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Attempts to explain the origin and diversification of vertebrates have commonly invoked the evolution of feeding ecology, contrasting the passive suspension feeding of invertebrate chordates and larval lampreys with active predation in living jawed vertebrates. Of the extinct jawless vertebrates that phylogenetically intercalate these living groups, the feeding apparatus is well-preserved only in the early diverging stem-gnathostome heterostracans. However, its anatomy remains poorly understood. Here, we use X-ray microtomography to characterize the feeding apparatus of the pteraspid heterostracan Rhinopteraspis dunensis (Roemer, 1855). The apparatus is composed of 13 plates arranged approximately bilaterally, most of which articulate from the postoral plate. Our reconstruction shows that the oral plates were capable of rotating around the transverse axis, but likely with limited movement. It also suggests the nasohypophyseal organs opened internally, into the pharynx. The functional morphology of the apparatus in Rhinopteraspis precludes all proposed interpretations of feeding except for suspension/deposit feeding and we interpret the apparatus as having served primarily to moderate the oral gape. This is consistent with evidence that at least some early jawless gnathostomes were suspension feeders and runs contrary to macroecological scenarios that envisage early vertebrate evolution as characterized by a directional trend towards increasingly active food acquisition.

The origin of vertebrate paired appendages is one of the most investigated and debated examples of evolutionary novelty 1-7 . Paired appendages are widely considered as key innovations that enabled new opportunities for controlled swimming and gill ventilation and were prerequisites for the eventual transition from water to land. The past 150 years of debate 8-10 has been shaped by two contentious theories 4,5 : the ventrolateral fin-fold hypothesis 9,10 and the archipterygium hypothesis 8 . The latter proposes that fins and girdles evolved from an ancestral gill arch. Although studies in animal development have revived interest in this idea 11-13 , it is apparently unsupported by fossil evidence. Here we present palaeontological support for a pharyngeal basis for the vertebrate shoulder girdle. We use computed tomography scanning to reveal details of the braincase of Kolymaspis sibirica 14 , an Early Devonian placoderm fish from Siberia, that suggests a pharyngeal component of the shoulder. We combine these findings with refreshed comparative anatomy of placoderms and jawless outgroups to place the origin of the shoulder girdle on the sixth branchial arch. These findings provide a novel framework for understanding the origin of the pectoral girdle. Our evidence clarifies the location of the presumptive head-trunk interface in jawless fishes and explains the constraint on branchial arch number in gnathostomes 15 . The results revive a key aspect of the archipterygium hypothesis and help reconcile it with the ventrolateral fin-fold model., (© 2023. The Author(s).)

The neurocranium is an integral part of the vertebrate head, itself a major evolutionary innovation 1,2 . However, its early history remains poorly understood, with great dissimilarity in form between the two living vertebrate groups: gnathostomes (jawed vertebrates) and cyclostomes (hagfishes and lampreys) 2,3 . The 100 Myr gap separating the Cambrian appearance of vertebrates 4-6 from the earliest three-dimensionally preserved vertebrate neurocrania 7 further obscures the origins of modern states. Here we use computed tomography to describe the cranial anatomy of an Ordovician stem-group gnathostome: Eriptychius americanus from the Harding Sandstone of Colorado, USA 8 . A fossilized head of Eriptychius preserves a symmetrical set of cartilages that we interpret as the preorbital neurocranium, enclosing the fronts of laterally placed orbits, terminally located mouth, olfactory bulbs and pineal organ. This suggests that, in the earliest gnathostomes, the neurocranium filled out the space between the dermal skeleton and brain, like in galeaspids, osteostracans and placoderms and unlike in cyclostomes 2 . However, these cartilages are not fused into a single neurocranial unit, suggesting that this is a derived gnathostome trait. Eriptychius fills a major temporal and phylogenetic gap in our understanding of the evolution of the gnathostome head, revealing a neurocranium with an anatomy unlike that of any previously described vertebrate., (© 2023. The Author(s).)

In modern vertebrates, the craniofacial skeleton is complex, comprising cartilage and bone of the neurocranium, dermatocranium and splanchnocranium (and their derivatives), housing a range of sensory structures such as eyes, nasal and vestibulo-acoustic capsules, with the splanchnocranium including branchial arches, used in respiration and feeding. It is well understood that the skeleton derives from neural crest and mesoderm, while the sensory elements derive from ectodermal thickenings known as placodes. Recent research demonstrates that neural crest and placodes have an evolutionary history outside of vertebrates, while the vertebrate fossil record allows the sequence of the evolution of these various features to be understood. Stem-group vertebrates such as Metaspriggina walcotti (Burgess Shale, Middle Cambrian) possess eyes, paired nasal capsules and well-developed branchial arches, the latter derived from cranial neural crest in extant vertebrates, indicating that placodes and neural crest evolved over 500 million years ago. Since that time the vertebrate craniofacial skeleton has evolved, including different types of bone, of potential neural crest or mesodermal origin. One problematic part of the craniofacial skeleton concerns the evolution of the nasal organs, with evidence for both paired and unpaired nasal sacs being the primitive state for vertebrates., (© 2022 Wiley Periodicals LLC.)

Fossil fish from the Silurian of China continue to surprise. These so-called 'maxillate placoderms', including the newly described Bianchengichthys micros, show a range of anatomical features that question our picture of vertebrate evolution and diversification., (Crown Copyright © 2021. Published by Elsevier Inc. All rights reserved.)

Chondrichthyan dentitions are conventionally interpreted to reflect the ancestral gnathostome condition but interpretations of osteichthyan dental evolution in this light have proved unsuccessful, perhaps because chondrichthyan dentitions are equally specialized, or else evolved independently. Ischnacanthid acanthodians are stem-Chondrichthyes; as phylogenetic intermediates of osteichthyans and crown-chondrichthyans, the nature of their enigmatic dentition may inform homology and the ancestral gnathostome condition. Here we show that ischnacanthid marginal dentitions were statodont, composed of multicuspidate teeth added in distally diverging rows and through proximal superpositional replacement, while their symphyseal tooth whorls are comparable to chondrichthyan and osteichthyan counterparts. Ancestral state estimation indicates the presence of oral tubercles on the jaws of the gnathostome crown-ancestor; tooth whorls or tooth rows evolved independently in placoderms, osteichthyans, ischnacanthids, other acanthodians and crown-chondrichthyans. Crown-chondrichthyan dentitions are derived relative to the gnathostome crown-ancestor, which possessed a simple dentition and lacked a permanent dental lamina, which evolved independently in Chondrichthyes and Osteichthyes.

In classical theory, teeth of vertebrate dentitions evolved from co-option of external skin denticles into the oral cavity. This hypothesis predicts that ordered tooth arrangement and regulated replacement in the oral dentition were also derived from skin denticles. The fossil batoid ray Schizorhiza stromeri (Chondrichthyes; Cretaceous) provides a test of this theory. Schizorhiza preserves an extended cartilaginous rostrum with closely spaced, alternating saw-teeth, different from sawfish and sawsharks today. Multiple replacement teeth reveal unique new data from micro-CT scanning, showing how the 'cone-in-cone' series of ordered saw-teeth sets arrange themselves developmentally, to become enclosed by the roots of pre-existing saw-teeth. At the rostrum tip, newly developing saw-teeth are present, as mineralized crown tips within a vascular, cartilaginous furrow; these reorient via two 90° rotations then relocate laterally between previously formed roots. Saw-tooth replacement slows mid-rostrum where fewer saw-teeth are regenerated. These exceptional developmental data reveal regulated order for serial self-renewal, maintaining the saw edge with ever-increasing saw-tooth size. This mimics tooth replacement in chondrichthyans, but differs in the crown reorientation and their enclosure directly between roots of predecessor saw-teeth. Schizorhiza saw-tooth development is decoupled from the jaw teeth and their replacement, dependent on a dental lamina. This highly specialized rostral saw, derived from diversification of skin denticles, is distinct from the dentition and demonstrates the potential developmental plasticity of skin denticles., (© 2015 The Authors.)

The outer armour of fossil jawless fishes (Heterostraci) is, predominantly, a bone with a superficial ornament of dentine tubercles surrounded by pores leading to flask-shaped crypts (ampullae). However, despite the extensive bone present in these early dermal skeletons, damage was repaired almost exclusively with dentine. Consolidation of bone, by dentine invading and filling the vascular spaces, was previously recognized in Psammolepis and other heterostracans but was associated with ageing and dermal shield wear (reparative). Here, we describe wound repair by deposition of dentine directly onto a bony scaffold of fragmented bone. An extensive wound response occurred from massive deposition of dentine (reactionary), traced from tubercle pulp cavities and surrounding ampullae. These structures may provide the cells to make reparative and reactionary dentine, as in mammalian teeth today in response to stimuli (functional wear or damage). We suggest in Psammolepis, repair involved mobilization of these cells in response to a local stimulatory mechanism, for example, predator damage. By comparison, almost no new bone is detected in repair of the Psammolepis shield. Dentine infilling bone vascular tissue spaces of both abraded dentine and wounded bone suggests that recruitment of this process has been evolutionarily conserved over 380 Myr and precedes osteogenic skeletal repair.

The adhesion disc of living remoras (Echeneoidea: Echeneidae) represents one of the most remarkable structural innovations within fishes. Although homology between the spinous dorsal fin of generalized acanthomorph fishes and the remora adhesion disc is widely accepted, the sequence of evolutionary-rather than developmental-transformations leading from one to the other has remained unclear. Here, we show that the early remora †Opisthomyzon (Echeneoidea: †Opisthomyzonidae), from the early Oligocene (Rupelian) of Switzerland, is a stem-group echeneid and provides unique insights into the evolutionary assembly of the unusual body plan characteristic of all living remoras. The adhesion disc of †Opisthomyzon retains ancestral features found in the spiny dorsal fins of remora outgroups, and corroborates developmental interpretations of the homology of individual skeletal components of the disc. †Opisthomyzon indicates that the adhesion disc originated in a postcranial position, and that other specializations (including the origin of pectination, subdivision of median fin spines into paired lamellae, increase in segment count and migration to a supracranial position) took place later in the evolutionary history of remoras. This phylogenetic sequence of transformation finds some parallels in the order of ontogenetic changes to the disc documented for living remoras.

Teeth and jaws constitute a model of the evolutionary developmental biology concept of modularity and they have been considered the key innovations underpinning a classic example of adaptive radiation. However, their evolutionary origins are much debated. Placoderms comprise an extinct sister clade or grade to the clade containing chondrichthyans and osteichthyans, and although they clearly possess jaws, previous studies have suggested that they lack teeth, that they possess convergently evolved tooth-like structures or that they possess true teeth. Here we use synchrotron radiation X-ray tomographic microscopy (SRXTM) of a developmental series of Compagopiscis croucheri (Arthrodira) to show that placoderm jaws are composed of distinct cartilages and gnathal ossifications in both jaws, and a dermal element in the lower jaw. The gnathal ossification is a composite of distinct teeth that developed in succession, polarized along three distinct vectors, comparable to tooth families. The teeth are composed of dentine and bone, and show a distinct pulp cavity that is infilled centripetally as development proceeds. This pattern is repeated in other placoderms, but differs from the structure and development of tooth-like structures in the postbranchial lamina and dermal skeleton of Compagopiscis and other placoderms. We interpret this evidence to indicate that Compagopiscis and other arthrodires possessed teeth, but that tooth and jaw development was not developmentally or structurally integrated in placoderms. Teeth did not evolve convergently among the extant and extinct classes of early jawed vertebrates but, rather, successional teeth evolved within the gnathostome stem-lineage soon after the origin of jaws. The chimaeric developmental origin of this model of modularity reflects the distinct evolutionary origins of teeth and of component elements of the jaws.

Cartilaginous vertebrate skeletons leave few records as fossils, unless mineralized. Here, we report outstanding preservation of early stages of cartilage differentiation, present in the Devonian vertebrate Palaeospondylus gunni. In large specimens of Palaeospondylus, enlarged, hypertrophic cell spaces (lacunae) are dominant in the cartilage matrix, each defined by thin mineralized matrix, where phosphorus and calcium co-occur. This is comparable to living endochondral cartilage, where cell hypertrophy and matrix mineralization mark the end of an ontogenetic process of cell growth and division before bone formation. New information from small individuals of Palaeospondylus demonstrates that the skeleton comprises mostly unmineralized organic matrix with fewer hypertrophic cell spaces, these occurring only in the central regions of each element. Only here has the surrounding matrix begun to mineralize, differing from the larger specimens in that phosphorus is dominant with little associated calcium at these earlier stages. This reflects cellular control of mineralization in living tissues through phosphate accumulation around hypertrophic cells, with later increase in calcium in the cartilaginous matrix. These features are always associated with endochondral bone development, but in the Palaeospondylus skeleton, this bone never develops. This skeletal state is thus far unique among vertebrates, with two alternative explanations: either later stages of endochondral bone development have been lost in Palaeospondylus, or, in a stepwise acquisition of the mineralized skeleton, these late stages have not yet evolved., (Copyright © 2011 Wiley Periodicals, Inc., A Wiley Company.)

Palaeospondylus gunni (Devonian, Scotland) is an enigmatic vertebrate, assigned to various jawless and jawed groups since its original description. New sections through the whole body allow description of a novel skeletal tissue for Palaeospondylus, comprising the entire skeleton. This tissue is mineralized cartilage and is characterized by large cell spaces embedded in minimal matrix. Bone is completely absent. Calcium phosphate mineralization has a differential topography of deposition within the cartilage that reflects a biogenic origin, despite subsequent diagenetic modification. This combination of hypertrophied cell spaces surrounded by regionalized mineralized matrix differs from all other cartilage in fossil and extant vertebrates. However, it compares most closely to gnathostome endochondral bone in early developmental stages. For example, Palaeospondylus skeletal histology differs from the Devonian agnathan Euphanerops and extant lamprey cartilage. Comparison with mineralized cartilage of armored fossil agnathans and placoderms shows the histology is not comparable to globular calcified cartilage. It also differs from that in extant chondrichthyan mineralized tesserae, which is restricted to a subperichondral zone. Amongst this diversity of calcified cartilage types we discuss various interpretations, including one that implicates tissue either in developmental stasis, before osteoblasts can deposit bone, or at a phylogenetic stage when this step has not evolved. These very different hypotheses highlight difficulties in interpreting fossil ontogenies when phylogenetic relationships are uncertain. Nevertheless, we propose that the composition of the Palaeospondylus skeleton represents a fossilized ontogenetic stage of endochondral bone, a type of bone characteristic of osteichthyan vertebrates., ((c) 2009. Published by Elsevier Ltd. All rights reserved.)

Evidence of reproductive biology is extremely rare in the fossil record. Recently the first known embryos were discovered within the Placodermi, an extinct class of armoured fish, indicating a viviparous mode of reproduction in a vertebrate group outside the crown-group Gnathostomata (Chondrichthyes and Osteichthyes). These embryos were found in ptyctodontids, a small group of placoderms phylogenetically basal to the largest group, the Arthrodira. Here we report the discovery of embryos in the Arthrodira inside specimens of Incisoscutum ritchiei from the Upper Devonian Gogo Formation of Western Australia (approximately 380 million years ago), providing the first evidence, to our knowledge, for reproduction using internal fertilization in this diverse group. We show that Incisoscutum and some phyllolepid arthrodires possessed pelvic girdles with long basipterygia that articulated distally with an additional cartilaginous element or series, as in chondrichthyans, indicating that the pelvic fin was used in copulation. As homology between similar pelvic girdle skeletal structures in ptyctodontids, arthrodires and chondrichthyans is difficult to reconcile in the light of current phylogenies of lower gnathostomes, we explain these similarities as being most likely due to convergence (homoplasy). These new finds confirm that reproduction by internal fertilization and viviparity was much more widespread in the earliest gnathostomes than had been previously appreciated.

The enigmatic Devonian fossil Palaeospondylus gunni was identified as a larval form, metamorphosing into the lungfish Dipterus valenciennesi. Morphological features used to identify P. gunni as a larval lungfish include enlarged cranial ribs, rudimentary limb girdles, and absence of teeth. However, this combination of features does not characterize the extant lungfish Neoceratodus forsteri, even at very young stages, nor early stages of Devonian and younger fossil lungfish. Absence of teeth is problematic because early ontogenetic stages of fossil and living lungfish possess full dentitions including marginal teeth. Also problematic are cranial ribs as a defining character of lungfish, as these also occur in certain actinopterygians. It is argued that Neoceratodus is an obligate neotene (reproductively mature larva), with the implication that metamorphosis was a feature of the ontogeny of early lungfish. Pedomorphic characters have been recognized in Neoceratodus and other post-Devonian lungfish, including large cells and correspondingly large genome size; these latter characters correlate with neoteny in salamanders. Small cells preserved in fossil bone suggest that Devonian lungfish had a smaller genome than post-Devonian lungfish, implying that they were not neotenic. As fossil lungfish cell sizes (and genomes) increased in the late Paleozoic, the diversity of lungfish morphologies decreased, so that taxa like Sagenodus and Conchopoma show morphological similarity to Neoceratodus, marking a point in phylogeny at which metamorphosis was potentially lost. Since ancestral larval characters are retained in neotenic adults, we predict that Devonian larvae should resemble these post-Devonian taxa, a prediction which Palaeospondylus does not fulfill., ((c) 2006 Wiley-Liss, Inc.)

Although the lungfish (Dipnoi) belong within the Osteichthyes, their dentitions are radically different from other osteichthyans. Lungfish dentitions also show a uniquely high structural disparity during the early evolution of the group, partly owing to the independent variation of odontogenic and odontoclastic processes that are tightly and stereotypically coordinated in other osteichthyans. We present a phylogenetic analysis of early lungfishes incorporating a novel approach to coding these process characters in preference to the resultant adult dental morphology. The results only partially resolve the interrelationships of Devonian dipnoans, but show that the widely discussed hypothesis of separate tooth-plated, dentine-plated, and denticulated lineages is unlikely to be true. The dipnoan status of Diabolepis is corroborated. Lungfish dentitions seem to have undergone extensive and nonparsimonious evolution during the early history of the group, but much of the resulting disparity can be explained by a modest number of evolutionary steps in the underlying developmental processes, those for dental formation (odontogenic) and those for the remodeling of dentine tissue (odontoclastic). Later in lungfish evolution, this disparity was lost as the group settled to a pattern of dental development that is just as stereotypic as, but completely different from, that of other osteichthyans.

The correlation of the origin of teeth with jaws in vertebrate history has recently been challenged with an alternative to the canonical view of teeth deriving from separate skin denticles. This alternative proposes that organized denticle whorls on the pharyngeal (gill) arches in the fossil jawless fish Loganellia are precursors to tooth families developing from a dental lamina along the jaw, such as those occurring in sharks, acanthodians, and bony fishes. This not only indicates that homologs of tooth families were present, but also illustrates that they possessed the relevant developmental controls, prior to the evolution of jaws. However, in the Placodermi, a phylogenetically basal group of jawed fishes, the state of pharyngeal denticles is poorly known, tooth whorls are absent, and the presence of teeth homologous to those in extant jawed fishes (Chondrichthyes + Osteichthyes) is controversial. Thus, placoderms would seem to provide little evidence for the early evolution of dentitions, or of denticle whorls, or tooth families, at the base of the clade of jawed fishes. However, organized denticles do occur at the rear of the placoderm gill chamber, but are associated with the postbranchial lamina of the anterior trunkshield, assumed to be part of the dermal cover. Significantly, these denticles have a different organization and morphology relative to the external dermal trunkshield tubercles. We propose that they represent a denticulate part of the visceral skeleton, under the influence of pharyngeal patterning controls comparable to those for pharyngeal denticles in other jawed vertebrates and Loganellia., (Copyright 2003 Wiley-Liss, Inc.)

Placoderms are extinct jawed fishes of the class Placodermi and are basal among jawed vertebrates. It is generally thought that teeth are absent in placoderms and that the phylogenetic origin of teeth occurred after the evolution of jaws. However, we now report the presence of tooth rows in more derived placoderms, the arthrodires. New teeth are composed of gnathostome-type dentine and develop at specific locations. Hence, it appears that these placoderm teeth develop and are regulated as in other jawed vertebrates. Because tooth development occurs only in derived forms of placoderms, we suggest that teeth evolved at least twice, through a mechanism of convergent evolution.

The Holocephali is a major group of chondrichthyan fishes, the sister taxon to the sharks and rays (Elasmobranchii). However, the dentition of extant holocephalans is very different from that of the elasmobranchs, lacking individual tooth renewal, but comprising dental plates made entirely of self-renewing dentine. This renewal of all tissues occurs at the postero-lingual plate surface, as a function of their statodont condition. The fossil record of the holocephalans illuminates multiple different trends in the dentition, including shark-like teeth through to those with dentitions completely lacking individual teeth. Different taxa illustrate developmental retention of teeth but with fusion in their serial development. Dentine of different varieties comprises these teeth and composite dental plates, whose histology includes vascularized tubes within coronal dentine, merging with basal trabecular dentine. In this coronal vascularized dentine, extensive hypermineralization forms a wear resistant tissue transformed into a variety of morphologies. Through evolution, hypermineralized dentine becomes enclosed within the trabecular dentine, and specialized by reduction into specific zones within a composite dental plate, with these increasing in morphological disparity, all reflecting loss of defined teeth but retention of dentine production from the inherited developmental package. [ABSTRACT FROM AUTHOR]

The Onychodontiformes is a poorly known sarcopterygian fish group, with four genera currently described, predominantly from the Middle-Late Devonian. A new onychodont, Bukkanodus jesseni n. gen. and sp., from the Fairy Formation, Victoria, Australia. is of Early Devonian (mid-late Pragian) age, representing one of the oldest known occurrences of this group. Other Pragian onychodonts are represented by a single lower jaw from China, while older occurrences (Lochkovian) include a lower jaw also from China and isolated teeth from Nevada. The Australian material, though disarticulated, includes skull, jaw, and palatal and dental specimens. These specimens share characteristics with younger onychodont taxa, including a laterally compressed tooth whorl with main and accessory rows containing an equal number of teeth, an anteriorly arched shape of the preniaxilla. a lateral rostral bone participating in the orbital margin, parietals separated in the midline by small bones, the insertion of the basisphenoid region of the braincase into a posterior area on the parasphenoid, and an ethmoidal sensory canal running along the dorsal margin of the premaxilla rather than through the bone. Bukkanodus jesseni differs from other onychodonts in the presence of a distinct vomer, coronoids with fang pairs, a restricted herringbone pattern of ribs on enamel ridges of teeth, and the presence of a cluster of large pores on certain skull and jaw bones. The course of the ethmoid sensory canal and this cluster of pores are plesiomorphic features for the Onychodontiformes, also occurring in primitive sarcopterygian taxa such as Youngolepis, Powichthys, and Kenichthys. The opening of this cluster on the internal surface of the premaxilla resembles the rostral organ of derived coelacanths. [ABSTRACT FROM AUTHOR]

The fossil group Placodermi is the most phylogenetically basal of the clade of jawed vertebrates but lacks a marginal dentition comparable to that of the dentate Chondrichthyes, Acanthodii and Osteichthyes (crown-group Gnathostomata). The teeth of crown-group gnathostomes are part of an ordered dentition replaced from, and patterned by, a dental lamina, exemplified by the elasmobranch model. A dentition recognised by these criteria has been previously judged absent in placoderms, based on structural evidence such as absence of tooth whorls and typical vertebrate dentine. However, evidence for regulated tooth addition in a precise spatiotemporal order can be observed in placoderms, but significantly, only within the group Arthrodira. In these fossils, as in other jawed vertebrates with statodont, non-replacing dentitions, new teeth are added at the ends of rows below the bite, but in line with biting edges of the dentition. The pattern is different on each gnathal bone and probably arises from single odontogenic primordia on each, but tooth rows are arranged in a distinctive placoderm pattern. New teeth are made of regular dentine comparable to that of crown-gnathostomes, formed from a pulp cavity. This differs from semidentine previously described for placoderm gnathalia, a type present in the external dermal tubercles. The Arthrodira is a derived taxon within the Placodermi, hence origin of teeth in placoderms occurs late in the phylogeny and teeth are convergently derived, relative to those of other jawed vertebrates. More basal placoderm taxa adopted other strategies for providing biting surfaces and these vary substantially, but include addition of denticles to the growing gnathal plates, at the margins of pre-existing denticle patches. These alternative strategies and apparent absence of regular dentine have led to previous interpretations that teeth were entirely absent from the placoderm dentition. A consensus view emerged that a dentition, as developed... [ABSTRACT FROM AUTHOR]