PHYLOGENETIC INTERRELATIONSHIPS AND REDUCTIVE EVOLUTION IN NEOTROPICAL CHARACIDIIN FISHES (CHARACIFORMES, OSTARIOPHYSI)

Abstract— Miniaturization, which results in the presence of numerous apparently paedomorphic characters associated with reduced size, is a common phenomenon among neotropical fishes, with over 85 miniature species distributed among the five major ordinal groups. Eleven species are recognized as miniatures within the Characidiinae, a monophyletic subunit of Characiformes. A reconstruction of characidiin phylogeny is used to analyze the history of miniaturization events. Former hypotheses of origin of miniaturization among characidiins are rejected, underscoring the need for phylogenetic frameworks in the study of ontogenetic changes associated with the phenomenon of miniaturization. The 11 instances of miniature species can be most parsimoniously attributed to three independent miniaturization events within the Characidiinae.
Reductive characters comprise a large proportion of phylogenetically informative characters within the Characidiinae. In the largest group of miniatures, reductive characters represent more than half of the character state transformations affecting supraspecific relationships among Elachocharax, Klausewitzia, Odontocharacidium and Microcharacidium . An analysis of patterns of character state distributions fails to reject the null hypothesis of character independence. A distinction is made between the concepts of character independence, defined as the origination of character states from different (non-simultaneous) evolutionary events, and character correlation, defined as the association of character states in terminal taxa. Character correlation is not a sufficient criterion to reject Hennig's auxiliary principle, according to which the "presence of apomorphous characters in different species is always reason for suspecting kinship, and their origin by convergence should not be assumed a priori". High values of character correlation are the expected result of congruent patterns of character distribution.     

The radiation of characiform fishes and the limits of resolution of mitochondrial ribosomal DNA sequences

Phylogenetic relationships among fishes from ostariophysan orders, characiform families, and serrasalmin genera (e.g., "piranhas") were examined using partial mitochondrial ribosomal DNA sequences of the 12S and 16S genes. Phylogenetic information content of these sequences was assessed at three levels of taxonomic inclusiveness by analyzing the patterns of nucleotide substitution using secondary structure models. Conserved and variable regions were identified, mapped onto the structural models, and compared at increasing levels of taxonomic divergence. In general, loop regions (unpaired) exhibited a higher level of variation than did stem regions (paired). A high proportion of compensatory substitutions was observed in stem regions in three data sets, suggesting strong selection to maintain the secondary structure. Saturation due to multiple substitutions was indicated by decreasing transition/transversion ratios and strong structural constraints on variation in comparisons among orders of Ostariophysi but was not obvious among families of Characiformes and was not detected among serrasalmin genera. Reliable phylogenetic signal successfully reconstructed relationships among serrasalmin genera. However, aside from a few well-supported clades, relationships could not be reconstructed with confidence among characiform families and ostariophysan orders. The reciprocal monophyly of African and Neotropical characiform lineages was rejected (based on maximum likelihood ratio tests), and some support for previous hypotheses based on morphology was provided by the molecular data. The radiation of characiform fishes is discussed in a historical biogeographic context.     

Monophyly of the Citharidae (Pleuronectoidei: Pleuronectiformes: Teleostei) with considerations of pleuronectoid phylogeny

The monophyly of Citharidae, the pleuronectoid family thought to be a transitional group between the Psettodoidei and other typical pleuronectoids, has been in question mainly because of the lack of recognized synapomorphies for the family. In this study, the citharid skeleton is described, and the monophyly of the family and its phylogenetic position within the Pleuronectoidei are reassessed following a phylogenetic analysis based on 45 osteological, myological, and external characters. The Citharidae was found to represent a monophyletic group, supported by six synapomorphies (e.g., first dorsal proximal radials firmly wedged into notch of blind side lateral ethmoid; anterior dorsal proximal radials tightly mutually attached; blind side posterior nostril considerably enlarged). Two previously recognized subfamilies, sinistral Citharinae and dextral Brachypleurinae, are invalid because of their nonmonophylies. Interrelationships of the pleuronectoids shown herein are discussed. Received: February 19, 2001 / Revised: July 10, 2001 / Accepted: July 23, 2001     

Digest: Resolving phylogenomic conflicts in characiform fishes†

How can taxonomists best resolve the challenge of curating and analyzing large phylogenomic datasets that produce incongruent but highly supported topologies? Betancur‐R et al. used a recently established hypothesis‐testing procedure on a large dataset of genes and species to study the evolutionary relationships of characiform fishes, finding that past conclusions of non‐monophyly may have been problematic and establishing monophyly with high confidence. The new findings highlight the importance of using dense taxon sampling to resolve conflicting relationships with phylogenomic data.     

Interrelationships of Gasterosteiformes (Actinopterygii,Percomorpha)

Phylogenetic relationships of Gasterosteiformes were studied using osteological examination of representatives of 11 families of gasterosteiform fishes, as ingroups, and 5 families of other smegmamorph fishes (Atheriniformes, Elassomatiformes, and Synbranchiformes), as outgroups. Based on phylogenetic analysis of 110 informative osteological characters, nine synapomorphies were found to unite all Gasterosteiformes and support was provided to the hypothesis that the order Gasterosteiformes (including Hypoptychidae and Indostomidae) is a monophyletic group. Furthermore, based on the synapomorphies provided for the subgroups, three suborders in Gasterosteiformes are recognized: Hypoptychoidei, Gasterosteoidei, and Syngnathoidei.     

Anatomy and evolution of the mandibular, hyopalatine, and opercular muscles in characiform fishes (Teleostei: Ostariophysi)

The Characiformes are distributed throughout large portions of the freshwaters of Africa and America. About 90% of the almost 2000 characiform species inhabit the American rivers, with their greatest diversity occurring in the Neotropical region. As in most other groups of fishes, the current knowledge about characiform myology is extremely poor. This study presents the results of a survey of the mandibular, hyopalatine, and opercular musculature of 65 species representing all the 18 traditionally recognized characiform families, including the 14 subfamilies and several genera incertae sedis of the Characidae, the most speciose family of the order. The morphological variation of these muscles across the order is documented in detail and the homologies of the characiform adductor mandibulae divisions are clarified. Accordingly, the mistaken nomenclature previously applied to these divisions in some characiform taxa is herein corrected. Contradicting some previous studies, we found that none of the examined characiforms lacks an A3 section of the adductor mandibulae, but instead some taxa have an A3 continuous with A2. Derived myological features are identified as new putative synapomorphies for: the Characoidei; the clade composed of the Alestidae, Characidae, Gasteropelecidae, Cynodontoidea, and Erythrinoidea; the clade Cynodontoidea plus Erythrinoidea; the clade formed by Ctenoluciidae and Erythrinidae; the Serrasalminae; and the Triportheinae. Additionally, new myological data seems to indicate that the Agoniatinae might be more closely related to cynodontoids and erythrinoids than to other characids.     

Are monophyly and synapomorphy the same or different? Revisiting the role of morphology in phylogenetics

Species are groups of organisms, marked out by reproductive (replicative) properties. Monophyletic taxa are groups of species, marked out by synapomorphies. In Nelson’s analysis, monophyly and synapomorphy are identical relations. Monophyly and synapomorphy, however, are not equivalent relations. Monophyly is epistemically not accessible, whereas synapomorphy is epistemically accessible through character analysis. Monophyly originates with speciation, the two sister‐species that come into being through the splitting of the ancestral species lineage forming a monophyletic taxon at the lowest level of inclusiveness. Synapomorphy provides the empirical evidence for monophyly, inferred from character analysis in the context of a three‐taxon statement. If synapomorphy and monophyly were equivalent, phylogenetic systematists should find a single tree, instead of multiple equally parsimonious trees. Understanding synapomorphy as the relevant evidence for phylogenetic inference reveals a category mistake in contemporary phylogenetics: the treatment of morphological characters mapped onto molecular trees as synapomorphies and homoplasies. The mapping of morphological characters onto nodes of a molecular tree results in an empirically empty procedure for synapomorphy discovery. Morphological synapomorphies and homoplasies can only be discovered by morphological and combined analyses. The use of morphology in phylogenetic inference in general is defended by examples from Laurales and Squamata in particular. To make empirical evidence scientifically relevant in order to search for concordance, or dis‐concordance, of phylogenetic signal, is certainly more fruitful for phylogenetics than the uncritical mapping of morphological traits on a molecular scaffold. © The Willi Hennig Society 2010.     

Phylogenetic position of the family Trichodontidae (Teleostei: Perciformes), with a revised classification of the perciform suborder Cottoidei

The phylogenetic relationships of the family Trichodontidae and suborder Cottoidei (and zoarcoid Bathymasteridae) are reconstructed morphologically. The monophyly of the Trichodontidae, Cottoidei, and Zoarcoidei is unambiguously supported by 14 synapomorphies, including 1 newly recognized (and also 4 synapomorphies when ACCTRAN is accepted). It is assumed that the Trichodontidae is deeply nested within the Cottoidei, and the family and Cottoidea plus Cyclopteroidea have a sister relationship, supporting a previously inferred molecular phylogenetic hypothesis. We propose that the Trichodontidae is placed into the cottoid superfamily Trichodontoidea.     

The Phylogeny of the Myrmecophagidae (Mammalia, Xenarthra, Vermilingua) and the Relationship of Eurotamandua to the Vermilingua

A cladistic investigation of the phylogenetic relationships among the three extant anteater genera and the three undoubted extinct myrmecophagid genera is performed based upon osteological characteristics of the skull and postcranial skeleton. One hundred seven discrete morphological characters are analyzed using the computer program PAUP. Characters are polarized via comparison to the successive xenarthran outgroups Tardigrada (represented by the living sloth Bradypus) and Cingulata (represented by the recent armadillos Dasypus and Euphractus). The analysis results in a single most-parsimonious tree (TL = 190, CI = 0.699, RI = 0.713). The tree corroborates the monophyly of the subfamilies Cyclopinae and Myrmecophaginae, the former including the extant Cyclopes and the Pliocene genus Palaeomyrmidon. Within the Myrmecophaginae the Miocene genus Protamandua is the sister taxon to a clade including the remaining three genera. The recent Tamandua is in turn the sister taxon to the extant Myrmecophaga plus the Pliocene genus Neotamandua. Contrary to the suggestions of recent authors, weak support is provided for the taxonomic distinctiveness of the latter genus from the recent Myrmecophaga. The monophyly of the Myrmecophagidae is supported by 15 unequivocal synapomorphies. The monophyly of the Cyclopinae and Myrmecophaginae is supported by 3 and 13 unambiguous synapomorphies, respectively. The enigmatic Eocene genus Eurotamandua, from the Messel fauna of Germany, is coded for the 107 morphological characters above and included in two subsequent PAUP analyses. The palaeanodont Metacheiromys is also added to these two analyses as a nonxenarthran outgroup to test for the possibility that Eurotamandua lies outside the Xenarthra. In the first analysis, Eurotamandua is constrained a priori to membership in the Vermilingua. The single most-parsimonious tree (TL = 224, CI = 0.618) that results places Eurotamandua as the sister group to the remaining anteater genera, contra Storch and Habersetzer's (1991) assignment of Eurotamandua to the vermilinguan subfamily Myrmecophaginae. Eurotamandua shares six unequivocal synapomorphies with other anteaters, including the absence of teeth and the presence of a lateral tuberosity on the fifth metatarsal. The remaining vermilinguans are united by 11 unequivocal synapomorphies, plus an additional 10 ambiguous synapomorphies. In the second analysis, the position of Eurotamandua is unconstrained. The resulting single most-parsimonious tree (TL = 219, CI = 0.632) places Eurotamandua outside Vermilingua as the sister group to the Pilosa (Vermilingua plus Bradypus). The monophyly of this node is supported by four unambiguous synapomorphies in the unconstrained analysis. Further manipulation of this second analysis shows that placement of Eurotamandua as the sister group to the Xenarthra or to the Palaeanodonta adds three steps to the shortest tree but is more parsimonious than its placement as a sister group to the Vermilingua is the previous analysis. The addition of pangolins to the analysis does little to alter the major phylogenetic conclusions of the study. The allocation of Eurotamandua to the Xenarthra, but as a sister group to the Pilosa, is a novel arrangement which leaves open the biogeographic question of how a xenarthran reached Western Europe during the Eocene.     

Neogastropod phylogenetic relationships based on entire mitochondrial genomes

Background  

The Neogastropoda is a highly diversified group of predatory marine snails (Gastropoda: Caenogastropoda). Traditionally, its monophyly has been widely accepted based on several morphological synapomorphies mostly related with the digestive system. However, recent molecular phylogenetic studies challenged the monophyly of Neogastropoda due to the inclusion of representatives of other caenogastropod lineages (e.g. Littorinimorpha) within the group. Neogastropoda has been classified into up to six superfamilies including Buccinoidea, Muricoidea, Olivoidea, Pseudolivoidea, Conoidea, and Cancellarioidea. Phylogenetic relationships among neogastropod superfamilies remain unresolved.     

Study on the systematic position of Metriini based on characters of the larval head (Coleoptera: Carabidae)

Abstract. Characters of the head of larvae of Metrius contractus Eschscholtz, Ozaenini and Paussini are interpreted phylogenetically. The monophyly of Metriini + Ozaenini + Paussini is substantiated by several synapomorphies such as hyperprognathism and strong constriction of the neck. Ozaenini and Paussini together form the sister-group of Metriini. Ozaenini are paraphyletic. The monophyly of Paussini + Ozaenini excluding Pachyteles is indicated by two possible synapomorphies. Several synapomorphies are shared by Physea + Paussini. Secondary prognathism, large membranous submento-mental area and other derived features are considered autapomorphies of Paussini. Paussini excluding Platyrhopalopsis are characterized by the loss of the palpifer. The monophyly of a group which comprises Geadephaga excluding Trachypachini is suggested by several synapomorphic features. A very basal position of the metriine—paussine lineage within Carabidae is indicated by several plesiomorphic features. A hypopharyngeal filter apparatus with a dense fringe of well-arranged, long hairs is a possible autapomorphy of Anisochaeta. The results of this study do not indicate a close relationship between the metriine—paussine lineage and the tribes Brachinini and Crepidogastrini as has been suggested in recent works.     

Morphological evidence supporting the monophyly of the family Polynemidae (Teleostei: Perciformes) and its sister relationship with Sciaenidae

The monophyly of Polynemidae was evaluated and its sister relationship with Sciaenidae discussed, based on osteological and myological characters from 24 polynemid species in eight genera, with comparisons with acanthomorph fishes from literature and 86 species in 8 orders and 63 families examined. Polynemidae was inferred as a monophyletic group, strongly supported by 19 synapomorphies, including four unique characters (unnamed bone present on cephalic sensory canal extending from supratemporal, third actinost not supporting pectoral-fin rays, section A1 comprising lateral and medial elements, and division of obliquus inferioris present between lower postcleithrum and rod-like process on coracoid) in percoids. In addition, seven pectoral girdle characters were recognized, with the girdle possessing filament-like sensory rays, an adaptation to benthic life in muddy water. The sister relationship of Polynemidae and Sciaenidae was supported by six synapomorphies, including two rather rare (a single branchiostegal ray suspended by epihyal and posterior portions of pelvic bones on both sides interdigitated) and two unique characters (metapterygoid and quadrate interdigitated medially and anterior extension of the nasal canal).     

Patterns of variation in life history among South American fishes in seasonal environments

Summary Ten traits related to life history theory were measured or estimated for 71 freshwater fish species from two locations in the Venezuelan llanos. Multivariate statistics and cluster analysis revealed three basic endpoint patterns bounding a two-dimensional continuum. A suite of attributes associated with parental care and aseasonal reproduction appeared to correspond to an equilibrium strategy. A second group of small fishes was distinguished by traits associated with rapid colonizing ability: early maturation, continuous reproduction, and small clutches. The third basic pattern was associated with synchronized reproduction during the early wet season, high fecundity, absence of parental care, and breeding migrations. A subset of mostly small fishes exhibiting little or no parental care, small clutches, and two to four month reproductive seasons was intermediate between the opportunistic (rapidly colonizing) and seasonal strategies. All ten life history variables showed significant effects of phylogeny. The cluster of species corresponding to the equilibrium group was dominated by siluriform fishes and perciforms of the Cichlidae. The opportunistic cluster was dominated by cyprinodontiform and characiform fishes, whereas the seasonal cluster contained primarily characiform and siluriform fishes. Seven of nine traits were significantly correlated with body length. The three reproductive patterns are interpreted as being adaptative with respect to relative intensity and predictability of temporal and spatial variation in abiotic environmental parameters, food availability, and predation pressure.     

Phylogenetic relationships within the speciose family Characidae (Teleostei: Ostariophysi: Characiformes) based on multilocus analysis and extensive ingroup sampling

Background

With nearly 1,100 species, the fish family Characidae represents more than half of the species of Characiformes, and is a key component of Neotropical freshwater ecosystems. The composition, phylogeny, and classification of Characidae is currently uncertain, despite significant efforts based on analysis of morphological and molecular data. No consensus about the monophyly of this group or its position within the order Characiformes has been reached, challenged by the fact that many key studies to date have non-overlapping taxonomic representation and focus only on subsets of this diversity.

Results

In the present study we propose a new definition of the family Characidae and a hypothesis of relationships for the Characiformes based on phylogenetic analysis of DNA sequences of two mitochondrial and three nuclear genes (4,680 base pairs). The sequences were obtained from 211 samples representing 166 genera distributed among all 18 recognized families in the order Characiformes, all 14 recognized subfamilies in the Characidae, plus 56 of the genera so far considered incertae sedis in the Characidae. The phylogeny obtained is robust, with most lineages significantly supported by posterior probabilities in Bayesian analysis, and high bootstrap values from maximum likelihood and parsimony analyses.

Conclusion

A monophyletic assemblage strongly supported in all our phylogenetic analysis is herein defined as the Characidae and includes the characiform species lacking a supraorbital bone and with a derived position of the emergence of the hyoid artery from the anterior ceratohyal. To recognize this and several other monophyletic groups within characiforms we propose changes in the limits of several families to facilitate future studies in the Characiformes and particularly the Characidae. This work presents a new phylogenetic framework for a speciose and morphologically diverse group of freshwater fishes of significant ecological and evolutionary importance across the Neotropics and portions of Africa.     

白鱼属鱼类的系统发育(鲤形目:鲤科)

白鱼属(Anabarilius Cockerell)是我国特有的鲤科鱼类,主要分布于云南东部、中部和四川南部。分布范围不大,但种的分化强烈。 Regan(1904,1908, 1918)最先报道四种白鱼,但均置于Barilius属。尔后,Cockerell(1923)以Barilius andersoni Regan为模式种建立白鱼属;Nichols(1927)、张孝威(1944)、易伯鲁、吴清江(1964)、刘振华、何纪昌(1983)先后记述了部分种和亚种。陈银瑞、褚新洛(1980)在系统整理的基础上,澄清了白鱼属属级分类上的混乱,记述了三新种和一新亚种。何纪昌、王重光(1984)用数值分类法探讨了白鱼属的分类,报道了二新种和一新亚种。至此,白鱼属共包括15个种和亚种。     

Grundsätze der phylogenetischen Systematik (Eine praxisorientierte Übersicht)

The essential elements of phylogenetic systematics in the sense of Hennig are emphasized: The search for synapomorphies based on a special method of comparative morphology, and the aim of an exclusive use of synapomorphies for kinship proof and the basis of systematics. Special aspects of comparative morphology are: “Directed comparisons” steady reciprocal reflection between comparative morphological result and the system methodical efforts for the realization of distinctive details, comprehensive documentation and functional interpretation. This is equally true for recent and fossil forms. Most suitable for the method (in the sense defined above) are groups with numerous differentiated morphological characters, which can also be preserved in the fossil state. The less this is the case the less is the chance for achieving necessary numbers of well proven synapomorphies. Even so, it is not permitted—for those who want to perform phylogenetic systematics in the sense of Hennig—to use convergences, parallelisms or symplesiomorphies in the sense of “synapomorphies” as phylogenetic arguments for kinship relations. Numerous examples and diagrams demonstrate the methodological proceeding, and differences towards other methods of phylogenetical reconstruction and interpretation. Special attention is paid to direct and indirect conclusions drawn from fossils: Time of origin of characters, stem groups and *groups; predictions concerning the appearance (set of characters) of fossils and simultaneous existence of “neighbour groups” (sister groups, and more distantly related taxa).     

Phylogeny of the Cocculinoidea (Mollusca, Gastropoda)

Abstract. The superfamily Cocculinoidea is a group of marine, deep-water, limpet-like gastropods. Recent speculation surrounding their affinities has concentrated on their placement within the Gastropoda. However, phylogenetic relationships within the Cocculinoidea, especially the monophyly of families and genera within the group, remain poorly understood. Phylogenetic analysis of 31 morphological characters for 15 cocculinoidean taxa and 2 outgroups resulted in a single most parsimonious tree, length=70, CI=0.62, and RI=0.71. Monophyly of the Cocculinoidea, Cocculinidae, and the genera Cocculina and Coccopigya was supported; Paracocculina and Coccocrater were found to be paraphyletic. Character optimization demonstrates that many characters often cited as diagnostic of various taxa, are often homoplastic and/or synapomorphies at different hierarchical levels.     

Phylogenetic analysis of the family Ariidae (Ostariophysi: Siluriformes), with a hypothesis on the monophyly and relationships of the genera

Ariid monophyly and intrafamilial relationships are investigated based on cladistic analysis of 230 morphological characters. Terminal taxa examined include whenever possible type‐species, or the most morphologically similar species to the type‐species of the nominal genera, and the largest possible number of species, including cleared and stained specimens, available in zoological collections. Previous hypotheses about monophyly of the Ariidae are strongly corroborated by new synapomorphies discovered in the present study. The subfamily Galeichthyinae and the remaining ariids are strongly supported by new morphological characters. The monotypic subfamily Bagreinae is recognized as the sister group to all nongaleichthyin ariids, supported by a large series of exclusive synapomorphies. A new concept of Ariinae is presented: the subfamily is found to be unequivocally monophyletic and includes all ariid genera, except Galeichthys and Bagre. New data supporting the monophyly of the genera included in the Ariinae are introduced and previous hypotheses of monophyly, species composition, morphological definition, and relationships are reviewed and discussed.     

A retroposon analysis of Afrotherian phylogeny

Recent comprehensive studies of DNA sequences support the monophyly of Afrotheria, comprising elephants, sirenians (dugongs and manatees), hyraxes, tenrecs, golden moles, aardvarks, and elephant shrews, as well as that of Paenungulata, comprising elephants, sirenians, and hyraxes. However, phylogenetic relationships among paenungulates, as well as among nonpaenungulates, have remained ambiguous. Here we applied an extensive retroposon analysis to these problems to support the monophyly of aardvarks, tenrecs, and golden moles, with elephant shrews as their sister group. Regarding phylogenetic relationships in Paenungulata, we could characterize only one informative locus, although we could isolate many insertions specific to each of three lineages, namely, Proboscidea, Sirenia, and Hyracoidea. These data prompted us to reexamine phylogenetic relationships among Paenungulata using 19 nuclear gene sequences resulting in three different analyses, namely, short interspersed element (SINE) insertions, nuclear sequence analyses, and morphological cladistics, supporting different respective phylogenies. We concluded that these three lineages diverged very rapidly in a very short evolutionary period, with the consequence that ancestral polymorphism present in the last common ancestor of Paenungulata results in such incongruence. Our results suggest the rapid fixation of many large-scale morphological synapomorphies for Tethytheria; implications of this in relation to the morphological evolution in Paenungulata are discussed.     

The adductor mandibulae muscle complex in lower teleostean fishes (Osteichthyes: Actinopterygii): comparative anatomy,synonymy, and phylogenetic implications

Bony fishes of the morphologically diverse infraclass Teleostei include more than 31 000 species, encompassing almost one‐half of all extant vertebrates. A remarkable anatomical complex in teleosts is the adductor mandibulae, the primary muscle in mouth closure and whose subdivisions vary in number and complexity. Difficulties in recognizing homologies amongst adductor mandibulae subdivisions across the Teleostei have hampered the understanding of the evolution of this system and consequently its application in phylogenetic analyses. The adductor mandibulae in representatives of all lower teleost orders is described, illustrated, and compared based on broad taxonomic sampling complemented by extensive literature information. Muscle division homologies are clarified via the application of a standardized homology‐driven anatomical terminology with synonymies provided to the myological terminologies of previous studies. Phylogenetic implications of the observed variations in the adductor mandibulae are discussed and new possible synapomorphies are proposed for the Notacanthiformes, Ostariophysi, Cypriniformes, Siluriphysi, Gymnotiformes, and Alepocephaloidei. New characters corroborate the putative monophyly of the clades Albuliformes plus Notacanthiformes (Elopomorpha), Argentinoidei plus Esocoidei plus Salmonoidei (Protacanthopterygii) and Hemiodontidae plus Parodontidae (Characiformes). We further confirm the validity of characters from the adductor mandibulae previously proposed to support the monophyly of the Esocoidei and the gonorynchiform clade Gonorynchoidei plus Knerioidei. © 2014 The Linnean Society of London