PUBLISHED VERSION Liping Dong, Zbyněk Roček, Yuan Wang, Marc E H. Jones Anurans from the Lower Cretaceous Jehol Group of western Liaoning, China PLoS One, 2013; 8(7):e69723-1-e69723-17 Copyright: © 2013 Dong et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. PERMISSIONS http://www.plosone.org/static/license http://creativecommons.org/licenses/by/3.0/ 17 November, 2014 http://hdl.handle.net/2440/87016 Anurans from the Lower Cretaceous Jehol Group of Western Liaoning, China Liping Dong1*, Zbyneˇk Rocˇek2, Yuan Wang1, Marc E. H. Jones3 1 Key Laboratory of Vertebrate Evolution and Human Origin of Chinese Academy of Sciences, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, China, 2 Institute of Geology, Department of Palaeobiology, Academy of Sciences of the Czech Republic, Prague, Czech Republic, 3 Research Department of Cell and Developmental Biology, University College London, London, United Kingdom Abstract Background: To date, the Lower Cretaceous Jehol Group of western Liaoning, China has yielded five monotypic genera of anurans, including Liaobatrachus grabaui, Callobatrachus sanyanensis, Mesophryne beipiaoensis, Dalianbatrachus mengi, and Yizhoubatrachus macilentus. However, the validity and distinctness of these taxa have been questioned. Methodology/Principal Finding: We provide a comprehensive analysis of the Jehol frogs that includes a re-examination of the published taxa as well as an examination of a number of new specimens that have been collected over the past 10 years. The results show that the five previously named taxa can be referred to three species of one genus–Liaobatrachus grabaui, L. beipiaoensis comb. nov. and L. macilentus comb. nov.. The diagnosis of Liaobatrachus is revised, and a new diagnosis is provided for each species of this genus. We also establish Liaobatrachus zhaoi sp. nov., on the basis of a dozen well-preserved specimens from a new locality. This taxon is distinguished by a unique combination of characteristics, including relatively long hind limbs, a rounded rather than triangular acetabulum, and a gradually-tapering cultriform process of the parasphenoid. In addition, an unnamed frog from a higher horizon, which has narrow sacral diapophyses and particularly long legs, is different from Liaobatrachus and represents another form of anuran in the Jehol Biota. Conclusion/Significance: Comparisons with other Mesozoic and extant anurans and the primary phylogenetic analysis both suggest that Liaobatrachus is a member of the anuran crown-group and forms a polytomy with leiopelmatids (Ascaphus and Leiopelma) and the remaining crown-group anurans (Lalagobatrachia). Citation: Dong L, Rocˇek Z, Wang Y, Jones MEH (2013) Anurans from the Lower Cretaceous Jehol Group of Western Liaoning, China. PLoS ONE 8(7): e69723. doi:10.1371/journal.pone.0069723 Editor: Andrew A. Farke, Raymond M. Alf Museum of Paleontology, United States of America Received January 7, 2013; Accepted June 6, 2013; Published July 26, 2013 Copyright: ß 2013 Dong et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Funding: This research was supported by grants from the National Natural Science Foundation of China (KA211102, to YW) and the Ministry of Science and Technology of China (KZ129739 and KQ207420, to YW). It was also supported by the Sino-Czech academic exchange program. ZR was a visiting professor at the Institute of Vertebrate Paleontology and Paleoanthropology (IVPP), Chinese Academy of Sciences, Beijing, while completing this research; he was also partly supported by the Institute of Geology, Academy of Sciences of the Czech Republic, Prague (Institutional Project AV0Z30130516). MEHJ thanks the British Council for travel funds associated with Darwin Now that facilitated a visit to the IVPP. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Competing Interests: The authors have declared that no competing interests exist. * E-mail: [email protected] Introduction Mesophryne beipiaoensis Gao and Wang 2001 [10], Dalianbatrachus mengi Gao and Liu 2004 [11], and Yizhoubatrachus macilentus Gao The Jehol Biota represents an Early Cretaceous lacustrine and Chen 2004 [12]. Because each taxon was based on a single terrestrial ecosystem with a high degree of diversity and endemism specimen, some anatomical characters were ambiguous or even [1,2]. It contains a variety of Mesozoic taxa including the early incorrectly interpreted. For example, Liaobatrachus grabaui was bird Confuciusornis and its relatives [3], the feathered tyrannosaur- reported to have no ribs [8], the centra of Mesophryne beipiaoensis oid Yutyrannus huali [4], and the early eutherian Acristatherium were reported to be procoelous [10], and Dalianbatrachus mengi was yanensis [5]. The Jehol Biota is best known from the Jehol Group, considered to have fused frontoparietals [11]. Some revisions have which is exposed in western Liaoning, northern Hebei and been made [13], but many problems and uncertainties remain southeastern Inner Mongolia. The Jehol Group consists of, in unresolved. Based on new fossil discoveries and a re-examination ascending order, the Dabeigou Formation, the Yixian Formation of all reported taxa, this paper provides the first comprehensive and the Jiufotang Formation. Its age has been estimated as study of the Jehol anurans, including revisions of established taxa 129.760.5–122.160.3 Ma [6] or 131–120 Ma [2] by different and erection of a new taxon. recent studies, indicating that the biota lasted about 7–9 Ma (early Barremian to early Aptian). Materials and Methods Amphibians are an important component of the Jehol Biota, with 4 urodele [7] and 5 anuran taxa having been formally Institutional Abbreviations reported [8–12]. The anurans include Liaobatrachus grabaui Ji and Ji CYH, Chaoyang Bird Fossil National Geopark, Chaoyang, 1998 [8], Callobatrachus sanyanensis Wang and Gao 1999 [9], Liaoning; DNM D, Dalian Natural History Museum, Dalian, PLOS ONE | www.plosone.org 1 July 2013 | Volume 8 | Issue 7 | e69723 Jehol Anurans Liaoning; GMV, Geological Museum of China, Beijing; IVPP V, appendicular skeleton and nearly all of the vertebral column Institute of Vertebrate Paleontology and Paleoanthropology, preserved), IVPP V13238 (a partial skeleton with part of the Chinese Academy of Sciences, Beijing; LPM, Liaoning Paleontol- vertebral column, the pelvis and the hind limbs preserved), ogy Museum, Shenyang, Liaoning; MV, Nanjing Institute of IVPP V13379 (a partial skeleton with most of the cranial bones Geology and Palaeontology, Chinese Academy of Sciences, missing), and IVPP V14608 (a partial skeleton consisting of the Nanjing, Jiangsu; ZMNH M, Zhejiang Museum of Natural posterior portion of the vertebral column, the pelvis and the History, Hangzhou, Zhejiang. hind limbs). Specimen IVPP V14979, which represents several frogs preserved on two associated slabs, was also recovered Localities and Materials from this locality. Some of the skeletons are nearly complete, We examined 25 specimens from eight fossil localities in the with only a few bones disarticulated and displaced. The first area around Yixian, Beipiao and Chaoyang in western Liaoning slab bears two adult skeletons (IVPP V14979.1 and IVPP (Fig. 1). The anuran-bearing horizons are the Lujiatun, Jian- V14979.2) and two juvenile skeletons (IVPP V14979.3 and shangou and Dawangzhangzi Beds of the Yixian Formation, and IVPP V14979.4), whereas the second slab (not considered in the Jiufotang Formation (Fig. 2) [14]. There are not anurans the paper as they are the young individuals of IVPP V14979.1) currently known from the Dabeigou Formation. The localities and bears two frogs: IVPP V14979.5 and IVPP V14979.6 [15]. their anuran fossil content are as follows (also see Table 1): 3. Sihetun Locality: 1.5 km southwest of Sihetun Cunminzu (a ‘‘Cunminzu’’ is a subdivision of a village), Chaomidianzi 1. Lujiatun Locality: near Lujiatun Village, 25 km northwest of Village, Shangyuan Town, Beipiao City, Liaoning Province; Beipiao City, Liaoning Province; Lujiatun Bed, Yixian Jianshangou Bed, Yixian Formation; Barremian/Aptian. Formation; Barremian. Specimen IVPP V13245 (a three- Specimens found here include GMV2126 (see Ji and Ji, 1998 dimensionally preserved incomplete skeleton) was excavated [8]), IVPP V11525 (see Wang and Gao, 1999 [9]), MV 77 (a here. nearly complete skeleton with a slightly displaced pelvis), IVPP 2. Qianyangou Locality: about 1.5 km west of Qianyangou V12717 (a nearly complete skeleton with part of the vertebral Village, Shangyuan Town, Beipiao City, Liaoning Province; column displaced and the limbs not preserved), and CYH 004 Lujiatun Bed, Yixian Formation; Barremian. All of the (a nearly complete skeleton). specimens from this locality are three-dimensionally preserved. 4. Heitizigou Locality: 0.5 km east of Libalanggou Village, They include IVPP V13236 (a partial skeleton with part of the Zhangjiying Township, Beipiao City, Liaoning Province; skull and most of the postcranial bones preserved), IVPP Jianshangou Bed, Yixian Formation; Barremian/Aptian. V13239 (a partial skeleton