The three-dimensionally articulated oral apparatus of a Devonian heterostracan sheds light on feeding in Palaeozoic jawless fishes.

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.

Fossil evidence for a pharyngeal origin of the vertebrate pectoral girdle.

The origin of vertebrate paired appendages is one of the most investigated and debated examples of evolutionary novelty1-7. 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 debate8-10 has been shaped by two contentious theories4,5: the ventrolateral fin-fold hypothesis9,10 and the archipterygium hypothesis8. The latter proposes that fins and girdles evolved from an ancestral gill arch. Although studies in animal development have revived interest in this idea11-13, 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 sibirica14, 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 gnathostomes15. The results revive a key aspect of the archipterygium hypothesis and help reconcile it with the ventrolateral fin-fold model.

The oldest three-dimensionally preserved vertebrate neurocranium.

The neurocranium is an integral part of the vertebrate head, itself a major evolutionary innovation1,2. 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 vertebrates4-6 from the earliest three-dimensionally preserved vertebrate neurocrania7 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, USA8. 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 cyclostomes2. 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.

Tales of the tongue.

Since first evolving 350 million years ago, the tongue has taken myriad forms, unlocking new niches and boosting the diversity of life.

A high latitude Gondwanan species of the Late Devonian tristichopterid Hyneria (Osteichthyes: Sarcopterygii).

We describe the largest bony fish in the Late Devonian (late Famennian) fossil assemblage from Waterloo Farm near Makhanda/Grahamstown, South Africa. It is a giant member of the extinct clade Tristichopteridae (Sarcopterygii: Tetrapodomorpha) and most closely resembles Hyneria lindae from the late Famennian Catskill Formation of Pennsylvania, USA. Notwithstanding the overall similarity, it can be distinguished from H. lindae on a number of morphological points and is accordingly described as a new species, H. udlezinye sp. nov. The preserved material comprises most of the dermal skull, lower jaw, gill cover and shoulder girdle. The cranial endoskeleton appears to have been unossified and is not preserved, apart from a fragment of the hyoid arch adhering to a subopercular, but the postcranial endoskeleton is represented by an ulnare, some semi-articulated neural spines, and the basal plate of a median fin. The discovery of H. udlezinye shows that Hyneria is a cosmopolitan genus extending into the high latitudes of Gondwana, not a Euramerican endemic. It supports the contention that the derived clade of giant tristichopterids, which alongside Hyneria includes such genera as Eusthenodon, Edenopteron and Mandageria, originated in Gondwana.

Organization of the body wall in lampreys informs the evolution of the vertebrate paired appendages.

Vertebrate paired appendages are one of the most important evolutionary novelties in vertebrates. During embryogenesis, the skeletal elements of paired appendages differentiate from the somatic mesoderm, which is a layer of lateral plate mesoderm. However, the presence of the somatic mesoderm in the common ancestor of vertebrates has been controversial. To address this problem, it is necessary but insufficient to understand the developmental process of lateral plate mesoderm formation in lamprey (jawless vertebrates) embryos. Here, I show the presence of the somatic mesoderm in lamprey (Lethenteron camtschaticum) embryos using plastic sectioning and transmission electron microscopy analysis. During the early pharyngeal stages, the somatic mesoderm transforms from the lateral plate mesoderm in the trunk region. Soon after, when the cardiac structures were morphologically distinct, the somatic mesoderm was recognized through the cardiac to more caudal regions. These findings indicated that the somatic mesoderm evolved before the emergence of paired appendages. I also discuss the developmental changes in the body wall organization in the common ancestor of vertebrates, which is likely related to the evolution of the paired appendages.

Evidence for high-performance suction feeding in the Pennsylvanian stem-group holocephalan Iniopera.

The Carboniferous (358.9 to 298.9 Ma) saw the emergence of marine ecosystems dominated by modern vertebrate groups, including abundant stem-group holocephalans (chimaeras and relatives). Compared with the handful of anatomically conservative holocephalan genera alive today-demersal durophages all-these animals were astonishingly morphologically diverse, and bizarre anatomies in groups such as iniopterygians hint at specialized ecological roles foreshadowing those of the later, suction-feeding neopterygians. However, flattened fossils usually obscure these animals' functional morphologies and how they fitted into these important early ecosystems. Here, we use three-dimensional (3D) methods to show that the musculoskeletal anatomy of the uniquely 3D-preserved iniopterygian Iniopera can be best interpreted as being similar to that of living holocephalans rather than elasmobranchs but that it was mechanically unsuited to durophagy. Rather, Iniopera had a small, anteriorly oriented mouth aperture, expandable pharynx, and strong muscular links among the pectoral girdle, neurocranium, and ventral pharynx consistent with high-performance suction feeding, something exhibited by no living holocephalan and never clearly characterized in any of the extinct members of the holocephalan stem-group. Remarkably, in adapting a distinctly holocephalan anatomy to suction feeding, Iniopera is more comparable to modern tetrapod suction feeders than to the more closely related high-performance suction-feeding elasmobranchs. This raises questions about the assumed role of durophagy in the evolution of holocephalans' distinctive anatomy and offers a rare glimpse into the breadth of ecological niches filled by holocephalans in a pre-neopterygian world.

Vertebrate cranial evolution: Contributions and conflict from the fossil record.

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.

Patterns of vertebrate biodiversity in a tropical dry and mangrove forest matrix / Patrones de biodiversidad de vertebrados en una matriz de bosque seco tropical y manglar

Introduction: Tropical dry forests and mangroves, two of the world's most endangered ecosystems, each host a different set of environmental conditions which may support unique assemblages of species. However, few studies have looked at the unique vertebrate biodiversity in regions where both habitats occur side-by-side. Objective: To assess the vertebrate diversity and patterns of habitat usage in a mangrove and tropical dry forest matrix in an unprotected region of Northwestern Costa Rica. Methods: The study was conducted in a 7 km2 matrix of mangrove and tropical dry forests between Cabuyal and Zapotillal bays in Northwestern Costa Rica, south of Santa Rosa National Park. From September 2017 to March 2018, we used 13 automatic camera traps over 1 498 trap days to capture species utilizing the region and assess their patterns of habitat usage both spatially and temporally. Results: Seventy vertebrate species from 42 families in 27 orders were detected, including several globally threatened species. Over half of all species were detected in only one habitat, particularly amongst avian (78 %) and mammalian (42 %) species. Tropical dry forests hosted the greatest number of unique species and supported a greater percentage of herbivores than mangrove or edge habitats, which were dominated by carnivorous and omnivorous species. Mean detections per camera trap of all species increased significantly from the coldest and wettest month (Oct) to the hottest and driest months (Jan & Feb) in tropical dry forests. Sample-based rarefaction analysis revealed that survey length was sufficient to sample the tropical dry forest and edge habitats, though mangroves require further sampling. Conclusions: Taxa found to utilize different forest types may utilize each for different stages of their life cycle, moving between areas as environmental conditions change throughout the year. General patterns of global biodiversity favoring carnivore and omnivore usage of mangrove forests was confirmed in our study.

Introducción: Los bosques secos tropicales y los manglares, dos de los ecosistemas más amenazados del mundo, albergan cada uno un grupo de condiciones ambientales que pueden albergar conjuntos únicos de especies. Sin embargo, pocos estudios han analizado la biodiversidad única de vertebrados en regiones donde ambos hábitats se encuentran uno al lado del otro. Objetivo: Evaluar la diversidad de vertebrados y los patrones de uso del hábitat en una matriz de manglar y bosque seco tropical en una región no protegida del noroeste de Costa Rica. Métodos: El estudio se realizó en una matriz de 7 km2 de manglares y bosques secos tropicales en las bahías de Cabuyal y Zapotillal en el noroeste de Costa Rica, al sur del Parque Nacional Santa Rosa. De septiembre 2017 a marzo 2018, utilizamos 13 cámaras trampa automáticas durante 1 498 días trampa para capturar especies que utilizan la región y evaluar sus patrones de uso espacial y temporal del hábitat. Resultados: Se detectaron 70 especies de vertebrados de 42 familias y 27 órdenes, incluidas varias especies amenazadas a nivel mundial. Más de la mitad de todas las especies se encontraron en un solo hábitat, particularmente aves (78 %) y mamíferos (42 %). Los bosques secos tropicales albergan el mayor número de especies únicas y sustentan un mayor porcentaje de herbívoros que los hábitats de borde de manglares, que estaban dominados u hospedados por especies carnívoras y omnívoras. Las detecciones promedio por cámara trampa de todas las especies aumentaron significativamente desde el mes más frío y húmedo (octubre) hasta los meses más cálidos y secos (enero y febrero) en los bosques secos tropicales. El análisis de rarefacción basado en muestras reveló que la duración del estudio fue suficiente para muestrear los hábitats de bosque seco tropical y de borde, aunque los manglares requieren más muestreo. Conclusiones: Se encontró que los taxones pueden usar varios tipos de bosque en las diferentes etapas de su ciclo de vida, moviéndose entre áreas a medida que las condiciones ambientales cambian a lo largo del año. En nuestro estudio se confirmaron patrones generales de la biodiversidad global que favorecen el uso de los bosques de manglar por parte de carnívoros y omnívoros.

The oldest gnathostome teeth.

Mandibular teeth and dentitions are features of jawed vertebrates that were first acquired by the Palaeozoic ancestors1-3 of living chondrichthyans and osteichthyans. The fossil record currently points to the latter part of the Silurian period4-7 (around 425 million years ago) as a minimum date for the appearance of gnathostome teeth and to the evolution of growth and replacement mechanisms of mandibular dentitions in the subsequent Devonian period2,8-10. Here we provide, to our knowledge, the earliest direct evidence for jawed vertebrates by describing Qianodus duplicis, a new genus and species of an early Silurian gnathostome based on isolated tooth whorls from Guizhou province, China. The whorls possess non-shedding teeth arranged in a pair of rows that demonstrate a number of features found in modern gnathostome groups. These include lingual addition of teeth in offset rows and maintenance of this patterning throughout whorl development. Our data extend the record of toothed gnathostomes by 14 million years from the late Silurian into the early Silurian (around 439 million years ago) and are important for documenting the initial diversification of vertebrates. Our analyses add to mounting fossil evidence that supports an earlier emergence of jawed vertebrates as part of the Great Ordovician Biodiversification Event (approximately 485-445 million years ago).

Galeaspid anatomy and the origin of vertebrate paired appendages.

Paired fins are a major innovation1,2 that evolved in the jawed vertebrate lineage after divergence from living jawless vertebrates3. Extinct jawless armoured stem gnathostomes show a diversity of paired body-wall extensions, ranging from skeletal processes to simple flaps4. By contrast, osteostracans (a sister group to jawed vertebrates) are interpreted to have the first true paired appendages in a pectoral position, with pelvic appendages evolving later in association with jaws5. Here we show, on the basis of articulated remains of Tujiaaspis vividus from the Silurian period of China, that galeaspids (a sister group to both osteostracans and jawed vertebrates) possessed three unpaired dorsal fins, an approximately symmetrical hypochordal tail and a pair of continuous, branchial-to-caudal ventrolateral fins. The ventrolateral fins are similar to paired fin flaps in other stem gnathostomes, and specifically to the ventrolateral ridges of cephalaspid osteostracans that also possess differentiated pectoral fins. The ventrolateral fins are compatible with aspects of the fin-fold hypothesis for the origin of vertebrate paired appendages6-10. Galeaspids have a precursor condition to osteostracans and jawed vertebrates in which paired fins arose initially as continuous pectoral-pelvic lateral fins that our computed fluid-dynamics experiments show passively generated lift. Only later in the stem lineage to osteostracans and jawed vertebrates did pectoral fins differentiate anteriorly. This later differentiation was followed by restriction of the remaining field of fin competence to a pelvic position, facilitating active propulsion and steering.

The oldest complete jawed vertebrates from the early Silurian of China.

Molecular studies suggest that the origin of jawed vertebrates was no later than the Late Ordovician period (around 450 million years ago (Ma))1,2. Together with disarticulated micro-remains of putative chondrichthyans from the Ordovician and early Silurian period3-8, these analyses suggest an evolutionary proliferation of jawed vertebrates before, and immediately after, the end-Ordovician mass extinction. However, until now, the earliest complete fossils of jawed fishes for which a detailed reconstruction of their morphology was possible came from late Silurian assemblages (about 425 Ma)9-13. The dearth of articulated, whole-body fossils from before the late Silurian has long rendered the earliest history of jawed vertebrates obscure. Here we report a newly discovered Konservat-Lagerstätte, which is marked by the presence of diverse, well-preserved jawed fishes with complete bodies, from the early Silurian (Telychian age, around 436 Ma) of Chongqing, South China. The dominant species, a 'placoderm' or jawed stem gnathostome, which we name Xiushanosteus mirabilis gen. et sp. nov., combines characters from major placoderm subgroups14-17 and foreshadows the transformation of the skull roof pattern from the placoderm to the osteichthyan condition10. The chondrichthyan Shenacanthus vermiformis gen. et sp. nov. exhibits extensive thoracic armour plates that were previously unknown in this lineage, and include a large median dorsal plate as in placoderms14-16, combined with a conventional chondrichthyan bauplan18,19. Together, these species reveal a previously unseen diversification of jawed vertebrates in the early Silurian, and provide detailed insights into the whole-body morphology of the jawed vertebrates of this period.

Ultrastructure reveals ancestral vertebrate pharyngeal skeleton in yunnanozoans.

Pharyngeal arches are a key innovation that likely contributed to the evolution of the jaws and braincase of vertebrates. It has long been hypothesized that the pharyngeal (branchial) arch evolved from an unjointed cartilaginous rod in vertebrate ancestors such as that in the nonvertebrate chordate amphioxus, but whether such ancestral anatomy existed remains unknown. The pharyngeal skeleton of controversial Cambrian animals called yunnanozoans may contain the oldest fossil evidence constraining the early evolution of the arches, yet its correlation with that of vertebrates is still disputed. By examining additional specimens in previously unexplored techniques (for example, x-ray microtomography, scanning and transmission electron microscopy, and energy dispersive spectrometry element mapping), we found evidence that yunnanozoan branchial arches consist of cellular cartilage with an extracellular matrix dominated by microfibrils, a feature hitherto considered specific to vertebrates. Our phylogenetic analysis provides further support that yunnanozoans are stem vertebrates.

"Arch"-etyping vertebrates.

Cellular details of gill arches in Cambrian fossils reignite a centuries-old debate.

Squamation and scale morphology at the root of jawed vertebrates.

Placoderms, as the earliest branching jawed vertebrates, are crucial to understanding how the characters of crown gnathostomes comprising Chondrichthyes and Osteichthyes evolved from their stem relatives. Despite the growing knowledge of the anatomy and diversity of placoderms over the past decade, the dermal scales of placoderms are predominantly known from isolated material, either morphologically or histologically, resulting in their squamation being poorly understood. Here we provide a comprehensive description of the squamation and scale morphology of a primitive taxon of Antiarcha (a clade at the root of jawed vertebrates), Parayunnanolepis xitunensis, based on the virtual restoration of an articulated specimen by using X-ray computed tomography. Thirteen morphotypes of scales are classified to exhibit how the morphology changes with their position on the body in primitive antiarchs, based on which nine areas of the post-thoracic body are distinguished to show their scale variations in the dorsal, flank, ventral, and caudal lobe regions. In this study, the histological structure of yunnanolepidoid scales is described for the first time based on disarticulated scales from the type locality and horizon of P. xitunensis. The results demonstrate that yunnanolepidoid scales are remarkably different from their dermal plates as well as euantiarch scales in lack of a well-developed middle layer. Together, our study reveals that the high regionalization of squamation and the bipartite histological structure of scales might be plesiomorphic for antiarchs, and jawed vertebrates in general.

Many vertebrates have an outer skeleton covering their body. Some, like crocodiles, have large bony plates of armor, while others, like fish, have small slippery scales. The type, shape, and arrangement of these structures can tell scientists a lot about how different species evolved. Most modern fish are completely covered in scales, but this has not always been the case. Over 400 millions of years ago in the Earth's oceans lived a major group of armored fish called antiarch placoderms which had a combination of bony plates, scales and naked skin. These ancient fish are particularly interesting to scientists because they were one of the first jawed vertebrates to evolve. However, much of what is known about this group has come from isolated materials, which has made it difficult to study the organization and shape of their scales. To overcome this, Wang and Zhu used a specialized x-ray imaging procedure to create a three-dimensional model of one of the best-preserved antiarch placoderm species, Parayunnanolepis xitunensis. The model showed that the fish had thirteen types of scales, found in nine distinct regions on its body. To better understand the structure of these scales, Wang and Zhu looked at the fossils of other extinct jawed fish which where were found in the region where P. xitunensis once lived. The scales of these ancient fish were very different from their bony plates, and from the scales of modern fish. Understanding the skin armor of ancient fish could help to explain how the scales of modern vertebrates evolved. The next step is to look in more detail at the scales of other placoderms to see how they changed over time.

Morphology of Palaeospondylus shows affinity to tetrapod ancestors.

Palaeospondylus gunni, from the Middle Devonian period, is one of the most enigmatic fossil vertebrates, and its phylogenetic position has remained unclear since its discovery in Scotland in 1890 (ref. 1). The fossil's strange set of morphological features has made comparisons with known vertebrate morphotype diversity difficult. Here we use synchrotron radiation X-ray micro-computed tomography to show that Palaeospondylus was a sarcopterygian, and most probably a stem-tetrapod. The skeleton of Palaeospondylus consisted solely of endoskeletal elements in which hypertrophied chondrocyte cell lacunae, osteoids and a small fraction of perichondral bones developed. Despite the complete lack of teeth and dermal bones, the neurocranium of Palaeospondylus resembles those of stem-tetrapod Eusthenopteron2 and Panderichthys3, and phylogenetic analyses place Palaeospondylus in between them. Because the unique features of Palaeospondylus, such as the cartilaginous skeleton and the absence of paired appendages, are present in the larva of crown tetrapods, our study highlights an unanticipated heterochronic evolution at the root of tetrapods.

Embryonic muscle splitting patterns reveal homologies of amniote forelimb muscles.

Limb muscles are remarkably complex and evolutionarily labile. Although their anatomy is of great interest for studies of the evolution of form and function, their homologies among major amniote clades have remained obscure. Studies of adult musculature are inconclusive owing to the highly derived morphology of modern amniote limbs but correspondences become increasingly evident earlier in ontogeny. We followed the embryonic development of forelimb musculature in representatives of six major amniote clades and found, contrary to current consensus, that these early splitting patterns are highly conserved across Amniota. Muscle mass cleavage patterns and topology are highly conserved in reptiles including birds, irrespective of their skeletal modifications: the avian flight apparatus results from slight early topological modifications that are exaggerated during ontogeny. Therian mammals, while conservative in their cleavage patterns, depart drastically from the ancestral amniote musculoskeletal organization in terms of topology. These topological changes occur through extension, translocation and displacement of muscle groups later in development. Overall, the simplicity underlying the apparent complexity of forelimb muscle development allows us to resolve conflicting hypotheses about homology and to trace the history of each individual forelimb muscle throughout the amniote radiations.

Early Origins of Divergent Patterns of Morphological Evolution on the Mammal and Reptile Stem-Lineages.

The origin of amniotes 320 million years ago signaled independence from water in vertebrates and was closely followed by divergences within the mammal and reptile stem lineages (Synapsida and Reptilia). Early members of both groups had highly similar morphologies, being superficially "lizard-like" forms with many plesiomorphies. However, the extent to which they might have exhibited divergent patterns of evolutionary change, with the potential to explain the large biological differences between their living members, is unresolved. We use a new, comprehensive phylogenetic dataset to quantify variation in rates and constraints of morphological evolution among Carboniferous-early Permian amniotes. We find evidence for an early burst of evolutionary rates, resulting in the early origins of morphologically distinctive subgroups that mostly persisted through the Cisuralian. Rates declined substantially through time, especially in reptiles. Early reptile evolution was also more constrained compared with early synapsids, exploring a more limited character state space. Postcranial innovation in particular was important in early synapsids, potentially related to their early origins of large body size. In contrast, early reptiles predominantly varied the temporal region, suggesting disparity in skull and jaw kinematics, and foreshadowing the variability of cranial biomechanics seen in reptiles today. Our results demonstrate that synapsids and reptiles underwent an early divergence of macroevolutionary patterns. This laid the foundation for subsequent evolutionary events and may be critical in understanding the substantial differences between mammals and reptiles today. Potential explanations include an early divergence of developmental processes or of ecological factors, warranting cross-disciplinary investigation. [Amniote; body size; constraint; phylogeny; rate.].

Forebrain Architecture and Development in Cyclostomes, with Reference to the Early Morphology and Evolution of the Vertebrate Head.

The vertebrate head and brain are characterized by highly complex morphological patterns. The forebrain, the most anterior division of the brain, is subdivided into the diencephalon, hypothalamus, and telencephalon from the neuromeric subdivision into prosomeres. Importantly, the telencephalon contains the cerebral cortex, which plays a key role in higher order cognitive functions in humans. To elucidate the evolution of the forebrain regionalization, comparative analyses of the brain development between extant jawed and jawless vertebrates are crucial. Cyclostomes - lampreys and hagfishes - are the only extant jawless vertebrates, and diverged from jawed vertebrates (gnathostomes) over 500 million years ago. Previous developmental studies on the cyclostome brain were conducted mainly in lampreys because hagfish embryos were rarely available. Although still scarce, the recent availability of hagfish embryos has propelled comparative studies of brain development and gene expression. By integrating findings with those of cyclostomes and fossil jawless vertebrates, we can depict the morphology, developmental mechanism, and even the evolutionary path of the brain of the last common ancestor of vertebrates. In this review, we summarize the development of the forebrain in cyclostomes and suggest what evolutionary changes each cyclostome lineage underwent during brain evolution. In addition, together with recent advances in the head morphology in fossil vertebrates revealed by CT scanning technology, we discuss how the evolution of craniofacial morphology and the changes of the developmental mechanism of the forebrain towards crown gnathostomes are causally related.