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).

Spiny chondrichthyan from the lower Silurian of South China.

Modern representatives of chondrichthyans (cartilaginous fishes) and osteichthyans (bony fishes and tetrapods) have contrasting skeletal anatomies and developmental trajectories1-4 that underscore the distant evolutionary split5-7 of the two clades. Recent work on upper Silurian and Devonian jawed vertebrates7-10 has revealed similar skeletal conditions that blur the conventional distinctions between osteichthyans, chondrichthyans and their jawed gnathostome ancestors. Here we describe the remains (dermal plates, scales and fin spines) of a chondrichthyan, Fanjingshania renovata gen. et sp. nov., from the lower Silurian of China that pre-date the earliest articulated fossils of jawed vertebrates10-12. Fanjingshania possesses dermal shoulder girdle plates and a complement of fin spines that have a striking anatomical similarity to those recorded in a subset of stem chondrichthyans5,7,13 (climatiid 'acanthodians'14). Uniquely among chondrichthyans, however, it demonstrates osteichthyan-like resorptive shedding of scale odontodes (dermal teeth) and an absence of odontogenic tissues in its spines. Our results identify independent acquisition of these conditions in the chondrichthyan stem group, adding Fanjingshania to an increasing number of taxa7,15 nested within conventionally defined acanthodians16. The discovery of Fanjingshania provides the strongest support yet for a proposed7 early Silurian radiation of jawed vertebrates before their widespread appearance5 in the fossil record in the Lower Devonian series.

Nostolepis scale remains (stem Chondrichthyes) from the Lower Devonian of Qujing, Yunnan, China.

Based initially on microfossils, Nostolepis is one of the first known 'acanthodians', which constitute a paraphyletic assemblage of plesiomorphic members of the total group Chondrichthyes. Its wide distribution has potential implications for stratigraphic comparisons worldwide. Six species of Nostolepis have been reported in China, including one species from the Xitun Formation (Lochkovian, Lower Devonian) of Qujing, eastern Yunnan. Acid preparation of rock samples from the Xitun Formation has yielded abundant acanthodian remains. Based on both morphological and histological examinations, here we identify five species of Nostolepis, including two new species. N. qujingensis sp. nov. is characterized by thin scales devoid of the neck anteriorly and the dentine tubules rarely present in the anterior part of the crown. N. digitus sp. nov. is characterized by parallel ridges on anterior and lateral margins of the crown, and the neck constricted and ornamented with pore openings. We extend the duration of N. striata in China from the Pridoli of Silurian (Yulungssu Formation) to the Lower Devonian in Qujing and report the first occurrences of N. amplifica and N. consueta in this region. This study increases the diversity of the Lower Devonian Xitun Fauna and provides a better understanding of the paleogeographic distribution of Nostolepis.

A new Silurian fish close to the common ancestor of modern gnathostomes.

The Silurian Period occupies a pivotal stage in the unfolding of key evolutionary events, including the rise of jawed vertebrates.1-4 However, the understanding of this early diversification is often hampered by the patchy nature of the Silurian fossil record,5 with the articulated specimens of jawed vertebrates only known in isolated localities, most notably Qujing, Yunnan, China.6-9 Here, we report a new Silurian maxillate placoderm, Bianchengichthys micros, from the Ludlow of Chongqing, with a near-complete dermatoskeleton preserved in articulation. Although geographically separated, the new taxon resembles the previously reported Qilinyu in possessing a unique combination of dermatoskeletal characters. However, the dermal bone of the mandible in Bianchengichthys unexpectedly differs from those in both Qilinyu and Entelognathus and displays a broad oral lamina carrying a line of tooth-like denticles, in addition to the marginal toothless flange. The external morphology of the pectoral fin is preserved and reveals an extensively scale-covered lobate part, flanked by a fringe of lepidotrichia-like aligned scales. The phylogenetic analysis reveals that Bianchengichthys is positioned immediately below Entelognathus plus modern gnathostomes. The discovery significantly widens the distribution of Silurian placoderm-grade gnathostomes in South China and provides a range of morphological disparity for the outgroup comparison to the earliest evolution of jaws, dentitions, and pectoral fins in modern gnathostomes. We also demonstrate that the previously reported Silurian placoderms from central Vietnam10 are maxillate placoderms close to Qilinyu, Silurolepis, and Bianchengichthys, corroborating the paleogeographic proximity between the Indochina and South China blocks during the Middle Paleozoic.11.