure and a first instalment of the Zoocode Glossary. The Zoocode regulates the status of zoological nomina and nomenclatural acts (onomatergies). Its aim is to provide an explicit, precise and objective nomenclatural system for the unambiguous and universal naming of all zoological taxa recognised by taxonomists, so that, in the frame of a given classification, the nomen of each taxon is unique and distinct. It relies on a Nomenclatural Process consisting in four main stages: nomenclatural assignment and availability, taxonomic allocation, nomenclatural validity and correctness, and registration of nomina and onomatergies. Whereas the Code currently in force is based on six stated Principles, the Zoocode recognises 17 distinct ones. We here submit these documents to the consideration of the international community of zootaxonomists, in the perspective of the incorporation of these proposals into the next version of the Code.
The largest published morphological data matrix for analysis of the phylogeny of Paleozoic limbed vertebrates (Ruta M, Coates MI. 2007. Journal of Systematic Palaeontology 5:69–122) has supported a number of phylogenetic hypotheses that are, to varying degrees, controversial; for example, it has recovered Seymouriamorpha, Diadectomorpha and (in some trees) Caudata as paraphyletic and found the “temnospondyl hypothesis” on the origin of the modern amphibians (TH) to be more parsimonious than the “lepospondyl hypothesis” (LH) – although we show that the difference in tree length between these hypotheses, given this matrix, is only one step. A thorough investigation of the matrix reveals thousands of suboptimal scores, of which many are clearly due to typographic and similar errors, and many questionable coding decisions: there are logically linked (redundant) characters, binary characters of which only one state is described (the “nonadditive binary coding” approach to inapplicable scores), and even characters where many taxa were scored after their presumed relatives. Furthermore, all characters – even the most obviously continuous ones – were unordered, the effects of ontogeny were not sufficiently taken into account, and data published after 2001 were mostly excluded despite occasional overlaps in authors between those publications and the matrix discussed here.
After these issues are improved – we document and justify all changes to the matrix –, our revised version of the matrix yields much longer trees with a different topology, for instance monophyletic Caudata, Diadectomorpha and (in some trees) Seymouriamorpha, Ichthyostega more rootward than Acanthostega, Anthracosauria more rootward than Temnospondyli which includes Caerorhachis, and the modern amphibians nested among “lepospondyls”; the LH is 10 steps more parsimonious than the TH and 15 steps more than the “polyphyly hypothesis” (PH), as determined by comparing an unconstrained to two constrained analyses. Bootstrap values, however, are low; only the LH and the PH are distinguishable at p = 0.05, and even that only according to one of three statistical tests.
For a separate set of (again three) analyses, we added 48 OTUs to the original 102. This has a destabilizing effect on some parts of the tree, such as the interrelationships of Anthracosauria, Temnospondyli and Caerorhachis. Yet, many of the added taxa have a fully resolved position or nearly so; this concerns the well known Chroniosaurus (representing Chroniosuchia), which is found as the sister-group to a clade that contains Solenodonsaurus, Seymouriamorpha, Diadectomorpha, Amniota, the “lepospondyls” and the modern amphibians, but at the other extreme is also true for isolated lower-jaw material (notably Metaxygnathus, Densignathus, Sigournea and Doragnathus). Despite the addition of the temnospondyls Gerobatrachus, Micropholis and Tungussogyrinus and the extremely peramorphic salamander Chelotriton, the difference between the LH and the TH only shrinks to 9 steps, and that between the LH and the PH to 13 steps. Among the “lepospondyls”, interestingly, Brachydectes (Lysorophia) is not found lying next to Lissamphibia. However, bootstrap values plummet, and all three hypotheses become indistinguishable at p = 0.05.
We then duplicated all six analyses after coding all losses of bones as irreversible. The impact on the results is modest. Anthracosauria is consistently placed more rootward than Temnospondyli; given the original taxon sample, the LH is 12 steps shorter than the TH and 17 steps shorter than the PH, which alone is statistically distinguishable from the LH, while the analyses of the expanded taxon sample recover the LH as 10 steps shorter than the TH and only 12 steps shorter than the PH, all three being statistically indistinguishable.
We think that more robust results could be obtained by adding the many characters used in other analyses or discussed in the literature. We discuss the phylogenetic relationships of certain taxa, approaches to coding, and certain character complexes, in particular the supposed middle ear of temnospondyls.
The absence of rules in the International Code of Zoological Nomenclature for nomenclature of taxa above superfamily is a source of instability and confusion, especially with the recent increase in number of higher taxa following multiplication of phylogenetic analyses. A recent proposal concerning such rules, submitted elsewhere, is briefly presented here, and its consequences regarding nomenclature of higher taxa of recent amphibians are summarised. The class nomen AMPHIBIA should be credited to DE BLAINVILLE (1816) instead of LINNAEUS (1758). The nomen LISSAMPHIBIA Haeckel, 1866 is an invalid junior synonym of BATRACHIA Brongniart, 1800, that applies to one of the superorders of the subclass including all recent amphibians. The valid nomen of this subclass is NEOBATRACHI Sarasin & Sarasin, 1890. The three orders of recent amphibians should be known as ANURA Duméril, 1806, URODELA Duméril, 1806 and GYMNOPHIONA Rafinesque-Schmaltz, 1814. The nomina SAUENTIA Laurenti, 1768, CAUDATA Scopoli, 1777, APODA Oppel, 1811, ARCHAEOBATRACHIA Reig, 1958 and NEOBATRACHIA Reig, 1958 are invalid and should no longer be used.
I recently published a survey of the different meanings of the nomen Amphibia in taxonomic publications since 1758 (Dubois 2015). The 'meaning' of a nomen in zoological nomenclature depends on the system used for the allocation of nomina to taxa, and several such systems can be used (see e.g. Dubois 2006a-b). In the paper at stake, I used the 'orostensional nomenclatural system' (OONS) for class-series nomenclature. In this system, a class-series nomen-i.e., a nomen above the rank superfamily, therefore one whose taxonomic allocation is not regulated by the Code (Anonymous 1999)-applies, in a given classification, to the most inclusive class-series taxon that includes all its originally expressly included nominal genera (conucleogenera) and excludes all its originally expressly excluded nominal genera (alienogenera)-if such a taxon indeed exists in this classification. However, if one a least of the alienogenera is now part of the most inclusive taxon including all the conucleogenera, the nomen cannot be taxonomically allocated and qualifies as an anaptonym in the classification used as reference (Dubois 2006a-b, 2011), although it may not be so under another taxonomic frame.
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.
After the critical study of a number of concepts and problems of zoological nomenclature and the proposal of terminological novelties, 24 formal Rules for nomenclature of higher-ranked taxa (i.e. for taxa above superfamily) are formulated, that are compatible with the current Code and could possibly be incorporated in the latter. These Rules concern: 1) the availability of nomina; 2) their allocation to taxa, through a set of generic nomina that together con-stitute the onomatophore of the nomen, and in some cases a set of generic nomina expressly excluded from the taxon designated by the nomen; and 3) their validity, based usually on priority, but in some cases taking into account widespread use of a nomen outside the field of systematics. RÉSUMÉ Proposition de Règles pour l'incorporation des nomina de taxons de rangs supérieurs d'animaux dans le Code international de nomenclature zoologique. 2. Les Règles proposées et leur justification théorique. Après l'examen critique d'un certain nombre de concepts et problèmes de la nomenclature zoologique et la proposition de nouveaux termes, 24 Règles formelles pour la nomenclature zoologique des taxons de rangs supérieurs (c'est-à-dire au-dessus du rang de superfamille) sont formulées. Celles-ci sont compatibles avec le Code actuel et pourraient être incorporées dans celui-ci. Ces règles concernent : 1) la disponibilité des nomina ; 2) leur attribution aux taxons, au moyen d'un ensemble de nomina génériques qui constituent ensemble l'onomatophore du nomen, et dans certains cas d'un ensemble de nomina génériques expressément exclus du taxon désigné par le nomen ; et 3) leur validité, qui repose en règle générale sur la priorité, mais qui dans certains cas prend en compte l'usage répandu d'un nomen en dehors du domaine de la systématique.
In zoological taxonomy, unambiguous and universal designation of a given taxon at a given rank in a given classification requires to establish its valid nomen and the correct spelling of the latter. Consequently, a need to resolve the situations of conflict that may exist between available nomina and spellings arises. These situations of nomenclatural conflict arehere designated as zygoidy. They are of three kinds: zygonymy (conflicts of homonymy or synonymy between nomina for validity), zygography (conflicts of parography between spellings for correctness) and zygophory (conflicts of airetophory between onomatophore designations for validity). The resolution of these conflicts relies on several basic Principles already formulated in the Code (the Principles of Homonymy, Priority and “First-Reviser”, here renamed “Airesy”), as well as additional ones not yet recognized in this text: the Principles of Zygoidy, Proedry, Nomography and
Sozoidy. A detailed analysis of these situations is provided. It allows to identify in which cases and under which conditions these Principles apply. This complete inventory of possible situations and solutions will be of great usefulness for a forecoming automatisation and computerization of the Rules of the Code in forthcoming databases and Web applications.
INTRODUCTION During a recent collecting trip, financed by grant W 956-2 from WOTRO (Netherlands Foundation for the Advancement of Tropical Research), I spent 47 days (22 August-7 October 1968) on the Sipaliwini savannah in southern Surinam. The Sipaliwini savannah is part of an extensive savannah that covers the watershed between the Sipaliwini River in Surinam and the Paroe River in Brazil. Though the watershed is very low, the waters of Sipaliwini and Paroe Rivers never seem to make contact, as is the case with other rivers on the Rupununi savannah and in southern French Guiana. About one tenth of this savannah is situated in Surinam, the remainder in Brazil (see inset fig. 3). G. Mann (in Fittkau et al., 1968) calls this a tropical wet savannah and describes the ecology of this habitat extensively. The soil of the Sipaliwini savannah is classified by Beek & Bramao (in Fittkau et al., 1968: 92) as belonging to Soil Region Β 1-Guiana Uplands, and is composed of Latosols and Red-Yellow Podzolic soils. This difference in soil composition with the coastal savannahs (with a sandy soil) is evident in the vegetation, while also there are differences in physiography. In the coastal savannahs the ground is covered with a sparse cover of grasses, but there are also many shrubs, up to two metres in height. The Sipaliwini savannah is nearly completely covered with grasses, only in the wet valleys Mauritia palms and on the hills a few widely spaced, low, crooked trees, never forming a closed canopy, are present. Shrubs are absent. In some places extensive bare rockslates are exposed and the top and slopes of quite a few hills are covered with large rocks. In contrast with the flat coastal savannahs the Sipaliwini savannah is very hilly, the highest point on Surinam
Determination of the proximate cause of sexual dimorphism remains difficult, especially when trying to discriminate between sexual selection and ecological causation. A clear rejection of one of these hypotheses would advance the direction of future investigations, especially for cryptic and/or subterranean species. Sexual dimorphism in head size, but not body size, is confirmed for Schistometopum thomense, a fossorial caecilian from the island of São Tomé, in the Gulf of Guinea. However, a quantitative and qualitative study of diet reveals no significant difference between males and females sampled from three sites on the island. Females appear to take larger and heavier prey (principally earthworms) than males, despite having a significantly smaller head size. We tentatively reject the ecological causation hypothesis, and discuss several testable hypotheses for evidence of sexual selection.
Based on recent morphological and molecular cladistic hypotheses, a
new ergotaxonomy (provisional classification) of the family Megophryidae is
presented, including three subfamilies, four tribes, ten genera and six
subgenera.
Five new species of Nectophrynoides Noble 1926 are described from the forests of the Eastern Arc Mountains of Tanzania. Two of the new species were already recognised as such, but not formally described. Two more were recog-nised as undescribed in the collection of the Natural History Museum, London. The fifth was collected during a field survey of the Udzungwa Scarp Forest Reserve in early 2003. A description of the advertisement call and some ecologi-cal information are provided for the latter species. Little is known of the ecology and behaviour of any of the other new species. An updated key to the species of Nectophrynoides is given. The high species diversity and pattern of endemism of Nectophrynoides suggests the genus is closely associated with the geological and climatic history of the Eastern Arc Mountains. Given the limited distribution of the new species and continuing habitat loss in the Eastern Arc Mountains (a globally recognised biodiversity hotspot), a rapid conservation assessment of the status of these species is necessary.
Muséum national d'Histoire naturelle, Département Systématique et Évolution, Reptiles et Amphibiens, USM 0602, case postale 30, 25 rue Cuvier, F-75231 Paris cedex 05 (France) ohler@mnhn.fr. 2006. — Phylogenetic relationships and generic taxonomy of the tribe Paini (Amphibia, Anura, Ranidae, Dicroglossinae), with diagnoses of two new genera. Zoosystema 28 (3) : 769-784. ABSTRACT A preliminary cladistic analysis of the relationships between 26 frog species of the tribe Paini (Ranidae, Dicroglossinae) was carried out on the basis of 31 morphological characters, mainly from external morphology of adults. Combined with the results of a molecular analysis published elsewhere, these data 1) confi rm that, after exclusion of the species Rana delacouri Angel, 1928, the Paini are a homophyletic group, and 2) allow to redefi ne the genera of this tribe, which are now six in number, including two new ones described herein. RÉSUMÉ Relations phylogénétiques et taxonomie générique de la tribu Paini (Amphibia, Anura, Ranidae, Dicroglossinae), avec diagnoses de deux nouveaux genres. Une analyse cladistique préliminaire des relations entre 26 espèces de grenouilles de la tribu des Paini (Ranidae, Dicroglossinae) a été menée en utilisant 31 caractères morphologiques, principalement de la morphologie externe des adultes. Combinées avec les résultats d'une analyse moléculaire publiée ailleurs, ces données 1) confi rment qu'après exclusion de l'espèce Rana delacouri Angel, 1928, les Paini constituent un groupe homophylétique, et 2) permettent de redéfi nir les genres de cette tribu, qui sont maintenant au nombre de six, dont deux genres nouveaux décrits ici.
The known diversity of dart-poison frog species has grown from 70 in the 1960s to 247 at present, with no sign that the discovery of new species will wane in the foreseeable future. Although this growth in knowledge of the diversity of this group has been accompanied by detailed investigations of many aspects of the biology of dendrobatids, their phylogenetic relationships remain poorly understood. This study was designed to test hypotheses of dendrobatid diversification by combining new and prior genotypic and phenotypic evidence in a total evidence analysis. DNA sequences were sampled for five mitochondrial and six nuclear loci (approximately 6,100 base pairs [bp]; x¯ = 3,740 bp per terminal; total dataset composed of approximately 1.55 million bp), and 174 phenotypic characters were scored from adult and larval morphology, alkaloid profiles, and behavior. These data were combined with relevant published DNA sequences. Ingroup sampling targeted several previously unsampled species, including Aromobates nocturnus, which was hypothesized previously to be the sister of all other dendrobatids. Undescribed and problematic species were sampled from multiple localities when possible. The final dataset consisted of 414 terminals: 367 ingroup terminals of 156 species and 47 outgroup terminals of 46 species. Direct optimization parsimony analysis of the equally weighted evidence resulted in 25,872 optimal trees. Forty nodes collapse in the strict consensus, with all conflict restricted to conspecific terminals. Dendrobatids were recovered as monophyletic, and their sister group consisted of Crossodactylus, Hylodes, and Megaelosia, recognized herein as Hylodidae. Among outgroup taxa, Centrolenidae was found to be the sister group of all athesphatanurans except Hylidae, Leptodactyidae was polyphyletic, Thoropa was nested within Cycloramphidae, and Ceratophryinae was paraphyletic with respect to Telmatobiinae. Among dendrobatids, the monophyly and content of Mannophryne and Phyllobates were corroborated. Aromobates nocturnus and Colostethus saltuensis were found to be nested within Nephelobates, and Minyobates was paraphyletic and nested within Dendrobates. Colostethus was shown to be rampantly nonmonophyletic, with most species falling into two unrelated cis- and trans-Andean clades. A morphologically and behaviorally diverse clade of median lingual process-possessing species was discovered. In light of these findings and the growth in knowledge of the diversity of this large clade over the past 40 years, we propose a new, monophyletic taxonomy for dendrobatids, recognizing the inclusive clade as a superfamily (Dendrobatoidea) composed of two families (one of which is new), six subfamilies (three new), and 16 genera (four new). Although poisonous frogs did not form a monophyletic group, the three poisonous lineages are all confined to the revised family Dendrobatidae, in keeping with the traditional application of this name. We also propose changes to achieve a monophyletic higher-level taxonomy for the athesphatanuran outgroup taxa. Analysis of character evolution revealed multiple origins of phytotelm-breeding, parental provisioning of nutritive oocytes for larval consumption (larval oophagy), and endotrophy. Available evidence indicates that transport of tadpoles on the dorsum of parent nurse frogs—a dendrobatid synapomorphy—is carried out primitively by male nurse frogs, with three independent origins of female transport and five independent origins of biparental transport. Reproductive amplexus is optimally explained as having been lost in the most recent common ancestor of Dendrobatoidea, with cephalic amplexus arising independently three times.
The extant amphibians are one of the most diverse radiations of terrestrial vertebrates (>6800 species). Despite much recent focus on their conservation, diversification, and systematics, no previous phylogeny for the group has contained more than 522 species. However, numerous studies with limited taxon sampling have generated large amounts of partially overlapping sequence data for many species. Here, we combine these data and produce a novel estimate of extant amphibian phylogeny, containing 2871 species (∼40% of the known extant species) from 432 genera (∼85% of the ∼500 currently recognized extant genera). Each sampled species contains up to 12,712 bp from 12 genes (three mitochondrial, nine nuclear), with an average of 2563 bp per species. This data set provides strong support for many groups recognized in previous studies, but it also suggests non-monophyly for several currently recognized families, particularly in hyloid frogs (e.g., Ceratophryidae, Cycloramphidae, Leptodactylidae, Strabomantidae). To correct these and other problems, we provide a revised classification of extant amphibians for taxa traditionally delimited at the family and subfamily levels. This new taxonomy includes several families not recognized in current classifications (e.g., Alsodidae, Batrachylidae, Rhinodermatidae, Odontophrynidae, Telmatobiidae), but which are strongly supported and important for avoiding non-monophyly of current families. Finally, this study provides further evidence that the supermatrix approach provides an effective strategy for inferring large-scale phylogenies using the combined results of previous studies, despite many taxa having extensive missing data.
In recent years, avian systematics has been characterized by a diminished reliance on morphological cladistics of modern taxa, intensive palaeornithogical research stimulated by new discoveries and an inundation by analyses based on DNA sequences. Unfortunately, in contrast to significant insights into basal origins, the broad picture of neornithine phylogeny remains largely unresolved. Morphological studies have emphasized characters of use in palaeontological contexts. Molecular studies, following disillusionment with the pioneering, but non-cladistic, work of Sibley and Ahlquist, have differed markedly from each other and from morphological works in both methods and findings. Consequently, at the turn of the millennium, points of robust agreement among schools concerning higher-order neornithine phylogeny have been limited to the two basalmost and several mid-level, primary groups. This paper describes a phylogenetic (cladistic) analysis of 150 taxa of Neornithes, including exemplars from all non-passeriform families, and subordinal representatives of Passeriformes. Thirty-five outgroup taxa encompassing Crocodylia, predominately theropod Dinosauria, and selected Mesozoic birds were used to root the trees. Based on study of specimens and the literature, 2954 morphological characters were defined; these characters have been described in a companion work, approximately one-third of which were multistate (i.e. comprised at least three states), and states within more than one-half of these multistate characters were ordered for analysis. Complete heuristic searches using 10 000 random-addition replicates recovered a total solution set of 97 well-resolved, most-parsimonious trees (MPTs). The set of MPTs was confirmed by an expanded heuristic search based on 10 000 random-addition replicates and a full ratchet-augmented exploration to ascertain global optima. A strict consensus tree of MPTs included only six trichotomies, i.e. nodes differing topologically among MPTs. Bootstrapping (based on 10 000 replicates) percentages and ratchet-minimized support (Bremer) indices indicated most nodes to be robust. Several fossil Neornithes (e.g. Dinornithiformes, Aepyornithiformes) were placed within the ingroup a posteriori either through unconstrained, heursitic searches based on the complete matrix augmented by these taxa separately or using backbone-constraints. Analysis confirmed the topology among outgroup Theropoda and achieved robust resolution at virtually all levels of the Neornithes. Findings included monophyly of the palaeognathous birds, comprising the sister taxa Tinamiformes and ratites, respectively, and the Anseriformes and Galliformes as monophyletic sister-groups, together forming the sister-group to other Neornithes exclusive of the Palaeognathae (Neoaves). Noteworthy inferences include: (i) the sister-group to remaining Neoaves comprises a diversity of marine and wading birds; (ii) Podicipedidae are the sister-group of Gaviidae, and not closely related to the Phoenicopteridae, as recently suggested; (iii) the traditional Pelecaniformes, including the shoebill (Balaeniceps rex) as sister-taxon to other members, are monophyletic; (iv) traditional Ciconiiformes are monophyletic; (v) Strigiformes and Falconiformes are sister-groups; (vi) Cathartidae is the sister-group of the remaining Falconiformes; (vii) Ralliformes (Rallidae and Heliornithidae) are the sister-group to the monophyletic Charadriiformes, with the traditionally composed Gruiformes and Turniciformes (Turnicidae and Mesitornithidae) sequentially paraphyletic to the entire foregoing clade; (viii) Opisthocomus hoazin is the sister-taxon to the Cuculiformes (including the Musophagidae); (ix) traditional Caprimulgiformes are monophyletic and the sister-group of the Apodiformes; (x) Trogoniformes are the sister-group of Coliiformes; (xi) Coraciiformes, Piciformes and Passeriformes are mutually monophyletic and closely related; and (xii) the Galbulae are retained within the Piciformes. Unresolved portions of the Neornithes (nodes having more than one most-parsimonious solution) comprised three parts of the tree: (a) several interfamilial nodes within the Charadriiformes; (b) a trichotomy comprising the (i) Psittaciformes, (ii) Columbiformes and (iii) Trogonomorphae (Trogoniformes, Coliiformes) + Passerimorphae (Coraciiformes, Piciformes, Passeriformes); and (c) a trichotomy comprising the Coraciiformes, Piciformes and Passeriformes. The remaining polytomies were among outgroups, although several of the highest-order nodes were only marginally supported; however, the majority of nodes were resolved and met or surpassed conventional standards of support. Quantitative comparisons with alternative hypotheses, examination of highly supportive and diagnostic characters for higher taxa, correspondences with prior studies, complementarity and philosophical differences with palaeontological phylogenetics, promises and challenges of palaeogeography and calibration of evolutionary rates of birds, and classes of promising evidence and future directions of study are reviewed. Homology, as applied to avian examples of apparent homologues, is considered in terms of recent theory, and a revised annotated classification of higher-order taxa of Neornithes and other closely related Theropoda is proposed. (c) 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 149, 1-95.
This edited volume is provides an authoritative synthesis of knowledge about the history of life. All the major groups of organisms are treated, by the leading workers in their fields. With sections on: The Importance of Knowing the Tree of Life; The Origin and Radiation of Life on Earth; The Relationships of Green Plants; The Relationships of Fungi; and The Relationships of Animals. This book should prove indispensable for evolutionary biologists, taxonomists, ecologists interested in biodiversity, and as a baseline sourcebook for organismic biologists, botanists, and microbiologists. An essential reference in this fundamental area.
Although currently most taxonomists claim to adhere to the concept of ‘phylogenetic taxonomy’, in fact most of the zoological classifications currently published are only in part ‘phylogenetic’ but include also phenetic or gradist approaches, in their arbitrary choices of the nodes formally recognised as taxa and in their attribution of ranks to these taxa. We here propose a new approach to ‘phylogenetic taxonomy and nomenclature’, exemplified by a phylogenetic classification or cladonomy of the extant amphibians (subclass Lissamphibia of the class Amphibia) derived from a supermatrix-based phylogenetic analysis using 4060 amphibian species, i.e. about half of the 8235 species recognised on 31 October 2020. These taxa were represented by a mean of 3029 bp (range: 197–13849 bp) of DNA sequence data from a mean of 4 genes (range: 1‒15). The cladistic tree thus generated was transferred into a classification according to a new taxonomic and nomenclatural methodology presented here, which allows a bijective or isomorphic relationship between the phylogenetic hypothesis and the classification through a rigorous use of suprageneric ranks, in which their hierarchy mirrors the structure of the tree. Our methodology differs from all previous ones in several particulars: [1] whereas the current International Code of Zoological Nomenclature uses only three ‘groups of names’ (species, genus and family), we recognise four nominal-series (species, genus, family and class); [2] we strictly follow the Code for the establishment of the valid nomen (scientific name) of taxa in the three lower nominal-series (however, in a few situations, we suggest improvements to the current Rules of the Code); [3] we provide precise and unambiguous Criteria for the assignment of suprageneric nomina to either the family- or the class-series, excluding nomina proposed expressly under unranked or pseudoranked nomenclatural systems; [4] in the class-series, for which the Code provides only incomplete Rules concerning availability, we provide precise, complete and unambiguous Criteria for the nomenclatural availability, taxonomic allocation and nomenclatural validity and correctness of nomina; [5] we stress the fact that nomenclatural ranks do not have biological definitions or meanings and that they should never be used in an ‘absolute’ way (e.g., to express degrees of genetic or phenetic divergence between taxa or hypothesised ages of cladogeneses) but in a ‘relative’ way: two taxa which are considered phylogenetically as sister-taxa should always be attributed to the same nomenclatural rank, but taxa bearing the same rank in different ‘clades’ are by no means ‘equivalent’, as the number of ranks depends largely on the number of terminal taxa (species) and on the degree of phylogenetic resolution of the tree; [6] because of this lack of ‘equivalence’, some arbitrary criteria are necessary to fix a starting point for assigning a given suprageneric rank to some taxa, from which the ranks of all other taxa will automatically derive through a simple implementation of the hierarchy of ranks: for this purpose we chose the rank family and we propose a ‘Ten Criteria Procedure’ allowing to fix the position of this rank in any zoological classification. As a result of the implementation of this set of Criteria, we obtained a new ranked classification of extant lissamphibians using 25 suprageneric ranks below the rank class (11 class-series and 14 family-series ranks), and including 34 class-series and 573 family-series taxa, and where the 575 genera we recognise are referred to 69 families and 87 subfamilies. We provide new nomina and diagnoses for 10 class-series taxa, 171 family-series taxa, 14 genus-series taxa and 1 species. As many new species of amphibians are permanently described, this classification and its nomenclature will certainly have to change many times in the future but, using the clear, explicit, complete, automatic and unambiguous methodology presented here, these changes will be easy to implement, and will not depend on subjective and arbitrary choices as it has too often been the case in the last decades. We suggest that applying this methodology in other zoological groups would improve considerably the homogeneity, clarity and usefulness of zoological taxonomy and nomenclature.
This paper explores two aspects of the question of the allocation of nomina to taxa in zoological nomenclature. The widespread belief that this allocation is effected only through onomatophores (‘types’) is shown to be wrong: if onomatophores only were involved, each zoological nomen would apply to all the taxa including the specimen(s) designated as onymophoront of each species-series nomen, up to the animal kingdom. Onomatophores do not provide the limit(s) of the taxon/taxa to which they apply. These limits are provided by two other pieces of information: the nominal-series to which the nomen of a taxon is assigned and the onomatostasis of this nomen, i.e. the onomatophore of the sister-taxon of the taxon under consideration. In species-, genus- and family-series nomenclature, the onomatostasis of a nomen is not fixed but depends on the ergotaxonomy (working classification) adopted, which is ideally based on at least one accepted phylogenetic hypothesis. The situation is more complex in the class-series nomenclature. So far, the stages of taxonomic allocation and nomenclatural validity in this nomenclature have not been regulated by formal Rules shared by all zootaxonomists, so that chaos and miscommunication are in order regarding the Criteria to be used in these domains, which is detrimental to zoological nomenclature and its perception by the international scientific community. The adoption of clear and strict Rules in this respect appears as an urgent need. These Rules should allow both to provide clear objective Criteria for the allocation of class-series nomina to taxa, but also to validate the few very well known and consensually adopted class-series nomina (pansozonyms). For this purpose, five possible nomenclatural systems of allocation of nomina are compared. The best solution appears to have two different systems, one for almost all nomina (distagmonyms), and one only for pansozonyms. The latter system relies on a special kind of onomatostases, which are fixed and do not depend on the classification adopted: this allows to attach permanently these nomina to some well-known higher taxa, whatever changes are brought to the cladistic hypotheses and ergotaxonomic frames.
After discussing some aspects of the biogeography of Plethodontidae, the authors conclude that these may have reached western Europe from northeastern North America by the end of Mesozoic, in agreement with the hypothesis proposed by Laurent (1975). The morphological, biochemical, caryological and ethological differences occurring between the Californian species of Hydromantes (sensu lato) and the European ones appear to justify the erection for the American species of the new genus Hydromantoides. This one shall include the three species platycephalus (Camp, 1916) (type-species), shastae (Gorman & Camp, 1953) and brunus (Gorman, 1954).
A few additions, comments and corrections are here provided on the Duplostensional Nomenclatural System (DONS) for higher zoological nomenclature recently proposed. They concern the concept of hypnokyronym and related concepts, the quantitative requirements for awarding a nomen the status of sozomorph, the nomen LISSAMPHIBIA and related nomina, as well as the correction of a few errata in the original publication.
Three years monitoring of the French populations of Rana pyrenaica Serra-Cobo, 1993 (Amphibia: Ranidae): first data on tadpole abundance. The Pyrenean frog Rana pyrenaica Serra-Cobo, 1993 is an endemic species from the western Pyrenees. The discovery of this species is recent, and its French distribution has long been restricted to the southern side of the Pyrenees and a few streams. In 2009, the situation changed following the discovery of several localities on the northern side of the Pyrenees. These findings prompted the conservation N.G.O. 'Cistude Nature' to implement a conservation programme of the species, to 1 – raise awareness about that species, 2 – continue to search for new populations, 3 – implement a suitable population monitoring. This paper presents the first results of a three-year survey based on repeated counts of Rana pyrenaica tadpoles on their breeding site, from an adapted protocol of 'PopAmphibiens' launched by the French Herpetological Society (SHF) and the French National Museum of Natural History (MNHN). N-mixture models for estimating population size from spatially replicated counts used in this study provide a first overview on abundance of tadpoles of this species and bring first ever published information on its detection probabilities. They depict strong disparities between sites and years with estimated densities ranging between 0.2 and 12.4 tadpoles/m². On the basis of those first results and considering the regional conservation status of the French population of this species, we argue that monitoring of these French populations should be continued in the upcoming years, to act as an ecological surveillance of this remarkable species for which the responsibility of France and the Aquitaine region is very high.
The almost complete absence in the Code of Rules for the
nomenclature of higher zoological taxa is a severe nuisance for
scientific communication about biodiversity and should urgently be
remedied for. After a brief survey of previous proposals in this
respect, a new system of class-series nomenclature is presented: the
Duplostensional Nomenclatural System (DONS). Being ostensional
and not intensional as some recently proposed alternative
nomenclatural systems, DONS is compatible with the system of the
Code for the nomina of taxa from the rank subspecies to the rank
superfamily, although it differs from it in several respects,
particularly in being monosemic and in not using the Principle of
Coordination. This system proposes precise, automatic, universal
and stringent Criteria for the nomenclatural availability of nomina,
their nominal-series assignment, their taxonomic allocation, and
their nomenclatural validity and correctness. It has several
advantages over the other systems of higher nomenclature so far
proposed, including three previous Ostensional Nomenclatural
Systems (ONS). One of its particularities is that it uses two different
systems of taxonomic allocation of nomina, one for well-known
nomina and one for more obscure ones. This allows to maintain as
valid the few really well-known higher zoological nomina (such as
AVES, COLEOPTERA or MOLLUSCA), even when they were not the
first ones to have been proposed for the taxa to which they apply or
when these taxa are modified through the inclusion of more basal,
often fossil, taxa. The other nomina follow simpler Criteria and
require less work for their allocation and validation. Adoption by
the international community of taxonomists of this system, with or
without incorporation of these Criteria into the Code, would allow
higher zoological nomenclature to abandon its current nonscientific
practices based on arbitrariness, subjectivity and
lobbying, and to be at last based on a clear, explicit, repeatable
methodology that would allow non-ambiguous international
communication about taxa. The use of the proposed Criteria is
exemplified through a detailed analysis of the steps allowing to
establish under DONS the valid nomina of the higher nomina of
recent amphibians (frogs, salamanders and cecilians).
Alkaline phosphatase (ALP) and aminopeptidase (AMP) distribution has been investigated histochemically in the digestive tract of Eutardigrada of the genera Macrobiotus and Hypsibius, in relation to cell features, to pH values inside the gut and to different life conditions. In the salivary glands, ALP seems to play a role in secretory activity, rather than in digestion. ALP and AMP are most active in the midgut epithelium, localized in the brush border and in the cytoplasm, but have different distributions. In the brush border, ALP may be involved mainly in absorption, and AMP both in secretion of proteolytic enzymes for extracellular digestion, and in the final breakdown and absorption of proteins cytoplasmic ALP and AMP may participate in intracellular digestion. A peritrophic membrane-like structure is often found. In the anterior hindgut, ALP at the cell membrane infoldings of the rectal pads may play a role in active transport related to osmoregulation. Distinct cell types cannot be identified in the midgut but the cardia and the pyloric region are somewhat different from the middle midgut and are characterized by a stronger AMP activity; the cardia may be a compartment of cell proliferation and differentiation. Variations of single cells may depend on a repeated morpho-functional cycle, while changes of the whole epithelium of the midgut may be related to the feeding period and to general physiological conditions.
We describe a new genus of fossil frog from the Miocene, Valentine Formation, Barstovian (13 - 14.5 million years ago) of northern Nebraska in the northern Great Plains USA. This frog was originally described as Pseudacris nordensis in 1964. The large, diverse genus Pseudacris is composed of 18 other species, all of which are extant. A total of 164 specimens (7 fossils, 114 dry skeletons, 43 cleared and stained specimens) from 17 museums and collections was examined for the new genus and the 18 extant species of Pseudacris . Diagnostic characteristics of the ilia differentiate the distinct new fossil genus from all extant species of Pseudacris . The new genus is known from three locations in the basin of the Niobrara River. Two extant species ( P. clarkii and P. crucifer ) are also known from several fossils of the same stratigraphic horizon as the new genus. Co-occurrence of the new genus with P. clarkii and P. crucifer in the same horizon clearly indicates that they were markedly distinct 13-14.5 million years ago. We propose that the northern Great Plains may have been a center of diversity for terrestrial vertebrates during the Miocene, and discuss models for speciation, the first step in the evolution of a new genus.
To understand the taxonomic and phylogenetic relationships of Alsodes montanus and A. pehuenche the species were compared with species of the genera Alsodes and Telmatobius. Comparisons were based on internal and external, qualitative and quantitative characters. Information was analysed phenetically and phylogenetically. Phenetic analysis reveals for A. montanus and A. pehuenche an independent position with respect to other Alsodes and intermediate between species of Abodes and Telmatobius. Phylogenetic analysis reveals that A. montanus and A. pehuenche have more ancestral states of characters than the species of Abodes and Telmatobius analysed.A new genus, containing A. montanus and A. pehuenche is proposed.
Torrent frogs of the genus Petropedetes Reichenow, 1874 as currently understood have a disjunct distribution with species endemic to West, Central or East Africa. We herein present a phylogenetic analysis including all but one of the currently described 12 species of the genus. Maximum Likelihood and Bayesian analyses of combined nuclear (rag1, SIA, BDNF) and mitochondrial (16S, 12S, cytb) genes of more than 3500 base pairs, revealed clades corresponding to the three sub-Saharan regions. Molecular results are confirmed by morphological differences. Surprisingly, the three geographic clades do not form a monophyletic group with respect to closely related families Pyxicephalidae and Conrauidae and therefore require taxonomic changes. We resurrect ArthroleptidesNieden, 1911 for the East African taxa. The Central African taxa remain in the genus Petropedetes. The West African members are placed in the new genus Odontobatrachus gen. nov. The phylogenetic position of the new genus remains incertae sedis as it was not assigned to any of the four families included in our analyses. Potential new species have been detected within all three major clades, pointing to a still not fully clarified diversity within African torrent frogs.
Marmorerpeton gen. nov. represented by M. kermackisp. nov. and M. freemani sp. nov., is the earliest known genus of fossil salamander from the Upper Bathonian, Middle Jurassic of Kirtlington, Oxfordshire, England. Marmorerpeton is more primitive than any other known salamander in the absence of intravertebral spinal nerve foramina in the atlantal centrum, but in other features it resembles members of the family Scapherpetontidae, neotenous salamanders otherwise known from the Upper Cretaceous and Palaeocene. The Kirtlington herpetofauna is a unique freshwater assemblage of Middle Jurassic small amphibians and reptiles, several of which represent the earliest known occurrences of their respective groups.RésuméMarmorerpeton gen. nov., représenté par M. kermacki sp. nov. et M. freemani sp. nov., provenant du Bathonien supérieur, Jurassique moyen de Kirtlington, Oxfordshire, Angleterre, est le plus ancien genre de caudate fossile. Il est plus primitif que les autres caudates connus à cause de l'absence des foramens intravertébraux dans le centrum atlantal, mais il ressemble par d'autres caractères aux membres de la famille des Scapherpetontidae, caudates néoténiques du Crétacé supérieur et du Paléocène. L'herpétofaune de Kirtlington constitue un ensemble unique d'amphibiens et de reptiles dulçaquicoles du Jurassique moyen, certains de ses éléments représentent les premiers spécimens connus de leurs groupes respectifs.
All living and fossil salamanders known till now are primarily Laurasian and have amphicoelous or opisthocoelous vertebrae. But Cretaceous salamanders from Gondwanan continents (South America and Africa) have procoelous vertebrae. One of them, from the Maastrichtian of Bolivia, is here described as Noterpeton bolivianum gen. and sp. nov. and it is referred to a family of its own, Noterpetontidae fam. nov. It is suggested that the procoelous Gondwanan forms could represent the sister group of all the other salamanders, but no detailed phyletic study can be undertaken on the basis of the available material.RésuméTous les urodèles, actuels et fossiles, connus jusqu'à présent sont laurasiatiques ou ont pénétré sur des territoires gondwaniens à partir de la Laurasie; en outre, ils possèdent des vertèbres amphicoeles ou opisthocoeles. Mais des urodèles provenant du Crétacé de continents gondwaniens (Amérique du Sud et Afrique) ont des vertèbres procoeles. L'un d'entre eux, du Maastrichtien de Bolivie, est décrit sous le nom de Noterpeton bolivianum gen. et sp. nov. et il est rapporté à une famille nouvelle, les Noterpetontidae. Les urodèles procoeles du Gondwana pourraient représenter le groupe frère de tous les autres urodèles; cependant, aucune étude phylétique ne peut être entreprise sur la base du matériel disponible.
Six larval forms of the bufonid genus Ansonia from Borneo share the following synapomorphies: cup-like, ventral oral disc; an expanded post-dental portion of the lower lip, which has a papillate margin; upper keratinized jaw sheath divided; body markedly flattened ventrally; eyes set relatively far back on the body. All of these tadpoles live on the bottom in strong currents, except for larval Ansonia leptopus, which lives in drifts of dead leaves that accumulate in eddies within streams. These larval forms differ among themselves in body shape, development of inframarginal papillae on the lower lip, size of the gap between the keratinized parts of the upper jaw sheath, width of the post-dental portion of the lower lip, relative lengths of upper and lower rows of labial teeth, and arrangement of the gut coils. One form has an abdominal sucker. Changes in these characters are not correlated; the derived condition in one character is not always associated with the derived state in another. Consequently, these tadpoles cannot be arranged in a simple morphocline from least to most derived, again with the exception of A. leptopus, which is the least modified in all respects. Although tadpoles of Ansonia resemble those of the neotropical bufonid genus Atelopus in general specialization for benthic life in flowing water, they differ from that group in body form and details of the oral disc.