Postcranial disparity of galeaspids and the evolution of swimming speeds in stem-gnathostomes.

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.

New data on the cranial anatomy of Pterogonaspis (Tridensaspidae, Galeaspida) from the lower Devonian of Yunnan, China and its evolutionary implications.

New material of the tridensaspid Pterogonaspis yuhaii is described from the Xujiachong Formation in Qujing City, Yunnan Province, revealing the morphology of the ventral side of the headshield for the first time. Most significantly, the cranial anatomy of Pterogonaspis provides the first fossil evidence for the position of the esophagus in galeaspids. The esophagus and dorsal aortae display a central placement in the postbranchial region and do not share a common canal as in osteostracans. Moreover, the paired and symmetrically extending dorsal aortae in galeaspids strikingly resemble those of jawed vertebrates, probably representing a plesiomorphic condition for vertebrates because they are also present in amphioxus and hagfishes. In contrast, the unpaired and asymmetrically extending dorsal aorta in osteostracans is similar to that of lampreys, probably resulting from convergent evolution. In addition, our new data also exhibit some characters that galeaspids uniquely share with osteostracans, such as the complete endoskeletal postbranchial wall and the extensive lateral expansion of the headshield with the marginal arteries and veins.

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.

A new genus of sinogaleaspids (Galeaspida, stem-Gnathostomata) from the Silurian Period in Jiangxi, China.

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.

New polybranchiaspiform fishes (Agnatha: Galeaspida) from the Middle Palaeozoic of China and their ecomorphological implications.

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 Evolution of the Zygomatic Bone From Agnatha to Tetrapoda.

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.

Fossil jawless fish from China foreshadows early jawed vertebrate anatomy.

Most living vertebrates are jawed vertebrates (gnathostomes), and the living jawless vertebrates (cyclostomes), hagfishes and lampreys, provide scarce information about the profound reorganization of the vertebrate skull during the evolutionary origin of jaws. The extinct bony jawless vertebrates, or 'ostracoderms', are regarded as precursors of jawed vertebrates and provide insight into this formative episode in vertebrate evolution. Here, using synchrotron radiation X-ray tomography, we describe the cranial anatomy of galeaspids, a 435-370-million-year-old 'ostracoderm' group from China and Vietnam. The paired nasal sacs of galeaspids are located anterolaterally in the braincase, and the hypophyseal duct opens anteriorly towards the oral cavity. These three structures (the paired nasal sacs and the hypophyseal duct) were thus already independent of each other, like in gnathostomes and unlike in cyclostomes and osteostracans (another 'ostracoderm' group), and therefore have the condition that current developmental models regard as prerequisites for the development of jaws. This indicates that the reorganization of vertebrate cranial anatomy was not driven deterministically by the evolutionary origin of jaws but occurred stepwise, ultimately allowing the rostral growth of ectomesenchyme that now characterizes gnathostome head development.