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Late Jurassic Salamandroid from Western Liaoning, China

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Late Jurassic Salamandroid from Western Liaoning, China Correction EVOLUTION, GEOLOGY Correction for “Late Jurassic salamandroid from western Liaon- ing, China,” by Ke-Qin Gao and Neil H. Shubin, which appeared in issue 15, April 10, 2012, of Proc Natl Acad Sci USA (109:5767– 5772; first published March 12, 2012; 10.1073/pnas.1009828109). The authors note that on page 5772, left column, first full para- graph, lines 17–20 to lines 1–2 of the adjacent right column, “This evidence, in keeping with our previous report of an early crypto- branchid from Inner Mongolia (9), rejects the purported timing of 140 Ma for this cladogenetic event (16), but supports a more recent study based on complete mitochondrial genomes for a Middle Ju- rassic divergence of the Salamandroidea from Cryptobranchoidea (8)” should instead appear as “This evidence, in keeping with our previous report of an early cryptobranchid from Inner Mongolia (9), supports molecular calibrations that imply Middle Jurassic di- vergence of the Salamandroidea from Cryptobranchoidea (8, 16).” www.pnas.org/cgi/doi/10.1073/pnas.1210191109 12260 | PNAS | July 24, 2012 | vol. 109 | no. 30 www.pnas.org Downloaded by guest on September 26, 2021 Late Jurassic salamandroid from western Liaoning, SEE COMMENTARY China Ke-Qin Gaoa,1 and Neil H. Shubinb aSchool of Earth and Space Sciences, Peking University, Beijing 100871, China; and bDepartment of Organismal Biology and Anatomy, University of Chicago, Chicago, IL 60637 Edited by David B. Wake, University of California, Berkeley, CA, and approved January 25, 2012 (received for review July 14, 2010) A Jurassic salamander, Beiyanerpeton jianpingensis (gen. et sp. known record of the Salamandroidea, predating a previously nov.), from a recently found site in western Liaoning Province, Early Cretaceous record (Valdotriton, 116–114 Ma) by some 40 China is the earliest known record of Salamandroidea. As a Late Ma (14). In contrast to the salamander fossils from the nearby Jurassic record of the group, it extends the range of the clade by Daohugou site, specimens from Jianping are all preserved with ~40 Ma. The Late Jurassic taxon is neotenic and represented by genuinely fossilized skeletons rather than preserved as natural exceptionally preserved specimens, including fully articulated cra- molds or impressions resulting from postmortem dissolution of nial and postcranial skeletons and bony gill structures close to the the skeletons. The superb preservation of skeletal details sheds cheek region. The fossil beds, consisting of dark-brown volcanic new light on the evolution of key characters, including palatal ash shales of the Upper Jurassic Tiaojishan (Lanqi) Formation structures and marginal teeth. In addition, a phylogenetic study (Oxfordian), underlie trachyandesite rocks that have yielded incorporating this important fossil taxon supports the hypothesis a SHRIMP zircon U-Pb date of 157 ± 3 Ma. The fossiliferous beds that the divergence of the Salamandroidea from their sister clade are substantially older than the Jehol Group, including the Yixian Cryptobranchoidea likely took place before the Late Jurassic on Formation (40Ar/39Ar dates of 122–129 Ma), but slightly younger the Eurasian continent (8). 40 39 than the Middle Jurassic Daohugou horizon ( Ar/ Ar date of Systematic Paleontology 164 ± 4 Ma). The early fossil taxon shares with extant salaman- droids derived character states, including: separated nasals lacking Amphibia Linnaeus, 1758; Lissamphibia Haeckel, 1866; Caudata a midline contact, angular fused to the prearticular in the lower Scopoli, 1777; Urodela Dumèril, 1806; Suborder Salamandroidea Beiyanerpeton jianpingensis jaw, and double-headed ribs on the presacral vertebrae. In con- Noble, 1931; gen. et sp. nov. trast to extant salamandroids, however, the salamander shows Etymology. a discrete and tooth-bearing palatine, and unequivocally nonpedi- Beiyan (ancient place name, meaning Northern Yan cellate and monocuspid marginal teeth in large and presumably State) + herpeton (Greek: creeping animal); Jianping, county mature individuals. The finding provides insights into the evolu- town nearby the type locality. tion of key characters of salamanders, and also provides direct Holotype. Peking University Paleontological Collection (PKUP) evidence supporting the hypothesis that the split between Cryp- V0601, a nearly complete skeleton exposed in ventral view. tobranchoidea and Salamandroidea had taken placed before the Late Jurassic Oxfordian time. In this aspect, both paleontological Referred Specimens. PKUP V0602-0606, and several unnumbered and molecular data now come to agree. specimens; all preserved as articulated cranial and postcranial skeletons. Urodela | phylogeny | fossil record | molecular clock Type Locality and Horizon. Guancaishan, near Jianping, Liaoning alamanders are a diverse group of modern amphibians, with Province, China; Upper Jurassic Tiaojishan (Lanqi) Formation. Shigh ecological diversity and extreme morphological conserva- tism (1–4). Among the 10 extant families, the Hynobiidae and Diagnosis. A Late Jurassic neotenic salamandroid diagnosed by Cryptobranchidae are classified in the suborder Cryptobranchoidea, the following unique combination of character states: Skull rel- and the remaining families (except for the Sirenidae) in another atively wider than long, both anterodorsal and anteromedial suborder, Salamandroidea. Phylogenetic relationships of the Sir- fenestrae developed; sensory canal present on external surface enidae remain contentious, and recent phylogenetic analyses place of premaxilla and maxilla; prefrontal present; lacrimal retained; the family in either basal or crownward positions (5–8). The palatine present as a discrete and dentate element; marginal Cryptobranchoidea have their earliest record from the Middle teeth nonpedicellate, simple peg-like and significantly shorter Jurassic in Asia (9–13), and the Salamandroidea have an Early than vomerine teeth; posterolateral wings of parasphenoid Cretaceous record from Europe (14). Iridotriton was described as grooved for internal carotid artery; operculum separate from a putative salamandroid from the Upper Jurassic Morrison For- stapes; hypobranchial I and II well ossified; basibranchial I and II mation (15), but our evaluation of the available data suggest that it co-ossified; angular lost by fusion with prearticular; presacral is probably a cryptobranchoid (see SI Appendix for details). Within vertebrae 15 in number; vertebral centrum short, strongly con- the phylogenetic framework of the Urodela, the split of the Sala- stricted, with prominent subcentral keel; phalangeal 2-2-3-2/3 in mandroidea from the Cryptobranchiodea is a major cladogenetic manus, and 2-2-3-4-3 in pes. event in the evolution of salamanders, but the timing for this major – event is still in dispute (8, 16 18), largely because of lack of early EVOLUTION fossil evidence. New molecular data (8) point to a substantially Author contributions: K.-Q.G. and N.H.S. designed research; K.-Q.G. and N.H.S. performed earlier age for this split than does the existing fossil record (14) research; K.-Q.G. analyzed data; and K.-Q.G. and N.H.S. wrote the paper. fi before the nding of the fossils reported in this article. The authors declare no conflict of interest. The salamandroid described herein is based on the specimens collected from the Guancaishan site near the county town of This article is a PNAS Direct Submission. Jianping in western Liaoning Province, China (Fig. 1). With a ra- Freely available online through the PNAS open access option. diometric date of 157 ± 3 Ma from the overlaying rocks (12), the See Commentary on page 5557. 1 fossil beds of the Upper Jurassic Tiaojishan (Lanqi) Formation To whom correspondence should be addressed. E-mail: [email protected]. GEOLOGY consist of volcanic ash deposited in a lake during Oxfordian time This article contains supporting information online at www.pnas.org/lookup/suppl/doi:10. (19). Thus, this Late Jurassic fossil taxon documents the earliest 1073/pnas.1009828109/-/DCSupplemental. www.pnas.org/cgi/doi/10.1073/pnas.1009828109 PNAS | April 10, 2012 | vol. 109 | no. 15 | 5767–5772 Fig. 1. Map and geologic section showing the geographic location and stratigraphic position of the fossil site (Tiaojishan Formation) in re- lation to Daohugou (Haifanggou Formation) locality in Inner Mongolia (geologic section modified from ref. 12). The nearby Lower Cre- taceous Xintaimen site is also shown in the map. Solid diamond denoting fossil localities. Description and Comparison. As best shown by the holotype the parietal, and a relatively narrow lateral end above the quad- (PKUP V0601), the salamandroid is a medium-sized salamander rate as part of the cranio-mandibular joint. with a standard snout-vent length of ~100 mm (Fig. 2). Bony The maxilla is short, with a pointed posterior extremity free structures of gill rakers (branchial denticles) are preserved in all from articulation with any bony elements. Similar to the pre- large specimens known for the taxon, giving clear indication of maxilla, the lateral surface of the maxilla also has a deep groove the gill structure for suction feeding as a neotenic feature of the running horizontally towards the external narial opening. This Late Jurassic salamander. The tail is roughly the same length groove may indicate the anterior extension of the infraorbital as the trunk, as shown in PKUP V0602. Caudal vertebrae are sensory canal as a branch of the Cranial Nerve V (22). The dorsoventrally expanded, reflecting a laterally compressed tail, maxilla develops
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