Behavioural evolution pattern and biogeographical migration of the Paleozoic psammichnitids
ZHANG Lijun1, PANG Zhichao1, YANG Qiqi1, FAN Liu1, WEI Fan2,3, SONG Huibo1, NIU Yongbin1
1 Institute of Resources and Environment,Key Laboratory of Biogenic Traces and Sedimentary Minerals of Henan Province, Henan Polytechnic University,Henan Jiaozuo 454003,China; 2 Institute of Palaeontology,MEC International Joint Laboratory for Palaeobiology and Palaeoenvironment, and Yunnan Key Laboratory for Palaeobiology,Yunnan University,Kunming 650500,China; 3 State Key Laboratory of Paleobiology and Stratigraphy,Nanjing Institute of Geology and Paleontology, Chinese Academy of Sciences,Nanjing 210008,China
Abstract：Psammichnites and Dictyodora are grouped into psammichnitids,representing the typical slug-like bulldozers with a snorkel living in the marine environments during the Palaeozoic. Through the morphological and functional analysis of Paleozoic Psammichnites and Dictyodora,combining with their spatio-temporal distribution characteristics,it is found: (1)Psammichnites generally has short and shallow grooves,and the wall height of Dictyodora gradually increased with time. (2)Psammichnites is mainly distributed in the shallow-marine environment,whereas Dictyodora is mainly distributed in deep-marine environment with occasional occurrences of certain ichnospecies in the shallow-marine environment. (3)The ichnodiversity of Psammichnites ichnospecies rapidly increased in the Carboniferous shallow-marine environment,belonging to the top-heavy pattern. Psammichnites gigas is distributed widely in the Early Cambrian strata,which is of biostratigraphic importance. The ichnodiversity of Dictyodora ichnospecies rapidly expanded in the Ordovician,representing the bottom-heavy pattern. (4)Psammichnites and Dictyodora were mainly distributed in the Iapetus and Rheic oceans in the middle and high latitudes of the southern hemisphere during the Cambrian to Ordovician. Due to the formation of Laurussia and the closure of Iapetus and Rheic oceans,Psammichnites and Dictyodora migrated to the new empty ecological niches from the Carboniferous to Permian,indicating that the adaptation and migration of trace makers to new ecological niches might be strongly influenced by environmental changes. Until now Psammichnites and Dictyodora have not been found in the post-Permian strata,possibly suggesting that their trace markers went extinct in the end-Permian mass extinction. On the basis of the above systematic summary and induction of the behavior habits,biogeographic migration characteristics and evolutionary characteristics of Psammochnites and Dictyodora,it can provide new data and new understanding of the evolutionary ecology and biogeographic migration characteristics of trace groups reflected by trace fossils from a detailed and profound perspective.
ZHANG Lijun,PANG Zhichao,YANG Qiqi et al. Behavioural evolution pattern and biogeographical migration of the Paleozoic psammichnitids[J]. JOPC, 2023, 25(2): 382-391.
[1] 党志英,张立军. 2020a. 西秦岭甘南地区上二叠统毛毛隆组古环境演变: 基于遗迹网络分析. 古地理学报, 22(6): 1171-1180. [Dang Z Y,Zhang L J.2020a. Palaeoenvironmental evolution of the Upper Permian Maomaolong Formation in Gannan area of the West Qinling: based on ichnonetwork analysis. Journal of Palaeogeography(Chinese Edition), 22(6): 1171-1180] [2] 党志英,张立军. 2020b. 西秦岭甘南地区二叠纪—三叠纪遗迹化石及其古环境意义. 古生物学报, 59(4): 430-444. [Dang Z Y,Zhang L J.2020b. Trace fossils and their paleoenvironmental significance of the Permian-Triassic succession in the Gannan region of West Qinling. Acta Palaeontologica Sinica, 59(4): 430-444] [3] 胡世学,朱茂炎,罗惠麟,米歇尔斯坦纳,赵方巨,李国详,刘琦,张志飞. 2014. 关山生物群. 昆明: 云南科技出版社. [Hu S,Zhu M,Luo H-L,Steiner M,Zhao F,Li G-X,Liu Q,Zhang Z.2014. The Guanshan Biota. Kunming: Yunan Science and Technology Press] [4] 罗惠麟,李勇,胡世学,傅晓平,侯蜀光,刘兴尧,陈良忠,李锋军,庞纪院,刘琦. 2008. 云南东部早寒武世马龙动物群和关山动物群. 昆明: 云南科技出版社. [Luo H-L,Li Y,Hu S,Fu X,Hou S,Liu X,Chen L,Li F,Pan J,Liu Q.2008. The Malong and Guanshan Biota in the easter Yunnan. Kunming: Yunnan Science and Technology Press] [5] 戎嘉余,黄冰. 2014. 生物大灭绝研究三十年. 中国科学: 地球科学, 44(3): 377-404. [Rong J Y,Huang B.2014. Study of Mass Extinction over the past thirty years: a synopsis. Scientia Sinica Terrae, 44(3): 377-404] [6] 杨式溥,胡云绪. 1992. 西秦岭西段志留纪遗迹化石. 现代地质, 6(4): 384-391. [Yang S P,Hu Y X.1992. Silurian trace fossils from the western part of west Qinling. Geosience, 6(4): 384-391] [7] 左国朝,金松桥,朱伟元. 1984. 甘肃夏河县下卡加—完尕滩一带二叠系浊积岩及有关粗碎屑沉积物. 沉积学报, 2(3): 75-81. [Zuo G C,Jin S Q,Zhu W Y.1984. Permian turbidites and related coarse clastic sediments in Xiakajia-Wangatan area of Xiahe County,Gansu Province. Acta Sedimentologica Sinica, 2(3): 75-81] [8] Baucon A,Neto de Carvalho C.2008. From the river to the sea: Pramollo,a new ichnolagerstätte from the Carnic Alps(Carboniferous,Italy-Austria). Studi Trentini Science Naturali. Acta Geologica, 83: 87-114. [9] Benton M.1982. Dictyodora and associated trace fossils from the Palaeozoic of Thuringia. Lethaia, 15: 115-132. [10] Benton M,Trewin N H.1980. Dictyodora from the Silurian of Peebleshire,Scotland. Palaeontology, 23: 501-513. [11] Chlupáč I.1997. Palaeozoic ichnofossils in phyllites near Železný Brod,northern Bohemia. Journal of the Czech Geological Society, 42: 75-86. [12] Domeier M.2016. A plate tectonic scenario for the Iapetus and Rheic oceans. Gondwana Research, 36: 275-295. [13] Feng X,Chen Z Q,Zhao L,Lan Z.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. [14] Häntzschel W.1975. Trace fossils and problemmatica. In: Teichert G(ed). Treatise on Invertebrate Paleontology,part W,Miscellanea,Supplement 1. Kansas: Geological Society of America and University of Kansas Press,1-269. [15] Jensen S,Carvalho C,Palacios T.2016. Trace fossils from the Barrancos and Colorada formations,Ordovician,Ossa-Morena Zone,Portugal and Spain. Comunicações Geológicas, 103: 159-167. [16] Lang V,Pek I,Zapletal J.1979. Ichnofosilie kulmu jihovýchodnisti Drahanské vrchoviny. Acta Universitatis Palackianae Olomucensis,Geographica-Geologica, 62: 57-96. [17] Li L,Meng Q R,Pullen A,Garzione C,Wu G,Wang Y,Ma S,Duan L.2014. Late Permian-early Middle Triassic back-arc basin development in West Qinling,China. Journal of Asian Earth Sciences, 87: 116-129. [18] Llompart C,Wieczorek J.1997. Trace fossils from culm facies of Minorca Island. Prace: Panstwowego Instytutu Geologicznego,100-101. [19] Luca P,Basilici G.2013. A prodeltaic system controlled by hyperpycnal flows and storm waves: reinterpretation of the Punta Negra Formation(Lower-Middle Devonian,Argentine Precordillera). Brazilian Journal of Geology, 43: 673-694. [20] MacNaughton R B,Fallas K M,Finley T D.2021. Psammichnites gigas from the lower Cambrian of the Mackenzie Mountains,northwest Canada,and their biostratigraphic implications. Ichnos, 28: 164-175. [21] Mángano M G,Buatois L A,Rindsberg A K.2002. Carboniferous Psammichnites: systematic re-evalution,taphonomy and autecology. Ichnos, 9: 1-22. [22] Mángano M G,Buatois L A,MacNaughton R B,Jensen S,Gougeon R,Marcos A,Meek D,Piñuela L,García-Ramos J C.2022. The Psammichnites-Taphrhelminthopsis conundrum: implications for Calibrating the Cambrian explosion. Earth-Science Reviews, 227: 03971. [23] McIlroy D,Heys G R.1997. Palaeobiological significance of Plagiogmus arcuatus from the lower Cambrian of central Australia. Alcheringa, 21: 161-178. [24] Mikuláš R,Lehotský T,Bábek O.2002. Ichnofabric of the Culm facies: a case study of the Moravice Formation(Lower Carboniferous;Moravia and Silesia,Czech Republic). Geologica Carpathica, 53: 141-148. [25] Mikuláš R,Lehotský T,Bábek O.2004. Trace fossils of the Moravice Formation in the southern part of the Nízký Jeseník Mts.(Lower Carboniferous,Culm facies;Moravia,Czech Republic). Bulletin of Geosciences, 79: 81-98. [26] Orr P J,Pickerill R.1995. Trace fossils from early Silurian flysch of the Waterville Formation,Maine,USA. Northeastern Geology, 17: 394-414. [27] Orr P J.1996. The ichnofauna of the Skiddaw Group(Early Ordovician)of the Lake District,England. Geological Magazine, 133: 193-216. [28] Orr P,Benton M,Trewin N.1996. Deep marine trace fossil assemblages from the Lower Carboniferous of Menorca,Balearic Islands,western Mediterranean. Geological Journal, 31: 235-258. [29] Orr P,Howe M.1999. Macrofauna and ichnofauna of the Manx Group(Early Ordovician),Isle of Man. Geological Society,London,Special Publications, 160: 33-44. [30] Pazos P J,Gutiérrez C,Fernández D E,Heredia A M,Comerio M.2015a. The unusual record of Nereites,wrinkle marks and undermat mining trace fossils from the late Silurian-earliest Devonian of central-western margin of Gondwana(Argentina). Palaeogeography,Palaeoclimatology,Palaeoecology, 439: 4-16. [31] Pazos P J,Heredia A M,Fernandez D E,Gutierrez C,Comerio M.2015b. The ichnogenus Dictyodora from late Silurian deposits of central-western Argentina: ichnotaxonomy,ethology and ichnostratigrapical perspectives from Gondwana. Palaeogeography, Palaeoclimatology, Palaeoecology, 439: 27-37. [32] Pek I,Zapletal J.1990. The importance of ichnology in geologic studies of the eastern Bohemian Massif(lower Carboniferous),Czechoslovakia. Ichnos, 1: 147-149. [33] Pickerill R,Fyffe L,Forbes W.1987. Late Ordovician-early Silurian trace fossils from the Matapedia Group,Tobique River,western New Brunswick,Canada. Marine Sediments and Atlantic Geology, 23: 77-87. [34] Rong J Y,Fan J X,Arnold I. M,Li G X.2007. Dynamic patterns of latest Proterozoic-Palaeozoic-early Mesozoic marine biodiversity in South China. Geological Journal, 42: 431-454. [35] Seilacher-Drexler E,Seilacher A.1999. Undertraces of sea pens and moon snails and possible fossil counterparts. Neues Jahrbuch für Geologie und Paläontologie-Abhandlungen, 214: 195-210. [36] Seilacher A.1997. The meaning of the Cambrian Explosion. Bulletin of National Museum of Natural Science, 10: 1-9. [37] Seilacher A.2007. Trace Fossil Analysis. New York: Springer-Verlag. [38] Seilacher A,Gámez-Vintaned.1995. Psammichnites gigas: ichological expression of the Cambrian explosion. Proceedings Sixth Paleobenthos International Symposium,Alghero: 151-152. [39] Seilacher A,Buatois L A,Mángano M G.2005. Trace fossils in the Ediacaran-Cambrian transition: behavioral diversification,ecological turnover and environmental shift. Palaeogeography,Palaeoclimatology,Palaeoecology, 227: 323-356. [40] Servais T,Harper D A T.2018. The Great Ordovician Biodiversification Event(GOBE): definition,concept and duration. Lethaia, 51: 151-164. [41] Servais T,Owen A W,Harper D A T,Kröger B,Munnecke A.2010. The Great Ordovician Biodiversification Event(GOBE): the palaeoecological dimension. Palaeogeography,Palaeoclimatology,Palaeoecology, 294: 99-119. [42] Stigall A.2012. Speciation collapse and invasive species dynamics during the Late Devonian “Mass Extinction”. GSA Today, 22: 4-9. [43] Torsvik T H,Cocks L R M. 2016. Earth History and Palaeogeography. Cambridge: Cambridge University Press. [44] Uchman A.2004. Phanerozoic history of deep-sea trace fossils. Geological Society,London,Special Publications, 228: 125-139. [45] Uchman A,Hanken N-M,Binns R.2005. Ordovician bathyal trace fossils from metasiliciclastics in central Norway and their sedimentological and paleogeographical implications. Ichnos, 12: 105-133. [46] 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: 75-87. [47] Zhang L J,Buatois L A,Mángano M G.2022. Potential and problems in evaluating secular changes in the diversity of animal-substrate interactions at ichnospecies rank. Terra Nova, 34: 433-440. [48] Zhang L J,Fan R Y,Dang Z Y,Gong Y M.2020. The youngest known Dictyodora from the Late Permian(Lopingian)deep sea in West Qinling,central China. Palaeogeography,Palaeoclimatology,Palaeoecology, 558: 109948. [49] Neto de Carvalho C.2001. Implicações geométricas na etologia do produtor de Dictyodora tenuis(M’ Coy)no Arenigiano de Barrancos(Portugal): custos e benefícios de ser um pioneiro. Seminarios de Paleontologia de Zaragoza, 5: 369-378. [50] Stepanek, Geyer G.1989. Spurenfossilien aus dem Kulm(Unterkarbon)des Frankenwaldes. Beringeria, 1: 1-55.
