Early Actinopterygians Research Articles

A new, exceptionally well-preserved Permian actinopterygian fish from the Minnekahta Limestone of South Dakota, USA

The evolutionary history of Palaeozoic and early Mesozoic ray-finned fishes (Actinopterygii) is obscured by an incomplete fossil record and under-study of late Palaeozoic (Permo-Carboniferous) actinopterygians. To help address this issue, we describe a new species of ray-finned fish, Concentrilepis minnekahtaensis gen. et sp. nov., from the late early Permian (Kungurian) Minnekahta Limestone of South Dakota (USA). This taxon is represented by Field Museum of Natural History FMNH PF 3721, an exceptionally well-preserved partial three-dimensional head and trunk preserving the external anatomy and some internal elements from the skull, paired fins and scale cover. Concentrilepis is also represented by Yale Peabody Museum 18649, a laterally compressed individual that preserves the posterior portion of the body. This taxon has features of the paraphyletic ‘paleoniscoids’, including an immobile maxilla with a narrow suborbital process and a broad postorbital expansion, a heterocercal caudal fin, and rhombic ganoid scales. Concentrilepis is distinguished from similar species by the dual lateral expansions on its frontal (= parietal of sarcopterygians), the exclusion of the lacrimal from the oral margin, and lateral scales that are longer than deep and ornamented with straight, concentric ridges at their margins and diagonal ridges at their centres. We conducted a maximum parsimony analysis in PAUP and a Bayesian analysis with the Mkv model in MrBayes with 73 other taxa and 222 characters that placed this taxon in a polytomy with other late Palaeozoic and early Mesozoic ‘paleoniscoids’. We also found that the application of Bayesian and parsimony techniques in tandem provided the necessary caution in interpreting our results and focused direction for future study. Continued efforts to gather more morphological data from ‘paleoniscoids’ in concert with the development and testing of novel phylogenetic characters and inference techniques will be necessary to improve our understanding of the interrelationships of late Palaeozoic and early Mesozoic actinopterygians. http://zoobank.org/urn:lsid:zoobank.org:pub:9F12F99D-A33A-41C2-AC34-F4936786020B

Tanyrhinichthys mcallisteri, a long-rostrumed Pennsylvanian ray-finned fish (Actinopterygii) and the simultaneous appearance of novel ecomorphologies in Late Palaeozoic fishes

Abstract The Carboniferous radiation of fishes was marked by the convergent appearance of then-novel but now common ecomorphologies resulting from changes in the relative proportions of traits, including elongation of the front of the skull (rostrum). The earliest ray-finned fishes (Actinopterygii) with elongate rostra are poorly known, obscuring the earliest appearances of a now widespread feature in actinopterygians. We redescribe Tanyrhinichthys mcallisteri, a long-rostrumed actinopterygian from the Upper Pennsylvanian (Missourian) of the Kinney Brick Quarry, New Mexico. Tanyrhinichthys has a lengthened rostrum bearing a sensory canal, ventrally inserted paired fins, posteriorly placed median fins unequal in size and shape, and a heterocercal caudal fin. Tanyrhinichthys shares these features with sturgeons, but lacks chondrostean synapomorphies, indicating convergence on a bottom-feeding lifestyle. Elongate rostra evolved independently in two lineages of bottom-dwelling, freshwater actinopterygians in the Late Pennsylvanian of Euramerica, as well as in at least one North American chondrichthyan (Bandringa rayi). The near-simultaneous appearance of novel ecomorphologies among multiple, distantly related lineages of actinopterygians and chondrichthyans was common during the Carboniferous radiation of fishes. This may reflect global shifts in marine and freshwater ecosystems and environments during the Carboniferous favouring such ecomorphologies, or it may have been contingent on the plasticity of early actinopterygians and chondrichthyans.

A new species of Burguklia (Pisces, Actinopterygii) from the Middle Permian of the Volga Region (European Russia)

A new species of early actinopterygians, Burguklia minichorum sp. nov., is described based on the occurrence of numerous isolated scales from freshwater deposits of the Urzhumian and Severodvinian regional stages (Wordian–Capitanian) of the East European Platform. The type and only hitherto known species of the genus, Burguklia gdali Kazantseva-Selezneva, 1980, comes from the Early Permian of Siberia. The new discovery indicates the existence of non-marine palaeogeographic pathways between Siberia and European Russia during the Permian.

Cranial anatomy of the predatory actinopterygian Brazilichthys macrognathus from the Permian (Cisuralian) Pedra de Fogo Formation, Parnaíba Basin, Brazil

ABSTRACTBrazilichthys macrognathus is the only named actinopterygian from the Permian (Cisuralian) Pedra de Fogo Formation of northeastern Brazil, in which it is represented by a single three-dimensionally preserved, but incompletely described, skull of unclear systematic placement. We used X-ray micro-computed tomography (µCT) to redescribe its anatomy, in particular parts of the internal skeleton, and assess its phylogenetic affinities. We correct errors in the original description, including the number of infraorbital bones and the misidentification of the dermosphenotic as sclerotic ossifications. These reinterpretations of external anatomy are accompanied by new data on internal structure, including the palate, the parasphenoid, and the branchial and hyoid arches. A maximum parsimony analysis of anatomical data resolves Brazilichthys as a stem actinopterygian, crownward of all Devonian species. This placement is supported by the absence of a dermosphenotic posterior ramus and the presence of opercular process of the hyomandibula. A similar placement is suggested by a Bayesian analysis of the same data set, although relationships throughout the tree are less resolved. Our results reject previous interpretations of Brazilichthys as a relative of Birgeriidae, a Triassic group traditionally placed within the actinopterygian crown. Although Acrolepis is too poorly known to be included in our analysis, we also reject relationship between that taxon and Brazilichthys, because their only shared similarities appear to be broadly distributed among early actinopterygians.

A new actinopterygian from the Late Devonian Gogo Formation, Western Australia

AbstractThe study of early actinopterygians (ray‐finned fishes) from the Devonian has been hampered by imperfect preservation in the majority of taxa. The Late Devonian (early Frasnian) Gogo Formation of north‐western Western Australia is notable in producing complete fossil actinopterygians with exceptional three‐dimensional preservation of both the dermal and endoskeletal anatomy. Four taxa have been described and have proved invaluable in understanding the anatomy of early representatives of this clade. Here, we present a fifth Gogo taxon, based on a single exceptionally preserved specimen and representing a new genus and species of early ray‐finned fish. The neurocranium was CT scanned, permitting a detailed examination of the endocast. The new taxon possesses unusual features including a greatly enlarged spiracular opening and extensive spinose ornamentation on the dermal skull bones, median ridge scutes and lepidotrichia. The endocast displays a mosaic of characters, some of which are similar to Mimipiscis and non‐actinopterygian outgroups, while other features are more akin to Raynerius from the Late Devonian of France as well as several stratigraphically younger taxa. A phylogenetic analysis resolves the new form as sister taxon to fishes from the Late Devonian of the northern hemisphere and all post‐Devonian actinopterygians, confirming that the assemblage of fossil ray fins from the Gogo Formation is part of the wider Devonian actinopterygian radiation.

Internal cranial anatomy of Early Triassic species of \u2020Saurichthys (Actinopterygii: \u2020Saurichthyiformes): implications for the phylogenetic placement of \u2020saurichthyiforms

Background†Saurichthyiformes were a successful group of latest Permian–Middle Jurassic predatory actinopterygian fishes and constituted important, widely-distributed components of Triassic marine and freshwater faunas. Their systematic affinities have long been debated, with †saurichthyiforms often being aligned with chondrosteans, a group today comprising sturgeons and paddlefishes. However, their character-rich endocranial anatomy has not been investigated in detail since the first half of the 20th century. Since then, major advances have occurred in terms of our understanding of early actinopterygian anatomy, as well as techniques for extracting morphological data from fossils.ResultsWe used μCT to study the internal cranial anatomy of two of the stratigraphically oldest representatives of †Saurichthys, from the Early Triassic of East Greenland and Nepal. Our work revealed numerous previously unknown characters (e.g., cryptic oticooccipital fissure; intramural diverticula of braincase; nasobasal canals; lateral cranial canal; fused dermohyal), and permitted the reevalution of features relating to the structure of cranial fossae, basicranial circulation and opercular anatomy of the genus. Critically, we reinterpret the former †saurichthyiform opercle as an expanded subopercle. For comparison, we also produced the first digital models of a braincase and endocast of a sturgeon (A. brevirostrum). New information from these taxa was included in a broad phylogenetic analysis of Actinopterygii. †Saurichthyiforms are resolved as close relatives of †Birgeria, forming a clade that constitutes the immediate sister group of crown actinopterygians. However, these and other divergences near the actinopterygian crown node are weakly supported.ConclusionsOur phylogeny disagrees with the historically prevalent hypothesis favoring the chondrostean affinities of †saurichthyiforms. Previously-proposed synapomorphies uniting the two clades, such as the closure of the oticooccipital fissure, the posterior extension of the parasphenoid, and the absence of an opercular process, are all widespread amongst actinopterygians. Others, like those relating to basicranial circulation, are found to be based on erroneous interpretations. Our work renders the †saurichthyiform character complex adequately understood, and permits detailed comparisons with other stem and crown actinopterygians. Our phylogenetic scheme highlights outstanding questions concerning the affinity of many early actinopterygians, such as the Paleozoic–early Mesozoic deep-bodied forms, which are largely caused by lack of endoskeletal data.

Exceptional preservation reveals gastrointestinal anatomy and evolution in early actinopterygian fishes.

Current knowledge about the evolutionary morphology of the vertebrate gastrointestinal tract (GIT) is hindered by the low preservation potential of soft tissues in fossils. Exceptionally preserved cololites of individual †Saurichthys from the Middle Triassic of Switzerland provide unique insights into the evolutionary morphology of the GIT. The GIT of †Saurichthys differed from that of other early actinopterygians, and was convergent to that of some living sharks and rays, in exhibiting up to 30 turns of the spiral valve. Dissections and literature review demonstrate the phylogenetic diversity of GIT features and signs of biological factors that influence its morphology. A phylogenetically informed analysis of a dataset containing 134 taxa suggests that body size and phylogeny are important factors affecting the spiral valve turn counts. The high number of turns in the spiral valve of †Saurichthys and some recent sharks and rays reflect both energetically demanding lifestyles and the evolutionary histories of the groups.

An exceptionally preserved Late Devonian actinopterygian provides a new model for primitive cranial anatomy in ray-finned fishes.

Actinopterygians (ray-finned fishes) are the most diverse living osteichthyan (bony vertebrate) group, with a rich fossil record. However, details of their earliest history during the middle Palaeozoic (Devonian) ‘Age of Fishes' remains sketchy. This stems from an uneven understanding of anatomy in early actinopterygians, with a few well-known species dominating perceptions of primitive conditions. Here we present an exceptionally preserved ray-finned fish from the Late Devonian (Middle Frasnian, ca 373 Ma) of Pas-de-Calais, northern France. This new genus is represented by a single, three-dimensionally preserved skull. CT scanning reveals the presence of an almost complete braincase along with near-fully articulated mandibular, hyoid and gill arches. The neurocranium differs from the coeval Mimipiscis in displaying a short aortic canal with a distinct posterior notch, long grooves for the lateral dorsal aortae, large vestibular fontanelles and a broad postorbital process. Identification of similar but previously unrecognized features in other Devonian actinopterygians suggests that aspects of braincase anatomy in Mimipiscis are apomorphic, questioning its ubiquity as stand-in for generalized actinopterygian conditions. However, the gill skeleton of the new form broadly corresponds to that of Mimipiscis, and adds to an emerging picture of primitive branchial architecture in crown gnathostomes. The new genus is recovered in a polytomy with Mimiidae and a subset of Devonian and stratigraphically younger actinopterygians, with no support found for a monophyletic grouping of Moythomasia with Mimiidae.

Endoskeletal structure in Cheirolepis (Osteichthyes, Actinopterygii), An early ray-finned fish.

As the sister lineage of all other actinopterygians, the Middle to Late Devonian (Eifelian–Frasnian) Cheirolepis occupies a pivotal position in vertebrate phylogeny. Although the dermal skeleton of this taxon has been exhaustively described, very little of its endoskeleton is known, leaving questions of neurocranial and fin evolution in early ray‐finned fishes unresolved. The model for early actinopterygian anatomy has instead been based largely on the Late Devonian (Frasnian) Mimipiscis, preserved in stunning detail from the Gogo Formation of Australia. Here, we present re‐examinations of existing museum specimens through the use of high‐resolution laboratory‐ and synchrotron‐based computed tomography scanning, revealing new details of the neuro‐cranium, hyomandibula and pectoral fin endoskeleton for the Eifelian Cheirolepis trailli. These new data highlight traits considered uncharacteristic of early actinopterygians, including an uninvested dorsal aorta and imperforate propterygium, and corroborate the early divergence of Cheirolepis within actinopterygian phylogeny. These traits represent conspicuous differences between the endoskeletal structure of Cheirolepis and Mimipiscis. Additionally, we describe new aspects of the parasphenoid, vomer and scales, most notably that the scales display peg‐and‐socket articulation and a distinct neck. Collectively, these new data help clarify primitive conditions within ray‐finned fishes, which in turn have important implications for understanding features likely present in the last common ancestor of living osteichthyans.

Heart pigmentation in the gray bichir, Polypterus senegalus (Actinopterygii: Polypteriformes).

The occurrence of pigment cells in the heart is well documented in amphibians, birds and mammals. By contrast, information on heart pigmentation in fish is extremely sparse. The aim is to report the presence of pigment cells over the entire surface of the heart in the gray bichir, Polypterus senegalus. The sample consisted of 12 hearts, which, after gross anatomical examination, were studied using histochemical and immunohistochemical techniques for light microscopy, and transmission electron microscopy. The pigment cells were located in the subepicardium, showing a regular distribution pattern across the whole heart, except for the anterior end of the outflow tract, where the pigmentation was much more intense. The cells contained dark, ovoid-shaped organelles which was consistent with a melanosome cell identity. As in other vertebrates, the physiological role of the pigment cells in the heart of the gray bichir is unknown. The absence of such cells in hearts of other polypteriforms suggests that cells containing melanin are not essential for normal fish heart function. Basing on literature data concerning tetrapods, it can be inferred that the pigment cells of the heart of the gray bichir derive from the neural crest. If this were true, our findings would provide the first evidence for the presence of neural crest-derived cells in the subepicardium of adult hearts of early actinopterygians.

Heads or tails: staged diversification in vertebrate evolutionary radiations

Adaptive radiations, bouts of morphological divergence coupled with taxonomic proliferation, underpin biodiversity. The most widespread model of radiations assumes a single round, or 'early burst', of elevated phenotypic divergence followed by a decline in rates of change or even stasis. A vertebrate-specific model proposes separate stages: initial divergence in postcranial traits related to habitat use, followed by diversification in cranial morphology linked to trophic demands. However, there is little empirical evidence for either hypothesis. Here, we show that, contrary to both models, separate large-scale radiations of actinopterygian fishes proceeded through distinct cranial and later postcranial stages of morphological diversification. Early actinopterygians and acanthomorph teleosts dispersed in cranial morphospace immediately following the end-Devonian extinction and the Cretaceous origin of the acanthomorph clade, respectively. Significant increases in postcranial morphological variation do not occur until one interval after cranial diversification commenced. Therefore, our results question the universality of the 'general vertebrate model'. Based on the results of model-fitting exercises and application of the divergence order test, we find little evidence that the early onset of cranial diversification in these two radiations is due to elevated rates of cranial change relative to postcranial change early in their evolutionary histories. Instead, postcranial and cranial patterns are best fit by an Ornstein-Uhlenbeck model, which is characterized by constant evolutionary rates coupled with a strong central tendency. Other groups have been reported to show early saturation of cranial morphospace or tropic roles early in their histories, but it is unclear whether these patterns are attributable to dynamics similar to those inferred for our two model radiations.

Devonian actinopterygian phylogeny and evolution based on a redescription ofStegotrachelus finlayi

Basal actinopterygian phylogeny is in a state of flux. Discoveries of new Devonian and Carboniferous taxa have not only contributed to our knowledge of these animals, but have also complicated our understanding of their interrelationships. Over the preceding 50 years, there have been several new discoveries and redescriptions of early actinopterygians; however, Tegeolepis clarki, Kentuckia hlavini, and Stegotrachelus finlayi have never been fully redescribed. Although previous authors have re-examined the skull of S. finlayi, new material has recently been collected, in 1973 and 2002. This study synthesizes information from original and new material to fully redescribe S. finlayi, and incorporates these data into a cladistic analysis that examines the interrelationships of early actinopterygians. Phylogenetic systematic data returned support for a monophyletic Stegotrachelidae – an evolutionary systematic hypothesis generated over 40 years ago. Previous palaeoenvironmental and biogeographic hypotheses were tested, with results supporting the independent freshwater origins of several Devonian actinopts, although a regional endemic grade of Gondwanan taxa is not confirmed.

New interpretations of the skull of a primitive bony fish Erpetoichthys calabaricus (Actinopterygii: Cladistia)

Polypterid fishes are considered the basalmost group of extant actinopterygians and may be a direct link to understanding the systematics and evolution of the first bony fishes. Several investigations have been conducted on one member genus, Polypterus; however, since the first specimens of its sister taxon Erpetoichthys calabaricus were described, remarkably little work has been done on the species. We review terminology critical to understanding cranial morphology in polypterids and present a new description of the skull of E. calabaricus as observed through classical methods of skeletal preparation, X-radiographic microfocus computed tomography, and 3D-digital reconstruction. Differences among E. calabaricus and at least three species of Polypterus (P. bichir, P. senegalus, and P. endlicheri), besides the gross variation in size, include an overall elongation of the skull roof observable in most elements of E. calabaricus with a shortening of most associated processes. In addition, several elements present in species of Polypterus are absent in E. calabaricus. As a result, Polypterus should not be used as a proxy for the family Polypteridae to the exclusion of E. calabaricus in phylogenetic studies, which examine early actinopterygians. Each should be treated separately, to resolve inter- and intrarelationships of Polypteridae.

Neuronal organization of the melanin‐concentrating hormone system in primitive actinopterygians: Evolutionary changes leading to teleosts

Hypothalamic melanin-concentrating hormone (MCH) neurones occur in all vertebrates and have an apparent neuromodulatory role. In teleost fish, however, MCH is used also as a neurohypophysial hormone, controlling skin color, and as a hypophysiotrophic peptide. This work describes the central location of immunoreactive MCH perikarya and their projections to the pituitary in a range of ancestral fish to determine the phylogenetic stage when the peptide adopted these roles. In all actinopterygians examined, including polypteriformes, chondrosteans, holosteans, and teleosts, immunoreactive fibers were abundant in the median eminence or, in the case of teleosts, within the pars distalis itself, suggesting MCH acquired a hypophysial regulatory role early in vertebrate evolution. MCH fibers appeared to be absent from the posterior neurohypophysis of the polypteriform Calamoichthys but were evident in this region in the chondrostean Acipensor, the holosteans Lepisosteus and Amia, and all teleosts, suggesting its use as a neurohypophysial hormone. The ability of MCH to induce skin pallor seems to have arisen at a later stage, probably in the preholosteans. This hormonal role coincides with the migration of MCH perikarya away from the ventricular surface and their enlargement into magnocellular neurones. In the higher teleosts, magnocellular hypothalamo-neurohypophysial neurones predominate in size and number, whereas smaller periventricular MCH neurones associated with the paraventricular organ, that are prominent in lampreys, early actinopterygians and tetrapods, are reduced in teleosts. The data suggest that, in teleost fish, earlier functions of the peptide may have become subordinate to its novel pigmentary role.

Actinopterygians from the Namurian of Bearsden, Scotland, with comments on early actinopterygian neurocrania

Abstract Two new stem-actinopterans, Melanecta anneae gen. et sp. nov. and Woodichthys bearsdeni gen. et sp. nov. , are described from the Manse Burn Formation (basal Namurian) of Bearsden, Glasgow. Melanecta resembles Devonian taxa such as Howqualepis and Mimia . Significant features include a large head, long gape, well preserved otoliths, micromeric squamation, broad-based fins, and fringing fulcra restricted to the extended leading pectoral edge. New data on Erpetoichthys otoliths are presented for comparison with those of Melanecta and other early actinopterygians. The primitive actinopterygian condition probably includes partly vateritic otoliths, with a small sagitta and large asteriscus contained within a single chamber. The small scales of Melanecta may be a primitive characteristic, but the apparent independence of factors such as size, histology, and scale row to somite ratios, obscures clear evolutionary trends. Woodichthys resembles Moythomasia and Kentuckia , and includes a flattened but detailed neurocranium. A new neurocranial specimen of Pteronisculus magnus examined for comparison reveals new data on the otic labyrinth, lateral cranial canals, and postorbital wall. Haemopoietic organ accommodation is considered insufficient explanation for lateral cranial canal elaboration. Several new basisphenoidal and postorbital wall characters are identified which are associated with early stages in posterior myodome evolution. Most of these characters are present in Moythomasia durgaringa .

Actinopterygians from the Namurian of Bearsden, Scotland, with comments on early actinopterygian neurocrania

Two new stem-actinopterans,Melanecta anneaegen.etsp. nov.andWoodichthys bearsdenigen.etsp. nov., are described from the Manse Burn Formation (basal Namurian) of Bearsden, Glasgow.Melanectaresembles Devonian taxa such asHowqualepisandMimia. Significant features include a large head, long gape, well preserved otoliths, micromeric squamation, broad-based fins, and fringing fulcra restricted to the extended leading pectoral edge. New data onErpetoichthysotoliths are presented for comparison with those ofMelanectaand other early actinopterygians. The primitive actinopterygian condition probably includes partly vateritic otoliths, with a small sagitta and large asteriscus contained within a single chamber. The small scales ofMelanectamay be a primitive characteristic, but the apparent independence of factors such as size, histology, and scale row to somite ratios, obscures clear evolutionary trends.WoodichthysresemblesMoythomasiaandKentuckia, and includes a flattened but detailed neurocranium. A new neurocranial specimen ofPteronisculus magnusexamined for comparison reveals new data on the otic labyrinth, lateral cranial canals, and postorbital wall. Haemopoietic organ accommodation is considered insufficient explanation for lateral cranial canal elaboration. Several new basisphenoidal and postorbital wall characters are identified which are associated with early stages in posterior myodome evolution. Most of these characters are present inMoythomasia durgaringa.