Paleoenvironmental significance of ichnogenus Skolithos in the aftermath of the end-Permian mass extinction: a case study of the Lower Triassic Heshanggou Formation in western Henan Province
Xing Zhi-Feng1,2,3, Li Wan-Ying1, Qi Yong-An1,2,3, Zheng Wei1,2,3, Xu Xin1, Zhang Xiang-Yun1, Li Da1,2,3, Wu Pan-Pan1
1 College of Resources and Environment,Henan Polytechnic University,Henan Jiaozuo 454003,China; 2 Henan Province Key Laboratory of Biogenic Traces and Sedimentary Minerals,Henan Jiaozuo 454003,China; 3 Central Plains Economic Zone Coalbed Methane and Shale Gas Collaborative Innovation Center,Henan Jiaozuo 454003,China
Abstract The Lower Triassic ichnofossils in the western Henan record the evolution of trace-maker communities from remnant to recovery after the Permian-Triassic mass extinction(PTME). Skolithos is characterized by vertical or high angle inclination to the rock layer,and is widely preserved in the Lower Triassic Heshanggou Formation. Macroscopic and microscopic characteristics of Skolithos were studied on abundant,well-developed Skolithos specimens collected from the Heshanggou Formation in Dengfeng and Yiyang areas.We conducted a three dimensional occupation cubes of ecospace utilization and ecosystem engineering by analyzing the motility,feeding strategy,the interaction between organisms and substrate, and the way of biological modification of sediments. We also analyzed the paleoenvironmental significance of Skolithos and explored the high environmental pressure indicated by Skolithos ichnofossils. The results show that Skolithos is mainly affected by high environmental pressures such as high temperatures,dry and hot climate and giant monsoon in the early stage of recovery. As an opportunistic ichnogenus, Skolithos was widely developed in meandering river and shallow lacustrine deposits. The above results may provide a reference for the study of continental sedimentary environment and paleoecological background of the Lower Triassic in the western Henan.
Fund:Co-funded by the National Natural Science Foundation of China(Nos. 41772110,41872111,41102008,41902113),Key Research Project of Higher Education Institutions in Henan Province(No. 20A170010) and Key Scientific and Technological Projects in Henan Province(No. 202102310336)
Corresponding Authors:
Zheng Wei,born in 1979,is an associate professor in the college of resources and environment of Henan Polytechnic University. He is mainly engaged in researches of sedimentology,ichnology. E-mail: zhengw99@hpu.edu.cn.
About author: Xing Zhi-Feng,born in 1973,is an associate professor in the college of resources and environment of Henan Polytechnic University. She is mainly engaged in researches of sedimentology,paleontology and ichnology. E-mail: xingzhifeng925@hpu.edu.cn.
Cite this article:
Xing Zhi-Feng,Li Wan-Ying,Qi Yong-An et al. Paleoenvironmental significance of ichnogenus Skolithos in the aftermath of the end-Permian mass extinction: a case study of the Lower Triassic Heshanggou Formation in western Henan Province[J]. JOPC, 2022, 24(3): 461-478.
Xing Zhi-Feng,Li Wan-Ying,Qi Yong-An et al. Paleoenvironmental significance of ichnogenus Skolithos in the aftermath of the end-Permian mass extinction: a case study of the Lower Triassic Heshanggou Formation in western Henan Province[J]. JOPC, 2022, 24(3): 461-478.
[1] 常龙,王长征,胡斌,陈传浩. 2017. 东濮凹陷沙三段正常三角洲沉积中遗迹化石及意义. 沉积与特提斯地质, 37(3): 66-73. [Chang L,Wang C Z,Hu B,Chen C H.2017. Trace fossils from the normal delta deposits in the third member of the Shahejie Formation in the Dongpu depression and their significance. Sedimentary Geology and Tethyan Geology, 37(3): 66-73] [2] 陈浩,黄继新,常广发,武军昌,孙天建. 2018. 基于全岩心CT的遗迹化石识别及沉积环境分析: 以加拿大麦凯Ⅲ油砂区块为例. 古地理学报, 20(4): 703-712. [Chen H,Huang J X,Chang G F,Wu J C,Sun T J.2018. Ichnology identification and sedimentary environment analysis based on CT scanning: a case study from Mackay Ⅲ oil sands,Canada. Journal of Palaeogeography(Chinese Edition), 20(4): 703-712] [3] 陈璐,陆廷清,郭溦,李爽,陈泉均,张矩生,王广强. 2019. 川西南峨眉山地区上白垩统夹关组遗迹组构特征及其环境意义. 古生物学报, 58(2): 222-231. [Chen L,Lu T Q,Guo Z,Li S,Chen Q J,Zhang J S,Wang G Q.2019. Ichnofabrics and their environmental significance from the Upper Cretaceous Jiaguan Formation in Emei MT. area,Southwest Sichuan Basin. Acta Palaeontologica Sinica, 58(2): 222-231] [4] 陈秀艳,姜在兴,师晶,徐杰. 2012. 东辛地区沙三段遗迹化石特征与沉积环境分析. 断块油气田, 19(4): 462-466. [Chen X Y,Jiang Z X,Shi J,Xu J.2012. Ichnofossil characteristics and depositional environment of Es 3 reservoir in Dongxin area. Fault Block Oil & Gas Field, 19(4): 462-466] [5] 初庆春. 1988. 河北抚宁柳江盆地中寒武统徐庄组遗迹化石及沉积环境. 北京大学学报(自然科学版), 24(2): 220-234. [Chu Q C.1988. Trace fossils from the Xuzhuang Formation(Middle Cambrian)of Liujiang Basin(Funing,Hebei Province)and their depositional environments. Acta Scientiarum Naturalium Universitatis Pekinensis, 24(2): 220-234] [6] 杜圣贤,张园园,李越,张尚坤,刘凤臣,宋香锁,穆西南,周腾飞,周瑶琪,卢辉楠. 2020. 山东青岛早白垩世遗迹化石和皱饰构造的指相意义. 地层学杂志, 44(2): 174-180. [Du S X,Zhang Y Y,Li Y,Zhang S K,Liu F C,Song X S,Mu X N,Zhou T F,Zhou Y Q,Lu H N.2020. Ichnofossils and wrinkle structures as facies indicators from the Lower Cretaceous in Qingdao,Shandong,eastern China. Journal of Stratigraphy, 44(2): 174-180] [7] 高正海,李瑞云,王秋军,董重,闫德飞,孙柏年. 2013. 柴达木盆地东北缘中新统遗迹化石组合及其沉积环境. 沉积学报, 31(3): 413-420. [Gao Z H,Li R Y,Wang Q J,Dong Z,Yan D F,Sun B N.2013. Trace fossil assemblages in the Miocene from northeast margin of Qaidam Basin and its sedimentary environment. Acta Sedimentologica Sinica, 31(3): 413-420] [8] 龚一鸣,胡斌,卢宗盛,齐永安,张国成. 2009. 中国遗迹化石研究80年. 古生物学报, 48(3): 322-337. [Gong Y M,Hu B,Lu Z S,Qi Y A,Zhang G C.2009. Study of trace fossils in the past eighty years in China. Acta Palaeontologica Sinica, 48(3): 322-337] [9] 胡斌,张国成,郭卫星,齐永安,张利伟. 2004. 豫西济源谭庄组河流沉积中的Stipsellus遗迹组构. 煤田地质与勘探, 32(3): 1-5. [Hu B,Zhang G C,Guo W X,Qi Y A,Zhang L W.2004. The Stipsellus ichnofabric in the fluvial deposits of Upper Triassic Tanzhuang Formation in Jiyuan,western Henan. Coal Geology and Exploration, 32(3): 1-5] [10] 胡斌,芦旭辉,宋慧波. 2015. 豫西登封地区下三叠统中的遗迹化石及其沉积环境. 地层学杂志, 39(4): 454-465. [Hu B,Lu X H,Song H B.2015. Lower Triassic ichnofossils and sedimentary environments in the Dengfeng area,west Henan Province. Journal of Stratigraphy, 39(4): 454-465] [11] 胡斌,齐永安,宋慧波,牛永斌,张立军,郑伟,王长征. 2021. 中国遗迹学研究十年进展. 古地理学报, 23(2): 284-320. [Hu B,Qi Y A,Song H B,Niu Y B,Zhang L J,Zheng W,Wang C Z.2021. Progress of monomorphology in China in the past decade. Journal of Palaeogeography(Chinese Edition), 23(2): 284-320] [12] 金惠,张俊,马石玉. 2013. 四川盆地上三叠统遗迹化石及其沉积环境. 岩性油气藏, 25(1): 24-28. [Jin H,Zhang J,Ma S Y.2013. Trace fossils and sedimentary environment of the Upper Triassic Xujiahe Formation in Sichuan Basin. Lithologic Reservoir, 25(1): 24-28] [13] 荆锡贵,陈政安,李凤杰,文胜男,谢胜军. 2019. 龙门山地区中泥盆统养马坝组风暴沉积中的遗迹化石及其环境意义. 沉积学报, 37(4): 749-757. [Jing X G,Chen Z A,Li F J,Wen S N,Xie S J.2019. Ichnofossils and their environmental significance in storm deposit from the Middle Devonian Yangmaba Formation in Longmenshan area. Acta Sedimentologica Sinica, 37(4): 749-757] [14] 李妲,齐永安,代明月,王敏. 2016. 基于遗迹化石的寒武系馒头组潮间带食物网特征. 河南理工大学学报(自然科学版), 35(2): 194-201. [Li D,Qi Y A,Dai M Y,Wang M.2016. Characteristics of the intertidal food web and paleoecology based on trace fossils in Mantou Formation of Cambrian. Journal of Henan Polytechnic University(Natural Science), 35(2): 194-201] [15] 刘松,曾宪官,陈应权,闫德飞. 2017. 甘肃宝积山盆地上三叠统Skolithos遗迹化石研究. 甘肃地质, 26(2): 1-6. [Liu S,Zeng X G,Chen Y Q,Yan D F.2017. Skolithos from the Upper Triassic Baojishan basin of Gansu Province. Gansu Geology, 26(2): 1-6] [16] 罗茂,龚一鸣,张立军,殷亚飞. 2021. 遗迹化石: 探索生物与环境相互作用的重要信息载体. 古生物学报, 60(3): 347-356. [Luo M,Gong Y M,Zhang L J,Yin Y F.2021. Trace fossil: a significant agent for exploring organism environment interactions. Acta Palaeontologica Sinica, 60(3): 347-356] [17] 齐永安,苏现波,陈传诗. 1993. 河南济源晚三叠世曲流河沉积中的痕迹化石及其意义. 河南地质, 11(2): 119-122,161. [Qi Y A,Su X B,Chen C S.1993. Trace fossils in Late Triassic meandering river sediments in Jiyuan,Henan Province and their significance. Henan Geology, 11(2): 119-122,161] [18] 齐永安. 1998. 遗迹化石与潮控滨线海泛面的识别及准层序相组合: 以塔里木盆地下志留统塔塔埃尔塔格组为例. 矿物岩石, 37(4): 13-17. [Qi Y A.1998. Trace fossils and recognition of flooding surface and facies associations of parasequen,identification of tidal controlled shoreline flooding surface and parasequences in tide-controlled shoreline. Journal of Mineralogy and Petrology, 37(4): 13-17] [19] 齐永安. 1999. 生物扰动和遗迹组构的描述与分析. 河南地质, 17(4): 273-277. [Qi Y A.1999. The description and analysis of bioturbation and ichnofabric. Henan Geology, 17(4): 273-277] [20] 齐永安,胡斌,Bruno Mistiaen,Jean-Pierre Nicollin.2005. 粤北晚泥盆世天子岭组遗迹组构及其环境解释. 煤田地质与勘探, 33(2): 11-14. [Qi Y A,Hu B,Mistiaen B,Nicollin J P.2005. Ichnofabrics and their environmental interpretation from Tianziling Formation(Upper Devonian)of northern Guangdong. Coalfield Geology and Exploration, 33(2): 11-14] [21] 齐永安,胡斌,张国成,龚一鸣. 2007. 豫西济源地区中三叠世油房庄组遗迹组构及其环境解释. 沉积学报, 25(3): 372-379. [Qi Y A,Hu B,Zhang G C,Gong Y M.2007. Ichnofabrics and their environmental interpretation from Middle Triassic Youfangzhuang Formation,Jiyuan region,western Henan Province. Acta Sedimentologica Sinica, 25(3): 372-379] [22] 齐永安,王敏,李妲,代明月. 2012. 寒武纪底质革命: 从微生物席底到生物扰动混合底. 河南理工大学学报(自然科学版), 31(2): 159-164. [Qi Y A,Wang M,Li D,Dai M Y.2012. Cambrian substrate revolution: from matgrounds to bioturbated mixgrounds. Journal of Henan Polytechnic University(Natural Science), 31(2): 159-164] [23] 齐永安,姚倩,代明月,李妲. 2014a. 豫西登封寒武系馒头组碳酸盐岩中的风暴沉积及相关的生物成因构造. 地质调查与研究, 37(4): 241-248. [Qi Y A,Yao Q,Dai M Y,Li D.2014a. Storm deposits and biogenic structures from the carbonate of the Mantou Formation(Cambrian),Dengfeng area,western Henan. Geological Survey and Research, 37(4): 241-248] [24] 齐永安,孟瑶,代明月,李妲. 2014b. 豫西登封地区寒武系第二统朱砂洞组生物成因的豹斑构造. 地质科技情报, 33(5): 1-8. [Qi Y A,Meng Y,Dai M Y,Li D.2014b. Biogenic leopare patch structures from the Zhushadong Formation(Cambrian Series 2),Dengfeng area,western Henan. Geological Science and Technology Information, 33(5): 1-8] [25] 钱利军,时志强,欧莉华. 2010. 二叠纪—三叠纪古气候研究进展—泛大陆巨型季风气候: 形成、发展与衰退. 海相油气地质, 15(3): 52-58. [Qian L J,Shi Z Q,Ou L H.2010. Research advances in Permian-Traissic paleoclimate: formation,development and decline of pangaean megamonsoon. Marine Origin Petroleum Geology, 15(3): 52-58] [26] 王道生,张国成. 2005. 济源盆地中三叠世油房庄组的古土壤特征. 河南理工大学学报(自然科学版), 24(4): 278-282. [Wang D S,Zhang G C.2005. Features of paleosols from Middle Triassic Youfangzhuang Formation,Jiyuan Basin. Journal of Henan Polytechnic University(Natural Science), 24(4): 278-282] [27] 王长征. 2015. 豫西白垩纪盆地遗迹化石及沉积环境研究. 河南理工大学博士论文,1-158. [Wang C Z.2015. Study on trace fossils and sedimentary environment of Cretaceous Basin in western Henan. Doctoral dissertation of Henan Polytechnic University,1-158] [28] 王海邻. 2018. 我国东部沿海潮坪沉积中的现代生物遗迹研究. 河南理工大学博士论文,1-173. [Wang H L.2018. Study on modern biological relics in tidal flat sediments along the east coast of China. Doctoral dissertation of Henan Polytechnic University,1-173] [29] 王媛媛,王学芹,胡斌. 2019. 黄河三角洲潮坪环境中现代生物遗迹组成与分布特征. 沉积学报, 37(6): 1244-1257. [Wang Y Y,Wang X Q,Hu B.2019. The composition and distribution characteristics of biogenic sedimentary structures in tidal flat of Yellow River Delta. Acta Sedimentologica Sinica, 37(6): 1244-1257] [30] 吴灏,丁奕,金幸生,杜天明,郑文杰,李越. 2020. 浙江义乌上白垩统金华组滨湖相红层中的遗迹化石. 地层学杂志, 44(4): 428-438. [Wu H,Ding Y,Jin X S,Du T M,Zheng W J,Li Y.2020. Trace fossils from the lakeside red beds of the Upper Cretaceous Jinhua Formation in Yiwu,Zhejiang Province. Journal of Stratigraphy, 44(4): 428-438] [31] 邢智峰,周虎,林佳,王敏,郑伟. 2018. 河南宜阳下三叠统刘家沟组微生物成因沉积构造演化及其对古环境变化的响应. 古地理学报, 20(3): 191-206. [Xing Z F,Zhou H,Lin J,Wang M,Zheng W.2018. Evolution of Microbially Induced Sedimentary Structures and their response to palaeoenvironment variation of the Lower Triassic Liujiagou Formation in Yiyang area,Henan Province. Journal of Palaeogeography(Chinese Edition), 20(3): 191-206] [32] 邢智峰,林佳,周虎,李妲,齐永安,郑伟. 2019. 豫西宜阳地区晚二叠世晚期—早三叠世沉积记录及其对古气候变化的响应. 高校地质学报, 25(3): 401-411. [Xing Z F,Lin J,Zhou H,Li D,Qi Y A,Zheng W.2019. Palaeoclimate changes revealed by sedimentary records of the Upper Permian-Lower Triassic strata in Yiyang area,western Henan Province,China. Geological Journal of China Universities, 25(3): 401-411] [33] 杨式溥. 1983. 广西横县六景早泥盆世遗迹化石及其古生态学意义. 中国区域地质, 5(3): 11-22. [Yang S P.1983. Early Devonian trace fossils from Liujing,Hengxian County,Guangxi and their paleoecological significance. Regional Geology of China, 5(3): 11-22] [34] 杨式溥,张建平,杨美芳. 2004. 中国遗迹化石. 北京: 科学出版社. [Yang S P,Zhang J P,Yang M F.2004. Chinese Trace Fossils. Beijing: Science Press] [35] 于水情,邢智峰,周虎. 2015. 豫西下三叠统刘家沟组微生物成因沉积构造. 四川地质学报, 35(4): 483-486. [Yu S Q,Xing Z F,Zhou H.2015. Microbial induced sedimentary structure in the Lower Triassic Liujiagou Formation in West Henan. Acta Geologica Sichuan, 35(4): 483-486] [36] 曾羽. 1988. 贵州赫章早奥陶世湄潭期Skolithos遗迹相及其古环境. 贵州地质, 5(1): 66-71,103. [Zeng Y.1988. The significance of the Skolithos ichnofacies and palaeoenvironment in the Meitan period of the Early Ordovician in Hezhang,Guizhou. Guizhou Geology, 5(1): 66-71,103] [37] 张立军,龚一鸣,马会珍. 2011. 华南泥盆纪遗迹化石及遗迹相. 古地理学报, 13(4): 397-418. [Zhang L J,Gong Y M,Ma H Z.2011. The Devonian trace fossils and ichnofacies from South China. Journal of Palaeogeography(Chinese Edition), 13(4): 397-418] [38] 张立军,龚一鸣. 2013. 四川龙门山地区下泥盆统平驿铺组的遗迹化石. 古生物学报, 52(1): 86-95. [Zhang L J,Gong Y M.2013. Ichnocoenosis of the Lower Devonian Pingyipu Fornation,Longmen mountion,Sichuan. Acta Palaeontologica Sinica, 52(1): 86-95] [39] 张利伟,姚建新,张立军. 2017. 豫西地区早三叠世微生物成因沉积构造的微观特征. 微体古生物学报, 34(4): 418-427. [Zhang L W,Yao J X,Zhang L J.2017. Microscopic features of microbial induced sedimen tary structures in the Early Triassic terrestrial deposits in western Henan Province,China. Acta Micropalaeontologica Sinica, 34(4): 418-427] [40] 张喜林,朱筱敏,郭长敏,杨俊生,王玉秀. 2006. 苏北盆地高邮凹陷古近系戴南组滨浅湖沉积中的遗迹化石. 沉积学报, 24(1): 81-89. [Zhang X L,Zhu X M,Guo C M,Yang J S,Wang Y X.2006. Trace fossils of shore-shallow lacustrine Dainan Formation(Paleogene)in Gaoyou sag,Subei Basin,Jiangsu Province. Acta Sedimentologica Sinica, 24(1): 81-89] [41] 张欣平,卿上康. 1988. 新邵县马栏边下石炭统的遗迹化石及其沉积环境. 湖南地质, 7(2): 53-61,86-87. [Zhang X P,Qing S K.1988. Trace fossils and the sedimentary environments of the Lower Carboniferous in Malanbian,Xinshao County. Geology of Hunan, 7(2): 53-61,86-87] [42] 郑锋利. 2019. 华北盆地西部太原组碳酸盐岩中遗迹组合与古氧相的响应特征. 河南理工大学硕士论文,1-94. [Zheng F L.2019. Response characteristics of trace assemblages and paleooxygen facies in carbonate rocks of Taiyuan Formation in western North China Basin. Masteral thesis of Henan Polytechnic University,1-94] [43] 郑伟,齐永安,张忠慧,邢智峰. 2016. 豫西荥阳陆相二叠纪—三叠纪之交的微生物成因构造(MISS)及其地质意义. 地球科学进展, 31(7): 737-750. [Zheng W,Qi Y A,Zhang Z H,Xing Z F.2016. Characteristic and geological significance of Microbially Induced Sedimentary Structures(MISS)in terrestrial P-T boundary in Xingyang,western Henan Province. Advances in Earth Science, 31(7): 737-750] [44] 朱士兴,陈辉能,高建平. 2001. 山西灵丘云彩岭组的时代归属: Skolithos潜穴化石的研究和意义. 前寒武纪研究进展, 24(3): 163-174. [Zhu S X,Chen H N,Gao J P.2001. Pipe-Rocks-Skolithos from Lingqiu County of Shanxi Province,China. Progress in Precambrian Research, 24(3): 163-174] [45] Alpert S P.1974. Systematic review of the genus Skolithos. Journal of Paleontology, 48(4): 661-669. [46] Aswan,Rudyawan A A,Oo T Z.2018. Ichnofossils study of paleocene sediment source rock cores from Bintuni Basin,west Papua,eastern Indonesia. IOP Conference Series: Earth and Environmental Science, 162(1): 012034. [47] Buatois L A,Guillermo F A.1996. Trace fossils and sedimentary facies from a Late Cambrian-Early Ordovician tide-dominated shelf(Santa Rosita Formation,northwest Argentina): implications for ichnofacies models of shallow marine successions. Ichnos, 5(1): 53-88. [48] Buatois L A,Mángano M G.2011. Ichnology: organism substrate interactions in space and time. Cambridge University Press, 149(4): 750. [49] Cribb A T,Bottjer D J.2020. Complex marine bioturbation ecosystem engineering behaviors persisted in the wake of the End Permian Mass Extinction. Scientific Reports, 10(1): 1-8. [50] Ding Y,Liu J N,Chen F F.2020. Ichnology,palaeoenvironment and ecosystem dynamics of the Early Cambrian(Stage 4,Series 2)Guanshan biota,South China. Geological Journal, 55(1): 1-18. [51] Driese S G,Byers C W,Dott R H.1980. Tidal deposition in the basal Upper Cambrian Mt. Simon Formation in Wisconsin. Journal of Sedimentary Petrology, 51: 367-381. [52] Droser M L.1991. Ichnofabric of the Paleozoic Skolithos ichnofacies and the nature and distribution of the Skolithos piperock. Palaios, 69(3): 316-325. [53] Eisawi A A M,Babikir I A A,Salih K A O.2011. Paleoecological significance of newly discovered trace fossils near the Gedaref town,eastern Sudan. Journal of African Earth Sciences, 61(3): 233-237. [54] Ekdale A A.1985. Paleoecology of the marine endobenthos. Palaeogeography,Palaeoclimatology,Palaeoecology, 50(1): 63-81. [55] Ekwenye O C,Nichols G,Okogbue C O,Mode A W.2016. Trace fossil assemblages in the tide-dominated estuarine system: Ameki Group,south-eastern Nigeria. Journal of African Earth Sciences, 80(2): 189-214. [56] Gregory M R,Campbell K A,Zuraida R,Martin A J.2006. Plant traces resembling Skolithos. Ichnos, 13(4): 205-216. [57] Hallam A,Swett K.1966. Trace fossils from the Lower Cambrian pipe rock of the north-west Highlands. Scottish Journal of Geology, 2: 101-106. [58] Knaust D,Thomas R D K,Curran H A.2018. Skolithos linearis Haldeman,1840 at its Early Cambrian type locality,chickies rock,Pennsylvania: analysis and designation of a neotype. Earth-Science Reviews, 185: 5-31. [59] Lukman A M,Ayuba R,Alege T S.2018. Sedimentology and depositional environments of the Maastrichtian Mamu Formation,northern Anambra Basin,Nigeria. Advances in Applied Science Research, 9(2): 53-68. [60] Mángano M G,Buatois L A.2014. Decoupling of body-plan diversification and ecological structuring during the Ediacaran-Cambrian transition: evolutionary and geobiological feedbacks. Proceedings of the Royal Society B: Biological Sciences, 281(1780): 1-9. [61] McKie T.1990. Tidal and storm influenced sedimentation from a Cambrian transgressive passive margin sequence. Journal of the Geological Society of London, 147: 785-794. [62] Minter N J,Buatois L A,Mángano M G,Davies N S,Gibling M R,MacNaughton R B,Labandeira C C.2017 . Early bursts of diversification defined the faunal colonization of land. Nature Ecology and Evolution, 1(7): 175. [63] Nascimento D L,Netto R G.2019. Skolithos serratus in paleosols: paleobiological,paleoecological and paleobiogeographical insights. Palaeogeography,Palaeoclimatology,Palaeoecology, 530: 152-162. [64] Radley J D,Coram R A.2015. Derived Skolithos pipe rock in the Budleigh Salterton pebble beds(Early Triassic,East Devon,UK). Proceedings of the Geologists' Association, 126(2): 220-225. [65] Rygel M C,Fielding C R,Bann K L,Frank T D,Birgenheier L,Tye S C.2008. The Lower Permian Wasp Head Formation,Sydney basin: high-latitude,shallow marine sedimentation following the late Asselian to early Sakmarian glacial event in eastern Australia. Sedimentology, 55(5): 1517-1540. [66] Santos H P,Mángano M G,Soares J L,Afonso C R,Bandeira N J,Rudnitzki I D.2017. Ichnologic evidence of a Cambrian age in the southern Amazon craton: implications for the onset of the western Gondwana history. Journal of South American Earth Sciences, 76: 482-488. [67] Scotese C R.1994. Early Triassic paleogeographic map. In: Klein G D(ed). Pangea: paleoclimate,tectonics and sedimentation during accretion zenith and break up of a supercontinent. Geological Society of America Special Paper, 288: 7. [68] Seilacher A.1967. Bathymetry of trace fossils. Marine Geology, 5(5-6): 413-428. [69] Simpson E L,Eriksson K A.1990. Early Cambrian progradational and transgressive sedimentation patterns in Virginia: an example of the early history of a passive margin. Journal of Sedimentary Petrology, 60: 84-100. [70] Solórzano E J,Buatois L A,Rodríguez W J,Mángano M G.2017. From freshwater to fully marine: exploring animal substrate interactions along a salinity gradient(Miocene Oficina Formation of Venezuela). Palaeogeography,Palaeoclimatology,Palaeoecology, 482: 30-47. [71] Wang M,Qi Y A,Li D,Dai M Y,Chang Y G.2014. Ichnofabrics and their environmental interpretation from the fluvial deposits of the Middle Triassic Youfangzhuang Formation in western Henan,central China. Journal of Earth Science, 25(4): 648-661. [72] Yul K J,Hyun K J.2018. Trace fossils from the Late Pleistocene marginal marine deposits of Jeju Island,Korea: implications for the Psilonichnus and Skolithos ichnofacies. Journal of the Korean Earth Science Society, 39(1): 23-45. [73] Zonneveld J P.2016. Applications of experimental neoichnology to paleobiological and evolutionary problems. Palaios, 31(6): 275-279.