The characters of Palaeozoic jawed vertebrates

Newly discovered fossils from the Silurian and Devonian periods are beginning to challenge embedded perceptions about the origin and early diversification of jawed vertebrates (gnathostomes). Nevertheless, an explicit cladistic framework for the relationships of these fossils relative to the principal crown lineages of the jawed vertebrates (osteichthyans: bony fishes and tetrapods; chondrichthyans: sharks, batoids, and chimaeras) remains elusive. We critically review the systematics and character distributions of early gnathostomes and provide a clearly stated hierarchy of synapomorphies covering the jaw‐bearing stem gnathostomes and osteichthyan and chondrichthyan stem groups. We show that character lists, designed to support the monophyly of putative groups, tend to overstate their strength and lack cladistic corroboration. By contrast, synapomorphic hierarchies are more open to refutation and must explicitly confront conflicting evidence. Our proposed synapomorphy scheme is used to evaluate the status of the problematic fossil groups Acanthodii and Placodermi, and suggest profitable avenues for future research. We interpret placoderms as a paraphyletic array of stem‐group gnathostomes, and suggest what we regard as two equally plausible placements of acanthodians: exclusively on the chondrichthyan stem, or distributed on both the chondrichthyan and osteichthyan stems. © 2014 The Authors. Zoological Journal of the Linnean Society published by John Wiley & Sons Ltd on behalf of The Linnean Society of London     

Brief review of the stylophoran debate

The "calcichordate" theory interprets an extinct group of calcite-plated invertebrates, the stylophorans, as chordates. In this theory, cornute stylophorans are interpreted as stem chordates, whereas mitrate stylophorans are primitive members of the acraniates, tunicates, and craniates. However, this theory discounts major synapomorphies between cornutes and mitrates. These groups constitute a natural, monophyletic group which is here argued to lie within the echinoderm radiation. The "calcichordate" theory is, therefore, rejected because it relies on assumption-driven hypotheses of character transformation which are supported by ambiguous, poor, or missing fossil evidence. Stylophorans may lie at the base of the echinoderm clade and primitively lack pentameral symmetry, therefore casting light on the near-ancestral body organization of the phylum.     

Oldest Near-Complete Acanthodian: The First Vertebrate from the Silurian Bertie Formation Konservat-Lagerst?tte,Ontario

Background

The relationships between early jawed vertebrates have been much debated, with cladistic analyses yielding little consensus on the position (or positions) of acanthodians with respect to other groups. Whereas one recent analysis showed various acanthodians (classically known as ‘spiny sharks’) as stem osteichthyans (bony fishes) and others as stem chondrichthyans, another shows the acanthodians as a paraphyletic group of stem chondrichthyans, and the latest analysis shows acanthodians as the monophyletic sister group of the Chondrichthyes.

Methodology/Principal Findings

A small specimen of the ischnacanthiform acanthodian Nerepisacanthus denisoni is the first vertebrate fossil collected from the Late Silurian Bertie Formation Konservat-Lagerstätte of southern Ontario, Canada, a deposit well-known for its spectacular eurypterid fossils. The fish is the only near complete acanthodian from pre-Devonian strata worldwide, and confirms that Nerepisacanthus has dentigerous jaw bones, body scales with superposed crown growth zones formed of ondontocytic mesodentine, and a patch of chondrichthyan-like scales posterior to the jaw joint.

Conclusions/Significance

The combination of features found in Nerepisacanthus supports the hypothesis that acanthodians could be a group, or even a clade, on the chondrichthyan stem. Cladistic analyses of early jawed vertebrates incorporating Nerepisacanthus, and updated data on other acanthodians based on publications in press, should help clarify their relationships.     

CHARACTER DIAGNOSIS, FOSSILS AND THE ORIGIN OF TETRAPODS

I. The traditional view of the origin of tetrapod vertebrates is that they are descendants of fossil osteolepiform fish, of which Eusthenopteron is best known. In recent years both that conclusion and the methodology by which it has been reached have been challenged by practitioners of cladistic analysis. Particularly a recent review by Rosen et al. (1981) claims that Dipnoi (lungfish) are the sister-group of the Tetrapoda, that Osteolepiformes is a non-taxon and that Eusthenopteron is more distant from tetrapods than are Dipnoi, coelacanths and probably the fossil Porolepiformes. We attempt to refute all these concludions by use of the same cladistic technique. 2. We accept that all the above-mentioned groups, together with some less well-known taxa, can be united as Sarcopterygii by means of shared derived (apomorph) characters. We also agree that Porolepiformes and Actinistia (coelacanths) can be characterized as valid taxa. The primitive and enigmatic fossil fish Powichthys is accepted as representing the plesiomorph sister-group of true porolepiforms. 3. Only two apomorph features, the course of the jaw adductor muscles and the position of incurrent and excurrent nostrils, appear to unite all the fish, living and fossil, currently regarded as Dipnoi. The characteristic tooth plates and the presence of petrodentine both exclude important primitive fossil forms. 4. Contrary to the opinion of Rosen et al., Osteolepiformes can be characterized — by the arrangement of bones forming the cheek plate, the presence of basal scutes to the fins and by the unjointed radials of the median fins. However, if these are true autapomorphies they exclude any osteolepiform from direct tetrapod ancestry. 5. Tetrapoda is a monophyletic group characterized by ten or more autapomorphies, including the bones of the cheek plate, a stapes and fenestra ovalis, and a series of characters of the appendicular skeleton. 6. Tetrapods have a true choana (internal nostril). We accept that the posterior (excurrent) nostril of Dipnoi is the homologue of the tetrapod choana. However, we assert that the posterior nostril of all bony fish is the homologue of the choana. This assertion would be refuted if any fish showed separate posterior nostril and choana. We reject the claim that this ‘three nostril condition’ occurred in porolepiforms and osteolepiforms. The evidence for a choana in porolepiforms is inadequate. Osteolepiforms had a true choana, characterized as in tetrapods by its relationship to the bones of the palate, but no third nostril. Dipnoans are not choanate. 7. Following cladistic practice, the relationship of the extant taxa is established first. Dipnoi are thus shown to be the living sister-group of tetrapods, but only on ‘soft anatomy’ characters unavailable in fossils. Coelacanths are the living sister-group of the taxon so formed. 8. The relationship of the fossil taxa to the extant sarcopterygians is then considered. The synapomorphy scheme proposed by Rosen et al. is discussed at length. Virtually all the characters they use to exclude close relationship of Eusthenopteron (and hence all osteolepiforms) to tetrapods, in favour of coelacanths and dipnoans, are invalid. 9. A series of synapomorphies uniting osteolepiforms and tetrapods is proposed, including a true choana (hence the taxon Choanata), the histology of the teeth, and a number of characters of the humerus. The recently discovered fossil Youngolepis, which lacks a choana, represents the sister-group of the Choanata, and is not uniquely close to Powichthys. The latter, as a porolepiform (s.l.) is a member of the sister-group to Choanata plus Youngolepis. 10. Our cladistic analysis suggests that all the extinct taxa considered are more closely related to tetrapods than are the Dipnoi. Moreover fossil evidence suggests that Dipnoi, considered as an extant taxon, may not even be the living sister-group of Tetrapoda. Early fossil dipnoans appear to have been marine fish without specific adaptations for air breathing. If so the apparent synapomorphies of Dipnoi and Tetrapoda may be homoplastic — the insistence on grouping extant taxa first would then have yielded an invalid inference.     

Bystrow’s Paradox – gills,fossils, and the fish‐to‐tetrapod transition

Schoch, R.R. and Witzmann, F. 2011. Bystrow’s Paradox – gills, fossils, and the fish‐to‐tetrapod transition. —Acta Zoologica (Stockholm) 92 : 251–265. The issue of which breathing mechanism was used by the earliest tetrapods is still unsolved. Recent discoveries of stem tetrapods suggest the presence of internal gills and fish‐like underwater breathing. The same osteological features were used by Bystrow to infer a salamander‐like breathing through external gills in temnospondyl amphibians. This apparent contradiction – here called Bystrow’s Paradox – is resolved by reviewing the primary fossil evidence and the anatomy of the two gill types in extant taxa. Rather unexpectedly, we find that internal gills were present in a range of early crown tetrapods (temnospondyls), based on the anatomy of gill lamellae and location of branchial arteries on the ventral side of gill arch elements (ceratobranchials). Although it remains to be clarified which components are homologous in external and internal gills, both gill types are likely to have been present in Palaeozoic tetrapods – internal gills in aquatic adults of some taxa, and external gills in the larvae of these taxa and in larvae of numerous forms with terrestrial adults, which resorbed the external gills after the larval phase. Future developmental studies will hopefully clarify which mechanistic pathways are involved in gill formation and how these might have evolved.     

Fossil fishes from china provide first evidence of dermal pelvic girdles in osteichthyans

Background

The pectoral and pelvic girdles support paired fins and limbs, and have transformed significantly in the diversification of gnathostomes or jawed vertebrates (including osteichthyans, chondrichthyans, acanthodians and placoderms). For instance, changes in the pectoral and pelvic girdles accompanied the transition of fins to limbs as some osteichthyans (a clade that contains the vast majority of vertebrates – bony fishes and tetrapods) ventured from aquatic to terrestrial environments. The fossil record shows that the pectoral girdles of early osteichthyans (e.g., Lophosteus, Andreolepis, Psarolepis and Guiyu) retained part of the primitive gnathostome pectoral girdle condition with spines and/or other dermal components. However, very little is known about the condition of the pelvic girdle in the earliest osteichthyans. Living osteichthyans, like chondrichthyans (cartilaginous fishes), have exclusively endoskeletal pelvic girdles, while dermal pelvic girdle components (plates and/or spines) have so far been found only in some extinct placoderms and acanthodians. Consequently, whether the pectoral and pelvic girdles are primitively similar in osteichthyans cannot be adequately evaluated, and phylogeny-based inferences regarding the primitive pelvic girdle condition in osteichthyans cannot be tested against available fossil evidence.

Methodology/Principal Findings

Here we report the first discovery of spine-bearing dermal pelvic girdles in early osteichthyans, based on a new articulated specimen of Guiyu oneiros from the Late Ludlow (Silurian) Kuanti Formation, Yunnan, as well as a re-examination of the previously described holotype. We also describe disarticulated pelvic girdles of Psarolepis romeri from the Lochkovian (Early Devonian) Xitun Formation, Yunnan, which resemble the previously reported pectoral girdles in having integrated dermal and endoskeletal components with polybasal fin articulation.

Conclusions/Significance

The new findings reveal hitherto unknown similarity in pectoral and pelvic girdles among early osteichthyans, and provide critical information for studying the evolution of pelvic girdles in osteichthyans and other gnathostomes.     

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.     

More DNA support for a Cetacea/Hippopotamidae clade: the blood-clotting protein gene gamma-fibrinogen

Recent phylogenetic analyses of DNA sequences suggest that cetaceans (whales) and hippopotamid artiodactyls (hippos) are extant sister taxa. Consequently, the shared aquatic specializations of these taxa may be synapomorphies. This molecular view is contradicted by paleontological data that overwhelmingly support a monophyletic Artiodactyla (even-toed ungulates) and a close relationship between Cetacea and extinct mesonychian ungulates. According to the fossil evidence, molecular, behavioral, and anatomical resemblances between hippos and whales are interpreted as convergences or primitive retentions. In this report, competing interpretations of whale origins are tested through phylogenetic analyses of the blood-clotting protein gene gamma- fibrinogen from cetaceans, artiodactyls, perissodactyls (odd-toed ungulates), and carnivores (cats, dogs, and kin). In combination with published DNA sequences, the gamma-fibrinogen data unambiguously support a hippo/whale clade and are inconsistent with the paleontological perspective. If the phylogeny favored by fossil evidence is accepted, the convergence at the DNA level between Cetacea and Hippopotamidae is remarkable in its distribution across three genetic loci: gamma-fibrinogen, the linked milk casein genes, and mitochondrial cytochrome b.      

Divergence time estimation using fossils as terminal taxa and the origins of Lissamphibia

Were molecular data available for extinct taxa, questions regarding the origins of many groups could be settled in short order. As this is not the case, various strategies have been proposed to combine paleontological and neontological data sets. The use of fossil dates as node age calibrations for divergence time estimation from molecular phylogenies is commonplace. In addition, simulations suggest that the addition of morphological data from extinct taxa may improve phylogenetic estimation when combined with molecular data for extant species, and some studies have merged morphological and molecular data to estimate combined evidence phylogenies containing both extinct and extant taxa. However, few, if any, studies have attempted to estimate divergence times using phylogenies containing both fossil and living taxa sampled for both molecular and morphological data. Here, I infer both the phylogeny and the time of origin for Lissamphibia and a number of stem tetrapods using Bayesian methods based on a data set containing morphological data for extinct taxa, molecular data for extant taxa, and molecular and morphological data for a subset of extant taxa. The results suggest that Lissamphibia is monophyletic, nested within Lepospondyli, and originated in the late Carboniferous at the earliest. This research illustrates potential pitfalls for the use of fossils as post hoc age constraints on internal nodes and highlights the importance of explicit phylogenetic analysis of extinct taxa. These results suggest that the application of fossils as minima or maxima on molecular phylogenies should be supplemented or supplanted by combined evidence analyses whenever possible.     

Comments on hexanchiform phylogeny (Pisces, Neoselachii)

The Hexanchiformes (Cow Sharks) are regarded as a monophyletic taxon. Cladistic analysis shows that among the various neoselachian taxa proposed so far as the sister group of the Hexanchiformes a sister group relationship between the Hexanchiformes and a (still unnamed) taxon comprising the Squaliformes and Pristiophoriformes appears as the most probable hypothesis. In addition, MAISEY and WOLFRAM'S (1984) concept of hexanchiform interrelationships is critically reviewed. An alternative cladogram of hexanchiform interrelationships is developed which includes Recent as well as fossil hexanchiform taxa. In this cladogram the living genera Hexancbus and Notorynchus are sister groups and both taxa together form the sister group of the Recent Heptranchias. The fossil taxa +Notidanoides, +“Hexanchus” gracilis, +Notidanodon and +Weltonia are arranged in the stem lineage of recent Hexanchiformes.     

Squamate phylogeny, taxon sampling, and data congruence

To investigate the affinities of snakes, amphisbaenians and dibamids, the phylogenetic relationships among the major lineages (families) of extinct and extant squamates are assessed through a combined analysis of 248 osteological, 133 soft anatomical, and 18 ecological traits. The osteological data set represents a revision of previous data, taking into account recent criticism; the ecological data set is new. In addition, potentially critical fossil taxa (polyglyphanodontids and macrocephalosaurs) are included for the first time. The osteological and soft anatomical data sets each place snakes within anguimorphs, with dibamids and amphisbaenians near gekkotans. The putative primitive fossil amphisbaenian Sineoamphisbaena groups with macrocephalosaurs and polyglyphanodontids, together the sister group to scleroglossans. All three data sets are congruent, and these results are reinforced by combined analyses. In these, as in the osteological analyses, snakes are nested within marine lizards. However, exclusion of fossil taxa from the osteological data set results in a ‘limbless clade’ consisting of snakes, amphisbaenians and dibamids, and introduces significant conflict between osteology and soft anatomy. Also, deletion tests and character weighting reveal that the signal in the reduced osteological data set is internally contradictory. These results increase confidence in the arrangement supported by the all-taxon osteological, the soft anatomical, and the combined data, and suggest that exclusion of fossils confounds the signal in the osteological data set. Finally, the morphological data support the nesting of snakes within marine lizards, and thus a marine origin of snakes. This result still holds when relationships between living forms are constrained to the topology suggested by molecular sequences: if marine lizards are allowed to ‘float’ within this molecular framework, they form the stem group to snakes, and do not group with varanids as previously suggested.See also Electronic Supplement at: http://www.senckenberg.de/odes/05-04.htm.     

Phylogenetic systematics of leaffishes (Teleostei: Polycentridae,Nandidae)

The Asian (nandid) and Afro‐Neotropical (polycentrid) leaffishes represent two superficially similar, but historically poorly diagnosed families – a situation resulting in a convoluted systematic history. Here, and including for the first time in a molecular study all leaffish genera, we generate a hypothesis of the phylogenetic history of both groups. We analyse a multilocus molecular data set encompassing 257 acanthomorph taxa, carry out a survey and assessment of selected osteological characters for the polycentrid leaffishes and also provide a reanalysis of previously published morphological data. Our results confirm: (1) that the Polycentridae and Nandidae are only remotely related, and hence, the classic leaffishes are diphyletic; (2) that the Polycentridae is monophyletic, with new skeletal synapomorphies being congruent with molecular data in placing the enigmatic Afronandus – a taxon that thus far has never been included in any molecular study – as sistergroup to the remaining genera; (3) the monophyly of the Nandidae + Badidae and their inclusion into a larger monophyletic group – along with the Pristolepididae, Anabantoidei and Channoidei – comprising the Labyrinthici sensu Rosen & Patterson. We also review the morphological and molecular evidence for both the conflicting placement of Pristolepis and the putative sistergroup relationship between the labyrinth fishes (Anabantoidei) and snakeheads (Channoidei).     

Mass extinctions among tetrapods and the quality of the fossil record

The fossil record of tetrapods is very patchy because of the problems of preservation, in terrestrial sediments in particular, and because vertebrates are rarely very abundant. However, the fossil record of tetrapods has the advantages that it is easier to establish a phylogenetic taxonomy than for many invertebrate groups, and there is the potential for more detailed ecological analyses. The relative incompleteness of a fossil record may be assessed readily, and this can be used to test whether drops in overall diversity are related to mass extinctions or to gaps in our knowledge. Absolute incompleteness cannot be assessed directly, but a historical approach may offer clues to future improvements in our knowledge. One of the key problems facing palaeobiologists is paraphyly, the fact that many higher taxa in common use do not contain all of the descendants of the common ancestor. This may be overcome by cladistic analysis and the identification of monophyletic groups. The diversity of tetrapods increased from the Devonian to the Permian, remained roughly constant during the Mesozoic, and then began to increase in the late Cretaceous, and continued to do so during the Tertiary. The rapid radiation of 'modern' tetrapod groups--frogs, salamanders, lizards, snakes, turtles, crocodilians, birds and mammals--was hardly affected by the celebrated end-Cretaceous extinction event. Major mass extinctions among tetrapods took place in the early Permian, late Permian, early Triassic, late Triassic, late Cretaceous, early Oligocene and late Miocene. Many of these events appear to coincide with the major mass extinctions among marine invertebrates, but the tetrapod record is largely equivocal with regard to the theory of periodicity of mass extinctions.     

Phylogenetic relationships of fossil and Recent gonorynchiform fishes (Teleostei: Ostariophysi)

This paper represents the first cladistic analysis of the interrelationships of all nominal fossil and living gonorynchiform genera. Gonorynchiformes is the basal group of the superorder Ostariophysi, and is confirmed as monophyletic on the basis of 12 synapomorphies. The Gonorynchiformes is be subdivided into two monophyletic suborders, Chanoidei and Gonorynchoidei. The Chanoidei includes the family Chanidae, which in turn includes the Recent Chanos plus five fossil genera, grouped in two subfamilies: Chaninae (( Chanos +† Tharrhiai) + † Parachanos +† Dastilbe ) and † Rubiesichthyinae († Rubiesichthys +† Gordichthys ). † Aethalionopsis is the sister-group to the Chanidae. Gonorynchoidei includes two families Gonorynchidae and Kneriidae. Gonorynchidae is formed by ( Gonorynchus, † Notogoneus ) and four fossil taxa of uncertain definition and interrelationships: †Charitosomus, † Charitopsis, † Ramallichthys, and †fudeichthys. The last four genera were previously included in the families †Charitosomidae and †Judeichthyidae, which could not be supported as monophyletic in this analysis. Kneriidae consists of two subfamilies Phractolaeminae with one genus Phractolaemus, and Kneriinae which includes (( Kneria + Parakneria ) + ( Grasseichthys + Cromeria )), the latter two being paedomorphic forms. The Phractolaeminae and the Kneriinae are freshwater African taxa with no known fossil record. The order Gonorynchiformes is represented herein by 18 genera, extending back to the Early Cretaceous. More work is required to clarify the interrelationships of the Gonorynchidae and the paedomorphic characters that apparently played an important role in the evolution of this morphologically diverse group of fishes.     

SPERM WHALE PHYLOGENY REVISITED: ANALYSIS OF THE MORPHOLOGICAL EVIDENCE

Some recent analyses of three mitochondrial DNA regions suggest that sperm whales are the sister group to baleen whales and, therefore, the suborder Odontoceti (toothed whales) constitutes a paraphyletic group. I cladistically analyzed the available morphological data, including that from relevant fossil taxa, for all families of extant cetaceans to test this hypothesis. The results of this analysis unambiguously support a monophyletic Odontoceti including the sperm whales. All synapomorphies that support the Odontoceti node are decisive, not related to the evolution of highly correlated characters, and provide the same result regardless of what order of mammals is used as an outgroup. These numerous, anatomically diverse, and unambiguous characters make this clade one of the best-supported higher-level groupings among mammals. In addition, the fossil evidence refutes a sperm whale/baleen whale clade. Both the molecular and morphological data produce the same unrooted tree. The improper rooting of the molecular tree appears to be producing these seemingly incongruent phylogenies.     

Homologies in the fossil record: The middle ear as a test case

This paper examines the middle ear of fossil living animals in terms of the homologies which have been drawn between its parts in different vertebrate groups. Seven homologies are considered: 1, the middle ear cavity/spiracular pouch; 2, the stapes/hyomandibula; 3, the stapedial/hyomandibular processes; 4 the tympanic membrane; 5, the otic notch; 6, the fenestra ovalis; 7, and the stapedial/hyomandibular foramen. The reasons leading to assessments of homology are reviewed. Homologies 1 and 2, based largely on embryological evidence, are fairly robust, though there are arguments about the details. Homologies 3, 4 and 5 stem from ideas about early tetrapod evolution, and were influenced by contingent factors including the order and time of discovery of early fossil taxa, and perceptions of their phylogeny which resulted from this. They were also influenced by ideas of the evolution of terrestriality among tetrapods. Most of the conceptions have been overturned in recent years by new fossil discoveries and new ways of looking at old data. Homology 6 has been little considered. One possible hypothesis, placed in a strictly archetypal theoretical framework has been ignored but deserves consideration on other grounds. Homology 7 depends on how tetrapods are characterised, not a question which has posed difficulties until recently, but which is likely to with the discovery of intermediate fossil forms.     

Convergent evolution and character correlation in burrowing reptiles: towards a resolution of squamate relationships

The affinities of three problematic groups of elongate, burrowing reptiles (amphisbaenians, dibamids and snakes) are reassessed through a phylogenetic analysis of all the major groups of squamates, including the important fossil taxa Sineoamphisbaena, mosasauroids and Pachyrhachis; 230 phylogenetically informative osteological characters were evaluated in 22 taxa. Snakes (including Pachyrhachis) are anguimorphs, being related firstly to large marine mosasauroids, and secondly to monitor lizards (varanids). Scincids and cordylids are not related to lacertiforms as previously thought, but to anguimorphs. Amphisbaenians and dibamids are closely related, and Sineoamphisbaena is the sister group to this clade. The amphisbaenian-dibamid-Sineoamphisbaena clade, in turn, is related to gekkotans and xantusiids. When the fossil taxa are ignored, snakes, amphisbaenians and dibamids form an apparently well-corroborated clade nested within anguimorphs. However, nearly all of the characters supporting this arrangement are correlated with head-first burrowing (miniaturization, cranial consolidation, body elongation, limb reduction), and invariably co-occur in other tetrapods with similar habits. These characters are potentially very misleading because of their sheer number and because they largely represent reductions or losses. It takes very drastic downweighting of these linked characters to alter tree topology: if fossils are excluded from the analysis, a (probably spurious) clade consisting of elongate, fossorial taxa almost always results. These results underscore the importance of including all relevant taxa in phylogenetic analyses. Inferring squamate phylogeny depends critically on the inclusion of certain (fossil) taxa with combinations of character states that demonstrate convergent evolution of the elongate, fossorial ecomorph in amphisbaenians and dibamids, and in snakes. In the all-taxon analysis, the position of snakes within anguimorphs is more strongly-corroborated than the association of amphisbaenians and dibamids with gekkotans. When the critical fossil taxa are deleted, snakes ‘attract’ the amphisbaenian-dibamid clade on the basis of a suite of correlated characters. While snakes remain anchored in anguimorphs, the amphisbaenian-dibamid clade moves away from gekkotans to join them. Regardless of the varying positions of the three elongate burrowing taxa, the interrelationships between the remaining limbed squamates (‘lizards’) are constant; thus, the heterodox affinities of scincids, cordylids, and xantusiids identified in this analysis appear to be robust. Finally, the position of Pachyrhachis as a basal snake rather than (as recently suggested) a derived snake is supported on both phylogenetic and evolutionary grounds.     

Higher elasmobranch phylogeny and biostratigraphy

Living sharks, skates and rays share several derived skeletal characters that are absent in most extinct elasmobranchs, suggesting a monophyletic group of 'higher' elasmobranchs. Within this group opinions vary as to phylogenetic relationships, although three broad groups are generally recognized. Arguments for and against monophyly of these group (batoids; squalomorphs; galeomorphs) are examined. Many of their contained taxa are also of questionable validity. Cladistic analysis of living galeomorphs reveals a sequence of characters supporting monophyly of the group as whole, but not of its more generalized contained taxa. The temporal distribution of fossil galeomorphs corroborates the hypothesis of relationship suggested by neontological data; i.e. there is considerable stratigraphic harmony with Recent phylogenetic data.     

Fish and tetrapod communities across a marine to brackish salinity gradient in the Pennsylvanian (early Moscovian) Minto Formation of New Brunswick,Canada, and their palaeoecological and palaeogeographical implications

Euryhaline adaptations in Pennsylvanian vertebrates allowed them to inhabit the marine to freshwater spectrum. This is illustrated by new assemblages of fish and tetrapods from the early Moscovian Minto Formation of New Brunswick, Canada. Fish include chondrichthyans (xenacanthids and the enigmatic Ageleodus), acanthodians (gyracanthids and acanthodiforms), sarcopterygians (rhizodontids, megalichthyids and dipnoans), and actinopterygians (eurynotiforms). Tetrapods include small‐ to medium‐sized, and largely aquatic, stem tetrapods (colosteids) and anthracosaurs (embolomeres). A key finding is that the parautochthonous fossil assemblages are preserved across a salinity gradient, with diversity (measured by the Simpson Index) declining from open marine environments, through brackish embayments, and reaching a nadir in tidal estuaries. Chondrichthyans dominate the entire salinity spectrum (65% of fossils), a distribution that demonstrates a euryhaline mode of life, and one large predatory chondrichthyan, Orthacanthus, may have practised filial cannibalism in coastal nurseries because its heteropolar coprolites contain juvenile xenacanthid teeth. In contrast, other fish communities were more common in open marine settings while tetrapods were more common in coastal brackish waters. While all these faunas were also likely to have been euryhaline, their osmoregulation was, perhaps, less versatile. The demonstration of widespread euryhalinity among fish and aquatic tetrapods explains why Pennsylvanian faunas generally show a cosmopolitan biogeography because taxa were able to disperse via seaways. It also resolves the paradox of enriched strontium isotopic signatures observed in these faunas because organisms would have been, at times, exposed to continental water bodies as well. Therefore, our new findings contribute to the long‐running debate about the ecology of Pennsylvanian fishes and tetrapods.