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

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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the introduced to refer to this assemblage (Hong, 1983), but Daxinganling (Greater Khingan) Mountains, east of the was superseded by “Yanliao Fauna” after it became Hunshandak Sandland, and west of the Liao River.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

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). Shaded area enlarged and shown in detail. 1, Wuhuaxigou Locality; 2, Zhuanshanzi localities (Huang, 2015). Radiometric dates Jiangzhangzi Locality; 3, Chentazi Locality; 4, Daohugou Locality; 5, are available for some of these exposures. The Callovian Guancaishan Locality; 6, Wubaiding (Reshuitang) Locality; 7, Fangshen Jiulongshan Formation is probably exposed at the Locality; 8, Yujiagou Locality; 9, Liaotugou Locality; 10, Yipandaogou Locality; 11, Fanzhangzi Locality; 12, Bawanggou (Mutoudeng) Local- Wubaiding Locality, and samples collected from lavas ity; 13, Zhuanshanzi (Gangou or Nanshimen) Locality; 14, Yaolugou below the salamander-bearing beds at this site yielded a Locality; 15, Daxishan (Daxigou) Locality. 206Pb/238U SHRIMP age of 164±4 Ma (Liu et al., 2006b). However, the same sample also produced a 206Pb/238U localities are within the core geographical range of the TIMS age of 168±4.4 Ma (Liu Yongqing, personal Jehol Biota, and are widely accepted as sites where the communication). A minimum 206Pb/238U age of 157±3 Ma Haifanggou/Jiulongshan and/or Lanqi/Tiaojishan has been reported for the salamander-bearing beds at the formations crop out (Huang, 2015), but details remain Guancaishan Locality (Liu et al., 2006b). It is noteworthy poorly known. that controversy surrounds the stratigraphy at some of The Haifanggou/Jiulongshan Formation has been widely these localities. For example, the beds at the Zhuanshanzi accepted as Middle Jurassic in age based on Locality have been considered to pertain to the Tiaojishan paleontological data (Ren et al., 1995; Shen et al., 2003; Formation by some studies (Xu et al., 2015), but to the Xu et al., 2003). Isotopic radiometric data have supported Haifanggou/Jiulongshan Formation by others (Huang, this age assessment (Chen et al., 1997; Xu et al., 2003; Wu 2015). Similarly, either the Haifanggou/Jiulongshan et al., 2004) and further indicated that the formation is Formation (Huang, 2015) or Lanqi/Tiaojishan Formation partially Bathonian but mainly Callovian. For example, a (Liu et al., 2006b; Gao and Shubin, 2012) may crop out at 40Ar/39Ar age of 166.7+1.0 Ma was obtained from samples the Guancaishan Locality. collected from around the middle of the Haifanggou The Daxishan Locality, another major Yanliao site, Formation of Beipiao, western Liaoning Province (Chang exposes the Oxfordian Lanqi/Tiaojishan Formation. et al., 2013). However, the formation might extend into the Radiometric dates based on samples above, directly from, Upper Jurassic in some areas, such as in the Chengde Basin or below the Anchiornis-bearing bed at the Daxishan of northern Hebei Province (Liu et al., 2006a; Zhang et al., Locality in Linglongta, Jianchang, Liaoning Province 2008b). The Lanqi/Tiaojishan Formation is probably include U-Pb SHRIMP ages of 160.5±0.99 Ma and 2232 Vol. 90 No. 6 ACTA GEOLOGICA SINICA (English Edition) Dec. 2016 http://www.geojournals.cn/dzxben/ch/index.aspx http://mc.manuscriptcentral.com/ags

161.0±1.44 Ma (Liu et al., 2012), of 160.7±3.2 Ma (Peng Locality (Ji and Yuan, 2002; Zhang, 2002). Various ages et al., 2012), and of 160.7±1.7 Ma, 158.9±1.7 Ma, and have been proposed for the Daohugou exposures, ranging 159.5±2.3 Ma (Wang et al., 2013), suggesting that the from Middle Jurassic to Early Cretaceous (Wang et al., Anchiornis-bearing bed is Oxfordian in age. Exposures at 2000; Wang, 2000; Ji and Yuan, 2002; Ren and Yin, 2002; several other sites, including the Yaolugou and Zhang, 2002; Gao and Shubin, 2003; Wang et al., 2005; Xu Bawanggou localities, have been suggested to also and Zhang, 2005; Ji et al., 2006), but their assignment to represent the Lanqi/Tiaojishan Formation (Xu et al., the Bathonian-Callovian Haifanggou/Jiulongshan 2015). Volcanic rocks overlying the Tianyulong-bearing Formation is now widely accepted (Huang, 2015). The bed at the Yaolugou Locality have been radiometrically term ‘Linglongta Biota’ was introduced because the fossil dated to 158.5 ±1.6 Ma (Liu et al., 2012). assemblages from the Oxfordian Lanqi/Tiaojishan The most common elements of the Yanliao fossil Formation are best known from several localities, including assemblages include the plants Yanliaoia, Yimaia and the Daxishan Locality, that lie in Linglongta Town, Schmeissneria, the conchostracans Euestheria Jianchang County, Liaoning Province (Huang, 2015). luanpingensis and Qaidamestheria, the bivalves Ferganoconcha sibirica and Arguniella, the insect 3 Taphonomy, Paleoenvironment, and Yanliaocorixa chinensis, the fish Liaosteus, the Paleoclimate of the Yanliao Biota salamander Chunerpeton, the pterosaur Darwinopterus, and the dinosaur Anchiornis (Huang, 2015). Among these The Yanliao Biota bears a close taphonomic fossils, some are common in the lower Haifanggou/ resemblance to the Jehol Biota. For example, the majority Jiulongshan Formation (e.g., Euestheria luanpingensis, of the fossils recovered from the Yanliao Biota are Ferganoconcha sibirica, and Yanliaocorixa chinensis), complete and articulated; many fine-scale structures are some common in the upper Lanqi/Tiaojishan (e.g., preserved, including wing veins and mouthparts of insects Arguniella, Darwinopterus and Anchiornis), and some in (Zhang and Kluge, 2007; Ren et al., 2010; Wang and both (e.g., Chunerpeton and Yanliaoia). Only a small Zhang, 2010) and tiny branchiopod eggs (Shen and number of species are known in both the Haifanggou/ Huang, 2008); specimens often retain preserved soft Jiulongshan Formation and the Lanqi/Tiaojishan tissue, including setae and hairs in insects (Zhang et al., Formation, and only a few small clades are unique to the 2008c; Ren et al., 2010; Gao et al., 2012), skin Yanliao Biota. Among the vertebrate fossils, the theropod impressions in salamanders (Wang and Evans, 2006; dinosaur groups Scansoriopterygidae and Anchiornithinae Wang et al., 2010d) and lizards (Evans and Wang, 2007; (here defined as the most inclusive clade including Wang et al., 2010d), filamentous structures and Anchiornis but not Archaeopteryx, Gallus, Troodon, membranes in pterosaurs (Ji and Yuan, 2002; Wang et al., Dromaeosaurus, Unenlagia, or Epidexipteryx) are 2002), feathers and membranes in dinosaurs (Zhang et al., probably restricted to the Yanliao Biota, although the 2008a; Hu et al., 2009; Xu et al., 2015), and fur in juvenile holotype of Zhongornis haoae from the Jehol mammaliaforms (Meng et al., 2006; Zhou et al., 2013); Biota has been tentatively interpreted as a Cretaceous stomach contents are preserved in some fossils (Dong et scansoriopterygid (O'Connor and Sullivan, 2014).The al., 2012); and some rarely preserved organisms, such as scansoriopterygid and anchiornithine clades occur in both lichens, are plausibly represented in the biota (Wang et al., the Haifanggou/Jiulongshan Formation and the Lanqi/ 2010a). A few Yanliao fossils are even preserved in ways Tiaojishan Formation. However, scansoriopterygids come that provide behavioral information. For example, some mostly from the Haifanggou/Jiulongshan Formation, male-female pairs of the pachymeridiid Peregrinpa- whereas most anchiornithines (Pedopenna is likely to be chymeridium comitcola are preserved in a mating position, an anchiornithine, based on its pedal morphology) come with the head of the male near the tail of the female and from the Lanqi/Tiaojishan Formation. vice versa (Lu et al., 2011). Because of the compositional differences between the Recurrent volcanism probably explains the apparently fossil assemblages of the older Haifanggou/Jiulongshan sudden deaths, rapid burial, and excellent preservation of Formation and the younger Lanqi/Tiaojishan Formation, the Yanliao organisms (Chang et al., 2009; Sullivan et al., the Yanliao Biota has been divided into the geologically 2014). Other possible causal factors include severe older Daohugou Biota and the younger Linglongta Biota environmental and climatic changes, and even periodic (Huang, 2015). The term ‘Daohugou Biota’, although collapse of the aquatic ecosystem as argued for the Jehol sometimes used in reference to the entirety of the Yanliao Biota (Pan et al., 2011). Possible mechanisms of Biota (e.g. Sullivan et al., 2014), was originally introduced exceptional fossil preservation for the Yanliao Biota to refer to fossils from the exposed strata at the Daohugou include microbial processes (O'Brien et al., 2002; Jiang et Dec. 2016 ACTA GEOLOGICA SINICA (English Edition) Vol. 90 No. 6 2233 http://www.geojournals.cn/dzxben/ch/index.aspx http://mc.manuscriptcentral.com/ags al., 2011) and pyritization (Wang et al., 2008), although Jiang, 2006), bivalves (Wang et al., 1989; Chang and Sun, the mechanism may also have been an unknown one 1997; Jiang, 2006; Duan et al., 2009), conchostracans operating within an unusual fossilization microenvironment (Shen et al., 2003), ostracods (Duan et al., 2009) and produced by clay minerals (Briggs, 2003; Martínez- arachnids (Selden et al., 2008; Huang et al., 2009; Selden Delclòs et al., 2004). and Huang, 2010; Selden et al., 2011). In general, the Yanliao paleoenvironment and climate Among vertebrates (Fig. 5), fish from the Yanliao Biota are poorly known, though some tentative inferences are include acipenseriforms (Lu, 1995; Jin, 1999) and some possible. Study of some insect fossils suggested that the undescribed ptycholepids (Duan et al., 2009). Caudates of Daohugou area might have been a near–shore shallow uncertain systematic position are present (Wang, 2000; lacustrine basin with a warm, humid climate, diverse and Wang and Rose, 2005; Wang and Evans, 2006), in abundant vegetation, and highly aquiferous soil (Tan and addition to a cryptobranchid (Gao and Shubin, 2003), a Ren, 2002; Ren et al., 2010; Selden et al., 2011), though hynobiid (Wang, 2004) and salamandroids (Gao and several different microenvironments may have coexisted Shubin, 2012). Lizards are represented by a species of in the Daohugou paleolakes (Wang et al., 2008). In uncertain systematic position (Evans and Wang, 2012) and general, this inference is consistent with paleobotanical two possible scleroglossans (Evans and Wang, 2007; data from the Haifanggou Formation of western Liaoning Evans and Wang, 2009). Pterosaurs are diverse, and Province, which suggest warm but seasonal and/or moist include anurognathids (Wang et al., 2002; Lü and Hone, conditions (Sun et al., 2008). Some fossil evidence 2012), a rhamphorhynchine (Lü et al., 2012), supports a high paleoaltitude for the Daohugou area, scaphognathines (Czerkas and Ji, 2002; Lü et al., 2010b; including the presence of mountain insects (Ren, 2003; Lü and Bo, 2011; Cheng et al., 2012), wukongopterids Engel and Ren, 2008) and mountain salamanders (Wang, (Lü, 2009; Wang et al., 2009; Lü et al., 2010a; Wang et 2004). Most paleobotanical data suggest a hot, dry climate al., 2010c; Lü et al., 2011), and a possible istiodactylid in the Yanliao Area during the period when the Lanqi (Lü and Fucha, 2010; Martill and Etches, 2012). Formation was being deposited (Zheng, 2013), though Dinosaurs include several maniraptoran theropods some silicified coniferous wood fossils suggest a (Czerkas and Yuan, 2002; Zhang et al., 2002; Xu and subtropical, humid and seasonal climate in western Zhang, 2005; Zhang et al., 2008a; Xu et al., 2009; Xu et Liaoning Province (Jiang et al., 2008). Consequently, al., 2011; Godefroit et al., 2013a; Godefroit et al., 2013b) there seem to have been some temporal and spatial and the heterodontosaurid ornithischian Tianyulong (the variations in the Yanliao paleoenvironment and climate. holotype of Tianyulong confuciusi was originally claimed to be from the Lower Cretaceous Yixian Formation 4 Biodiversity and Paleoecology of the (Zheng et al., 2009), but later studies have considered the Yanliao Biota taxon to belong to the Yanliao Biota (Liu et al., 2012)). The mammaliaform assemblage is relatively diverse, and To date, at least 216 plant species (Jiang et al., 2008; includes the gliding mammal Volaticotherium (Meng et Wang, 2010a; Zheng, 2013), more than 500 insect and al., 2006), triconodonts (Yabe and Shikama, 1938; Zhou et other invertebrate species, and 36 vertebrate species have al., 1991), a yinotherian (Luo et al., 2007), docodonts (Ji been reported from the Yanliao Biota (Xu et al., 2016). et al., 2006; Luo et al., 2015b; Meng et al., 2015), Known plant fossils include spores, leaves, and wood haramiyidans (Zheng et al., 2013; Zhou et al., 2013; Bi et (Zheng, 2013), from lichens, horsetails, lycopsids, filicins, al., 2014), a multituberculate (Yuan et al., 2013), and a bennettitaleans, cycads, ginkgos, czekanowskialeans, eutherian (Luo et al., 2011). conifers, osmundaceans, dipterids, dicksonians, and Yanliao insect herbivores include various cicadomorphs possibly angiosperms (Fig. 3) (Pan, 1983; Wang, 2010a; (Huang and Cai, in press) and the prophalangopsid Wang and Wang, 2010; Zheng, 2013). Most of these Bacharaboilus lii, whose mouthparts appear suitable for fossils are from Liaoning, with 124 species from the feeding on vegetation (Gu et al., 2011). The biota also Haifanggou Formation and 92 species from the Lanqi includes scorpionflies that may have fed on pollination Formation (Zheng, 2013). drops secreted by gymnosperms (Ren, 1998; Ren et al., Insect fossils are represented by at least 500 species 2009; Shih et al., 2011). The inferred mutualistic from 24 major insect groups, among which coleopterans, relationship between the scorpionflies and the dipterans, homopterans, hemipterans and hymenopterans gymnosperms considerably preceded “the similar and are especially high in specific diversity (Fig. 4). Other independent coevolution of nectar-feeding flies, moths, and invertebrate groups (Fig. 4) include branchiopods (Huang beetles on angiosperms” (Ren et al., 2009). Direct evidence and Cai, in press), gastropods (Chang and Sun, 1997; for insect herbivory comes from stomach contents 2234 Vol. 90 No. 6 ACTA GEOLOGICA SINICA (English Edition) Dec. 2016 http://www.geojournals.cn/dzxben/ch/index.aspx http://mc.manuscriptcentral.com/ags

Fig. 3. Selected Yanliao plant fossils. (a), the possible angiosperm Schmeisseria; (b), the bennettitalean Anomozamites sp.; (c), the filicalean Coniopteris sp.; (d), the cycad Nilssonia sp.; (e), the conifer Yanliaoa sp.; (f), the ginkgoalean Yimaia capituliformis. Not to scale. preserved in some permopsocid fossils, which probably cockroach Graciliblatta bella (Liang et al., 2012), and the represent sporangia (Huang et al., 2016). Vertebrate flower bug Pumilanthocoris (Hou et al., 2012), all of herbivores include the haramiyidan mammal Megaconus which fed on other insects. Predatory spiders include the mammaliaformis (Zhou et al., 2013), the multituberculate insect-eating nephilid Nephila jurassica (Selden et al., mammal Rugosodon eurasiaticus (Yuan et al., 2013), and 2011), among others. Vertebrate predators are diverse. the heterodontosaurid dinosaur Tianyulong confuciusi Stomach contents show that the salamander Jeholotriton (Zheng et al., 2009). paradoxus fed on the conchostracan Euestheria Yanliao predatory insects include various odonatans luanpingensis (Dong et al., 2012), whereas the salamander (Huang and Cai, in press) such as the damsel-dragonfly Chunerpeton tianyiensis fed on the corixid insect Zygokaratawia reni (Nel et al., 2008), above-ground Yanliaocorixa chinensis (Dong et al., 2012). Several diurnal geinitziids (Cui et al., 2012), the raphidiomimid Yanliao pterosaurs were probably piscivorous (Sullivan et Dec. 2016 ACTA GEOLOGICA SINICA (English Edition) Vol. 90 No. 6 2235 http://www.geojournals.cn/dzxben/ch/index.aspx http://mc.manuscriptcentral.com/ags

Fig. 4. Selected Yanliao invertebrate fossils. (a), the orthopteran Allaboilus gigantus; (b), the palaeontinid Daohugoucossus shii; (c), the ephemeropteran Jurassonuru samoenus; (d), the sipho- napteran Pseudopulex jurassicus; (e), the corixid Yanliaocorixa chinensis; (f), the dipteran Strashila daohugouensis; (g), the harvestman Mesobunus martensi; (h), the bivalve Ferganoconcha sibirica; (i), the conchostracan Euestheria luanpingensis. Not to scale. al., 2014), but the anurognathid pterosaur Jeholopterus The most common aquatic insects include ningchengensis was probably an aerial insectivore Yanliaocorixa chinensis and ephemeropteran nymphs (Bennett, 2007; Sullivan et al., 2014) and the pterosaur (Huang and Cai, in press). Other aquatic insects include Darwinopterus robustodens has been suggested to have the chresmodid Jurachresmoda gaskelli, which might have fed on hard-carapaced coleopterans (Lü et al., 2011). The been able to skate across the surfaces of bodies of fresh theropod Anchiornis was probably carnivorous and/or water (Zhang et al., 2008c). Aquatic vertebrates include a insectivorous, and most Yanliao mammaliaforms were few fish, the salamanders Jeholotriton and Chunerpeton probably insectivorous (Luo, 2007). (Dong et al., 2012), and the mammaliaform Castorocauda 2236 Vol. 90 No. 6 ACTA GEOLOGICA SINICA (English Edition) Dec. 2016 http://www.geojournals.cn/dzxben/ch/index.aspx http://mc.manuscriptcentral.com/ags

Fig. 5. Selected Yanliao vertebrate fossils. (a), the ptycholepid fish PMOL-AF00743; (b), the cryptobranchid salamander Chunerpeton tianyiensis; (c), a juvenile lizard of uncertain systematic position; (d), the wukongopterid pterosaur Darwinopterus robustodens; (e), the anurognathid pterosaur Jeholopterus ningchengensis; (f), the deinonychosaurian theropod Anchiornis huxleyi; the scansoriopterygid theropods Epidexipteryx hui (g) and Yi qi (h); (i), the haramiyidan mammaliaform Shenshou lui; (j), the basal mammal Volaticotherium antiquus; (k), the eutherian mammal Juramaia sinensis. Not to scale. Dec. 2016 ACTA GEOLOGICA SINICA (English Edition) Vol. 90 No. 6 2237 http://www.geojournals.cn/dzxben/ch/index.aspx http://mc.manuscriptcentral.com/ags lutrasimilis (Ji et al., 2006). numerical dominance of salamanders among aquatic The majority of Yanliao mammaliaforms were probably vertebrates; and 6) numerical dominance of theropods, as terrestrial (Yuan et al., 2013; Zhou et al., 2013). However, opposed to ornithischians, among dinosaurs. The some may have been fossorial (Luo et al., 2007), or extremely unusual characteristics of this ecosystem, as perhaps even subterranean (Luo et al., 2015b). Similarly, preserved in the fossil record, are unquestionably due at several Yanliao mammaliaforms were scansorial (Luo et least in part totaphonomic filtering and collection bias al., 2011) or even arboreal (Meng et al., 2006; Zheng et (Sullivan et al., 2014; Xu et al., 2016). al., 2013; Meng et al., 2015). Arboreal vertebrate species probably also include several theropod dinosaurs (Zhang 5 Significance of the Yanliao Biota et al., 2008a; Hu et al., 2009; Xu et al., 2009) and at least one of the known lizards (Evans and Wang, 2009). The Yanliao fossils have significant implications for our Capable flyers include numerous insects and various understanding of many Mesozoic terrestrial and aquatic pterosaurs (Sullivan et al., 2014); vertebrates that probably groups. The earliest known definitive angiosperms are possessed a more limited capacity for aerial locomotion from the Early Cretaceous (Sun et al., 2002; Friis et al., include the mammaliaform Volaticotherium (Meng et al., 2005), considerably later than the predicted date of 2006), the theropod Anchiornis huxleyi and its close origination for angiosperms based on molecular clock relatives (Xu et al., 2011), and the bizarre studies (Sanderson et al., 2004; Smith et al., 2010). scansoriopterygid Yi qi, whose wings were membranous Several plants from the Yanliao Biota might be helpful in rather than feathered (Xu et al., 2015). understanding the origin of flowering plants (Wang et al., A few studies have identified evidence for specific 2007; Wang, 2010a; b; Wang and Wang, 2010) and/or the paleoecological interactions within the Yanliao Biota. evolution of key angiosperm features (Wang et al., 2010b), Several ectoparasites have been identified, including stem- though this suggestion has received little attention. The fleas which probably fed on dinosaurs, pterosaurs and Yanliao Biota contains the earliest known members of mammals (Gao et al., 2012; Huang et al., 2012). Several many insect groups, including fleas (Gao et al., 2012; species, including alloioscarabaeid insects, have been Huang et al., 2012), webspinners (Huang and Nel, 2009), suggested to have fed on decaying organic materials (Bai and mantophasmatods (Huang et al., 2008b), among et al., 2012), and Yanliao silphids are likely to have fed on others (Huang, 2014). These fossils provide significant carcasses of small vertebrates such as mammals (Cai et al., information on the origins of the groups they represent, 2014). The exceptionally well-preserved stridulatory and on patterns of early character evolution within those structures present in a particular basal katydid suggest that groups. For example, several stem-fleas from the Yanliao low-frequency insect song may have been a normal part of Biota support the hypothesis that fleas, which rank among the auditory background in the Yanliao Biota ecosystem the major ectoparasitic insect groups, are nested within the (Gu et al., 2012). scorpionflies, and these fossils additionally shed new light Yanliao Biota specimens provide evidence for on the evolution of parasite-host relationships in fleas interactions between insects and plants, beyond the (Gao et al., 2012; Huang et al., 2012). Yanliao fossils also presence of the probable insect herbivores mentioned reveal a high degree of evolutionary stasis in many insect above. The bennettitalean Anomozamites villosus groups (Huang et al., 2009; Ren et al., 2010), and in some frequently shows signs of herbivorous insect damage in spider lineages (Selden et al., 2008; Selden and Huang, the distal parts of the leaves, but the proximal parts bear 2010). Yanliao strashilid fossils deserve special note protective ‘hairs’ and generally remain intact (Pott et al., because they have helped to demonstrate that strashilids 2011). Some Yanliao insects appear to have mimicked are highly specialized flies (Diptera) rather than an parts of gymnosperms, just as many modern insects mimic enigmatic group of ectoparasites, and that paedomorphism parts of angiosperms. For example, two lacewings appear is present among endopterygote insects (Huang et al., to have mimicked pinnate cycadophyte leaves (Wang et 2013). In addition, there is evidence for parental care in al., 2010e), and a hangingfly seemingly mimicked a type carrion beetles from the Yanliao Biota (Cai et al., 2014). of ginkgo leaf (Wang et al., 2012). The Yanliao caudate fossils include the earliest known The Yanliao Biota displays several unusual members of three extant clades: cryptobranchids, paleoecological features (Sullivan et al., 2014; Xu et al., hynobiids, and salamandroids (Gao and Shubin, 2003; 2016): 1) a total absence of sauropodomorphs, and other Wang, 2004; Wang and Evans, 2006; Gao and Shubin, vertebrates larger than a few kg; 2) relative rarity of 2012; Jia and Gao, 2016). The Yanliao pterosaur fossils herbivorous vertebrates; 3) absence of definitively aquatic represent an important transitional stage in pterosaur or semi-aquatic reptiles; 4) relative rarity of fish; 5) evolution and demonstrate that modularity can play a key 2238 Vol. 90 No. 6 ACTA GEOLOGICA SINICA (English Edition) Dec. 2016 http://www.geojournals.cn/dzxben/ch/index.aspx http://mc.manuscriptcentral.com/ags role in major evolutionary transformations (Lü et al., consequence of environmental differences (a more closed 2010a). The Yanliao theropods strongly suggest that forest habitat for the Yanliao Biota vs a more open habitat several salient features, such as pennaceous feathers, the for the Shishugou and Shaximiao biotas). However, they four-winged condition, and even aerial locomotion, are may also suggest that some important clades genuinely primitive for Paraves and date back to approximately the originated and underwent their early diversification in the Middle Jurassic (Hu et al., 2009; Xu et al., 2015). Among Yanliao Area, whereas others were genuinely rare or mammaliaforms, Yanliao fossils extend the known record absent in northeast China during Yanliao times. In any of aquatic taxa (Ji et al., 2006), gliding taxa (Meng et al., case, the Yanliao Biota provides a unique window into the 2006) and eutherians (Luo et al., 2011) significantly poorly known terrestrial ecosystems of the Middle-Late backward in time. The holotype of the Yanliao Jurassic transitional period and advances our haramiyidan Megaconus is preserved with typical understanding of the early evolution of many clades. mammalian integument (Zhou et al., 2013), but the Although significant advances have been made in evolutionary significance of this feature is uncertain research on the Yanliao Biota over the last decade, our because of continuing controversy over the phylogenetic understanding of this important biota is still limited. In the position of haramiyidans. Several other Yanliao future, the following areas will be particularly worthy of haramiyidan fossils were reported to confirm that attention and investigation: 1) further analysis of the multituberculates are phylogenetically nested within anatomy, phylogenetic positions and functional Haramiyida and that haramiyidans lie within the morphology of key Yanliao taxa in order to achieve a mammalian crown group, a conclusion that would imply better understanding of important evolutionary events; 2) Triassic haramiyidans represent the oldest known crown- scientific as opposed to commercial excavation of Yanliao mammals (Zheng et al., 2013; Bi et al., 2014). However, a fossils, to make possible relatively unbiased recent study of Triassic haramiyidan fossils suggested reconstructions of the Yanliao ecosystem; 3) establishing a instead that haramiyidans are not closely related to high-precision temporal framework for the Yanliao Biota, multituberculates and fall outside the mammalian crown and accurately determining the geographical and (Luo et al., 2015a), which would imply that crown-group stratigraphic distributions of various Yanliao fossils; 4) mammals probably originated in the Jurassic as reconstruction of the Yanliao paleoenvironment; and 5) traditionally believed. If haramiyidans indeed fall outside analysis of the evolution of the Yaoliao ecosystem in a the crown-group, however, then Megaconus can be taken global context, in order to understand the evolution of the to demonstrate that a full pelage first appeared in non- wider earth-life system around the critical point in time crown mammaliaforms. represented by the Middle-Late Jurassic boundary. The Yanliao Biota shares some common faunal elements (e.g., haramiyid and docodont mammaliaforms, Acknowledgements and transitional pterosaurs) with contemporaneous continental fossil assemblages such as the Shishugou The authors thank Diying Huang, Zhexi Luo and Paul Biota of northwestern China (Clark et al., 2006; Hu et al., Barrett for constructive reviews of the manuscript, and Ge 2007; Martin et al., 2010; He et al., 2013; Andres et al., Sun, Shaolin Zheng, Diying Huang, Yongqing Liu, Keqin 2014) and the Shaximiao Biota of southwestern China Gao, Dongyu Hu, Changfu Zhou, Lijun Zhang, Xin Wang, (Peng et al., 2005), but is compositionally distinct from Haichun Zhang, David Hone, and Zhiyan Zhou for these more typical Jurassic assemblages in several discussion and for providing references and photographs. important respects. Relative to the Shishugou and This project was supported by the National Natural Shaximiao biotas, the Yanliao Biota contains much greater Science Foundation of China (41120124002) and the taxonomic diversity within some clades, such as the National Basic Research Program of China Pterosauria and Mammaliaformes, but much less diversity (2012CB821900). in other clades such as the Testudines, Crocodylomorpha, and Dinosauria; contains more crownward taxa within Manuscript received July 1, 2016 various clades such as the Theropoda and Mammalia; and accepted Aug. 10, 2016 contains many fewer herbivorous vertebrates, partly edited by Liu Lian reflecting a lack of tritylodontids, stegosaurian ornithischians, and sauropods. These significant References differences may partially represent a preservational artifact Andres, B., Clark, J., and Xu, X., 2014. The earliest (lacustrine deposition for the Yanliao Biota vs fluvial pterodactyloid and the origin of the group. Current Biology, 24(9): 1011–1016. deposition for the Shishugou and Shaximiao biotas) or a Dec. 2016 ACTA GEOLOGICA SINICA (English Edition) Vol. 90 No. 6 2239 http://www.geojournals.cn/dzxben/ch/index.aspx http://mc.manuscriptcentral.com/ags

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