Benthic animal response to the Paleozoic plant terrestrialization: insights from trace fossils

doi:10.7605/gdlxb.2026.135

Journal of Palaeogeography（Chinese Edition） ›› 2026, Vol. 28 ›› Issue (4): 1257-1274. doi: 10.7605/gdlxb.2026.135

• SPECIAL TOPIC ON THE 100TH ANNIVERSARY OF PROFESSOR FENG ZENGZHAO • Previous Articles Next Articles

Benthic animal response to the Paleozoic plant terrestrialization: insights from trace fossils

LI Feiyang(), ZHANG Lijun(), WANG Zongyan, CHAI Haiyan

School of Resources and Environment, Henan Key Laboratory of Coal Measure Unconventional Resources Accumulation and Exploitation, Henan Jiaozuo 454003, China

Received:2026-04-10 Revised:2026-05-28 Online:2026-08-01 Published:2026-08-14
Contact: ZHANG Lijun,born in 1982,is a professor. He is mainly engaged in teaching and researches on paleobiology and sedimentology. E-mail: Ljzhanghpu@gmail.com.
About author:
LI Feiyang,born in 1993,is a Ph.D. candidate. He is mainly engaged in research on ichnology. E-mail: 1169006628@qq.com.
Supported by:
Co-funded by the National Natural Science Foundation of China(42472037); Natural Science Foundation of Henan(252300421289)

底栖动物如何响应古生代植物陆地化进程: 遗迹化石的证据与启示^*

李飞洋(), 张立军(), 王宗延, 柴海艳

河南理工大学资源环境学院, 河南省煤系非常规资源成藏与开发重点实验室, 河南焦作 454003

通讯作者: 张立军,男,1982年生,教授,主要从事古生物学和沉积学的教学和科研工作。E-mail: Ljzhanghpu@gmail.com。
作者简介:
李飞洋,男,1993年生,博士研究生,从事遗迹学研究。E-mail: 1169006628@qq.com。
谨以此文纪念冯增昭先生100周年诞辰。
基金资助:
*国家自然科学基金项目(42472037); 河南省自然科学基金项目(252300421289)

Abstract

Abstract:

The Paleozoic plant terrestrialization(PPT)represents a pivotal point in the evolutionary history of life on Earth. This process not only reshaped surficial geomorphological landscapes but also exerted on others biotic evolution throughout the Phanerozoic. Until now,how benthic animals respond to PPT across different depositional environments,as well as their roles in the construction of benthic ecosystems,have remained insufficiently understood. Based on the compilation and synthesis of a global trace fossil dataset,this study establishes a time-depositional environment analytical framework. Within this framework,we systematically compare the evolutionary patterns of ichnodiversity,ichnodisparity,ichnotiering,and benthic ecosystem engineering across transitional coastal,fluvial,and lacustrine settings during the process of PPT. Furthermore,we quantitatively assess the behavioral responses of benthic animals to PPT. The results show that from the Cambrian to the Permian,the behavioral evolution of benthic animals exhibits pronounced time-limited and heterogeneity. During the Ordovician,benthic animals in transitional coastal environments had already begun to show initial responses to PPT,whereas those in fluvial and lacustrine environments display a clear temporal lag. This discrepancy is closely associated with the development of plant root systems,the progression of PPT,and subsequent forest expansion. Overall,with the progressive improvement of substrate stability and organic matter availability in both continental and transitional settings driven by terrestrial plants,benthic animals gradually expanded their ecospace utilization,accompanied by increased behavioral complexity and enhanced ecosystem engineering. In turn,benthic animals exerted a positive feedback on PPT through sediment reworking and the modulation of matter and energy fluxes. In addition to the influence of terrestrial vegetation,environmental factors such as climate change and tectonic activity,and sea-level fluctuations also played significant roles in shaping benthic evolutionary trajectories. This study provides a novel ichnological perspective for understanding Paleozoic ecosystem evolution and biotic-sediment interactions.

Key words: Paleozoic, terrestrialization, trace fossils, ecological feedback, benthic ecosystem

摘要：

古生代植物登陆是地球生命演化史上的重大事件,不仅重塑了地球表层的地貌景观,还对其他门类生物演化产生显著影响。截至目前,不同沉积环境中底栖动物如何响应植物登陆过程,以及对底栖生态系统构建方面的研究仍显薄弱。本研究在搜集和整理全球古生代底栖动物遗迹化石数据的基础上,建立以时间-沉积环境为核心的研究框架,系统对比在植物登陆演化过程中,海陆过渡环境、河流环境和湖泊环境中的遗迹多样性、遗迹歧异度、遗迹阶层和底栖生态系统工程的演化方式,评估底栖动物的行为演化对于植物登陆的响应。结果表明,古生代底栖动物向陆地环境的殖居具有显著的阶段性和差异性, 奥陶纪海陆过渡环境中的底栖动物已开始对植物登陆做出初步反馈,但在河流和湖泊环境中该响应却存在明显滞后,这一差异与植物根系的发育和森林化过程密切相关。总体来看,随着陆地植物对沉积环境中的底质稳定性和有机质等条件的改善,底栖动物逐渐扩展其生态位,生态空间利用和生态系统工程明显增强; 同时,底栖动物通过改造沉积物底质和物质能量流动,对植物陆地化过程产生正反馈作用。除陆地植物这一影响因素外,气候变化、构造运动和海平面波动等环境因素也对底栖动物行为习性演化产生深刻影响。研究成果对理解古生代地球生态系统演化和生物-沉积物相互作用提供了新的遗迹学视角。

关键词: 古生代, 陆地化, 遗迹化石, 生态反馈, 底栖生态系统

CLC Number:

P531
Q911.28

LI Feiyang, ZHANG Lijun, WANG Zongyan, CHAI Haiyan. Benthic animal response to the Paleozoic plant terrestrialization: insights from trace fossils[J]. Journal of Palaeogeography（Chinese Edition）, 2026, 28(4): 1257-1274.

李飞洋, 张立军, 王宗延, 柴海艳. 底栖动物如何响应古生代植物陆地化进程: 遗迹化石的证据与启示^*[J]. 古地理学报, 2026, 28(4): 1257-1274.

/ / Recommend / Download Citations

URL: http://www.gdlxb.cn/EN/10.7605/gdlxb.2026.135

http://www.gdlxb.cn/EN/Y2026/V28/I4/1257

Figures/Tables 4

Fig.1 A research framework on the impact of benthic animals on terrestrial ecological environments during plant terrestrialization

Fig.2 Typical trace fossils from transitional coastal and fluvial environments during the Cambrian-Permian

Fig.3 Time-environment matrix of benthic animal trace fossils under background of the Paleozoic plant terrestrialization

Fig.4 Ichnological metrics correlations with other parameters during the Silurian-Permian

References 104

[1]	崔智林. 1998. 新疆伊犁晚二叠世非海相Ophiomorpha. 地层学杂志, 22(3): 211-215,T002.
	[Cui Z L. 1998. The Upper Permian nonmarine Ophiomorpha of the Yily Basin,Xinjiang. Journal of Stratigraphy, 22(3): 211-215,T002]
[2]	郭文伟. 2022. 华北陆相晚二叠世—中三叠世磁性地层和遗迹化石演变. 中国地质大学博士学位论文.
	[Guo W W. 2022. Late Permian-Middle Triassic magnetostratigraphy and secular variation of terrestrial ichnofossils in North China. Doctoral dissertation of China University of Geosciences]
[3]	聂莎莎. 2023. 鄂尔多斯盆地东缘太原组遗迹化石及区域对比研究. 河南理工大学硕士学位论文.
	[Nie S S. 2023. Trace fossils of Taiyuan Formation in the eastern margin of Ordos Basin and their regional comparison. Masteral dissertation of Henan Polytechnic University]
[4]	牛永斌, 钟建华, 钟福平, 高玉飞, 禚元杰. 2008. 柴达木盆地南缘石炭系Chondrites遗迹化石的发现及其地质意义. 古地理学报, 10(5): 529-535.
	[Niu Y B, Zhong J H, Zhong F P, Gao Y F, Zhuo Y J. 2008. Discovery of trace fossil Chondrites in the Carboniferous in south margin of Qaidam Basin and its geological significance. Journal of Palaeogeography(Chinese Edition), 10(5): 529-535]
[5]	齐永安, 胡斌. 2001. 塔里木盆地下志留统遗迹组构及其环境解释. 古生物学报, 40(1): 116-126.
	[Qi Y A, Hu B. 2001. Lower Silurian ichnofabrics of Tarim Basin and their environmentary interpretation. Acta Palaeontologica Sinica, 40(1): 116-126]
[6]	王约. 2004. 贵州独山泥盆纪弗拉期—法门期生物灭绝后的生物遗迹. 古生物学报, 43(1): 131-141.
	[Wang Y. 2004. Some trace fossils after the Frasnian-Famennian extinct in Dushan area,southern Guizhou Province,China. Acta Palaeontologica Sinica, 43(1): 131-141]
[7]	薛进庄, 王嘉树, 李炳鑫, 黄璞, 刘乐. 2022. 陆地植物的起源、早期演化及地球环境效应. 地球科学, 47(10): 3648-3664.
	[Xue J Z, Wang J S, Li B X, Huang P, Liu L. 2022. Origin and early evolution of land plants and the effects on Earth’s environments. Earth Science, 47(10): 3648-3664]
[8]	薛进庄, 李炳鑫, 王嘉树, 刘乐, 黄璞, 熊聪慧, 沈冰. 2023. 早期维管植物辐射演化与长时间尺度水循环的耦合关系. 科学通报, 68(12): 1459-1472.
	[Xue J Z, Li B X, Wang J S, Liu L, Huang P, Xiong C H, Shen B. 2023. Coupling relationship between radiation of early vascular plants and the long-term water cycle. Chinese Science Bulletin, 68(12): 1459-1472]
[9]	薛进庄, 钟涛, 王徽, 李炳鑫, 杨琦琦, 王嘉树. 2026. 志留纪—泥盆纪维管植物的辐射演化及对陆地碳库的潜在影响. 古地理学报, 28(2): 708-723. doi: 10.7605/gdlxb.2025.078
	[Xue J Z, Zhong T, Wang H, Li B X, Yang Q Q, Wang J S. 2026. The Silurian-Devonian radiation of vascular plants and the potential effects on terrestrial carbon stocks. Journal of Palaeogeography(Chinese Edition), 28(2): 708-723]
[10]	杨式溥. 1999. 遗迹化石的古环境和古地理意义. 古地理学报, 1(1): 7-19.
	[Yang S P. 1999. Palaeoenvironmental and palaeogeographic significance of trace fossils. Journal of Palaeogeography(Chinese Edition), 1(1): 7-19]
[11]	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. 2004. Trace Fossils of China. Beijing: Science Press]
[12]	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]
[13]	张立军, 党志英, 李飞洋, 杨琦琦. 2021. 二叠纪—三叠纪之交深水海洋环境生态系统工程. 科学通报, 66(33): 4295-4306.
	[Zhang L J, Dang Z Y, Li F Y, Yang Q Q. 2021. Quantifying deep-water marine ecospace utilization and ecosystem engineering from Late Permian to Middle Triassic. Chinese Science Bulletin, 66: 4295-4306]
[14]	张小乐, 季兴开, 唐鹏, 王怿. 2023. 塔里木盆地西北缘志留系兰多维列统塔塔埃尔塔格组的遗迹化石. 地层学杂志, 47(4): 478-492.
	[Zhang X L, Ji X K, Tang P, Wang Y. 2023. Trace fossils from the Llandovery(Silurian)Tataaiertage Formation in the northwestern margin of Tarim Basin,Xinjiang. Journal of Stratigraphy, 47(4): 478-492]
[15]	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 their sedimentary environment of the Lower Carboniferous Malanbian,Xinshao County. Hunan Geology, 7(2): 53-61,86-87]
[16]	纵瑞文, 刘琦, 龚一鸣. 2011. 湖北武汉下志留统坟头组化石组合及沉积环境. 古地理学报, 13(3): 299-308. doi: 10.7605/gdlxb.2011.03.006
	[Zong R W, Liu Q, Gong Y M. 2011. Fossil association and sedimentary environments of the Lower Silurian Fentou Formation in Wuhan,Hubei Province. Journal of Palaeogeography(Chinese Edition), 13(3): 299-308]
[17]	Algeo T J, Scheckler S E, Scott A C. 1998. Terrestrial-marine teleconnections in the Devonian: links between the evolution of land plants,weathering processes,and marine anoxic events. Philosophical Transactions: Biological Sciences, 353: 113-130. doi: 10.1098/rstb.1998.0195 URL
[18]	Algeo T J, Scheckler S E, Maynard J B. 2001. Effects of the Middle to Late Devonian Spread of Vascular Land Plants on Weathering Regimes,Marine Biotas,and Global Climate. In: Gensel P G,Edwards D(eds). Plants Invade the Land. Columbia University Press: 213-236.
[19]	Banks H P. 1975. The oldest vascular land plants: a note of caution. Review of Palaeobotany and Palynology, 20(1-2): 13-25. doi: 10.1016/0034-6667(75)90004-4 URL
[20]	Bashforth A R, Cleal C J, Gibling M R, Falcon-Lang H J, Miller R F. 2014. Paleoecology of Early Pennsylvanian vegetation on a seasonally dry tropical landscape(Tynemouth Creek Formation,New Brunswick,Canada). Review of Palaeobotany & Palynology, 200: 229-263.
[21]	Bjerstedt T W. 1987. Trace fossils indicating estuarine deposystems for the Devonian-Mississippian Cloyd Conglomerate Member,Price Formation,Central Appalachians. Palaios, 2: 339-349. doi: 10.2307/3514759 URL
[22]	Briggs D E G, Miller M F, Isbell J L, Sidor C A. 2010. Permo-Triassic arthropod trace fossils from the Beardmore Glacier area,central Transantarctic Mountains,Antarctica. Antarctic Science, 22(2): 185-192. doi: 10.1017/S0954102009990708 URL
[23]	Buatois L A, Mángano M G. 1995. Post glacial lacustrine event sedimentation in an ancient mountain setting: Carboniferous Lake Malanzán(Western Argentina). Journal of Paleolimnology, 14: 1-22. doi: 10.1007/BF00682591 URL
[24]	Buatois L A, Mángano M G. 2004. Terminal Proterozoic-Early Cambrian ecosystems: Ichnology of the Puncoviscana Formation,Northwest Argentina. Trace Fossils in Evolutionary Palaeoecology. London: Taylor & Francis, 1-16.
[25]	Buatois L A, Mángano M G. 2011. Ichnology:Organism-substrate Interactions in Space and Time. Cambridge: Cambridge University Press,1-744.
[26]	Buatois L A, Mángano M G. 2013. Ichnodiversity and ichnodisparity: significance and caveats. Lethaia, 46: 281-292. doi: 10.1111/let.12018 URL
[27]	Buatois L A, Mangano M G, Maples C G, Lanier W P. 1998. Ichnology of an Upper Carboniferous fluvio-estuarine paleovalley: the tonganoxie sandstone,buildex quarry,eastern Kansas,USA. Journal of Paleontology, 72(1): 152-180. doi: 10.1017/S0022336000024094 URL
[28]	Buatois L A, Gingras M K, MacEachern J, Mángano M G, Zonneveld J P, Pemberton S G, Netto R G, Martin A. 2005. Colonization of brackish-water systems through time: evidence from the trace-fossil record. Palaios, 20(4): 321-347. doi: 10.2110/palo.2004.p04-32 URL
[29]	Buatois L A, Davies N S, Gibling M R, Krapovickas V, Labandeira C C, MacNaughton R B, Mángano M G, Minter N J, Shillito A P. 2022. The invasion of the land in deep time: integrating Paleozoic records of paleobiology,ichnology,sedimentology,and geomorphology. Integrative and Comparative Biology, 62: 297-331. doi: 10.1093/icb/icac059 URL
[30]	Capel E, Cleal C J, Gerrienne P, Servais T, Cascales-Miñana B. 2021. A factor analysis approach to modelling the early diversification of terrestrial vegetation. Palaeogeography,Palaeoclimatology,Palaeoecology, 566: 110170.
[31]	Chen B, Chen J T, Qie W K, Huang P, He T C, Joachimski M M, Regelous M, Pogge von Strandmann P A E, Liu J S, Wang X D, Montañez I P, Algeo T J. 2021. Was climatic cooling during the earliest Carboniferous driven by expansion of seed plants. Earth and Planetary Science Letters, 565: 116953.
[32]	Chen J Y, Liu J. 2024. Journey to the east: the oldest tetrapod fauna of east Pangea in Early Permian. National Science Review, 11(10): nwae249.
[33]	Cohen A S. 2003. Paleolimnology:The History and Evolution of Lake Systems. Oxford: Oxford University Press,1-485.
[34]	Corenblit D, Piégay H, Arrignon F, González-Sargas E, Bonis A, Davies N S, Ebengo D M, Garófano-Gómez V, Gurnell A M, Henry A L, Hortobágyi B, Martínez-Capel F, Steiger J, Tabacchi E, Tooth S, Vautier F, Walcker R. 2024. Interactions between vegetation and river morphodynamics. Part I: research clarifications and challenges. Earth-Science Reviews, 253: 104769. doi: 10.1016/j.earscirev.2024.104769 URL
[35]	Cramer B D, Jarvis I. 2020. Carbon isotope stratigraphy. IIn: Gradstein F M et al.(eds). Geologic Time Scale 2020. Amsterdam: Elsevier, 309-343.
[36]	Daeschler E B, Shubin N H, Jenkins F A. 2006. A Devonian tetrapod-like fish and the evolution of the tetrapod body plan. Nature, 440: 757-763. doi: 10.1038/nature04639
[37]	Davies N S, Gibling M R. 2010. Cambrian to Devonian evolution of alluvial systems: the sedimentological impact of the earliest land plants. Earth-Science Reviews, 98: 171-200. doi: 10.1016/j.earscirev.2009.11.002 URL
[38]	Davies N S, Gibling M R. 2011. Evolution of fixed-channel alluvial plains in response to Carboniferous vegetation. Nature Geoscience, 4(9): 629-633. doi: 10.1038/ngeo1237
[39]	Davies N S, Gibling M R. 2013. The sedimentary record of Carboniferous rivers: continuing influence of land plant evolution on alluvial processes and Palaeozoic ecosystems. Earth-Science Reviews, 120: 40-79. doi: 10.1016/j.earscirev.2013.02.004 URL
[40]	Davies N S, McMahon W J, Shillito A P, Veenma Y P, Craig J A. 2023. Ichnology of a Middle Devonian regression: environment,terrestrialization and true substrate controls on trace fossils of the Hangman Sandstone Formation,SW England. Palaeogeography,Palaeoclimatology,Palaeoecology, 625: 111671.
[41]	DiMichele W A, Pfefferkorn H W, Gastaldo R A. 2001. Response of Late Carboniferous and Early Permian plant communities to climate change. Annual Review of Earth and Planetary Sciences, 29(1): 461-487. doi: 10.1146/earth.2001.29.issue-1 URL
[42]	Ding Y, Liu J N, Liu Y, Chen F F. 2020. Trace fossils from Lower Cambrian Hongjingshao Formation,Yunnan,China: taxonomy,palaeoecology,palaeoenvironment. Palaeoworld, 29: 649-661. doi: 10.1016/j.palwor.2020.02.002
[43]	El-Refaiy A A, El-Sabbagh A M, El Hedeny M M, Mansour A S, El-Barkooky A N. 2023. Stratigraphical,microfacies,and ichnological characteristics and depositional environments of the Permo-Carboniferous Aheimer Formation,western side of the Gulf of Suez,Egypt. Facies, 69: 8. doi: 10.1007/s10347-023-00666-7
[44]	Falcon-Lang H J, Minter N J, Bashforth A R, Gibling M R, Miller R F. 2015. Mid-Carboniferous diversification of continental ecosystems inferred from trace fossil suites in the Tynemouth Creek Formation of New Brunswick,Canada. Palaeogeography,Palaeoclimatology,Palaeoecology, 440: 142-160. doi: 10.1016/j.palaeo.2015.09.002 URL
[45]	Fan R Y, Zong R W, Gong Y M. 2020. Fish swimming traces from the Upper Devonian of Wuhan,South China. Journal of Earth Science, 34: 1319-1322. doi: 10.1007/s12583-021-1509-9
[46]	Feng X Q, Chen Z Q, Zhao L S, Lan Z W. 2021. Middle Permian trace fossil assemblages from the Carnarvon Basin of Western Australia: implications for the evolution of ichnofaunas in wave-dominated siliciclastic shoreface settings across the Permian-Triassic boundary. Global and Planetary Change, 197: 103392.
[47]	Fillmore D L, Lucas S G, Simpson E L. 2010. Invertebrate trace fossils in semi-arid to arid braided-ephemeral-river deposits of the Mississippian middle member of the Mauch Chunk Formation,eastern Pennsylvania,USA. Palaeogeography,Palaeoclimatology,Palaeoecology, 292(1-2): 222-244. doi: 10.1016/j.palaeo.2010.03.047 URL
[48]	Flannery-Sutherland J T. 2021. Putative arthropod trace fossils from the Orcadian Basin at Achanarras Quarry(Middle Devonian of Scotland). Journal of the Geological Society, 178(4): jgs2020-jgs2233.
[49]	Foster G L, Royer D L, Lunt D J. 2017. Future climate forcing potentially without precedent in the last 420 million years. Nature Communications, 8: 14845.
[50]	Garvey J M, Hasiotis S T. 2008. An ichnofossil assemblage from the Lower Carboniferous Snowy Plains Formation,Mansfield Basin,Australia. Palaeogeography,Palaeoclimatology,Palaeoecology, 258: 257-276. doi: 10.1016/j.palaeo.2007.11.002 URL
[51]	Gensel P G. 1986. Diversification of land plants in the Early and Middle Devonian. Notes for a Short Course: Studies in Geology, 15: 64-80. doi: 10.1017/S0271164800001330 URL
[52]	Gensel P G. 2008. The earliest land plants. Annual Review of Ecology,Evolution,and Systematics, 39(1): 459-477. doi: 10.1146/ecolsys.2008.39.issue-1 URL
[53]	Gibling M R, Davies N S. 2012. Palaeozoic landscapes shaped by plant evolution. Nature Geoscience, 5(2): 99-105. doi: 10.1038/ngeo1376
[54]	Gouramanis C, McLoughlin S. 2016. Siluro-Devonian trace fossils from the Mereenie sandstone,Kings Canyon,Watarrka National Park,Amadeus Basin,northern Territory,Australia. Alcheringa: An Australasian Journal of Palaeontology, 40: 118-128. doi: 10.1080/03115518.2016.1099957 URL
[55]	Hao S G, Xue J Z. 2013. The early Devonian Posongchong Flora of Yunnan:A Contribution to An Understanding of The Evolution and Early Diversification of Vascular Plants. Beijing: Science Press,1-366.
[56]	Jin J S, Harper D A T, Rasmussen J A, Sheehan P M. 2012. Late Ordovician massive-bedded Thalassinoides ichnofacies along the palaeoequator of Laurentia. Palaeogeography,Palaeoclimatology,Palaeoecology, 367: 73-88.
[57]	Judd E J, Tierney J E, Lunt D J, Montañez I P, Huber B T, Wing S L, Valdes P J. 2024. A 485-million-year history of Earth’s surface temperature. Science, 385: eadk3705. doi: 10.1126/science.adk3705 URL
[58]	Keighley D G, Pickerill R K. 2003. Ichnocoenoses from the Carboniferous of eastern Canada and their implications for the recognition of ichnofacies in nonmarine strata. Atlantic Geology, 39(1): 1-22.
[59]	Kenrick P, Crane P R. 1997. The origin and early evolution of plants on land. Nature, 389: 33-39. doi: 10.1038/37918
[60]	Kenrick P, Wellman C H, Schneider H, Edgecombe G D. 2012. A timeline for terrestrialization: consequences for the carbon cycle in the Palaeozoic. Philosophical Transactions: Biological Sciences, 367(1588): 519-536. doi: 10.1098/rstb.2011.0271 URL
[61]	Lipp A G, Shorttle O, Sperling E A, Brocks J J, Cole D B, Crockford P W, Del Mouro L, Dewing K, Dornbos S Q, Emmings J F, Farrell U C, Jarrett A, Johnson B W, Kabanov P, Keller C B, Kunzmann M, Miller A J, Mills N T, O’Connell B, Peters S E, Planavsky N J, Ritzer S R, Schoepfer S D, Wilby P R, Yang J. 2021. The composition and weathering of the continents over geologic time. Geochemical Perspectives Letters, 17: 21-26. doi: 10.7185/geochemlet.2109
[62]	Lu J, Peng X, Yin L S, Ling Z Y, Yang M F, Zhang P X, Zhou K, Liu L, Dai S F, Shao L Y, Hilton J. 2026. Influence of plant terrestrialization on coal accumulation and deep time terrestrial carbon storage. Earth-Science Reviews, 274: 105390. doi: 10.1016/j.earscirev.2026.105390 URL
[63]	Lucas S G. 2021. Tetrapod origins. In: Alderton D,Elias S A(eds). Encyclopedia of Geology. Amsterdam: Elsevier,138-146.
[64]	MacNaughton R B, Cole J M, Dalrymple R W, Braddy S J, Briggs D E G, Lukie T D. 2002. First steps on land: Arthropod trackways in Cambrian-Ordovician eolian sandstone,southeastern Ontario,Canada. Geology, 30(5): 391-394. doi: 10.1130/0091-7613(2002)030<0391:FSOLAT>2.0.CO;2 URL
[65]	Mángano M G, Buatois L A. 2004. Reconstructing Early Phanerozoic intertidal ecosystems:ichnology of the Cambrian Campanario Formation in northwest Argentina. In: Webby B D,Mángano M G,Buatois L A(eds). Fossils and Strata. London: Taylor & Francis, 17-38.
[66]	Mángano M G, Buatois L A, Moya M C. 2021a. Trazas fósiles de trilobites de la Formación Mojotoro(Ordovícico Inferior-Medio de Salta,Argentina): implicancias paleoecológicas,paleobiológicas y bioestratigráficas. Spanish Journal of Palaeontology, 16(1): 9-28. doi: 10.7203/sjp.16.1.21577 URL
[67]	Mángano M G, Buatois L A, Waisfeld B G, Muñoz D F, Vaccari N E, Astini R A. 2021b. Were all trilobites fully marine Trilobite expansion into brackish water during the early Palaeozoic. Proceedings of the Royal Society B:Biological Sciences, 288(1944): 1-10.
[68]	Mángano M G, Waisfeld B G, Buatois L A, Vaccari N E, Muñoz D F. 2023. Evolutionary and ecologic controls on benthos distribution from an upper Cambrian incised estuarine valley: implications for the early colonization of marginal-marine settings. Palaeogeography,Palaeoclimatology,Palaeoecology, 626: 111692.
[69]	Marriott S B, Morrissey L B, Hillier R D. 2009. Trace fossil assemblages in Upper Silurian tuff beds: evidence of biodiversity in the Old Red Sandstone of southwest Wales,UK. Palaeogeography,Palaeoclimatology,Palaeoecology, 274(3-4): 160-172. doi: 10.1016/j.palaeo.2009.01.001 URL
[70]	McArthur J M, Howarth R J, Shields G A, Zhou Y. 2020. Strontium isotope stratigraphy. In: Gradstein F M et al(eds). Geologic Time Scale 2020. Amsterdam: Elsevier, 211-238.
[71]	Miller M F. 1984. Distribution of biogenic structures in Paleozoic nonmarine and marine-margin sequences: an actualistic model. Journal of Paleontology, 58(2): 550-570.
[72]	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 & Evolution, 1(7): 175.
[73]	Montañez I P, Poulsen C J. 2013. The Late Paleozoic Ice Age: an evolving paradigm. Annual Review of Earth and Planetary Sciences, 41(1): 629-656. doi: 10.1146/earth.2013.41.issue-1 URL
[74]	Morrissey L B, Braddy S J. 2004. Terrestrial trace fossils from the Lower Old Red Sandstone,southwest Wales. Geological Journal, 39(3-4): 315-336. doi: 10.1002/gj.v39:3/4 URL
[75]	Morrissey L B, Braddy S J, Dodd C, Higgs K T, Williams B P J. 2012a. Trace fossils and palaeoenvironments of the Middle Devonian Caherbla Group,Dingle Peninsula,southwest Ireland. Geological Journal, 47(1): 1-29. doi: 10.1002/gj.v47.1 URL
[76]	Morrissey L B, Hillier R D, Marriott S B. 2012b. Late Silurian and Early Devonian terrestrialisation: ichnological insights from the Lower Old Red Sandstone of the Anglo-Welsh Basin,U.K. Palaeogeography,Palaeoclimatology,Palaeoecology, 337: 194-215.
[77]	Netto R G, Balistieri P R M N, Lavina E L C, Silveira D M. 2009. Ichnological signatures of shallow freshwater lakes in the glacial Itararé Group(Mafra Formation,Upper Carboniferous-Lower Permian of Paraná Basin,southern Brazil). Palaeogeography,Palaeoclimatology,Palaeoecology, 272(3-4): 240-255. doi: 10.1016/j.palaeo.2008.10.028 URL
[78]	Pickerill R K. 1992. Carboniferous nonmarine invertebrate ichnocoenoses from southern New Brunswick,eastern Canada. Ichnos: An International Journal of Plant & Animal, 2(1): : 21-35.
[79]	Pollard J, Selden P A, Watts S. 2008. Trace fossils of the arthropod Camptophyllia from the Westphalian(Carboniferous)rocks of Lancashire,UK and their palaeoenvironmental context. Palaeogeography,Palaeoclimatology,Palaeoecology, 270(3-4): 399-406. doi: 10.1016/j.palaeo.2008.01.032 URL
[80]	Raven J A, Edwards D. 2013. Photosynthesis in early land plants:adapting to the terrestrial environment. In: Hanson D T,Rice S K(eds). Photosynthesis in Bryophytes and Early Land Plants. Dordrecht: Springer Netherlands, 29-58.
[81]	Ridgwell A, Zeebe R E. 2005. The role of the global carbonate cycle in the regulation and evolution of the Earth system. Earth and Planetary Science Letters, 234(3-4): 299-315. doi: 10.1016/j.epsl.2005.03.006 URL
[82]	Scotese C R, Song H J, Mills B J W, van der Meer D G. 2021. Phanerozoic paleotemperatures: the Earth’s changing climate during the last 540 million years. Earth-Science Reviews, 215: 103503. doi: 10.1016/j.earscirev.2021.103503 URL
[83]	Shillito A P, Davies N S. 2017. Archetypally Siluro-Devonian ichnofauna in the Cowie Formation, Scotland: implications for the myriapod fossil record and Highland Boundary Fault movement. Proceedings of the Geologists’ Association, 128(5-6): 815-828.
[84]	Shillito A P, Davies N S. 2021. The Silurian inception of inland desert ecosystems: trace fossil evidence from the Mereenie Sandstone,Northern Territory,Australia. Journal of the Geological Society, 178(4): jgs2020-jgs2243.
[85]	Stacey J, Wallace M W, Hood A V S, Shuster A M, Corlett H, Reed C P, Moynihan C. 2024. Ocean oxygenation and ecological restructuring caused by the Late Paleozoic evolution of land plants. Geology, 52: 948-952. doi: 10.1130/G52502.1 URL
[86]	Stein W E, Berry C M, Hernick L V, Mannolini F. 2012. Surprisingly complex community discovered in the mid-Devonian fossil forest at Gilboa. Nature, 483: 78-81. doi: 10.1038/nature10819
[87]	Toledo S, Bippus A C, Tomescu A M F. 2018. Buried deep beyond the veil of extinction: euphyllophyte relationships at the base of the spermatophyte clade. American Journal of Botany, 105(8): 1264-1285. doi: 10.1002/ajb2.1102 pmid: 29893501
[88]	Walker L J, Wilkinson B H, Ivany L C. 2002. Continental drift and Phanerozoic carbonate accumulation in shallow-shelf and deep-marine settings. The Journal of Geology, 110(1): 75-87. doi: 10.1086/324318 URL
[89]	Wang D M, Liu L, Meng M C, Xue J Z, Liu T, Guo Y. 2014. Cosmosperma polyloba gen. et sp. nov.,a seed plant from the Upper Devonian of South China. Naturwissenschaften, 101(8): 615-622. doi: 10.1007/s00114-014-1187-x URL
[90]	Wang J S, Zhang L J, Huang P, Liu L, Xue J Z. 2024. Sedimentology and ichnology of the Xiaxishancun Formation of Qujing,Yunnan: earliest Devonian marginal-marine environments and ecosystems in South China. Palaeogeography,Palaeoclimatology,Palaeoecology, 647: 112220.
[91]	Weber B, Braddy S J. 2003. A marginal marine ichnofauna from the Blaiklock Glacier Group(Lower Ordovician)of the Shackleton Range,Antarctica. Transactions of the Royal Society of Edinburgh: Earth Sciences, 94(1): 1-20. doi: 10.1017/S026359330000050X URL
[92]	Wellman C H, Cascales-Miñana B, Servais T. 2023. Terrestrialization in the Ordovician. Geological Society,London,Special Publications, 532: 171-190. doi: 10.1144/SP532-2022-92 URL
[93]	Wilmeth D T, Corsetti F A, Beukes N J, Awramik S M, Petryshyn V, Spear J R, Celestian A J. 2019. Neoarchean(2.7 Ga)lacustrine stromatolite deposits in the Hartbeesfontein Basin,Ventersdorp Supergroup,South Africa: implications for oxygen oases. Precambrian Research, 320: 291-302. doi: 10.1016/j.precamres.2018.11.009
[94]	Xue J Z, Deng Z Z, Huang P, Huang K J, Benton M J, Cui Y, Wang D M, Liu J B, Shen B, Basinger J F, Hao S G. 2016. Belowground rhizomes in paleosols: The hidden half of an Early Devonian vascular plant. Proceedings of the National Academy of Sciences of the United States of America, 113: 9451-9456.
[95]	Zhang L J, Zhao Z. 2015. Lower Devonian trace fossils and their palaeoenvironmental significance from the western Yangtze Plate,South China. Turkish Journal of Earth Sciences, 24(4): 325-343. doi: 10.3906/yer-1409-15 URL
[96]	Zhang L J, Qi Y A, Buatois L A, Mángano M G, Meng Y, Li D. 2017. The impact of deep-tier burrow systems in sediment mixing and ecosystem engineering in Early Cambrian carbonate settings. Scientific Reports, 7(1): 45773. doi: 10.1038/srep45773 URL
[97]	Zhang L J, Buatois L A, Mángano M G, Qi Y A, Tai C. 2019. Early Triassic trace fossils from South China marginal-marine settings: implications for biotic recovery following the end-Permian mass extinction. GSA Bulletin, 131: 235-251. doi: 10.1130/B31867.1 URL
[98]	Zhang L J, Buatois L A, Mángano M G, Fan R Y, Yang Q Q, Wei F, Wang Z, Ma X Y. 2024. Evolution and palaeoenvironmental migration of a Cambrian undermat miner. Palaeogeography,Palaeoclimatology,Palaeoecology, 655: 112559.
[99]	Zhang L J, Wang Z Y, Li Y, Chai H Y, Fan R Y, Wang J S, Wang Z. 2025a. How benthic animals responded to the Paleozoic plant terrestrialization: evidence from trace fossils and their ecosystem engineering reconstruction. Earth-Science Reviews, 270: 105236.
[100]	Zhang L J, Li Y, Yang Q Q, Wang Z Y, Chai H Y, Qie W K. 2025b. Dense Early Cambrian Psammichnites gigas circularis from South China and its palaeoecological significance for the Cambrian Information Revolution. Historical Biology, 38: 907-922. doi: 10.1080/08912963.2025.2522773 URL
[101]	Zhang Y L, Lai G M, Gong E P, Yuan D C, Wilson M A, Li Y. 2023. Characteristics of early Neoproterozoic stromatolites from southern Liaoning,North China: insights into the formation of stromatolites. Journal of Marine Science and Engineering, 11(9): 1709. doi: 10.3390/jmse11091709 URL
[102]	Zhao C X, Shi M, Feng Q L, Ye Y, Zareen Khan M, Feng F B. 2020. New study of microbial mats from the Mesoproterozoic Jixian Group,North China: evidence for photosynthesis and oxygen release. Precambrian Research, 344: 105734. doi: 10.1016/j.precamres.2020.105734 URL
[103]	Zhu M, Fan J H. 1995. Youngolepis from the Xishancun Formation(early Lochkovian)of Qujing,China. Geobios, 28: 293-299.
[104]	Zouicha A, Voigt S, Saber H, Marchetti L, Hminna A, El Attari A, Ronchi A, Schneider J W. 2021. First record of Permian continental trace fossils in the Jebilet Massif,Morocco. Journal of African Earth Sciences, 173: 104015.

Please choose a citation manager

Content to export