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A new palaeontinid (Insecta, Hemiptera, Cicadomorpha) from the Upper Jurassic Tiaojishan Formation of northeastern China and its biogeographic significance

Published online by Cambridge University Press:  11 December 2019

Yanzhe Fu Open the ORCID record for Yanzhe Fu [Opens in a new window] ,
Chenyang Cai  and
Diying Huang
Yanzhe Fu
Affiliation:
State Key Laboratory of Palaeobiology and Stratigraphy, Center for Excellence in Life and Paleoenvironment, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, China University of Chinese Academy of Science, Beijing 100049, China
Chenyang Cai
Affiliation:
Key Laboratory of Economic Stratigraphy and Palaeogeography, Center for Excellence in Life and Paleoenvironment, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, China
Diying Huang*
Affiliation:
State Key Laboratory of Palaeobiology and Stratigraphy, Center for Excellence in Life and Paleoenvironment, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, China
*
*Corresponding author
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Abstract

Cicadomorpha guancaishanensis new species, of the extinct family Palaeontinidae, is described from the Upper Jurassic Tiaojishan Formation at Guancaishan, Jianping County, Western Liaoning, northeastern China. It represents the first record of Palaeontinidae from the Tiaojishan Formation and highlights the palaeodiversity of the Late Jurassic palaeontinids. Cicadomorpha Martynov, 1926 previously included three species with extremely limited morphological differences from the Karabastau Formation of Kazakhstan, the Glushkovo Formation, and the Ukurei Formation of Chita, Transbaikalia, and Russia, indicating a correlated stratigraphic relationship. This discovery provides new evidence that closure of the Okhotsk Ocean began in the Middle Jurassic. Thus, there is no distinct geographical barrier between the southern and northern areas of Mongolia-Okhotsk Ocean in the Late Jurassic.

UUID: http://zoobank.org/64531de4-070c-43d4-ae01-106d4338027b.

Information

Type
Articles
Information
Journal of Paleontology , Volume 94 , Issue 3 , May 2020 , pp. 513 - 520
Copyright
Copyright © 2019, The Paleontological Society

Introduction

Palaeontinoidea (Insecta: Hemiptera: Cicadomorpha) is an extinct superfamily of hemipteran insects, comprising three families (i.e., Dunstaniidae Tillyard, Reference Tillyard1916, Mesogereonidae Tillyard, Reference Tillyard1921, and Palaeontinidae Handlirsch, Reference Handlirsch1906). Palaeontinidae are Mesozoic cicadomorphan hoppers, superficially resembling huge moths, which existed from the Triassic (Carnian) to the Cretaceous (Aptian), distributed in Europe, Central Asia, South Africa, China, South Korea, and Australia, especially in the Northern Hemisphere (Riek, Reference Riek1976; Shcherbakov and Popov, Reference Shcherbakov, Popov, Rasnitsyn and Quicke2002; Menon et al., Reference Menon, Heads and Martill2005; Wang et al., Reference Wang, Ren and Shih2007; Nam et al., Reference Nam, Wang, Ren, Kim and Szwedo2017; Szwedo, Reference Szwedo2018; Chen et al., Reference Chen, Beattie, Wang, Jiang, Zheng and Zhang2019; Li et al., Reference Li, Chen and Wang2019). The striking features of Palaeontinidae are the presence of a large body covered with numerous hairs; a small head with an extremely long rostrum reaching the end of the abdomen, which is suitable for sucking juice from a macrophyte; a wide pronotum and mesonotum; broad wings armed with dark bands to avoid predators; and a fully developed nodal line on the forewing, similar to that of tettigarctids (Carpenter, Reference Carpenter and Kaesler1992; Chen et al., Reference Chen, Zhang, Wang, Zheng and Wang2016; Nam et al., Reference Nam, Wang, Ren, Kim and Szwedo2017). Palaeontinids flourished during the Jurassic and the Early Cretaceous and became extinct in the middle Cretaceous, likely owing to the food crisis and emergence of various insectivorous mammals and birds in the Early Cretaceous (Menon et al., Reference Menon, Heads and Martill2005; Wang et al., Reference Wang, Zhang, Fang, Wang and Ji2008).

The first unequivocal palaeontinid in China was reported by Hong (Reference Hong1982) on the basis of an isolated hindwing. Palaeontinidae have a rich fossil record in northern China, including 57 species within 22 genera from the Early Jurassic to the Early Cretaceous. They have been reported from the Lower Jurassic Badaowan Formation of Karamai, Xinjiang, northwestern China; the Middle–Upper Jurassic Longmen Formation (= Jiulongshan Formation) of North Hebei, the Middle–Upper Jurassic Haifanggou Formation of West Liaoning, and the Daohugou beds (Haifanggou Formation), Inner Mongolia; and the Lower Cretaceous Chijinqiao Formation of Gansu and the Yixian Formation of Hebei, Liaoning, and Inner Mongolia (Fig. 1) (Wang and Ren, Reference Wang and Ren2009; B. Wang et al., Reference Wang, Zhang and Szwedo2009; Chen et al., Reference Chen, Zhang, Wang, Zheng and Wang2016). Among them, 17 genera and 42 species were collected from the Middle–Upper Jurassic of northeastern China, belonging to the ‘early assemblage’ of the Yanliao biota. Here we report a new species of Palaeontinidae from the Upper Jurassic Tiaojishan Formation of Guancaishan—Cicadomorpha guancaishanensis new species—representing the first fossil record of palaeontinids in the ‘late assemblage’ of the Yanliao biota and the third insect species described from the Tiaojishan Formation.

Figure 1.

Distribution of Mesozoic palaeontinids in China.

Geological settings

The new fossil palaeontinid preserved in black shale was collected by one of us (CC) in 2013 from the Upper Jurassic Tiaojishan Formation, at the upper layer of the lacustrine deposit at Guancaishan, near Muyingzi Village, Shahai Township, Jianping County, Chaoyang City, Liaoning Province, northeastern China (Fig. 2). This deposit can be generally assigned within the Linglongta beds, which are the lacustrine interbed clasolites of the large scale andesite of the Tiaojishan Formation mainly occurring in western Liaoning, northern Hebei, and western Beijing, including several exceptional fossil localities such as Daxishan, Nanshimen, Bawanggou, and Fanzhangzi (Huang, Reference Huang2015, Reference Huang2016; Huang et al., Reference Huang, Cai, Fu and Su2018a). The Linglongta biota contains numerous well-preserved fossils, e.g., insects, bivalves, conchostracans, ostracods, gastropods, charophytes, pollen, and spores (Duan et al., Reference Duan, Zhen, Hu, Zhang and Wang2009; Huang, Reference Huang2016; Huang et al., Reference Huang, Cai, Fu and Su2018a), as well as diverse vertebrate fossils including fishes (Huang, Reference Huang2015), salamanders (Gao and Shubin, Reference Gao and Shubin2012; Wang et al., Reference Wang, Dong and Evans2015; Jia and Gao, Reference Jia and Gao2016, Reference Jia and Gao2019), pterosaurs and their eggs (e.g., Lü, Reference Lü2009; Lü et al., Reference Lü, Unwin, Jin, Liu and Ji2010, Reference Lü, Unwin, Deeming, Jin, Liu and Ji2011; X. Wang et al., Reference Wang, Kellner, Jiang and Meng2009; Wang et al., Reference Wang, Kellner, Jiang, Cheng, Meng and Rodrigues2010; Cheng et al., Reference Cheng, Wang, Jiang and Kellner2012; Yang et al., Reference Yang, Jiang, McNamara, Kearns, Pittman, Kaye, Orr, Xu and Benton2019), feathered dinosaurs (e.g., Hu et al., Reference Hu, Lianhai, Zhang and Xu2009; Xu et al., Reference Xu, You, Du and Han2011; Godefroit et al., Reference Godefroit, Demuynck, Dyke, Hu, Escuillié and Claey2013a, Reference Godefroit, Cau, Hu, Escuillié, Wu and Dykeb), and mammals (e.g., Meng et al., Reference Meng, Hu, Wang, Wang and Li2006; Luo et al., Reference Luo, Yuan, Meng and Ji2011; Yuan et al., Reference Yuan, Ji, Meng, Tabrum and Luo2013; Zhou et al., Reference Zhou, Wu, Martin and Luo2013; Bi et al., Reference Bi, Wang, Guan, Sheng and Meng2014). However, the fossil insects from the Linglongta biota are poorly known. Only two species (i.e., Omma daxishanense Cai and Huang, Reference Cai and Huang2016 and Sinothemis difficilis Huang, Cai, and Nel, Reference Huang, Cai and Nel2018b) have been formally described from Daxishan and Guancaishan, respectively. Huang et al. (Reference Huang, Cai, Fu and Su2018a) illustrated abundant fossil insects with at least 16 orders from various localities of the Linglongta beds including Daxishan, Guancaishan (Muyingzi Village), and Zhuanshanzi (Nanshimen Village).

Figure 2.

(1) Stratigraphic column for the Tiaojishan Formation at Guancaishan, Jianping County, Liaoning Province, northeastern China; (2) map showing the Guancaishan locality; (3) the outcrop of the fossil, red arrow indicating the black shales from which the fossil was collected, and the white arrow indicating the volcanic ash layer slightly above the black shale.

Until now, the successive deposit of Yanliao biota contains both Daohugou and Linglongta beds only known from Guancaishan and Nanshan localities (Fig. 2). The Guancaishan deposits include the lower tuffaceous shale layers that have yielded very rich insects, bivalves, plants, and abundant salamanders (Gao and Shubin, Reference Gao and Shubin2012; Wang et al., Reference Wang, Dong and Evans2015), and the higher black insect layers (Fig. 2; Huang et al., Reference Huang, Cai, Fu and Su2018a), which are separated by a layer of intrusive rocks. The volcanic ash layer is located ~1 m above the fossil insect bedding plane dated as 157.3 ± 1.5 Ma (SIMS U-Pb zircon age), indicating the age of the fossil layer close to the boundary of Oxfordian and Kimmeridgian (i.e., 157.3 ± 1.0 Ma; Huang, Reference Huang2016).

Material and methods

The fossil specimen (NIGP 168220) was carefully prepared using a sharp knife under a stereo microscope. A photograph of general habitus was taken using a Canon EOS 5D Mark II camera with a Canon 100 mm macrolens. Photomicrographs were made using a Zeiss Discovery V16 stereoscope, in some cases moistened with 70% ethanol. The raw digital images were processed with Helicon Focus 6 software, and all images were modified and arranged in Adobe Photoshop CS6. Line drawings were prepared using a binocular Olympus SZX7 microscope and were modified with CorelDRAW X7 graphic software.

Wing venation terminology and cell nomenclature follow B. Wang et al. (Reference Wang, Zhang and Szwedo2009), Nel et al. (Reference Nel, Prokop, Nel, Grandcolas, Huang, Roques, Guilbert, Dostál and Szwedo2012), and Bourgoin et al. (Reference Bourgoin, Wang, Asche, Hoch, Soulier-Perkins, Stroinski, Yap and Szwedo2015). All measurements are given in mm.

Repository and institutional abbreviation

The specimen is deposited in the Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences (NIGP).

Systematic paleontology

Suborder Cicadomorpha Evans, Reference Evans1946
Superfamily Palaeontinoidea Handlirsch, Reference Handlirsch1906
Family Palaeontinidae Handlirsch, Reference Handlirsch1906
Genus Cicadomorpha Martynov, Reference Martynov1926

Type species

Cicadomorpha punctualata Martynov, Reference Martynov1926.

Other species

Cicadomorpha milva Shcherbakov, Reference Shcherbakov1988 and C. parula Shcherbakov, Reference Shcherbakov1988.

Cicadomorpha guancaishanensis new species
Figures 3–4

Holotype

NIGP 168220 (a, b), including part and counterpart, a nearly complete adult male with wings extended; stored at the Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing, China.

Diagnosis

Wings with distinct color bands and suborbicular spots; tegmen with Pc + CP emitting seven branches intersecting costal area; MP bifurcate, situated slightly after bifurcation point of CuA; MP1+2 as long as MP3+4; cross vein mp-cua with short, wedge-shaped vein; mp4-cua curved mediad; Pcu fused with A1 distinctly basad of bifurcation of MP; A1 nearly straight; hind wings with cross vein ir much longer than ScP + RA1.

Occurrence

Upper Jurassic Tiaojishan Formation at Guancaishan near Muyingzi Village, Jianping County, Chaoyang City, Liaoning Province, China; early Late Jurassic, near boundary of Oxfordian and Kimmeridgian.

Description

Body (Figs. 3.1, 4.1) ~32.1 mm length and 14.8 mm in width, with a pair of fore wings and hind wings, lacking only the legs.

Figure 3.

Cicadomorpha guancaishanensis n. sp. from the Upper Jurassic Tiaojishan Formation at Guancaishan: (1) holotype (NIGP 168220), male, showing general habitus moistened with 70% ethanol; (2–5) enlargements of (1), showing details of: (2) head and pronotum, moistened with 70% ethanol; (3) right tegmen; (4) discal cell and cross vein mp-cua; and (5) genitals. ce = compound eye; gs = genital style; pr = pronotum; ro = rostrum. Scale bars = 1 mm (4, 5); 2 mm (2, 3); 6 mm (1).

Figure 4.

Line drawings of Cicadomorpha guancaishanensis n. sp., holotype (NIGP 168220): (1) general habitus; (2) forewing; and (3) hind wing. A = anal vein; CP = costa posterior; CuA = cubitus anterior; CuP = cubitus posterior; MP = median posterior; Pc = precosta; Pcu = postcubitus; RA = radius anterior; RP = radius posterior; ScP = subcosta posterior. Scale bars = 4 mm (2, 3); 6 mm (1).

Head small (Fig. 3.2), length ~3.3 mm, width ~5.9 mm, rounded apically; compound eyes large, ovoid, positioned laterally; postclypeus swollen, 1.13 mm long, 0.55 mm wide; anteclypeus less swollen, nearly half length of postclypeus; clypellus triangularly elongated; rostrum long, as preserved ~3.1 mm, extending beyond thorax. Pronotum rounded (Fig. 3.2), nearly 2.3 times wider than head; anterior edge concave medially; anterolateral angles of pronotum notably expanded. Thorax partly obscured by extended mesonotum.

Tegmen subtriangular (Figs. 3.3, 4.2), 39.8 mm long, 12.6 mm wide, with length/width ratio ~3.2; tegmen armed with colored bands from base to end, dotted with suborbicular spots of varying sizes; costal margin slightly arched at base, then nearly straight; anteroapical angle rounded; posteroapical angle 148°; posteroapical margin slightly curved; postclaval margin almost straight, with position of bend connected with nodal line; basal cell wide; vein Pc + CP slightly curved, ending at level of nodal indentation, emitting seven branches intersecting costal area, these branches dispersed in increasing distance from each other; stem ScP separated from common stem R + MP at base, with terminal ScP obscure; stem R slightly curved, branching into RA and RP distinctly basad of nodal line, reaching one-third of tegmen length; RA and RP simple, with cross vein ir absent; RP connected with MP1; cross vein mp-cua segmenting discal cell, distinctly curved with short, wedge-shaped vein (Fig. 3.4), tapering toward end of tegmen; stem MP with four branches, branching into MP1+2 and MP3+4 slightly basad of nodal line and slightly after bifurcation of CuA; MP1+2 as long as MP3+4; cross vein im absent; mp4-cua curved mediad; CuA dispersed from stem CuA + CuP reaching 0.14 of tegmen length, branching into CuA1 and CuA2 reaching 0.38 of tegmen length; basal of CuA2 along posterior end of nodal line; CuP curved at base, then nearly straight; Pcu fused with A1 distinctly basad of bifurcation of stem MP, reaching one-quarter of tegmen length; A1 nearly straight.

Hind wings oval (Figs. 3.3, 4.3), ~25.4 mm long, with length/width ratio 1.7, dotted with colored bands and irregular suborbicular spots; costal margin distinctly arched; costal area long, almost half length of wing, with length/width ratio ~5.2; stem ScP + RA branching into ScP + RA1 and RA2, reaching 0.44 of wing length, connecting with stem RP + MP by cross vein ir; RA2 short; cross vein ir long, ~2.7 times longer than ScP + RA1; RP fused with MP1 for considerable distance before separating; MP with four branches, with MP1 and MP2 almost straight, MP3+4 branching MP3 and MP3 at same level of bifurcation of ScP + RA; cross vein mp-cua short; stem CuA branching into CuA1 and CuA2, reaching 0.30 of wing length; peripheral membrane narrow.

Abdomen with seven visible segments, clothed with dense hairs; pygofer short, wide, slightly longer than preceding abdominal sternite (Fig. 3.5).

Etymology

The species name is from Guancaishan, indicating the type locality of the species.

Remarks

The new species distinctly differs from the type species, Cicadomorpha punctualata from Kazakhstan, by following characters of the tegmen: (1) tegmen length near 40 mm (near 54 mm in C. punctualata); (2) stem R slightly shorter than stem MP (extremely short in C. punctualata); (3) MP branching into MP1+2 and MP3+4 slightly after bifurcation of CuA (branching distinctly basad of bifurcation of CuA in C. punctualata); and (4) cross vein mp4-cua long and curved (absent in C. punctualata).

The new species is distinguished from other species of Cicadomorpha from Chita, Transbaikalia by following characters: (1) Pc + CP of tegmen emitting seven branches intersecting the costal area (ScP emitting three or four branches in C. milva and C. parula); (2) tegmen with MP branching into MP1+2 and MP3+4 slightly after bifurcation of CuA (MP branching basad of bifurcation of CuA in C. milva and C. parula); (3) tegmen with MP1+2 as long as MP3+4 (slightly longer than MP3+4 in C. milva; nearly two times longer than MP3+4 in C. parula); (4) cross vein mp-cua of tegmen with a short, wedge-shaped vein (short vein absent in C. milva and C. parula); (5) cross vein mp4-cua of tegmen curved mediad (straight in C. parula); (6) tegmen with Pcu fused with A1 distinctly basad of bifurcation of MP (Pcu fused with A1 at same level of bifurcation of MP in C. milva; A1 isolated in C. parula); and (7) hind wings with cross vein ir ~2.7 times as long as ScP + RA1 (ir not longer than ScP + RA1 in C. milva and C. parula).

Discussion

Martynov (Reference Martynov1926) erected the genus Cicadomorpha and the single species C. punctualata based on an incomplete specimen from Kazakhstan. Shcherbakov (Reference Shcherbakov1988) placed two species in Cicadomorpha (C. milva and C. parula) from Chita, Transbaikalia, Russia. Cicadomorpha guancaishanensis n. sp. from the Upper Jurassic Linglongta beds can be firmly assigned to Cicadomorpha. The new species is more similar to C. milva from Chita than other species in terms of morphological characters, e.g., size of body and wings, stem MP of forewing distinctly longer than MP1+2 and MP3+4, cross vein mp4-cua of forewing curved mediad, and Pcu fused with A1. It is notable that all species of Cicadomorpha show extremely limited morphological differences. Therefore, these might indicate the biostratigraphic correlation of some well-known fossil localities: the Tiaojishan Formation (Yanliao area, northeastern China), the Karabastau Formation (Karatau, Kazakhstan), and the Glushkovo and the Ukurei Formations (Chita, Transbaikalia, Russia). There is some dispute about the ages of the Karabastau Formation, the Glushkovo Formation, and the Ukurei Formation. It has been suggested that the age of the Karabastau Formation is Middle–Late Jurassic (e.g., Kirichkova and Doludenko, Reference Kirichkova and Doludenko1996; Zhang, Reference Zhang2015); the age of Glushkovo Formation and the Ukurei Formation are poorly constrained, suggested as Late Jurassic (Sinitsa, Reference Sinitsa1993), Early Cretaceous (Sinitshenkova, Reference Sinitshenkova2005), or perhaps at the Jurassic/Cretaceous boundary (Rasnitsyn and Quicke, Reference Rasnitsyn and Quicke2002).

Our discovery provides new evidence suggesting that all of the above units are Late Jurassic, which is consistent with previous studies (e.g., Ponomarenko et al., Reference Ponomarenko, Sukatsheva and Vasilenko2009; Sinitsa, Reference Sinitsa2016). The geological range of the Tiaojishan Formation was limited to 161–153 Ma by recent analyses based on isotope dating (Huang, Reference Huang2019), which is obviously different from the view that the age of the Tiaojishan Formation is the Middle Jurassic (Deng et al., Reference Deng, Yao, Ye, Chen, Jin, Zhang, Xü, Zhao, Yuan and Zhang2003, Reference Deng, Lu, Zhao, Fan, Wang, Yang, Li and Sun2017; Duan et al., Reference Duan, Zhen, Hu, Zhang and Wang2009). Isotope dating of the volcanic ash layer at Guancaishan ~1 m above the fossil palaeontinid layer yielded an age of 157.3 ± 1.5 Ma (SIMS U-Pb zircon age), which is close to the boundary of Oxfordian and Kimmeridgian (157.3 ± 1.0 Ma; Huang, Reference Huang2016). Therefore, this study provides a relative age for the other three well-known localities, indicating that the insect probably lived during the early Late Jurassic.

The Late Jurassic (Tithonian) palaeontinids from Solnhofen, Germany and the Cretaceous palaeontinids constitute a monophyletic group, representing the late stage of palaeontinid evolution with the following combination characters: (1) forewing trigonal; (2) costal area and claval reduced; (3) veins RA, RP, and MP originating at a common point; (4) length of the hind wing nearly one half that of the forewing; and (5) RP of hind wing often fused with MP1 (B. Wang et al., Reference Wang, Zhang and Szwedo2009). Cicadomorpha completely differs from the Solnhofen palaeontinids and Cretaceous palaeontinids, but shares similarities with Palaeontiondes Martynov, Reference Martynov1937 and Daohugoucossus Wang, Zhang, and Fang, Reference Wang, Zhang and Fang2006 from the Middle–Upper Jurassic Haifanggou Formation at Daohugou, Inner Mongolia, northeastern China in having similar subtriangular forewings, costal area and claval wide, cross vein mp-cua segmenting the discal cell, and hind wing relatively large. However, Cicadomorpha differs from Palaeontiondes and Daohugoucossus by two distinct characters: costal area with some intersecting branches and cross vein r-m of hind wing absent, and RP fused with MP1 for a considerable distance. The cross vein r-m of the hind wing is present in Dunstaniidae and Mesogereonidae, and some genera of Palaeontinidae, e.g., Suljuktocossus Becker-Migdisova, Reference Becker-Migdisova1949, Ningchengia B. Wang et al., Reference Wang, Zhang and Szwedo2009, and Palaeontinodes, but it is absent from most Cretaceous palaeontinids (B. Wang et al., Reference Wang, Zhang and Szwedo2009).

Cicadomorpha appears to be descended from the early assemblage of the Yanliao biota (e.g., Daohugou) by sharing some groups, and then migrating throughout Eurasia in the Late Jurassic, suggesting their adaptability to surroundings to some extent. The age difference between the core fossil layers of the Daohugou and the Linglongta assemblages is < 5 Ma. Until now, no species common to both assemblages have been discovered, because of the strong tectonic deformation and the large-scale volcanic activity that occurred (Yanshan Movement, Phase A2), resulting in the ecosystem of Daohugou area being seriously collapsed (Huang, Reference Huang2015; Huang et al., Reference Huang, Cai, Fu and Su2018a).

Biogeographic implications

Siberia and Mongolia were separated by an enormous gulf in the Paleopacific, called the Mongolia-Okhotsk Ocean, which is located in the northern part of the Mongolian-North China Block, dated from the Triassic to the Jurassic (Zonenshain et al., Reference Zonenshain, Kuzmin and Natapov1990; Huang, Reference Huang2019). The Siberia Plate began a southward subduction and closure of the Mongolia-Okhotsk Ocean, to produce the Mongol-Okhotsk fold belt (Zorin, Reference Zorin1999; Kravchinsky et al., Reference Kravchinsky, Cogne, Harbert and Kuzmin2002). There is much debate about the beginning of the closure of the Mongolia-Okhotsk Ocean. Available data suggest that the closure of the Mongolia-Okhotsk Ocean occurred during the Middle–Late Jurassic (Zonenshain et al., Reference Zonenshain, Kuzmin and Natapov1990; Kravchinsky et al., Reference Kravchinsky, Cogne, Harbert and Kuzmin2002; Tomurtogoo et al., Reference Tomurtogoo, Windley, Kroner, Badarch and Liu2005; Huang, Reference Huang2019), although paleomagnetic studies have indicated that it was completely closed by the Early Cretaceous (~130 Ma; Zhao et al., Reference Zhao, Coe, Zhou, Wu and Wang1990; Enkin et al., Reference Enkin, Yang, Chen and Courtillot1992; Gilder and Courtillot, Reference Gilder and Courtillot1997; Metelkin et al., Reference Metelkin, Gordienko and Klimuk2007). A regional unconformity below the Tiaojishan Formation of Yanliao area, China, formed at 161 Ma (Huang et al., Reference Huang, Cai, Fu and Su2018a; Huang, Reference Huang2019). There is also a regional unconformity below the Toutunhe Formation—the Junggar Basin in northwestern China (correlated to the Tiaojishan Formation)—and the age of the bottom of the Toutunhe Formation is ~161.2 Ma (Fang et al., Reference Fang, Wu, Guo, Hou, Dong, Wang and Li2015). Huang (Reference Huang2019) suggested that a widespread unconformity in northern China is indicative of the Yanshan Movement, Phase A2, related to the closure of the Mongolia-Okhotsk Ocean. The closing of the Mongolia-Okhotsk Ocean probably occurred before the Yanshan Movement, Phase A1 (~168 Ma). The ancient landform north and northeast of the Yanliao area is uplift, and then a set of complex conglomerates is rapidly denuded.

All species of Cicadomorpha are confined to Eurasia (Fig. 5). During the early Late Jurassic, C. punctualata from Kazakhstan was located in the western Okhotsk Ocean, C. milva and C. parula from Chita, Transbaikalia were located in the north Okhotsk Ocean, and the new species from Guancaishan, northeastern China was located in the southern Okhotsk Ocean (Fig. 5). Considering the limited flight capacity of these early cicada-like insects, it was almost impossible to cross the vast ocean, and the similarities between the new species and C. milva from Chita instead of C. punctualata from Kazakhstan indicate that a biological connection emerged between Chita and northeastern China. Our discovery, therefore, inclines toward the global palaeogeographic reconstruction of Scotese (Reference Scotese2014; Fig. 5.1), and provides new evidence indicating that the closure of the Okhotsk Ocean began in the Middle Jurassic.

Figure 5.

Palaeogeographic distribution of Cicadomorpha during the Late Jurassic: (1) global palaeogeographic reconstruction, 158 Ma (after Scotese, Reference Scotese2014); (2) global palaeogeographic reconstruction, 150 Ma (after https://www2.nau.edu/rcb7/). 1 = Guancaishan, Liaoning Province, China; 2 = Karatau, Kazakhstan; 3 = Chita, Transbaikalia, Russia.

Acknowledgments

We sincerely thank three anonymous reviewers for their useful comments on the paper. This work was supported by the National Key R&D Program of China (2016YFC0600406), the Strategic Priority Research Program of the Chinese Academy of Sciences (XDB26000000 and XDB18000000), and the National Natural Science Foundation of China (41688103).

References

Becker-Migdisova, E., 1949, Mesozoic Homoptera of central Asia: Trudy Paleontologicheskogo Instituta, v. 22, p. 1–68. [in Russian]Google Scholar
Bi, S., Wang, Y., Guan, J., Sheng, X., and Meng, J., 2014, Three new Jurassic euharamiyidan species reinforce early divergence of mammals: Nature, v. 514, p. 579–584, doi:10.1038/nature13718.Google ScholarPubMed
Bourgoin, T., Wang, R., Asche, M., Hoch, H., Soulier-Perkins, A., Stroinski, A., Yap, S., and Szwedo, J., 2015, From micropterism to hyperpterism: recognition strategy and standardized homology-driven terminology of the forewing venation patterns in planthoppers (Hemiptera: Fulgoromorpha): Zoomorphology, v. 134, p. 63–77, doi:10.1007/s00435-014-0243-6.Google ScholarPubMed
Cai, C., and Huang, D., 2016, Omma daxishanense sp. nov., a fossil representative of an extant Australian endemic genus recorded from the Late Jurassic of China (Coleoptera: Ommatidae): Alcheringa, v. 41, p. 277–283, doi:10.1080/03115518.2016.1225251.Google Scholar
Carpenter, F.M., 1992, Superclass Hexapoda, in Kaesler, R.L., ed., Treatise on Invertebrate Paleontology, Part R, Arthropoda 4: Boulder, Colorado, and Lawrence, Kansas, Geological Society of America (and University of Kansas Press), 655 p.Google Scholar
Chen, J., Zhang, H., Wang, B., Zheng, X., and Wang, X., 2016, New Jurassic Sinopalaeocossus and related genera with notes on their evolutionary implications (Hemiptera, Palaeontinidae): Insect Systematics and Evolution, v. 47, p. 113–129, doi:10.1163/1876312X-47022136.Google Scholar
Chen, J., Beattie, R., Wang, B., Jiang, H., Zheng, Y., and Zhang, H., 2019, The first palaeontinid from the Late Jurassic of Australia (Hemiptera, Cicadomorpha, Palaeontinidae): Alcheringa, v. 43, p. 449–454, doi:10.1080/03115518.2019.1587787.Google Scholar
Cheng, X., Wang, X., Jiang, S., and Kellner, A.W., 2012, A new scaphognathid pterosaur from western Liaoning, China: Historical Biology, v. 24, p. 101–111, doi:10.1080/08912963.2011.635423.Google Scholar
Deng, S., Yao, Y., Ye, D., Chen, P., Jin, F., Zhang, Y., Xü, K., Zhao, Y., Yuan, X., and Zhang, S., 2003, The Jurassic System of North China (I): A Stratigraphic Overview: Beijing, Petroleum Industry Press, 399 p. [in Chinese with English abstract]Google Scholar
Deng, S., Lu, Y., Zhao, Y., Fan, R., Wang, Y., Yang, X., Li, X., and Sun, B., 2017, The Jurassic palaeoclimate regionalization and evolution of China: Earth Science Frontiers, v. 24, p. 106–142, doi:10.13745/j.esf.2017.01.007. [in Chinese with English abstract]Google Scholar
Duan, Y., Zhen, S., Hu, D., Zhang, L., and Wang, W., 2009, Preliminary report on Middle Jurassic strata and fossils from Linglongta area of Jianchang, Liaoning: Global Geology, v. 28, p. 143–147. [in Chinese with English abstract]Google Scholar
Enkin, R.J., Yang, Z., Chen, Y., and Courtillot, V., 1992, Paleomagnetic constraints on the geodynamic history of the major blocks of China from the Permian to the present: Journal of Geophysical Research Atmospheres, v. 971, p. 13953–13989, doi:10.1029/92JB00648.Google Scholar
Evans, J.W., 1946, A natural classification of leaf-hoppers (Homoptera, Jassoidea), part 2: Aetalionidae, Hylicidae, Eurymelidae: Transactions of the Royal Entomological Society of London, v. 97, p. 39–54.Google Scholar
Fang, Y., Wu, C., Guo, Z., Hou, K., Dong, L., Wang, L., and Li, L., 2015, Provenance of the southern Junggar Basin in the Jurassic: Evidence from detrital zircon geochronology and depositional environments: Sedimentary Geology, v. 315, p. 47–63, doi:10.1016/j.sedgeo.2014.10.014.Google Scholar
Gao, K., and Shubin, N.H., 2012, Late Jurassic salamandroid from western Liaoning, China: Proceedings of the National Academy of Sciences, v. 109, p. 5767–5772, doi:10.1073/pnas.1009828109.Google ScholarPubMed
Gilder, S., and Courtillot, V., 1997, Timing of the North-South China collision from new middle to late Mesozoic paleomagnetic data from the North China Block: Journal of Geophysical Research, v. 102, p. 17713–17727.Google Scholar
Godefroit, P., Demuynck, H., Dyke, G., Hu, D., Escuillié, F., and Claey, P., 2013a, Reduced plumage and flight ability of a new paravian theropod from China: Nature Communications, v. 4, p. 1394, doi:10.1038/ncomms2389.Google Scholar
Godefroit, P., Cau, A., Hu, D., Escuillié, F., Wu, W., and Dyke, G., 2013b, A Jurassic avialan dinosaur from China resolves the early phylogenetic history of birds: Nature, v. 498, p. 359–362, doi:10.1038/nature12168.Google Scholar
Handlirsch, B.A., 1906–1908, Die Fossilen Insekten und die Phylogenie der Rezenten Formen: Ein Handbuch für Paläontologen und Zoologen: Leipzig, Germany, Engelmann, 1430 pp.Google Scholar
Hong, Y., 1982, Mesozoic Fossil Insects of the Jiuquan Basin in Gansu Province: Beijing, Geological Publishing House, 223 p. [in Chinese]Google Scholar
Hu, D., Lianhai, H., Zhang, L., and Xu, X., 2009, A pre-Archaeopteryx troodontid theropod from China with long feathers on the metatarsus: Nature, v. 461, p. 640–643, doi:10.1038/nature08322.Google ScholarPubMed
Huang, D., 2015, Yanliao biota and Yanshan movement: Acta Palaeontologica Sinica, v. 54, p. 501–546. [in Chinese with English abstract]Google Scholar
Huang, D., ed., 2016, The Daohugou Biota: Shanghai, Shanghai Scientific and Technical Publishers, 332 pp. [in Chinese]Google Scholar
Huang, D., 2019, Jurassic integrative stratigraphy and timescale of China: Science China Earth Science, v. 62, p. 223–255, doi:10.1007/s11430-017-9268-7.Google Scholar
Huang, D., Cai, C., Fu, Y., and Su, Y., 2018a, The Middle-Late Jurassic Yanliao entomofauna: Palaeoentomology, v. 1, p. 3–31, doi:10.11646/palaeoentomology.1.1.2.Google Scholar
Huang, D., Cai, C., and Nel, A., 2018b, A new damsel-dragonfly of the small family Selenothemistidae from the Late Jurassic of China (Odonata: Isophlebioptera): Journal of Paleontology, v. 92, p. 1–5, doi:10.1016/j.cretres.2011.11.019.Google Scholar
Jia, J., and Gao, K., 2016, A new basal salamandroid (Amphibia, Urodela) from the Late Jurassic of Qinglong, Hebei Province, China: PLoS ONE, v. 11, p. e0153834, doi:10.1371.journal.pone.0153834.Google ScholarPubMed
Jia, J., and Gao, K., 2019, A new stem hynobiid salamander (Urodela, Cryptobranchoidea) from the Upper Jurassic (Oxfordian) of Liaoning Province, China: Journal of Vertebrate Paleontology, v. 39, p. e1588285, doi:10.1080/02724634.2019.1588285.Google Scholar
Kirichkova, A.I., and Doludenko, M.P., 1996, New data on phytostratigraphy of the Jurassic deposits of Kazakhstan: Stratigraphy and Geological Correlation, v. 4, p. 35–52.Google Scholar
Kravchinsky, V.A., Cogne, J.P., Harbert, W.P., and Kuzmin, M.I., 2002, Evolution of the Mongol-Okhotsk Ocean as constrained by new palaeomagnetic data from the Mongol-Okhotsk suture zone, Siberia: Geophysical Journal International, v. 148, p. 34–57, doi:10.1046/j.1365-246x.2002.01557.x.Google Scholar
Li, Y., Chen, J., and Wang, B., 2019, New Cretaceous palaeontinids (Insecta, Hemiptera) from northeast China: Cretaceous Research, v. 95, p. 130–137.Google Scholar
Lü, J., 2009, A new non-pterodactyloid pterosaur from Qinglong County, Hebei Province of China: Acta Geologica Sinica, v. 83, p. 189–199, doi:10.1111/j.1755-6724.2009.00062.x.Google Scholar
Lü, J., Unwin, D.M., Jin, X., Liu, Y., and Ji, Q., 2010, Evidence for modular evolution in a long-tailed pterosaur with a pterodactyloid skull: Proceedings of the Royal Society B, Biological Sciences, v. 277, p. 383–389, doi:10.1098/rspb.2009.1603.Google Scholar
Lü, J., Unwin, D.M., Deeming, D.C., Jin, X., Liu, Y., and Ji, Q., 2011, An egg-adult association, gender, and reproduction in pterosaurs: Science, v. 331, p. 321–324, doi:10.1126/science.1197323.Google ScholarPubMed
Luo, Z., Yuan, C., Meng, Q., and Ji, Q., 2011, A Jurassic eutherian mammal and divergence of marsupials and placentals: Nature, v. 476, p. 442–445, doi:10.1038/nature10291.Google ScholarPubMed
Martynov, A.V., 1926, To the knowledge of fossil insects from the Jurassic beds in Turkestan: Izvestiia Akademii nauk SSSR, v. 20, p. 1349–1366.Google Scholar
Martynov, A.V., 1937, Liassic insects from Shurab and Kizil-Kiya: Trudy Paleontologicheskogo Instituta, v. 7, p. 1–232. [in Russian with English summary]Google Scholar
Meng, J., Hu, Y., Wang, Y., Wang, X., and Li, C., 2006, A Mesozoic gliding mammal from northeastern China: Nature, v. 444, p. 889–893, doi:10.1038/nature05234.Google ScholarPubMed
Menon, F., Heads, S.W., and Martill, D.M., 2005, New Palaeontinidae (Insecta: Cicadomorpha) from the Lower Cretaceous Crato Formation of Brazil: Cretaceous Research, v. 26, p. 837–844, doi:10.1016/j.cretres.2005.05.005.Google Scholar
Metelkin, D.V., Gordienko, I.V., and Klimuk, V.S, 2007, Paleomagnetism of Upper Jurassic basalts from Transbaikalia: New data on the time of closure of the Mongol-Okhotsk Ocean and Mesozoic intraplate tectonics of Central Asia: Russian Geology & Geophysics, v. 48, p. 825–834, doi:10.1016/j.rgg.2007.09.004.Google Scholar
Nam, K.S., Wang, Y., Ren, D., Kim, J.H., and Szwedo, J., 2017, An extraordinary palaeontinid from the Triassic of Korea and its significance: Scientific Reports, v. 7, p. 40691, doi:10.1038/srep40691.Google ScholarPubMed
Nel, A., Prokop, J., Nel, P., Grandcolas, P., Huang, D., Roques, P., Guilbert, E., Dostál, O., and Szwedo, J., 2012, Traits and evolution of wing venation pattern in paraneopteran insects: Journal of Morphology, v. 273, p. 480–506, doi:10.1002/jmor.11036.Google ScholarPubMed
Ponomarenko, A.G., Sukatsheva, I.D., and Vasilenko, D.V., 2009, Some characteristics of the Trichoptera distribution in the Mesozoic of Eurasia (Insecta: Trichoptera): Paleontological Journal, v. 43, p. 282–295, doi:10.1134/S003103010903006X.Google Scholar
Rasnitsyn, A.P., and Quicke, D.L.J., 2002, History of Insects: Dordrecht, The Netherlands, Kluwer Academic Publisher, 513 p.Google Scholar
Riek, E.F., 1976, A new collection of insects from the Upper Triassic of South Africa: Annals of the Natal Museum, v. 22, p. 791–820.Google Scholar
Scotese, C.R., 2014, Atlas of Jurassic Paleogeographic Maps, PALEOMAP Atlas for ArcGIS, Volume 3, The Jurassic and Triassic: Evanston, Illinois, Mollweide Projection, PALEOMAP Project, maps 32–42, doi:10.13140/2.1.4850.4321.Google Scholar
Shcherbakov, D.E., 1988, New cicadas (Cicadina) from the late Mesozoic of Transbaikalia: Paleontological Journal, v. 4, p. 55–66.Google Scholar
Shcherbakov, D.E., and Popov, Y.A., 2002, Superorder Cimicidea Laicharting, 1781 order Hemiptera Linné, 1758: The bugs, cicadas, plantlice, scale insects, etc., in Rasnitsyn, A.P., and Quicke, D.L.J., eds., History of Insects: Dordrecht, The Netherlands, Kluwer Academic Publisher, p. 152–155.Google Scholar
Sinitsa, S.M., 1993, Jurassic and Lower Cretaceous of Central Mongolia (ostracodes, stratigraphy, and paleoreconstruction): Paleontological Journal, v. 42, p. 1–240.Google Scholar
Sinitsa, S.M., 2016, Jurassic dinosaurs of Transbaikalia and prospects of searching for them in Mongolia: Paleontological Journal, v. 50, p. 1401–1411, doi:10.1134/S0031030116120170.Google Scholar
Sinitshenkova, N.D., 2005, The oldest known record of an imago of Nemouridae (Insecta: Perlida = Plecoptera) in the late Mesozoic of eastern Transbaikalia: Paleontological Journal, v. 39, p. 38–40.Google Scholar
Szwedo, J., 2018, The unity, diversity and conformity of bugs (Hemiptera) through time: Earth and Environmental Science Transactions of the Royal Society of Edinburgh, v. 107, p. 109–128, doi:10.1017/S175569101700038X.Google Scholar
Tillyard, R.J., 1916, Mesozoic and Tertiary insects of Queensland and New South Wales: Queensland Geological Survey, Publication 253, p. 11–47.Google Scholar
Tillyard, R.J., 1921, Mesozoic insects of Queensland, no. 8, Hemiptera-Homoptera: Proceedings of the Linnean Society of New South Wales, v. 46, p. 270–284.Google Scholar
Tomurtogoo, T., Windley, B.F., Kroner, A., Badarch, G., and Liu, D.Y., 2005, Zircon age and occurrence of the Adaatsag ophiolite and Muron shear zone, Central Mongolia: Constraints on the evolution of the Mongol-Okhotsk ocean, suture and orogeny: Journal of the Geological Society, London, v. 162, p. 125–134, doi:10.1144/0016-764903-146.Google Scholar
Wang, B., Zhang, H., and Fang, Y., 2006, Some Jurassic Palaeontinidae (Insecta, Hemiptera) from Daohugou, Inner Mongolia, China: Palaeoworld, v. 15, p. 115–125, doi:10.1016/j.palwor.2006.03.006.Google Scholar
Wang, B., Zhang, H., Fang, Y., Wang, D., and Ji, S., 2008, New data on Cretaceous Palaeontinidae (Insecta: Hemiptera) from China: Cretaceous Research, v. 29, p. 551–560, doi:10.1016/j.cretres.2008.01.007.Google Scholar
Wang, B., Zhang, H., and Szwedo, J., 2009, Jurassic Palaeontinidae from China and the higher systematics of Palaeontinoidea (Insecta: Hemiptera: Cicadomorpha): Palaeontology, v. 52, p. 53–64, doi:10.1111/j.1475-4983.2008.00826.x.Google Scholar
Wang, X., Kellner, A.W., Jiang, S., and Meng, X., 2009, An unusual long-tailed pterosaur with elongated neck from western Liaoning of China: Anais da Academia Brasileira de Ciências, v. 81, p. 793–812, doi:10.1590/S0001-37652009000400016.Google ScholarPubMed
Wang, X., Kellner, A.W., Jiang, S., Cheng, X., Meng, X., and Rodrigues, T., 2010, New long-tailed pterosaurs (Wukongopteridae) from western Liaoning, China: Anais da Academia Brasileira de Ciências, v. 82, p. 1045–1062, doi:10.1590/S0001-37652010000400024.Google ScholarPubMed
Wang, Y., and Ren, D., 2009, New fossil palaeontinids from the Middle Jurassic of Daohugou, Inner Mongolia, China (Insecta, Hemiptera): Acta Geologica Sinica, v. 83, p. 33–38, doi:10.1111/j.1755-6724.2009.00004.x.Google Scholar
Wang, Y., Ren, D., and Shih, C., 2007, New discovery of palaeontinid fossils from the Middle Jurassic in Daohugou, Inner Mongolia (Homoptera, Palaeontinidae): Science in China, ser. D, Earth Sciences, v. 50, p. 481–486, doi:10.1007/s11430-007-0029-5.Google Scholar
Wang, Y., Dong, L., and Evans, S., 2015, Polydactyly and other limb abnormalities in the Jurassic salamander Chunerpeton from China: Palaeobiodiversity and Palaeoenvironments, v. 96, p. 49–59, doi:10.1007/s12549-015-0219-7.Google Scholar
Xu, X., You, H., Du, K., and Han, F., 2011, An Archaeopteryx-like theropod from China and the origin of Avialae: Nature, v. 475, p. 465–470, doi:10.1038/nature10288.Google ScholarPubMed
Yang, Z., Jiang, B., McNamara, M.E., Kearns, S.L., Pittman, M., Kaye, T.G., Orr, P.J., Xu, X., and Benton, M.J., 2019, Pterosaur integumentary structures with complex feather-like branching: Nature Ecology & Evolution, v. 3, p. 24–30, doi:10.1038/s41559-018-0728-7.Google ScholarPubMed
Yuan, C., Ji, Q., Meng, Q., Tabrum, A.R., and Luo, Z., 2013, Earliest evolution of multituberculate mammals revealed by a new Jurassic fossil: Science, v. 341, p. 779–783, doi:10.1126/science.1237970.Google ScholarPubMed
Zhang, J., 2015, Archisargoid flies (Diptera, Brachycera, Archisargidae and Kovalevisargidae) from the Jurassic Daohugou biota of China, and the related biostratigraphical correlation and geological age: Journal of Systematic Palaeontology, v. 13, p. 857–881, doi:10.1080/14772019.2014.960902.Google Scholar
Zhao, X., Coe, R.S., Zhou, Y., Wu, H., and Wang, J., 1990, New paleomagnetic results from northern China: Collision and suturing with Siberia and Kazakhstan: Tectonophysics, v. 184, p. 43–81.Google Scholar
Zhou, C., Wu, S., Martin, T., and Luo, Z., 2013, A Jurassic mammaliaform and the earliest mammalian evolutionary adaptations: Nature, v. 500, p. 163–167, doi:10.1038/nature12429.Google ScholarPubMed
Zonenshain, L.P., Kuzmin, M.I., and Natapov, L.M., 1990, Geology of the USSR: A plate-tectonic synthesis: American Geophysical Union, v. 21, p. 97–108.Google Scholar
Zorin, Y.A., 1999, Geodynamics of the western part of the Mongolia-Okhotsk collisional belt, Trans-Baikal region (Russia) and Mongolia: Tectonophysics, v. 306, p. 33–56.Google Scholar
Figure 0

Figure 1. Distribution of Mesozoic palaeontinids in China.

Figure 1

Figure 2. (1) Stratigraphic column for the Tiaojishan Formation at Guancaishan, Jianping County, Liaoning Province, northeastern China; (2) map showing the Guancaishan locality; (3) the outcrop of the fossil, red arrow indicating the black shales from which the fossil was collected, and the white arrow indicating the volcanic ash layer slightly above the black shale.

Figure 2

Figure 3. Cicadomorpha guancaishanensis n. sp. from the Upper Jurassic Tiaojishan Formation at Guancaishan: (1) holotype (NIGP 168220), male, showing general habitus moistened with 70% ethanol; (2–5) enlargements of (1), showing details of: (2) head and pronotum, moistened with 70% ethanol; (3) right tegmen; (4) discal cell and cross vein mp-cua; and (5) genitals. ce = compound eye; gs = genital style; pr = pronotum; ro = rostrum. Scale bars = 1 mm (4, 5); 2 mm (2, 3); 6 mm (1).

Figure 3

Figure 4. Line drawings of Cicadomorpha guancaishanensis n. sp., holotype (NIGP 168220): (1) general habitus; (2) forewing; and (3) hind wing. A = anal vein; CP = costa posterior; CuA = cubitus anterior; CuP = cubitus posterior; MP = median posterior; Pc = precosta; Pcu = postcubitus; RA = radius anterior; RP = radius posterior; ScP = subcosta posterior. Scale bars = 4 mm (2, 3); 6 mm (1).

Figure 4

Figure 5. Palaeogeographic distribution of Cicadomorpha during the Late Jurassic: (1) global palaeogeographic reconstruction, 158 Ma (after Scotese, 2014); (2) global palaeogeographic reconstruction, 150 Ma (after https://www2.nau.edu/rcb7/). 1 = Guancaishan, Liaoning Province, China; 2 = Karatau, Kazakhstan; 3 = Chita, Transbaikalia, Russia.