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Paired fins are a major innovation that evolved within the jawed vertebrate lineage after divergence from the living jawless vertebrates. Extinct jawless armored stem-gnathostomes exhibit a diversity of paired bodywall extensions, from skeletal processes to simple flaps. However, osteostracans (sister to jawed vertebrates) are interpreted to manifest the first true paired appendages in a pectoral position, with pelvic appendages evolving later in association with jaws. Here we show, based on articulated remains of Tujiaaspis vividus from the Silurian of China, that galeaspids (sister to osteostracans and jawed vertebrates) possessed three unpaired dorsal fins, an approximately symmetrical hypochordal tail and a pair of continuous, branchial to caudal ventro-lateral fins. The ventro-lateral fins compare to paired fin flaps in other stem-gnathostomes but specifically to the ventro-lateral ridges of cephalaspid osteostracans which also possess differentiated pectoral fins. The ventro-lateral fins as compatible with aspects of the fin-fold hypothesis for the origin of vertebrate paired appendages. Galeaspids manifest a precursor condition to osteostracans and jawed vertebrates where paired fins arose initially as continuous pectoral-pelvic lateral fins that our Computed Fluid Dynamic experiments demonstrate passively generate lift. Only later in the stem-lineage to osteostracans and jawed vertebrates did pectoral fins differentiate anteriorly. This was followed by restriction of the remaining field of fin competence to a pelvic position, facilitating active propulsion and steering.

Galeaspids are extinct jawless relatives of living jawed vertebrates whose contribution to understanding the evolutionary assembly of the gnathostome bodyplan has been limited by absence of postcranial remains. Here, we describe Foxaspis novemura gen. et sp. nov., based on complete articulated remains from a newly discovered Konservat-Lagerstätte in the Early Devonian (Pragian, ∼410 Ma) of Guangxi, South China. F. novemura had a broad, circular dorso-ventrally compressed headshield, slender trunk and strongly asymmetrical hypochordal tail fin comprised of nine ray-like scale-covered digitations. This tail morphology contrasts with the symmetrical hypochordal tail fin of Tujiaaspis vividus, evidencing disparity in galeaspid postcranial anatomy. Analysis of swimming speed reveals galeaspids as moderately fast swimmers, capable of achieving greater cruising swimming speeds than their more derived jawless and jawed relatives. Our analyses reject the hypothesis of a driven trend towards increasingly active food acquisition which has been invoked to characterize early vertebrate evolution.

A new genus and species of galeaspid, Falxcornus liui gen. et sp. nov., is described from the Xishancun Formation (Lochkovian, Lower Devonian) in Qujing, Yunnan, China. The new taxon displays a sui...

Galeaspida is an endemic clade of jawless stem-gnathostomes (‘ostracoderms’), known exclusively from the Silurian and Devonian of China and Vietnam. Among galeaspids, the family Sanqiaspidae is of particular interest for providing the first information on the galeaspid trunk and caudal fin; however, the cranial morphology of sanqiaspids remains poorly known. Here we report the first occurrence of Sanqiaspis in the Xujiachong Formation of Qujing, East Yunnan, and describe new material of Sanqiaspis from two other sites in South China. The new data corroborate the presence of the two median transverse canals in Sanqiaspis as a possible synapomorphy of the Sanqiaspidae. As a key member of the Xujiachong Assemblage, Sanqiaspis is known from the Xujiachong, Guanshanpo, Posongchong, and Si Ka formations. Stratigraphic correlation of eight Early Devonian sections in South China and North Vietnam indicates that the Sanqiaspis-bearing layers can be correlated with the transitional beds between the Lianhuashan and Nahkaoling formations in Liujing, Guangxi. Therefore, the middle-upper part of the Si Ka Formation in northern Vietnam and the upper part of the Lianhuashan Formation in Guangxi should be dated as Pragian, rather than Lochkovian. The fish assemblage from the Pragian of the South China Block documents the diversification of galeaspids, notably Huananaspidiformes, and exhibits much the same endemic pattern as that of early vascular plants and brachiopods. This endemism might be linked with the semi-closed epicontinental nature of the South China Sea during the Early Devonian, which formed a biodiversity hotspot as in the modern Red Sea.

A new genus and species of the Galeaspida Qushiaspis elaia gen. et sp. nov. is described from the Lower Devonian Xujiachong Formation of Qujing, Yunnan Province, southwestern China. The new genus d...

Micro-CT provides a means of nondestructively investigating the internal structure of organisms with high spatial resolution and it has been applied to address a number of palaeontological problems that would be undesirable by destructive means. This approach has been applied successfully to characterize the cranial anatomy of Shuyu, a 428 million-year-old galeaspid (jawless stem-gnathostome) from the Silurian of Changxing, Zhejiang Province, China. Here, we use the synchrotron X-ray tomographic microscopy (SRXTM) to further describe the circulatory system of the head of Shuyu. Our results indicate that the circulatory system of galeaspids exhibits a mosaic of primitive vertebrate and derived gnathostome characters, including a number of derived gnathostome characters that are absent from osteostracans — the group conventionally interpreted as the sister lineage of jawed vertebrates. Our study provides a rich source of information that can be used to infer and reconstruct the early evolutionary history of the vertebrate cardiovascular system.

Galeaspids are an endemic clade of jawless stem-gnathostomes known as ostracoderms. Their existence illuminates how specific characteristics developed in jawed vertebrates. Sinogaleaspids are of particular interest among the galeaspids but their monophyly is controversial because little is known about Sinogaleaspis xikengensis. Newly discovered sinogaleaspids from the Lower Silurian of Jiangxi, China provide a wealth of data and diagnostic features used to establish the new genus, Rumporostralis gen. nov., for Sinogaleaspis xikengensis. A morphological study showed that the sensory canal system of sinogaleaspids had mosaic features similar to those of three known galeaspids. There are 3–8 pairs of transverse canals in the Sinogaleaspidae, which suggests that the sensory canal system of galeaspid probably had a grid distribution with transverse canals arranged throughout the cephalic division. Phylogenetic analysis of Galeaspida supports the monophyly of the Sinogaleaspidae, consisting of Sinogaleaspis, Rumporostralis, and Anjiaspis. However, Shuyu and Meishanaspis form another monophyletic group, Shuyuidae fam. nov., which is outside all other eugaleaspidiforms. We propose a cladistically-based classification of Galeaspida based on our analysis.

Establishing the homology of the zygomatic or jugal bone and tracing its origin and early evolution represents a complex issue because of large morphological gaps between various groups of vertebrates. Using recent paleontological findings, we discuss the deep homology of the zygomatic or jugal bone in stem gnathostomes (placoderms) and examine its homology and modifications in crown gnathostomes (acanthodians, chondrichthyans and osteichthyans). The discovery of the placoderm Entelognathus from the Silurian of China (∼423 million years ago) established that the large dermal plates in placoderms and osteichthyans are homologous. In Entelognathus, the jugal was joined by a new set of bones (premaxilla, maxilla, and lachrymal), marking the first appearance of the typical vertebrate face found in tetrapods including humans. In non-Entelognathus placoderms, the jugal (homologized with the suborbital plate) occupied most of the cheek region and covered the palatoquadrate laterally. In antiarch placoderms (the most basal jawed vertebrates), the jugal (represented by the ventrally positioned mental plate) functioned as part of the upper jaw. In osteichthyans, the preopercular arose as a novel bone and separated the jugal from the opercular in piscine osteichthyans. A single bone in basal osteichthyans, the preopercular may have divided into two or three elements (the preopercular, the squamosal and/or the quadratojugal) in several later osteichthyan groups. Subsequent modifications of the jugal in the fish-tetrapod transition (its enlargement leading to its contact with the quadratojugal and the separation of the squamosal from the maxilla) brought the vertebrate face to the typical model we see in living tetrapods. Anat Rec, 300:16-29, 2017. © 2016 Wiley Periodicals, Inc.

Establishing the homology of the zygomatic or jugal bone and tracing its origin and early evolution represents a complex issue because of large morphological gaps between various groups of vertebrates. Using recent paleontological findings, we discuss the deep homology of the zygomatic or jugal bone in stem gnathostomes (placoderms) and examine its homology and modifications in crown gnathostomes (acanthodians, chondrichthyans and osteichthyans). The discovery of the placoderm Entelognathus from the Silurian of China (∼423 million years ago) established that the large dermal plates in placoderms and osteichthyans are homologous. In Entelognathus, the jugal was joined by a new set of bones (premaxilla, maxilla, and lachrymal), marking the first appearance of the typical vertebrate face found in tetrapods including humans. In non-Entelognathus placoderms, the jugal (homologized with the suborbital plate) occupied most of the cheek region and covered the palatoquadrate laterally. In antiarch placoderms (the most basal jawed vertebrates), the jugal (represented by the ventrally positioned mental plate) functioned as part of the upper jaw. In osteichthyans, the preopercular arose as a novel bone and separated the jugal from the opercular in piscine osteichthyans. A single bone in basal osteichthyans, the preopercular may have divided into two or three elements (the preopercular, the squamosal and/or the quadratojugal) in several later osteichthyan groups. Subsequent modifications of the jugal in the fish-tetrapod transition (its enlargement leading to its contact with the quadratojugal and the separation of the squamosal from the maxilla) brought the vertebrate face to the typical model we see in living tetrapods. Anat Rec, 300:16-29, 2017. © 2016 Wiley Periodicals, Inc.

The 438-370-million-year-old galeaspids, diversified armoured jawless vertebrates ('ostracoderms') from China and northern Vietnam, were assumed to have a benthic feeding habit in a coastal, marine environment. Here, we describe two new genera of galeaspid fishes, Platylomaspis gen. nov. and Nanningaspis gen. nov. from the Middle Palaeozoic of China. The two new forms are characterized by a rostral process and strikingly broad ventral rim, and clustered with Gumuaspis to form a new family, Gumuaspidae, which represents the most primitive clade of Polybranchiaspiformes. They extend the earliest occurrence of Polybranchiaspiformes backward about 19 million years, and expand its geographical distribution from southern China and northern Vietnam to the Tarim Basin, northwestern China. The new taxa exhibit many morphological convergences with modern rays, and might specify a new kind of lifestyle of galeaspids, the half burrowing habit. Probably benefiting from the new lifestyle, the Gumuaspidae has become the longest lasting galeaspid family. The new findings demonstrated that the demersal galeaspids had developed three different kinds of lifestyles: semi-infaunal benthic (half buried), epibenthic, and suprabenthic (nektonic) habits to accommodate to differentiated ecological niches, and reached the peak of their diversity by the Pragian of the Early Devonian.

The origin of the vertebrate jaw has been reviewed based on the molecular, developmental and paleontological evidences. Advances in developmental genetics have accumulated to propose the heterotopy theory of jaw evolution, i.e. the jaw evolved as a novelty through a heterotopic shift of mesenchyme-epithelial interaction. According to this theory, the disassociation of the nasohypophyseal complex is a fundamental prerequisite for the origin of the jaw, since the median position of the nasohypophyseal placode in cyclostome head development precludes the forward growth of the neural-crest-derived craniofacial ectomesenchyme. The potential impacts of this disassociation on the origin of the diplorhiny are also discussed from the molecular perspectives. Thus far, our study on the cranial anatomy of galeaspids, a 435-370-million-year-old ‘ostracoderm’ group from China and northern Vietnam, has provided the earliest fossil evidence for the disassociation of nasohypophyseal complex in vertebrate phylogeny. Using Synchrotron Radiation X-ray Tomography, we further show some derivative structures of the trabeculae (e.g. orbitonasal lamina, ethmoid plate) in jawless galeaspids, which provide new insights into the reorganization of the vertebrate head before the evolutionary origin of the jaw. These anatomical observations based on new techniques highlight the possibility that galeaspids are, in many respects, a better proxy than osteostracans for reconstructing the pre-gnathostome condition of the rostral part of the braincase. The cranial anatomy of galeaspids reveals a number of derived characters uniquely shared with gnathostomes. This raises the potential possibility that galeaspids might be the closest jawless relatives of jawed vertebrates. Our study provides an intriguing example of intersection between developmental biology-based model and fossil evidence.