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Late Jurassic Yanliao Biota: Chronology, Taphonomy, Paleontology and Paleoecology

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Late Jurassic Yanliao Biota: Chronology, Taphonomy, Paleontology and Paleoecology Vol. 90 No. 6 pp.2229–2243 ACTA GEOLOGICA SINICA (English Edition) Dec. 2016 An Updated Review of the Middle-Late Jurassic Yanliao Biota: Chronology, Taphonomy, Paleontology and Paleoecology XU Xing1, *, ZHOU Zhonghe1, Corwin SULLIVAN1, WANG Yuan1 and REN Dong2 1 Key Laboratory of Vertebrate Evolution and Human Origins, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing 100044, China 2 College of Life Sciences, Capital Normal University, Haidian District, Beijing 100048, China Abstract: The northeastern Chinese Yanliao Biota (sometimes called the Daohugou Biota) comprises numerous, frequently spectacular fossils of non-marine organisms, occurring in Middle-Upper Jurassic strata in western Liaoning, northern Hebei, and southeastern Inner Mongolia. The biota lasted for about 10 million years, divided into two phases: the Bathonian-Callovian Daohugou phase (about 168-164 million years ago) and the Oxfordian Linglongta phase (164-159 million years ago). The Yanliao fossils are often taphonomically exceptional (many vertebrate skeletons, for example, are complete and accompanied by preserved integumentary features), and not only are taxonomically diverse but also include the oldest known representatives of many groups of plants, invertebrates, and vertebrates. These fossils have provided significant new information regarding the origins and early evolution of such clades as fleas, birds, and mammals, in addition to the evolution of some major biological structures such as feathers, and have demonstrated the existence of a complex terrestrial ecosystem in northeast China around the time of the Middle-Late Jurassic boundary. Key words: Yanliao Biota, Daohugou phase, Linglongta phase, Middle-Late Jurassic, Yanliao area 1 Introduction 1983, when a rich insect assemblage was discovered from the Middle Jurassic Jiulongshan Formation in the Yanliao The Yanliao Area is a large region of northeast China, Area. The term “Yanliao Insect Fauna” was first situated north of the Yan Mountains, south of the introduced to refer to this assemblage (Hong, 1983), but Daxinganling (Greater Khingan) Mountains, east of the was superseded by “Yanliao Fauna” after it became Hunshandak Sandland, and west of the Liao River. apparent that a wide range of additional non-marine taxa Between roughly the middle of the Jurassic and the middle including conchostracans, bivalves, fish, reptiles, and of the Cretaceous, three major volcanic and sedimentary mammaliaforms coexisted with the insects (Ren et al., cycles in the Yanliao Area successively produced the 1995). Ren et al. (1995) further suggested that the fauna Middle-Upper Jurassic Haifanggou/Jiulongshan and should also be considered to include fossils from the Lanqi/Tiaojishan formations, the Upper Jurassic-Lower Mentougou, Tiaojishan, and Tuchengzi formations Cretaceous Tuchengzi/Houcheng and Zhangjiakou exposed in the Yanliao Area, which were then believed to formations, and the Lower Cretaceous Dabeigou, Yixian be Middle Jurassic. The diagnostic elements of the and Jiufotang formations. The famous and widely Yanliao Fauna were suggested to be the insect distributed Early Cretaceous Jehol fossil assemblages Yanliaocorixa, the fish Liaosteus, and the lizard formed during the third cycle. However, strata deposited ‘Yabeinosaurus’ (Ren et al., 1995), which are known only during the first cycle contain a comparable fossil biota of from the laterally equivalent Jiulongshan and Haifanggou Middle-Late Jurassic age that is known as the Yanliao formations (Hoffstetter, 1964; Zhou et al., 1991; Jin, 1999; Biota (Fig. 1) and is currently emerging as a source of Ren et al., 2010). It should be noted that ‘Yabeinosaurus’ paleontological discoveries that may rival the Jehol Biota here refers to ‘Yabeinosaurus’ youngi, which is known in importance. from the Jiulongshan/Haifanggou Formation of Lingyuan, The Yanliao fossil assemblages were first noticed in Liaoning. A recent study indicated that this species differs * Corresponding author. E-mail: [email protected] markedly in limb proportions from the type species of the © 2016 Geological Society of China 2230 Vol. 90 No. 6 ACTA GEOLOGICA SINICA (English Edition) Dec. 2016 http://www.geojournals.cn/dzxben/ch/index.aspx http://mc.manuscriptcentral.com/ags researchers and is preferred here. Sullivan et al. (2014) also defined the ‘Daohugou Biota’ more narrowly, restricting its scope to sites where the abundant salamander Chunerpeton tianyiensis is known to occur. Over the last decade, numerous significant fossils have been recovered from the Yanliao Biota, including plants that may represent the oldest known angiosperms (Wang, 2010a), the earliest known members of several insect clades (Huang et al., 2008a; Huang and Nel, 2009; Ren et al., 2010; Gao et al., 2012; Huang et al., 2012; Huang, 2014; Huang and Cai, in press), the best-preserved crown salamanders known from the Jurassic (Gao and Shubin, 2003; Gao and Shubin, 2012), transitional pterosaurs (Lü et al., 2010a; Wang et al., 2010c), some of the earliest known feathered dinosaurs such as the paravian Anchiornis and the heterodontosaurid Tianyulong (Xu and Zhang, 2005; Zhang et al., 2008a; Hu et al., 2009; Godefroit et al., 2013a; Godefroit et al., 2013b), the earliest known gliding and aquatic mammaliaforms (Ji et al., 2006; Meng et al., 2006), and the earliest known eutherian mammal (Luo et al., 2011). The Yanliao Biota has thus emerged as a terrestrial fossil assemblage of the Fig. 1. Stratigraphic distribution of the Middle-Upper Juras- greatest importance for understanding Mesozoic sic Yanliao Biota and the Lower Cretaceous Jehol Biota. ecosystems and the evolution of many major groups of Shading indicates the rock formations that contain the Yanliao and Jehol biotas. organisms, including the origins of some large modern clades. The present paper provides an updated review of genus, Yabeinosaurus tenuis from the Lower Cretaceous this important biota, building on previous reviews written Jehol Group, and its attribution to the genus is still from various perspectives (Liu et al., 2010; Zhou et al., equivocal (Evans and Wang, 2012). 2010; Guo et al., 2012; Zheng, 2013; Sullivan et al., 2014; During 2009 and 2010, the term ‘Yanliao Biota’ was Xu et al., 2016). independently introduced by multiple authors, but with different definitions. Sun et al. (2009) defined the Yanliao 2 Temporal and Geographical Distribution of Biota as the entire Jurassic flora and fauna of western the Yanliao Biota Liaoning Province and neighboring areas (Sun et al., 2009; Sun et al., 2011). Chang et al. (2009), by contrast, Like the Early Cretaceous Jehol Biota, the Yanliao used the term ‘Yanliao Biota’ to designate the fossil Biota is a collection of non-marine fossil assemblages content of the Haifanggou Formation of Liaoning mainly preserved in lacustrine deposits rich in volcanic Province and the laterally equivalent Jiulongshan ash. Major known Yanliao localities, many of which are Formation of Hebei Province, a definition concordant with referred to by more than one name in current literature, the original definition of the “Yanliao Insect Fauna”. Zhou include: the Fangshen (Zhou et al., 1991), Yujiagou (Lu, et al. (2010) considered the Yanliao Biota to include all 1995; Jin, 1999), Yipandaogou, Liaotugou (Wang et al., fossils from the Middle Jurassic Haifanggou/Jiulongshan 1989), Daxishan (or Daxigou), Yaolugou, Wubaiding (or Formation and the Upper Jurassic Lanqi/Tiaojishan Reshuitang) (Zhang and Wang, 2004; Wang et al., 2005; Formation. Because the Jiulongshan/Haifanggou Wang and Evans, 2006), Guancaishan (or Guancailing) Formation and the Lanqi/Tiaojishan Formation share (Liu et al., 2006b; Wang et al., 2010d; Gao and Shubin, strong similarities in terms of lithology, taphonomy and 2012; Liu et al., 2012), and Haifenggou localities in fossil content (Zhou et al., 2010; Sullivan et al., 2014), we western Liaoning Province; the Fanzhangzi, Bawanggou follow Zhou et al.’s (2010) definition of the Yanliao Biota. (or Mutoudeng) (Lü, 2009; Lü et al., 2012; Zheng et al., The Yanliao Biota has sometimes been referred to as the 2013), and Zhuanshanzi (or Nanshimen or Gangou) Daohugou Biota (e.g. Sullivan et al. 2014), after the localities in northern Hebei Province; and the Daohugou Locality in southeastern Inner Mongolia, but Wuhuaxigou, Chentaizi, Jiangzhangzi, and Daohugou the term ‘Yanliao Biota’ is gaining currency among localities in southeastern Inner Mongolia (Fig. 2). These Dec. 2016 ACTA GEOLOGICA SINICA (English Edition) Vol. 90 No. 6 2231 http://www.geojournals.cn/dzxben/ch/index.aspx http://mc.manuscriptcentral.com/ags Oxfordian. Two recent 40Ar/39Ar dates for the Lanqi Formation of western Liaoning are 159.5±0.6 Ma, based on samples from the top of the formation (Chang et al., 2013), and 161.8±0.4 Ma based on samples from the basal portion of the formation (Chang et al., 2009; Chang et al., 2013). The strata exposed at the Daohugou Locality belong mainly to the Bathonian-Callovian Haifanggou Formation. Radiometric dates based on samples collected from volcanic rocks overlying the fossil-bearing layers at the Daohugou Locality include two 206Pb/238U SHRIMP ages of 162±2 Ma and 152±2.3 Ma (Liu et al., 2006b), an 40Ar- 39Ar age of 159.8±0.8 Ma (He et al., 2004; Liu et al., 2006b), two 206Pb/238U SHRIMP ages of 166±1.5 Ma and 165±2.4 Ma, an 40Ar-39Ar age of 164±2.5 Ma (Chen et al., 2004), and two 206Pb/238U SHRIMP ages of 164±1.2 Ma and 165±1.2 Ma (Yang and Li, 2004). A 206Pb/238U SHRIMP age of 165.0 ±1.2 Ma was obtained based on samples collected from the fossil-bearing layers (Yang and Li, 2008), suggesting that the Daohugou beds are mostly Callovian. The beds at this locality have been informally termed the ‘Daohugou Formation’ (Zhang, 2002), but this usage has received little acceptance. Other sites at which strata of the Haifanggou/ Jiulongshan Formation are considered to be exposed Fig. 2. Geographic distribution of major Yanliao Biota lo- include the Wuhuaxigou, Chentaizi, Jiangzhangzi, calities (Daohugou Biota localities in red and Linglongta Wubaiding, Guancaishan, Haifenggou, Fanzhangzi, and Biota localities in blue).
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